Photoperiod- and temperature-dependent induction of larval diapause in a multivoltine bruchid, Bruchidius dorsalis

The wild bruchid beetle, Bruchidius dorsalis Fahraeus (Coleoptera: Bruchidae), has a multivoltine life cycle and overwinters in several developmental stages in the middle part of Japan. We investigated the incidence of diapause under different conditions of photoperiod (from L8:D16 to L16:D8) and temperature (at 20 °C and 24 °C). Our experiments revealed the following results: (1) B. dorsalis entered diapause at the final (late fourth) instar larva under short photoperiods, (2) the larval diapause incidence was dependent on temperature (critical photoperiods were 12.5 h at 20 °C and 12 h at 24 °C), (3) some individuals did not enter diapause under short-photoperiod conditions at 24 °C, and (4) the sensitive stages to the photoperiod were from the late egg stage to the early first instar larva. Based on these results, we discuss not only the evolution of a complex overwintering strategy inB. dorsalis but also the domestication process of stored-bean pests.     

Adaptation to the new land or effect of global warming? An age-structured model for rapid voltinism change in an alien lepidopteran pest

1. Hyphantria cunea Drury invaded Japan at Tokyo in 1945 and expanded its distribution gradually into northern and south-western Japan. All populations in Japan were bivoltine until the early 1970s, at which time trivoltine populations appeared in several southern regions. Presently, H. cunea exists as separate bivoltine and trivoltine populations divided around latitude 36 degrees . In the course of this voltinism change, the mean surface temperature in Japan rose by 1.0 degrees C. 2. To determine whether and how this temperature increase might be responsible for the voltinism change, we constructed an age-structured model incorporating growth speed driven by actual daily temperature and detailed mechanisms of diapause induction triggered by both daily photoperiod and temperature. 3. The simulation result suggests that both the acceleration of the growth speed and the prolongation of diapause induction are necessary to cause changes in voltinism, regardless of temperature increase. We concluded that the H. cunea population changed its life-history traits as an adaptation parallel with its invasion into the south-western parts of Japan. 4. Though the temperature increase had little effect on the fitness and heat stress in bivoltine and trivoltine populations, the trivoltine life cycle has become advantageous at least in marginal regions such as Tokyo.     

Divergent timing of egg-laying may maintain life history polymorphism in potentially multivoltine insects in seasonal environments

The length of the favourable season determines voltinism in insect populations. In some insects, there is variation in fecundity and timing of reproduction among females. If the length of the favourable season does not allow all offspring to develop into adults without diapause, the benefits of high early fecundity may outweigh the associated cost of low lifetime fecundity. We tested this by exploring mating frequencies of Pieris napi females along a latitudinal gradient in different generations. Pieris napi is a bivoltine butterfly, and genetically polyandrous females enjoy higher lifetime fecundity than monandrous ones. Polyandry is, however, coupled with a relatively low early fecundity. We found that monandrous females are more likely to produce an additional generation than polyandrous ones under conditions that allow production of only a partial summer generation. Monandrous females were also the first to emerge and slightly over-represented in the summer generation under conditions that allow the development of a complete summer generation. Further, a stochastic model shows that variation in the timing of reproduction between strategies is sufficient to explain the observed patterns. Thus, seasonality may counter-select against polyandry, or more generally against low early reproductive rate, and promote maintenance of polymorphism in life history strategies.     

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.     

Inclusion of host quality data improves predictions of herbivore phenology

Understanding the correspondence between ambient temperature and insect development is necessary to forecast insect phenology under novel environments. In the face of climate change, both conservation and pest control efforts require accurate phenological predictions. Here, we compare a suite of degree‐day models to assess their ability to predict the phenology of a common, oligophagous butterfly, the silver‐spotted skipper, Epargyreus clarus (Cramer) (Lepidoptera: Hesperiidae). To estimate model parameters, we used development time of eggs and larvae reared in the laboratory at six constant temperatures ranging from 8 to 38 °C and on two host plants of contrasting quality (kudzu and wisteria). We employed three approaches to determine the base temperature to calculate degree days: linear regression, modified reduced major axis regression, and application of a generic base temperature value of 10 °C, which is commonly used in the absence of laboratory data. To calculate the number of degree days required to complete a developmental stage, we used data from caterpillars feeding on high‐ and low‐quality hosts, both in the field and in the laboratory. To test model accuracy, we predicted development time of seven generations of larvae reared in the field on the same host plants across 3 years (2014–2016). To compare performance among models, we regressed predicted vs. observed development time, and found that r² values were significantly larger when accounting for host plant quality. The accuracy of development time predictions varied across the season, with estimates of the first two generations being more accurate than estimates of the third generation, when ambient temperatures dropped outside the range in which development rate and temperature have a linear relationship. Overall, we show that accounting for variation in host plant quality when calculating development time in the field is more important than the choice of the base temperature for calculating degree days.     

