Use of sperm precedence to infer the overwintering cost of insecticide resistance in the Colorado potato beetle

1 Resistance to insecticides is a model system for studying adaptation. Although selection for resistance is always strong in areas and seasons where populations are exposed to insecticides, costs of resistance, which may only be expressed in the absence of insecticide use, will shape how quickly resistance will evolve.
2 We used sperm precedence to measure the shifts in resistance to imidacloprid in a natural population during winter diapause in the Colorado potato beetle, Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae). Because adult beetles overwinter with viable autumn sperm, but sperm from spring matings take precedence over stored sperm, we used the difference in resistance of springmated and autumn-mated overwintered females to estimate the shift in resistance during the winter. Offspring of autumn-mated females were 2.7- or 2.5-fold more resistant compared with offspring from spring-mated females in two replicate trials.
3 We also measured the resistance of late summer and spring emergent adults in fields treated and untreated with imidacloprid in the first year. Adults from the treated field were 13.7-fold more resistant and adults from the untreated field were 2.6-fold more resistant compared with the next spring's emergers.
4 These large costs of resistance observed in the field and inferred from resistance declines during diapause help to explain the observation that imidacloprid resistance has increased only slowly over the decade of widespread use against this species, and how insecticide resistance in general can cycle annually.     

中华稻蝗长沙种群的生活史及其卵滞育的进化意义

中华稻蝗Oxya chinensis为重要的水稻害虫,在我国除青海、西藏、新疆、内蒙古等未见报道外,南起海南北至东北均有分布,在许多的分布区域1年发生1代。为探索中华稻蝗长沙种群的生活史及其季节适应特征,通过野外和实验室的研究,调查了其发生代数、孵化率和卵滞育率的季节变化及越冬卵的存活率。结果显示,中华稻蝗长沙种群为1年2代和1年1代混合发生:第1代卵产卵后大部分孵化为若虫而1年完成2代,但亦有19.4% -4.1%的卵不孵化而1年只能完成1世代。第1代成虫于6月上旬至8月上旬羽化,6月下旬至8月中旬产卵;第2代若虫于7月初开始孵化,9-10月羽化为成虫,10月上旬至11月下旬产卵。在室外自然条件下,中华稻蝗长沙种群6-8月(第1代)和10-11月(第2代)所产卵块均为部分滞育,滞育率为30%左右,皆无显著差异。然而,其卵滞育率在12月以后显著降低,仅为6.6%或以下,卵滞育快速地得以解除。因此,包括非滞育卵和滞育解除卵,中华稻蝗长沙种群的越冬卵皆以非滞育状态度过其后的寒冷季节。即使是遭遇长江流域2007年末至 2008年初异常寒冷的冬季,在长沙地区越冬后其卵的存活率亦在98%以上。非滞育状态的中华稻蝗长沙种群越冬卵完全能安全地越冬,其滞育的发生并非是为了提高其耐寒性而安全度过不适环境。并探讨了中华稻蝗长沙种群卵滞育的进化意义。     

Bi- and trivoltine complex life cycles in a Kanto (Japan) population of a wild bruchid Kytorhinus sharpianus

The partial trivoltinism and overwintering of Kytorhinus sharpianus Bridwell (Coleoptera: Bruchidae) was studied in the Kanto district, Japan. The later in the summer eggs were laid by the first-generation adults, the higher was the incidence of larval diapause in the second generation. The incidence of diapause also fluctuated between years, influencing the abundance of third-generation larvae. A relatively large proportion of third-generation larvae did not attain the diapause stage by the beginning of winter. The diapause development of larvae in diapause was completed by mid-January. Immature larvae of the third generation also overwintered and emerged as adults in the spring.     

