Historical and projected interactions between climate change and insect voltinism in a multivoltine species

Climate change can cause major changes to the dynamics of individual species and to those communities in which they interact. One effect of increasing temperatures is on insect voltinism, with the logical assumption that increases in surface temperatures would permit multivoltine species to increase the number of generations per year. Though insect development is primarily driven by temperature, most multivoltine insect species rely on photoperiodic cues, which do not change from year‐to‐year or in response to climate warming, to initiate diapause. Thus, the relationship between climate change and voltinism could be complex. We use a phenology model for grape berry moth, Paralobesia viteana (Clemens), which incorporates temperature‐dependent development and diapause termination, and photoperiod‐dependent diapause induction, to explore historical patterns in year‐to‐year voltinism fluctuations. We then extend this model to predict voltinism under varying scenarios of climate change to show the importance of both the quality and quantity of accumulated heat units. We also illustrate that increases in mean surface temperatures > 2 °C can have dramatic effects on insect voltinism by causing a shift in the ovipositional period that currently is subject to diapause‐inducing photoperiods.     

Geographic variation in diapause induction: the grape berry moth (Lepidoptera: Tortricidae)

Diapause in insects occurs in response to environmental cues, such as changes in photoperiod, and it is a major adaptation by which insects synchronize their activity with biotic resources and environmental constraints. For multivoltine agricultural insect pests, diapause initiation is an important consideration in management decisions, particularly toward the end of the growing season. The grape berry moth, Paralobesia viteana (Clemens), is the main insect pest affecting viticulture, and this insect responds to postsummer solstice photoperiods to initiate diapause. Because the range of grape berry moth extends from southern Canada to the southern United States, different populations are exposed to different photoperiodic regimes. We quantified the diapause response in grape berry moth populations from Arkansas, Michigan, New York, Pennsylvania, Texas, and Virginia, and observed latitudinal variation in diapause initiation. Populations from Michigan, New York, and Pennsylvania responded significantly different than those from Arkansas, Texas, and Virginia. We also observed, as a consequence of our experiments, that the timing of our laboratory studies influenced grape berry moth's response to photoperiod, ceteris paribus. Experiments that were conducted when grape berry moth would be naturally in diapause resulted in a significant higher proportion of diapausing pupae at photoperiods (i.e., >15 h) that generally do not induce diapause, suggesting that attention should be paid to the timing of behavioral and physiological experiments on insects. This relationship between photoperiod and diapause induction in grape berry moth across geographic regions will provide applicable knowledge to improve pest management decisions.     

昼夜变温幅度对小菜蛾不同发育阶段生活史性状的影响

【目的】昼夜变温幅度对昆虫的发育、存活、寿命、繁殖等核心生命活动有非常重要的影响。以往研究主要以恒温为主,温度设置不符合自然界中昼夜温度波动变化,无法明确温度波动幅度与恒温之间的生物效应差别。【方法】本研究采用了二步变温,模拟了不同的昼夜温度波动幅度(25±2℃,25±4℃,25±6℃,25±8℃,25±10℃和25±12℃)与相应恒温25℃,研究了不同变温幅度对十字花科世界性害虫小菜蛾Plutella xylostella不同生活史阶段中发育、存活、寿命与繁殖的影响。【结果】结果表明,变温幅度对小菜蛾不同阶段的发育、存活、寿命与繁殖影响存在显著差异。25±2℃,25±4℃和25±6℃对小菜蛾生活史性状的影响与恒温(25℃)相似,但25±10℃和25±12℃却产生了显著的负面影响。【结论】我们发现,较大的昼夜变温幅度显著影响小菜蛾不同阶段的发育、存活、寿命与繁殖,而适宜夜低温在一定程度上修复了日高温胁迫对小菜蛾的不利影响;并且认为昼夜变温幅度作为影响昆虫核心生命活动一种重要决定因素,必需纳入到昆虫种群数量预测模型中,才能真实地反映自然界中复杂变温模式对昆虫生态学效应的影响,才能提高昆虫田间发生预测预报的准确性。     

Food-mediated modulation of immunity in a phytophagous insect: An effect of nutrition rather than parasitic contamination

