Host‐plant quality adaptively affects the diapause threshold: evidence from leaf beetles in willow plantations

1. Voltinism of herbivorous insects can vary depending on environmental conditions. The leaf beetle Phratora vulgatissima L. is univoltine in Sweden but will sometimes initiate a second generation in short‐rotation coppice (SRC) willow plantations. 2. The study investigated whether increased voltinism by P. vulgatissima in plantations can be explained by (i) rapid life‐cycle development allowing two generations, or (ii) postponed diapause induction on coppiced willows. 3. In the field, no difference was found in the phenology or development of first‐generation broods between plantations (S. viminalis) and natural willow habitats (S. cinerea). However, the induction of diapause occurred 1–2 weeks later in SRC willow plantations. 4. Laboratory experiments indicated no genetic difference in the critical day‐length for diapause induction between beetles originating from plantations and natural habitats. Development time was unaffected by host‐plant quality but critical day‐length was prolonged by almost an hour when the beetles were reared on a non‐preferred willow species (S. phylicifolia). When reared on new leaves from re‐sprouting shoots of recently coppiced willow plants, diapause incidence was significantly less than when the beetles were reared on mature leaves from uncoppiced plants. 5. The study suggests that P. vulgatissima has a plastic diapause threshold influenced by host‐plant quality. The use of host‐plant quality as a diapause‐inducing stimulus is likely to be adaptive in cases where food resources are unpredictable, such as when new host‐plant tissue is produced after a disturbance. SRC willows may allow two beetle generations due to longer growing seasons of coppiced plants that grow vigorously.     

Phenology of brown marmorated stink bug described using female reproductive development

Temperature‐based degree‐day models describe insect seasonality and to predict key phenological events. We expand on the use of a temperature‐based process defining timing of reproduction through the incorporation of female reproductive physiology for the invasive pentatomid species Halyomorpha halys, the brown marmorated stink bug. A five‐stage ranking system based on ovary development was able to distinguish between the reproductive statuses of field‐collected females. Application of this ranking method described aspects of H. halys’ seasonality, overwintering biology, and phenology across geographic locations. Female H. halys were collected in the US from NJ, WV, NC, OR, and two sites in PA in 2006–2008 (Allentown, PA only) and 2012–2014. Results identify that H. halys enters reproductive diapause in temperate locations in the fall and that a delay occurs in developmental maturity after diapause termination in the spring. Modification of the Snyder method to identify biofix determined 12.7‐hr photoperiod as the best fit to define initiation of reproduction in the spring. Applying the biofix, we demonstrated significant differences between locations for the rate at which the overwintering generation transition into reproductive status and the factors contributing to this difference require further study. For example, after including abiotic variables influencing development such as temperature and photoperiod (critical diapause cue), reproduction occurred earlier in OR and for an extended period in NJ. This data describe a method to investigate insect seasonality by incorporating physiological development across multiple regions that can clarify phenology for insects with overlapping generations.     

Differences in photoperiod‐induced diapause plasticity among different populations of the bark beetle Ips typographus and its predator Thanasimus formicarius

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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.     

The forgotten season: the impact of autumn phenology on a specialist insect herbivore community on oak

1. Variation in spring phenology – like tree budburst – affects the structure of insect communities, but impacts of autumn phenology have been neglected. Many plant species have recently delayed their autumn phenology, and the timing of leaf senescence may be important for herbivorous insects. 2. This study explored how an insect herbivore community associated with Quercus robur is influenced by variation in autumn phenology. For this, schools were asked to record, across the range of oak in Sweden, the autumn phenology of oaks and to conduct a survey of the insect community. 3. To tease apart the relative impacts of climate from that of tree phenology, regional tree phenology was first modelled as a function of regional climate, and the tree‐specific deviation from this relationship was then used as the metric of relative tree‐specific phenology. 4. At the regional scale, a warmer climate postponed oak leaf senescence. This was also reflected in the insect herbivore community: six out of 15 taxa occurred at a higher incidence and five out of 18 taxa were more abundant, in locations with a warmer climate. Similarly, taxonomic richness and herbivory were higher in warmer locations. 5. Trees with a relatively late autumn phenology had higher abundances of leaf miners (Phyllonorycter spp.). This caused lower community diversity and evenness on trees with later autumn phenology. 6. The findings of the present study illustrate that both regional climate‐driven patterns and local variation in oak autumn phenology contribute to shaping the insect herbivore community. Community patterns may thus shift with a changing climate.     