Phenology and life history of the blowfly Calliphora vicina in stockfish production areas

Calliphora vicina Robineau‐Desvoidy (Diptera: Calliphoridae) causes yearly losses of 1–2 million Euros to the stockfish industry in Lofoten, Norway. To develop an efficient management program, knowledge of its life cycle and phenology in production areas is needed. Cohort studies in a simulated Lofoten climate showed that field abundance peaks of adults in early spring and midsummer can be explained by a cohort originating from stockfish and its subsequent generations. Laboratory simulations with normal, increased, and decreased Lofoten temperatures indicate that C. vicina overwinter as a mix of larvae, pupae, and adults, and a temperature change of ± 2 °C significantly influences reproductive timing, reproductive output, and female mortality. Flies originating from stockfish reproduced during the first summer when temperatures were increased 2 °C above normal. At lower temperatures, the reproductive investment was low or absent during the first summer and the adult flies entered the winter in a diapausing state. Most offspring produced during the first summer and autumn developed continuously without maternally induced diapause, pupated during the winter, and hatched in the early spring to co‐occur with their parent generation during stockfish production. Calliphora vicina showed flexibility in reproductive efforts and overwintering strategies. The high proportion of adults overwintering compared with the commonly used larval diapause strategy might be interpreted as an adaptation to exploit the stockfish resource. The majority of female C. vicina that cause damage to stockfish likely developed on fish dried the previous year, and a continuous year‐long trapping is recommended to decimate the population.     

How does Drosophila melanogaster overwinter?

In temperate regions low temperatures seem to be the most restrictive factor for survival of Drosophila natural populations, which depends on the capacity of one or more developmental stages to resist unfavourable winter conditions. In this study we have attempted to answer the question of how D. melanogaster overwinters under natural temperature conditions. Only adults overwintered and no diapause was observed in any developmental stage. Thus, developmental duration becomes a decisive component with respect to overwintering potential and, therefore, the preadult stages are unlikely to overwinter. Possible evolutionary steps in adaptation to cold regions are discussed.     

Presence of three diapauses in a subtropical cockroach: control mechanisms and adaptive significance

Abstract.  Diapause is common among insects and is regarded as an adaptive response to periodic occurrence of adverse conditions. It occurs at a particular developmental stage, typically only once in a lifetime. However, little is known about the details of the control mechanism of life cycles with multiple diapauses in insects. In this study, a complex 2-year life cycle with three types of diapause is reported in a subtropical cockroach, Symploce japonica : a winter diapause in mid-nymphal instars, a summer diapause in later nymphal instars, and a reproductive diapause is reported in the adult stage. Nymphal development was extremely slow either at short days (winter diapause) or long days (summer diapause). Nymphs in summer diapause matured rapidly when transferred from long days to short days, indicating that seasonal changes in day-length are the pivotal factor in the control of this life cycle. It is proposed that the main significance of winter diapause in this subtropical species is to enable the nymphs to survive the mild winter successfully with reduced energy demand, and that of summer diapause is to delay adult emergence until late in the autumn for successful induction of the following adult diapause. Adults do not emerge until shortly before winter, yet the presence of diapause in the adult stage does not simply appear to be a response to cope with the winter conditions but, instead, ensures that reproduction will occur early the next year, before summer, because reproduction is greatly hampered at high temperature.     

Seasonal development and variation in abundance among four annual flight periods in a butterfly: a 20‐year study of the speckled wood (Pararge aegeria)

Insects typically spend the winter in a species‐specific diapause stage. The speckled wood butterfly, Pararge aegeria, is unique in having two alternative diapause stages, hibernating as larvae or pupae. In southern Sweden this creates a seasonal flight pattern with four annual adult flight periods: the first in May (pupal diapause), the second in June (larval diapause), and the third and fourth directly developing offspring generations in July and August, respectively. We address the raison d'être of the two diapause pathways by (1) outdoor rearing of cohorts, and (2) performing transect censuses throughout the season for 20 years. We contend that an early start of next season provides a benefit accruing to pupal diapause; conversely, a large proportion of the offspring from adults of the fourth flight peak are unable to reach the pupal stage before winter, providing a benefit accruing to larval winter diapause. The results obtained show that the two hibernation pathways are unlikely to be genetically distinct because of a strong overlap between the two offspring generations, and because sibling offspring from the third and fourth flight periods are likely to choose either of the two hibernation pathways, thereby resulting in a genetic mixing of the pathways. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 635–649.     