Inherent to the cost of immunity, the immune system itself can exhibit tradeoffs between its arms. Phytophagous insects face a wide range of microbial and eukaryotic parasites, each activating different immune pathways that could compromise the activity of the others. Feeding larvae are primarily exposed to microbes, which growth is controlled by antibiotic secondary metabolites produced by the host plant. The resulting variation in abundance of microbes on plants is expected to differentially stimulate the insect antimicrobial immune defenses. Under the above tradeoff hypothesis, stimulation of the insect antimicrobial defenses is expected to compromise immune activity against eukaryote parasites. In the European grape berry moth, Eupoecilia ambiguella, immune effectors directed towards microbes are negatively correlated to those directed towards eukaryotic parasites among host plants. Here, we hypothesize this relationship is caused by a variable control of the microbial community among host plants by their antibiotic metabolites. To test this hypothesis, we first quantified antimicrobial activity in berries of several grape varieties. We then measured immune defenses of E. ambiguella larvae raised on artificial diets in which we mimicked levels of antimicrobial activity of grape berries using tetracycline to control the abundance of growing microbes. Another group of larvae was raised on artificial diets made of berry extracts only to control for the effect of nutrition. We found that controlling microbe abundance with tetracycline in diets did not explain variation in the immune function whereas the presence of berry extracts did. This suggests that variation in immune defenses of E. ambiguella among grape varieties is caused by nutritional difference among host plants rather than microbe abundance. Further study of the effects of berry compounds on larval immune parameters will be needed to explain the observed tradeoff among immune system components.     

An improved model for determining degree-day values from daily temperature data

Although using hourly weather data offers the greatest accuracy for estimating growing degree-day values, daily maximum and minimum temperature data are often used to estimate these values by approximating the diurnal temperature trends. This paper presents a new empirical model for estimating the hourly mean temperature. The model describes the diurnal variation using a sine function from the minimum temperature at sunrise until the maximum temperature is reached, another sine function from the maximum temperature until sunset, and a square-root function from then until sunrise the next morning. The model was developed and calibrated using several years of hourly data obtained from five automated weather stations located in California and representing a wide range of climate conditions. The model was tested against an additional data-set at each location. The temperature model gave good results, the root-mean-square error being less than 2.0 °C for most years and locations. The comparison with published models from the literature showed that the model was superior to the other methods. Hourly temperatures from the model were used to calculate degree-day values. A comparison between degree-day estimates determined from the model and those obtained other selected methods is presented. The results showed that the model had the best accuracy in general regardless of the season. Received: 25 October 2000 / Revised: 2 July 2001 / Accepted: 2 July 2001     

Effects of Fluctuating Daily Temperatures at Critical Thermal Extremes on Aedes aegypti Life-History Traits

Background

The effect of temperature on insect biology is well understood under constant temperature conditions, but less so under more natural, fluctuating conditions. A fluctuating temperature profile around a mean of 26°C can alter Aedes aegypti vector competence for dengue viruses as well as numerous life-history traits, however, the effect of fluctuations on mosquitoes at critical thermal limits is unknown.

Methodology/Principal Findings

We investigated the effects of large and small daily temperature fluctuations at low (16°C) and high (35–37°C) mean temperatures, after we identified these temperatures as being thresholds for immature development and/or adult reproduction under constant temperature conditions. We found that temperature effects on larval development time, larval survival and adult reproduction depend on the combination of mean temperature and magnitude of fluctuations. Importantly, observed degree-day estimates for mosquito development under fluctuating temperature profiles depart significantly (around 10–20%) from that predicted by constant temperatures of the same mean. At low mean temperatures, fluctuations reduce the thermal energy required to reach pupation relative to constant temperature, whereas at high mean temperatures additional thermal energy is required to complete development. A stage-structured model based on these empirical data predicts that fluctuations can significantly affect the intrinsic growth rate of mosquito populations.

Conclusions/Significance

Our results indicate that by using constant temperatures, one could under- or over-estimate values for numerous life-history traits compared to more natural field conditions dependent upon the mean temperature. This complexity may in turn reduce the accuracy of population dynamics modeling and downstream applications for mosquito surveillance and disease prevention.     