Effect of photoperiod and allatectomy on reproduction and (second) diapause induction in post-diapause adult females of the Colorado potato beetle,Leptinotarsa decemlineata Say

Summary

The effect of photoperiod and allatectomy on the rates of oviposition and juvenile hormone (JH) synthesis was studied in female beetles after the break of adult diapause. The results indicate that after diapause, photoperiod has no effect on oviposition rates. However, the photoperiodic effect on diapause induction is restored within a few days after the break of the first diapause. Allatectomy performed within one day after diapause did not induce a second diapause, whereas about 40% of the beetles allatectomized after 3 or more days responded by entering a second diapause. Oogenesis in the operated animals was never completely prevented under long and short day conditions. These preliminary results suggest a change in the sensitivity of post-diapause beetles to photoperiodic and endocrine manipulation as compared with pre-diapause insects.     

Large‐scale simulation of temperature‐dependent phenology in wintering populations of Bactrocera oleae (Rossi)

To implement Area‐Wide Pest Management protocols at a regional scale (Liguria, northern Italy), egg deposition and adult flight of olive fruit fly, Bactrocera oleae, were monitored during 2009, 2010 and 2011. The consequence of complete generation in late winter – early spring was also examined. The reliability of a degree‐day model was tested to simulate the insect cycle, starting from October oviposition and considering a 379.01°C cumulative degree‐day (CDD) needed to complete development. The model was validated and then used to simulate olive fruit fly phenology in the region of Liguria, using a GIS approach and the agrometeorological network in the region. The output of the CDD model was mapped with two different spatialization modelling techniques, geostatistical autocorrelation and regression correlation, and altitude, aspect and distance from the sea were assessed as elements of variability. The regression correlation model provided a more accurate indication of B. oleae diversity at the local scale than the geostatistical autocorrelation model and identified the high spatial climatic variability of Liguria. The potential application of the distribution of days after oviposition and prediction error maps in support of pest management planning is discussed.     

Variation in two phases of post‐winter development of a butterfly

The temporal aspects of life cycle characteristics, such as diapause development, are under strong selection in seasonal environments. Fine‐tuning of the life cycle may be particularly important to match the phenology of potential mates and resources as well as for optimizing abiotic conditions at eclosion. Here, we experimentally study the spring phenology of the orange tip butterfly, Anthocharis cardamines, by analysing post‐winter pupal development in three populations along a latitudinal cline in each of Sweden and the United Kingdom. These countries differ substantially in their seasonal temperature profile. By repeatedly recording pupal weights, we established that post‐winter development has two separate phases, with a more rapid weight loss in the second phase than in the first, likely corresponding to a ramping up of the rate of development. Variation in the duration of the first phase contributed more strongly than the second phase to the differences in phenology between the localities and sexes. We found that insects from Sweden had a faster overall rate of development than those from the United Kingdom, which is consistent with countergradient variation, as Sweden is colder during the spring than the United Kingdom. Similar trends were not observed at the within‐country scale, however. A cogradient pattern was found within Sweden, with populations from the north developing more slowly, and there was no clear latitudinal trend within the United Kingdom. In all localities, males developed faster than females. Our results point to the importance of variation in the progression of post‐winter development for spring phenology.     

Seasonal changes in the reproductive and life-history traits of Chrysomela populi L. (Coleoptera: Chrysomelidae)

Seasonal changes in the reproductive and life‐history traits of Chrysomela populi L. were investigated in Shobara (34°52′N, 133°01′E), Japan, in 2004 and 2005. Larvae were hatched and reared under natural photoperiod and temperature in 8 different periods between late April and early September in 2004. The incidence of adult diapause increased with progression of the season in 2004. The critical timing of diapause induction, as defined by 50% of females entering diapause, was estimated to occur between late July and early August. No effects of season on the survival rate from hatching to adult emergence, percentage ovipositing females and percentage females that deposited viable eggs were clearly detected. Adult weight at emergence fluctuated seasonally, which was probably caused by air temperature. No effect of season on the oviposition period was observed among females that averted diapause. Females that entered diapause in 2004 started oviposition from the first day of May in 2005. Reproductive output was significantly greater in diapause females than in non‐diapause females. This increase in the reproductive output of diapause females was caused by elongation of the oviposition period and consequently by enlargement of the number of egg clutches deposited. These results suggest that the seasonal timing of diapause induction in females of C. populi would be affected by various risks and benefits to reproduction and survival.     