Seasonal development of the lacewing Chrysopa dorsalis Burmeister (Neuroptera, Chrysopidae) in the forest-steppe zone of Russia

Seasonal adaptations of Chrysopa dorsalis Burmeister were studied in Belgorod Province of Russia (50°N, 36°E). It was found that in the forest-steppe zone two generations may be produced, but from 76 to 100% of the prepupae of the first generation entered diapause, about 40% of them having an obligate diapause. In the rest of the population, seasonal development was controlled by photoperiod and temperature. The critical day length was about 17 h at 20°C. Some prepupae were found to require two or three years to complete the diapause. This prolonged diapause seems to be a permanent element of the life cycle of the studied species. The proportion of the prepupae that remain in diapause after the first winter positively correlated with the percentage of diapausing prepupae of the first generation in the previous season (r = 0.8). The offspring from eggs laid by females simultaneously collected under the natural conditions emerged in successive years, and the tendency to enter a long-term diapause was not inherited. The rate of the larvae development and the prepupae weight were not different in individuals which had one-year-long and prolonged diapause. However, the weight loss due to dehydration and respiration during the first winter was slightly lower in the prepupae with the prolonged diapause. The expression of the prolonged diapause was independent of the photoperiod during the diapause induction, but it was increased at a high temperature (28°C). Probably, the prolonged diapause increases polymorphism of local Ch. dorsalis populations, ensuring their survival under unpredictable conditions.     

Diapause and implications for control of Penthaleus species and Halotydeus destructor (Acari: Penthaleidae) in southeastern Australia

Blue oat mites, Penthaleus spp., and redlegged earth mites, Halotydeus destructor (Tucker) are major winter pests of a variety of crops and pastures. In southern Australia earth mites exhibit a facultative egg diapause to survive unfavorable summer conditions. The initiation of diapause egg production in earth mites was investigated using field and shade-house experiments. Species differed in their timing of diapause. H. destructor mainly produced diapausing eggs towards the end of the active mite season in spring, although small numbers were also produced in winter. In contrast, Penthaleus major (Dugés) produced diapause eggs almost immediately after emergence in autumn and continued producing these eggs throughout the season. Penthaleus falcatus (Qin and Halliday) also produced diapause eggs in early winter, although the first appearance of these eggs was slightly later in the season than for P. major. The diapause response of an undescribed species was also somewhat later than in P. major and P. falcatus, but earlier than in H. destructor. Electrophoresis of P. major samples indicated that clones of this parthenogenic species may differ in their timing of diapause egg production, providing another potential selective factor contributing to the maintenance of clonal diversity within this group. The results highlight the importance of determining species composition when devising control strategies for earth mite outbreaks.     

Over-wintering stage polymorphism of a bruchine beetle: geographical variation in optimal diapause strategy

Abstract.  1. Most temperate insects undergo diapause at a specific developmental stage to overcome severe winters. The bruchine beetle Bruchidius dorsalis in a warmer region in Japan, however, has diverse over-wintering stages – non-diapausing larvae, diapausing larvae, and diapausing adults, whereas in a cooler region, it over-winters only as the final instar larva or adult in diapause.
2. The geographical pattern of over-wintering stages in 12 populations was investigated over a wide range along the mainland of Japan. It revealed that a distinct geographical pattern of over-wintering stages exists in relation to temperature conditions. In regions with warmer climates, the proportion of non-diapausing larvae increased and B. dorsalis had a more complex over-wintering stage structure.
3. Life cycles were also compared between two areas of Japan by field experiments. In the cooler area, the first generation over-wintered in the diapausing larval or adult stage. Conversely, in the warmer area, diapause was induced later and some of the first-generation adults produced second-generation offspring before over-wintering.
4. Based on the geographical cline of climates and the differences in cold hardiness among stages, we can demonstrate that the over-wintering stage variation among and within populations results from an adaptive timing of diapause induction in each region, because the late larval or adult diapauses protect pupae or eggs – which unlike other stages are not cold hardy – from being produced late in the season.     

Optimal switching to diapause in relation to the onset of winter

A model is presented that specifies optimal switching times for the induction of diapause relative to the onset of winter in a deterministic environment. Fitness is defined as an individual's contribution to the overwintering population in diapause at the time of the first hard frost. The fitness of a nondiapausing female is determined by the switching times of her offspring. If age-specific fecundity and survivorship do not change significantly from generation to generation, the optimal switching time precedes the onset of winter by a constant amount of time equal to the age of first reproduction plus the time to produce one offspring plus the difference in ages between the sensitive age and the diapause age. This result is independent of the time at which the original female began to reproduce. However, if fecundity or survivorship decreases toward the end of the season, the optimal switching time depends upon the time when she began to reproduce and should be more conservative by an amount of time that can be computed. Possible tests of the model are proposed.     