A physiologically based approach for degree-day calculation in pest phenology models: the case of the European Corn Borer (<Emphasis Type="BoldItalic">Ostrinia nubilalis</Emphasis> Hbn.) in Northern Italy

Phenological models based on degree-day accumulation have been developed to support the integrated pest management of many insects. Most of these models are based on linear relationships between temperature and development, and on daily time step simulations using daily minimum and maximum temperatures. This approach represents an approximation that does not take into account the insect physiological response to temperature, and daily temperature fluctuations. The objective of this work has been to develop a phenological model for the European corn borer (ECB) based on the insect physiological response to temperature and running at an hourly time step. Two modeling solutions based on the same generic compartmental system have been compared: the first based on a physiologically based relationship between temperature and development, and using hourly derived temperatures as input (HNL modeling solution); and the second based on a linear relationship between temperature and degree-day accumulation and using daily temperature (DL modeling solution). The two approaches have been compared using ECB moth capture data from the Piemonte region in Northern Italy. The HNL modeling solution showed the best results for all the accuracy indicators. The DL modeling solution showed a tendency to anticipate ECB phenological development too early. This tendency is attributable to the linear relationship between temperature and development, which does not take into account (1) the decline of this relationship at high temperatures, and (2) the daily fluctuation of temperature. As a consequence, degree-days accumulation is accelerated in the DL modeling solution and the phenological development anticipated.     

Effects of climate change scenarios on Tempranillo grapevine (Vitis vinifera L.) ripening: response to a combination of elevated CO2 and temperature, and moderate drought

Greenhouse experiments were conducted to investigate the impact of predicted climate change (elevated CO₂, 700 μmol CO₂ mol^?1 air vs. ambient; elevated temperature, 28/18°C vs. 24/14°C, day/night; and partial irrigation, 40% of field capacity vs. well-irrigated) on grape berry quality characteristics during ripening. Grapevine (Vitis vinifera L. cv. Tempranillo) fruiting cuttings were used as experimental plant material. Climate change shortened the time between grape veraison and full maturity. At harvest time, many of the grape quality parameters determined were affected by the different grape maturity. The data were re-grouped according to total soluble solids to factor out changes due to the shortened time to maturity, and the effects on grape quality were then re-examined. Under current CO₂ and temperature conditions, partial irrigation decreased berry malic acid concentration and facilitated anthocyanins extractability. Elevated CO₂ and temperature decreased berry malic acid and total anthocyanins potential in well-irrigated plants and increased tonality index, irrespective of water availability. In partial irrigation conditions, elevated CO₂ and temperature hindered the anthocyanins extractability. In summary, results indicate that climate change (elevated CO₂, high temperature and partial irrigation) affects phenology and berry quality.     

Projecting insect voltinism under high and low greenhouse gas emission conditions

We develop individual-based Monte Carlo methods to explore how climate change can alter insect voltinism under varying greenhouse gas emissions scenarios by using input distributions of diapause termination or spring emergence, development rate, and diapause initiation, linked to daily temperature and photoperiod. We show concurrence of these projections with a field dataset, and then explore changes in grape berry moth, Paralobesia viteana (Clemens), voltinism that may occur with climate projections developed from the average of three climate models using two different future emissions scenarios from the International Panel of Climate Change (IPCC). Based on historical climate data from 1960 to 2008, and projected downscaled climate data until 2099 under both high (A1fi) and low (B1) greenhouse gas emission scenarios, we used concepts of P. viteana biology to estimate distributions of individuals entering successive generations per year. Under the low emissions scenario, we observed an earlier emergence from diapause and a shift in mean voltinism from 2.8 to 3.1 generations per year, with a fraction of the population achieving a fourth generation. Under the high emissions scenario, up to 3.6 mean generations per year were projected by the end of this century, with a very small fraction of the population achieving a fifth generation. Changes in voltinism in this and other species in response to climate change likely will cause significant economic and ecological impacts, and the methods presented here can be readily adapted to other species for which the input distributions are reasonably approximated.     