The life cycle of the giant water bug of northwestern Patagonian wetlands: the effect of hydroperiod and temperature regime

We analyzed the effect of hydroperiod and water temperature on the life cycle of the giant water bug Belostoma bifoveolatum in two wetlands of northwestern Patagonia, Argentina. In each wetland, we estimated adult and nymph abundance and monitored water depth and temperature throughout the study period. We determined the age structure of the giant water bug population in each wetland, and estimated the cumulative degree‐days (DD) needed for eggs to hatch and for nymphs to complete their development. Individuals of B. bifoveolatum colonized temporary wetlands at the beginning of spring when daylight lasts 12 h. The breeding period varied with hydroperiod length and showed both univoltine and bivoltine strategies, with a relatively constant breeding season. Egg‐bearing males appeared in October, carrying between 35 and 144 eggs per individual. Hatching success was high (~80% of eggs) and cumulative temperature for the hatching event was between 250 and 300 DD (which represents 3–4 weeks in nature), while complete development occurred between 800 and 1220 DD (~7–8 weeks). Individuals were more abundant in shallow and sunny patches of the wetlands, where the temperature was comparatively high, than in deeper or shaded sites. These results showed that hydroperiod duration and temperature could be good regulators of voltinism and development in B. bifoveolatum, driving the population structure of this giant water bug at the southern end of its distribution range.     

Tell me what you eat and I'll tell you when you fly: diet can predict phenological changes in response to climate change

Changes in phenology are correlated with climate change. However, we still struggle to understand the traits making species susceptible to climate change, and the implications of species' reactions for communities and food webs. Butterflies and moths are an ecologically important group that have shown pronounced phenological changes over the last decades. Tests using a > 150-year dataset from 566 European butterfly and moth species demonstrated that variation in phenological change was strongly related to traits describing plant-herbivore interactions (larval diet breadth, diet composition), and the life cycle. The results indicate that climate change related shifts in phenology are correlated with the seasonal availability and palatability of food plants. Lepidopterans feeding on herbaceous plants showed smaller shifts in flight periods but larger increases in voltinism than lepidopterans feeding on woody plants. Consequently, the effect of herbivorous lepidopterans may increase in herb-rich grassland ecosystems under warmer conditions, and not in forest ecosystems.     

Effect of temperature on the reproductive success,developmental rate and brood characteristics of Ips sexdentatus (Boern.)

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How do host plant use and seasonal life cycle relate to insect body size: A case study on European geometrid moths (Lepidoptera: Geometridae)

We used European geometrid moths (>630 species) as a model group to investigate how life history traits linked to larval host plant use (i.e., diet breadth and host-plant growth form) and seasonal life cycle (i.e., voltinism, overwintering stage and caterpillar phenology) are related to adult body size in holometabolous insect herbivores. To do so, we applied phylogenetic comparative methods to account for shared evolutionary history among herbivore species. We further categorized larval diet breadth based on the phylogenetic structure of utilized host plant genera. Our results indicate that species associated with woody plants are, on average, larger than herb feeders and increase in size with increasing diet breadth. Obligatorily univoltine species are larger than multivoltine species, and attain larger sizes when their larvae occur exclusively in the early season. Furthermore, the adult body size is significantly smaller in species that overwinter in the pupal stage compared to those that overwinter as eggs or caterpillars. In summary, our results indicate that the ecological niche of holometabolous insect herbivores is strongly interrelated with body size at maturity.     

Population consequences of diapause in a model system: the European corn borer

The diapause biology of the European corn borer (ECB), Ostrinia nubilalis (Hübn.), is described based on natural and controlled environment studies of feral and lab-reared ECB's in North Carolina (NC). The diapause response is described as a function of photophase (h of light/day) as well as a function of larval age (instar) at onset of diapause-inducing conditions. A critical photophase of 14.4 h and a critical mean larval instar of 3.3 is found in the lab studies and supported by three years of insectary studies. Seven years of black light trapping of ECB moths in Goldsboro, NC, revealed the likelihood of up to four moth flights/year.Information about the diapause biology of this insect is used to explain both the number of flights and the relative magnitude of the final moth flights. On average, the majority of ECB lineages pass through three generations/year with early maturing ECB's producing a significant and predictable fourth generation. The timing and magnitude of the fourth flight can be partly explained on the basis of the critical photophase and the timing and age structure of previous ECB generations. In most years, the fourth flight is smaller than the third due to the majority of the fourth generation's predisposition towards diapause. However, in at least one case (1977), the fourth flight was unusually large and could be predicted by slight temporal shifts in the previous three flights resulting in the majority of the fourth generation larvae averting diapause. The value of the ECB-diapause interaction as a model system for the explanation and prediction of dynamic phenological events is discussed.

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Résumé La diapause d'O. nubilalis Hubn. est décrite d'après des études en conditions naturelles et programmées de souches sauvages ou élevées au laboratoire en Caroline du Nord. La diapause est décrite comme une fonction de la photophase (heures de jour/24 heures) et du stade larvaire au début des conditions inductrices de la diapause. Une photophase critique de 14,4 h et un stade critique larvaire de 3,3 ont été établis au laboratoire après 3 ans d'études en insectarium. 7 ans de piégeage à la lumière noire à Goldsboro, ont montré la vraisemblance de l'existence de 4 vols par an. Les données sur la diapause de cet insecte sont utilisées pour expliquer tant le nombre de vols que l'importance relative des derniers vols. En moyenne, la majorité des lignées ont 3 générations par an, O. nubilalis précoces produisant une quatrième génération conséquente et prédictible. La data et l'importance du 4ème vol peuvent être partiellement expliquées d'après la photophase critique, et la data et la structure en âge des générations précédentes. La plupart des années, le 4ème vol est moins important que le 3ème par suite de la prédisposition à la diapause de la majorité de la 4ème génération. Cependant, dans un cas au moins, en 1977, le 4ème était anormalement important et pouvait être prédit par de faibles changements temporels dans les 3 précédents vols, détournant de la diapause la majorité des chenilles de 4ème génération. La discussion porte sur la valeur du modèle fourni par la diapause de O. nubilalis pour expliquer et prédire la dynamique des évéments phénologiques.