Seasonal variation in embryonic diapause of the striped ground cricket, Allonemobius fasciatus

ABSTRACT. The incidence and intensity of embryonic diapause in the striped ground cricket, Allonemobius fasciatus DeGeer (Gryllidae), is influenced by the date of oviposition and the incubation temperature. At 20C, embryos develop normally until the end of the appendage-formation stage when diapause occurs. The intensity of 'winter' diapause at this stage is greatest in eggs laid early in the season. Incubation at higher temperature induces a different and earlier 'summer' diapause. The higher the temperature the earlier the stage at which this developmental suppression is imposed. At 27 and 30C, some individuals develop without any interruption and hatch within 20 days. The proportion of these fast-developing eggs is low in summer but increases towards the end of the laying season. In nature, this species is univoltine and precocious hatching before winter is unlikely.
When embryos diapausing at an early stage at 27C are transferred to low temperatures, they resume development and enter winter diapause. The time required to reach the winter-diapause stage after transfer is inversely related to the temperature (30-30C). At 30C, sunmer diapause is neither induced nor maintained.     

Patterns of diapause frequency and emergence in swede midges of southern Ontario

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High cold tolerance through four seasons and all free-living stages in an ectoparasite

Off-host stages of temperate parasites must cope with low temperatures. Cold tolerance is often highest in winter, as a result of diapause and cold acclimation, and low during the active summer stages. In some blood-feeding ectoparasites, offspring provisioning determines cold tolerance through all the non-feeding, off-host stages. Large size increases survival in the cold, but so far seasonal variation in within-female offspring size has not been associated with offspring cold tolerance. The deer ked (Lipoptena cervi) reproduces on cervids from autumn to spring. Newborn pupae drop off the host, facing frosts without any acclimation. We examined cold tolerance through 4 seasons and from birth to adulthood by means of short- and long-term frost exposure. We expected females to produce more tolerant offspring in winter than in spring. Large spring pupae survived prolonged frosts better than did small winter pupae. Thus more tolerant offspring were not produced when the temperature outside the host is at its lowest. Unexpectedly, the freezing points were -20 °C or below all year round. We showed that high cold tolerance is possible without acclimation regardless of life stage, which presumably correlates with other survival characteristics, such as the starvation resistance of free-living ectoparasites.     

Interpopulation variation of hibernal-aestival-diapause in the egg parasitoid wasp Anagrus takeyanus: adaptation to seasonal host-plant alternation of the tingid host, Stephanitis takeyai

The egg parasitoid wasp, Anagrus takeyanus Gordh & Dunbar (Hymenoptera: Mymaridae), is one of the few natural enemies of the andromeda lace bug, Stephanitis takeyai Drake & Maa (Heteroptera: Tingidae). Studying this wasp in two habitats, I found that at Kyoto, where the bug alternates its host plants seasonally, most wasp individuals in the overwintering eggs of the bug entered summer diapause immediately after winter diapause. However, at Nara, where the bug does not alternate its host plant, the wasp proved to have no summer diapause. Changes in the parasitization rate between generations differed at the two study sites. In Kyoto, the overwintered generation suffered the highest parasitism among the three generations of the year. In contrast, the first and second generations suffered higher parasitization rates in Nara. This difference in level of parasitization between the two study sites corresponded to the different diapause regime of the wasp and the seasonal population trends of the lace bug. Laboratory experiments showed that short photoperiod in combination with low temperature could terminate the long diapause at the time the non-aestivating individuals terminate winter diapause. As a life cycle without host plant alternation is likely to be an ancestral character of the bug, the aestivation of the wasp is thought to have been appended to overwintering as an adaptation to the evolution of seasonal host plant alternation in the lace bug.     