Predicting germination response of four cool-season range grasses to field-variable temperature regimes

Germination of non-dormant seeds under variable-temperature conditions can be predicted from constant-temperature germination data if it is assumed that instantaneous germination rate is independent of thermal history. Thermal-response models of this type have not been validated under simulated field-variable temperature conditions that vary in diurnal pattern, diurnal range and longer-term trends in mean–daily temperature. The purpose of this experiment was to evaluate germination response of thickspike wheatgrass (Elymus lanceolatus), bluebunch wheatgrass (Pseudoroegneria spicata), Sandberg bluegrass (Poa sandbergii) and bottlebrush squirreltail (Elymus elymoides) under both constant and field-variable temperature regimes in the laboratory. It was hypothesized that the thermal history assumption was valid and that constant-temperature data could be used to accurately estimate field-variable temperature response. Seeds were germinated at seven constant temperatures between 5 and 35°C, and under 18 variable-temperature regimes simulating six planting dates at three field sites. Predictions of germination time under variable-temperature conditions were accurate to within a fraction of 1 day up to a cumulative germination percentage of 70% for thickspike wheatgrass, 60% for bluebunch wheatgrass, 55% for Sandberg bluegrass and 70% for bottlebrush squirreltail. It was concluded that, for the variable-temperature regimes tested in this experiment, the thermal-history assumption was valid for earlier-germinating subpopulations.     

Carbaryl resistance in populations of grape berry moth (Lepidoptera: Tortricidae) in New York and Pennsylvania

We collected grape berry moth, Endopiza viteana (Clemens) (from cultivated and wild Vitis along Lake Erie in Pennsylvania and New York), and measured carbaryl susceptibility in first instars. A model of susceptibility was based on the concentration-mortality curve of laboratory-maintained colonies originating from wild Vitis with no prior history of carbaryl exposure, and a noncommercial vineyard with modest previous exposure to carbaryl. We estimated LC50 and LC90 for susceptible grape berry moth larvae at 45.4 and 2319 microg/ml, respectively. Bioassays on field-collected larvae from commercial vineyards in both states, where grape growers were abiding by current pest management guidelines for carbaryl use, revealed carbaryl resistance ratios from 7 to 71 at the LC50 level. With the loss or restriction of alternative chemical control tactics in the Food Quality Protection Act era, resistance management programs for grape berry moth should be immediately developed and implemented to regain the efficacy of this once effective insecticide and other related chemical compounds.     

Temperature data for phenological models

In an arid environment, the effect of evaporation on energy balance can affect air temperature recordings and greatly impact on degree-day calculations. This is an important consideration when choosing a site or climate data for phenological models. To our knowledge, there is no literature showing the effect of the underlying surface and its fetch around a weather station on degree-day accumulations. In this paper, we present data to show that this is a serious consideration, and it can lead to dubious models. Microscale measurements of temperature and energy balance are presented to explain why the differences occur. For example, the effect of fetch of irrigated grass and wetting of bare soil around a weather station on diurnal temperature are reported. A 43-day experiment showed that temperature measured on the upwind edge of an irrigated grass area averaged 4% higher than temperatures recorded 200 m inside the grass field. When the single-triangle method was used with a 10°C threshold and starting on May 19, the station on the upwind edge recorded 900 degree-days on June 28, whereas the interior station recorded 900 degree-days on July 1. Clearly, a difference in fetch can lead to big errors for large degree-day accumulations. Immediately after wetting, the temperature over a wet soil surface was similar to that measured over grass. However, the temperature over the soil increased more than that over the grass as the soil surface dried. Therefore, the observed difference between temperatures measured over bare soil and those over grass increases with longer periods between wettings. In most arid locations, measuring temperature over irrigated grass gives a lower mean annual temperature, resulting in lower annual cumulative degree-day values. This was verified by comparing measurements over grass with those over bare soil at several weather stations in a range of climates. To eliminate the effect of rainfall frequency, using temperature data collected only over irrigated grass, is recommended for long-term assessment of climate change effects on degree-day accumulation. In high evaporative conditions, a fetch of at least 100 m of grass is recommended. Our results clearly indicate that weather stations sited over bare soil have consistently higher degree-day accumulations. Therefore, especially in arid environments, phenology models based on temperature collected over bare soil are not transferable to those based on temperature recorded over irrigated grass. At a minimum, all degree-day-based phenology models reported in the literature should clearly describe the weather station site. Received: 25 October 2000 / Revised: 10 July 2001 / Accepted: 10 July 2001     