Effects of photoperiod and temperature on reproductive diapause in Ophraella communa (Coleoptera: Chrysomelidae), a potential biocontrol agent against Ambrosia artemisiifolia

Abstract To investigate the seasonal adaptation strategies of Ophraella communa to new habitats, the effects and regulation mechanisms of photoperiod and temperature on the reproductive diapause in a population collected from Changsha, Hunan were examined. Adults showed obvious reproductive diapause, which was regulated by photoperiod and temperature. At 30°C, there was no adult diapause occurring under either long‐day or short‐day conditions; at 25°C the pre‐oviposition period was short and fecundity was high in adult females under L : D 16 : 8 h, whereas under L : D 12 : 12 h, a few females entered reproductive diapause; at 20°C under short‐day conditions, all female adults entered diapause. The pre‐oviposition period was significantly prolonged when the pupae and adults were transferred from long‐days to short‐days, but the day length influence was not obvious when they were transferred only in the adult stage. However, the fecundity dropped greatly no matter whether the photoperiod shifted to short‐days only in the adult stage or whether the shift occurred in both the pupal and adult stage. The fecundity was extremely low when photoperiod shifted from long‐days to short‐days in both pupal and adult stages. This was an indication that the pupal and adult stages were the photoperiod‐sensitive stage for adult reproductive diapause. This was especially true for the photoperiod in the pupal stage, which has a distinctly significant regulative effect on reproductive diapause. Additionally, this article also addresses the reason for different photoperiodic response patterns in reproductive diapause induction between the Changsha strain and the Tsukuba strain (Japan) of O. communa.     

Spatio-temporal impact of climate change on the activity and voltinism of the spruce bark beetle, Ips typographus

The spruce bark beetle Ips typographus is one of the major insect pests of mature Norway spruce forests. In this study, a model describing the temperature-dependent thresholds for swarming activity and temperature requirement for development from egg to adult was driven by transient regional climate scenario data for Sweden, covering the period of 1961–2100 for three future climate change scenarios (SRES A2, A1B and B2). During the 20th century, the weather supported the production of one bark beetle generation per year, except in the north-western mountainous parts of Sweden where the climate conditions were too harsh. A warmer climate may sustain a viable population also in the mountainous part; however, the distributional range of I. typographus may be restricted by the migration speed of Norway spruce. Modelling suggests that an earlier timing of spring swarming and fulfilled development of the first generation will significantly increase the frequency of summer swarming. Model calculations suggest that the spruce bark beetle will be able to initiate a second generation in South Sweden during 50% of the years around the mid century. By the end of the century, when temperatures during the bark beetle activity period are projected to have increased by 2.4–3.8 °C, a second generation will be initiated in South Sweden in 63–81% of the years. The corresponding figures are 16–33% for Mid Sweden, and 1–6% for North Sweden. During the next decades, one to two generations per year are predicted in response to temperature, and the northern distribution limit for the second generation will vary. Our study addresses questions applicable to sustainable forest management, suggesting that adequate countermeasures require monitoring of regional differences in timing of swarming and development of I. typographus , and planning of control operations during summer periods with large populations of bark beetles.     

Development characteristics of the box‐tree moth Cydalima perspectalis and its potential distribution in Europe

The box‐tree moth Cydalima perspectalis (Walker) is an invasive pest causing severe damage to box trees (Buxus spp.). It is native to Japan, Korea and China, but established populations have been recorded in a number of locations across Europe since 2007 and the spread of the insect continues. The developmental investigations suggest that larvae overwinter mainly in their 3rd instar in Europe and that diapause is induced by a day length of about 13.5 h. One and a half to 2 months in the cold are necessary to terminate diapause. Threshold temperatures for development and number of degree‐days to complete a generation are slightly different from those calculated in previous studies in Japan. A bioclimatic (CLIMEX^®) model for C. perspectalis in Europe was developed, based on climate, ecological and developmental parameters from the literature and new field and laboratory studies on diapause termination, thermal requirements and phenology. The model was then validated with actual distribution records and phenology data. The current distribution and life history of C. perspectalis in Europe were consistent with the predicted distribution. The climate model suggests that C. perspectalis is likely to continue its spread across Europe, except for Northern Fenno‐Scandinavia, Northern Scotland and high mountain regions. The northern distribution of C. perspectalis is expected to be limited by a number of degree‐days above the temperature threshold insufficient to complete a generation, whereas its southern range is limited by the absence of a cold period necessary to resume diapause. The model predicts relatively high Ecoclimatic Indices throughout most of Europe, suggesting that the insect has the potential of becoming a pest in most of its predicted range. However, damage is likely to be higher in Southern and Central Europe where the moth is able to complete at least two generations per year.     