Asymmetric life-history decision-making in butterfly larvae

In temperate environments, insects appearing in several generations in the growth season typically have to decide during the larval period whether to develop into adulthood, or to postpone adult emergence until next season by entering a species-specific diapause stage. This decision is typically guided by environmental cues experienced during development. An early decision makes it possible to adjust growth rate, which would allow the growing larva to respond to time stress involved in direct development, whereas a last-minute decision would instead allow the larva to use up-to-date information about which developmental pathway is the most favourable under the current circumstances. We study the timing of the larval pathway decision-making between entering pupal winter diapause and direct development in three distantly related butterflies (Pieris napi, Araschnia levana and Pararge aegeria). We pinpoint the timing of the larval diapause decision by transferring larvae from first to last instars from long daylength (inducing direct development) to short daylength conditions (inducing diapause), and vice versa. Results show that the pathway decision is typically made in the late instars in all three species, and that the ability to switch developmental pathway late in juvenile life is conditional; larvae more freely switched from diapause to direct development than in the opposite direction. We contend that this asymmetry is influenced by the additional physiological preparations needed to survive the long and cold winter period, and that the reluctance to make a late decision to enter diapause has the potential to be a general trait among temperate insects.     

Diapausing fertilized adults

Summary A lake population of the cyclopoid copepod Cyclops strenuus Fischer had a 1-year life cycle including a diapause from June to January. Dormant stages consisted of fertilized females, not previously described as a diapause stage, and a small fraction of cop V. The longevity of the sperm in a diapausing female was at least 6–7 months. Pre-fertilization is assumed to be an adaptive mechanism to assure fertilization when male abundance is low. This life cycle pattern also gives a shorter time-lag between diapause break and offspring development, and also the low total adult population exerts a lower predation pressure on their offspring. The flexibility of life cycle patterns of copepods is emphasized.     

SEASONAL DECLINES IN OFFSPRING FITNESS AND SELECTION FOR EARLY REPRODUCTION IN NYMPH-OVERWINTERING GRASSHOPPERS

In this study, I examine the effects of natural and experimentally induced variation in life cycle timing on offspring fitness in Arphia sulphurea and Chortophaga viridifasciata, to understand the selective pressures shaping phenology in these two species of nymph-overwintering grasshoppers. Because these species lack embryonic diapause, hatching varies over a two month range under natural conditions. I used a cold treatment to delay hatching of some egg pods and extend the natural range of hatching dates. Due to the shorter time for growth and poorer growing conditions late in the fall, late-hatching nymphs of both species grew to a smaller size before winter and suffered higher overwinter mortality, compared to early nymphs. In addition, late nymphs that did survive the winter became reproductive later in the following year's breeding season. Size- dependent mortality of offspring during the winter is a strong selective pressure favoring early reproduction in these species. Female adult life history traits appear responsive to the seasonal declines in offspring fitness, in that late-maturing females began reproducing sooner after adult maturation and reproduced at a more rapid rate, even at the expense of having shorter adult longevity and producing fewer total egg pods. Experimental manipulations were crucial in understanding the fitness consequences of intrapopulation variation in the timing of specific life-cycle events for these species.     

Environmental control of the seasonal life cycle of a zoophytophagous mirid, <Emphasis Type="Italic">Adelphocoris triannulatus</Emphasis> (Hemiptera: Miridae)

Adelphocoris triannulatus (Stål) (Heteroptera: Miridae) is a zoophytophagous bug that inhabits grasslands and crop fields, and is widely distributed in eastern Asia. In this study, the seasonal occurrence and environmental factors controlling the seasonal life cycle of this bug were investigated in a local population in southern Kyushu, Japan. When insects were reared under different photoperiods at 20 and 25 °C, they produced diapause or nondiapause eggs depending on the photoperiod: when reared under short-day photoperiods (≤?13 h/day), females laid diapause eggs, whereas they laid nondiapause eggs when reared under long-day photoperiods (≥?14 h/day). Because the developmental duration of the pre-adult stages and maturation (minimum duration of one generation) is estimated to be approximately 30 days at summer temperatures, these results suggest that this bug has a multivoltine seasonal life cycle. In agreement with the laboratory results, only nondiapause eggs were produced until mid-August in the field, and an increasing proportion of diapause eggs were produced from early September. In addition to these findings, field sampling from spring to autumn in 2009 indicated that three or four generations were produced in one year. These results suggest that egg diapause plays a role in preventing nymphs of A. triannulatus from developing in winter, and thereby ensures the survival of the population in adverse seasons.