Genotype-environment interactions for insect growth in constant and fluctuating temperature regimes

Experiments on life history genetics are usually performed using constant temperature environments in the laboratory. However, the dynamics of insect growth can be influenced profoundly by daily fluctuations in temperature such as those which characterize field environments. We report here on experiments using different stocks and selected lines of a tropical butterfly, Bicyclus anynana, to examine whether genotype-environment interactions occur for three traits describing pre-adult growth. These traits were measured over two pairs of environments differing in mean temperature, each of which had a constant, and a cycling temperature regime. Development time, pupal weight and growth rate show genotype-environment interactions, especially at comparatively low average temperatures. Researchers should, therefore, take care when extrapolating from the form of genetic covariance matrices and ''trade-offs'' among life history traits found in constant temperature environments to those likely to occur in nature. <br>     

Potential effects of climate warming on the survivorship of adult Monochamus galloprovincialis

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On the canopy structure manipulation to buffer climate change effects on insect herbivore development

Insect pest development is often linearly related to air temperature, without taking into account the multiple interactions between the particular host plant and pest, the microclimatic conditions actually experienced by the insect, and the non-linear response of insect development rate to temperature. In this study, using an integrative biophysical model, we have investigated effects of both climatic and tree structure changes on the development of a phytophagous leaf mining moth (Phyllonorycter blancardella), taking into account the heterogeneous microclimatic conditions provided by its host plant, the domestic apple (Malus domestica), the larval body temperature rather than the ambient air temperature, and a non-linear development rate model. Hourly body temperature dynamics of larvae homogeneously dispersed in tree canopies were simulated from hourly meteorological conditions (medium IPCC climate change scenario) within the canopy of apple trees. To analyse the effect of tree architecture on leaf miner development, both pruned and unpruned trees, and one, two and three scaffold branched trees were used. Body temperature dynamics was used to compute larval development time and mortality following the non-linear developmental model for this insect. The results showed that tree pruning influences significantly larval development time and mortality. Nevertheless, the effects of manipulating tree structure on larval development and survival were relatively weak compared with the impact of chosen climate variations. This survey also showed that the variability in insect development time within a year and insect mortality change markedly with climatic variations, and highlights the importance of using non-linear rate curves and insect body temperatures instead of air temperature in forecasting models of climate-related insect pest outbreaks.     

Development and survival of the diamondback moth, Plutella xylostella (L.) (Lepidoptera: Yponomeutidae) as a function of temperature: effect on the number of generations in tropical and subtropical regions

The diamondback moth, Plutella xylostella (L.), is the most important pest of brassicaceous crops worldwide. Since temperature is the major abiotic factor influencing insect development and thermal requirements may vary among insect populations, it is important to know the effect of temperature on development and survival of a subtropical strain of P. xylostella. Development and survival of the diamondback moth was evaluated under seven constant temperatures ranging from 10°C to 35°C. Development was completed between 10°C and 32.5°C, but at 35°C all individuals died in the larval stage. Data were fitted to one linear and five nonlinear models. Considering as criteria the goodness of fit and the ability to estimate parameters of biological significance, the models Briere-1 and Briere-2 were the most adequate to describe the relationship between temperature and development of P. xylostella. The linear model demonstrated that P. xylostella required 312.5 degree-days above a lower threshold of 6.3°C to complete development. The degree-day model showed that the number of diamondback moth generations in the tropical region of Brazil is nearly twice the number in the subtropical region of the country. This result explains, at least in part, the higher population levels of this species in the tropical region of Brazil, and also demonstrates that P. xylostella is tolerant to a wide range of temperatures (6.1-32.5°C). Therefore, temperature cannot be considered a limiting factor for the occurrence of diamondback moth throughout the year in most regions of Brazil.     

Stage-specific control of grape berry moth, Endopiza viteana (Clemens) (Lepidoptera: Tortricidae), by selective and broad-spectrum insecticides

The insect growth regulators (IGRs) tebufenozide and methoxyfenozide and the broad-spectrum insecticides azinphosmethyl, carbaryl, and fenpropathrin were compared for their activity against adult, egg, and larval stages of the grape berry moth, Endopiza viteana (Clemens) (Lepidoptera: Tortricidae), under laboratory and vineyard conditions. Adult mortality was not affected by exposure to field-equivalent rates of tebufenozide or methoxyfenozide on grape clusters, whereas all the broad-spectrum compounds significantly reduced adult survival, compared with the untreated controls. Surviving adult moths laid significantly more eggs on berries treated with the IGRs than on berries treated with any of the broad-spectrum insecticides. Survival of these eggs through to late larval and pupal stages was significantly lower on methoxyfenozide-treated grapes than on untreated grapes, and no pupae were found when grapes were treated with azinphosmethyl or fenpropathrin. Neither of the growth regulator insecticides limited egg eclosion or larval development by E. viteana when insecticides were applied before egg laying, whereas broad-spectrum insecticides were effective against both eggs and neonates at this timing. When applied after egg eclosion, all insecticide treatments significantly reduced survival of grape berry moth larvae. Under vineyard conditions, berries with 1-d-old residues of tebufenozide or methoxyfenozide received more E. viteana eggs than berries treated with broad-spectrum compounds. After aging for 7 or 14 d, no significant effects on E. viteana survival were detected among treatments. Whereas broad-spectrum insecticides provide control of multiple life stages of E. viteana, integration of tebufenozide or methoxyfenozide into vineyard management programs for control of this pest will be most successful if applications are timed for egg hatch.     

Seasonal pattern of oviposition by the North American grape berry moth (Lepidoptera: Tortricidae)

The seasonal patterns of oviposition by the North American grape berry moth, Paralobesia viteana (Clemens) (Lepidoptera: Tortricidae) were monitored in juice grape (Vitis labrusca) vineyards in southwest Michigan. Egg deposition was recorded throughout the growing season at two vineyards in 2006, and at four vineyards from 2007 to 2009. In each vineyard, a random sample of 100 grape clusters was visually inspected twice‐weekly and the number of newly laid eggs was counted. We found that oviposition was continuous but variable throughout the season. Egg deposition started in early June coinciding with early grape bloom, continued at low level until mid‐ to late July, intensified in August close to veraison, and ended in September often before harvest. There were no consistent periods without oviposition that would indicate distinct generations. To determine the contribution of moth immigration into the vineyard to the pattern of oviposition, six grape plants located at the edge of a vineyard next to woods were covered with field cages and stocked with infested fruit. Oviposition and berry infestation were followed weekly on covered and exposed plants. Although higher numbers of eggs and infested berries were found on fruit of exposed vines than enclosed vines, egg deposition and berry infestation followed the same pattern in both treatments. This result indicates that the seasonal pattern of egg deposition is not dependent on immigration of grape berry moth of wild grape origin. The pattern of oviposition by grape berry moth described here contributes to the difficulty of controlling this pest using conventional insecticides with short residual activity.     

Sex ratios in naturally incubating Macrochelys temminckii nests,a species with type II temperature-dependent sex determination

The alligator snapping turtle, Macrochelys temminckii, exhibits type II temperature-dependent sex determination (TSD), wherein females are produced at high and low incubation temperatures. This TSD pattern is well studied at constant temperatures, but little work has focused on sex ratios in natural nests that experience daily and seasonal temperature fluctuations. We monitored nesting activity of reintroduced Macrochelys temminckii at Tishomingo National Wildlife Refuge in 2010–2011. Nests located prior to predation were excavated to determine clutch size and the eggs were reburied with a temperature data logger to collect nest temperatures. Overall, 24% of nests were protected with wire mesh prior to predation, and the average clutch size in intact nests was 22.4 eggs. Nest predation rates in the study population will likely approach 100% if nest protection efforts do not continue. Temperature profiles were used to compare estimated sex ratios using two methods—mean nest temperature during middle third of incubation and the degree-day model—to actual sex ratios in naturally incubated Macrochelys temminckii nests. The sex ratio in all 2010 recruits was female-biased (91.8% female); 2011 nests did not produce any hatchlings, likely the result of severe drought. The predicted sex ratios based on mean nest temperature and the degree-day model matched actual sex ratios in the warmer nests (0% male), but the degree-day model estimate proved more accurate in the cooler nest. A strongly skewed population sex ratio could become a threat to this reintroduced population if the strongly female-biased sex ratio in 2010 reflects a long-term trend.