Effects of nutrition and methoprene treatment upon reproductive diapause in Caloptilia fraxinella (Lepidoptera: Gracillariidae)

Abstract Female Caloptilia fraxinella exhibit a prolonged reproductive diapause immediately post adult emergence in mid‐summer until the next spring when mating, egg development and oviposition on fresh Fraxinus spp. leaflets occur. Factors that effect the termination of reproductive diapause are investigated in this species. Caloptilia fraxinella diapausing adults held in overwintering conditions (2 °C, LD 0 : 24 h) for 24 weeks terminate diapause after placement for 2 weeks in simulated summer conditions (24 °C, LD 16 : 8 h) only if they are provided with 10% sugar water. Exogenous application of the Juvenile Hormone (JH) analogue methoprene to moths in both early‐ (summer) and mid‐ (autumn) reproductive diapause demonstrates that JH affects diapause termination but a carbohydrate nutrition source also mediates mating and vitellogenesis. Mating between moth pairs early in diapause occurs only after treatment with methoprene and provision with sugar water. However, there is no impact of mating on the propensity of females to produce vitellogenic oöcytes. Moths collected in the autumn in mid‐diapause respond in a dose‐dependent fashion to methoprene treatment and the response is greater than that of moths early in diapause collected in the summer. Treatment with methoprene and access to sugar water results in vitellogenic oöcytes in 18.75% of females from mid‐diapause moth pairs treated with 0.01 μg methoprene per insect and in all females from pairs treated at the two highest doses of methoprene (0.1 and 1 μg per insect). Mating occurs only between moths in mid‐diapause treated with the two highest doses of methoprene and these doses induce 91% and 100% mating, respectively. Both control and methoprene‐treated males in mid‐diapause held under summer conditions mate successfully and pass a spermatophore to their methoprene‐treated female partner. These data demonstrate that female C. fraxinella undergo a prolonged reproductive diapause in which termination is dependent on JH and further mediated by a carbohydrate nutrition source. The production of vitellogenic oöcytes is independent of mating. These data also provide evidence that response of moths in diapause to exogenous applications of methoprene differs throughout the diapause period and between male and female C. fraxinella.     

Differences in the pre‐diapause and pre‐oviposition accumulation of critical nutrients in adult females of the beetle Colaphellus bowringi

In many insect species, the differentiation of development between diapause and reproduction first becomes obvious during the diapause preparation (pre‐diapause) and pre‐oviposition phases. However, the differentiation of nutrient accumulation between these two phases remains unclear. We compared the weights of pre‐diapause and reproductive adult female Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), and measured their triacylglycerol (TAG), protein, and carbohydrate content from emergence until they had fed for 4 days post‐eclosion. We also compared the ovarian development and accumulation of lipid droplets between pre‐diapause and reproductive adult females in order to determine whether we could visually detect differences in nutrient allocation. The weights of both pre‐diapause and reproductive females increased with duration of feeding. The fresh weight and water content of pre‐diapause females was significantly lower than that of reproductive females after feeding for 3 days post‐eclosion. Pre‐diapause females channeled their reserves into TAG in the fat body, whereas reproductive females converted nutrients into proteins and carbohydrates for egg development. These results quantify differences in nutrient accumulation between pre‐diapause and reproductive adult female C. bowringi, and provide clues for understanding the molecular mechanisms underlying differences in the allocation of nutrients between diapause and reproduction in insects.     

Insects in a warmer world: ecological, physiological and life-history responses of true bugs (Heteroptera) to climate change

Focusing on the southern green stink bug, Nezara viridula (Pentatomidae), in central Japan the effects of climate change on true bugs (Insecta: Heteroptera) are reviewed. In the early 1960s, the northern edge of the species's distribution was in Wakayama Prefecture (34.1°N) and distribution was limited by the +5°C coldest month (January) mean temperature isothermal line. By 2000, N. viridula was recorded 70 km further north (in Osaka, 34.7°N). Historical climate data were used to reveal possible causes of the northward range expansion. The increase of mean and lowest winter month temperatures by 1–2°C in Osaka from the 1950s to the 1990s improved potential overwintering conditions for N. viridula. This promoted northward range expansion of the species. In Osaka, adult diapause in N. viridula is induced after mid‐September, much later than in other local seed‐feeding heteropterans. This late diapause induction results in late‐season ineffective reproduction: some females start oviposition in autumn when the progeny have no chance of attaining adulthood and surviving winter. Both reproductive adults and the progeny die. A period from mid‐September to early November represents a phenological mismatch: diapause is not yet induced in all adults, but it is already too late to start reproduction. Females that do not start reproduction but enter diapause in September have reduced postdiapause reproductive performance: they live for a shorter period, have a shorter period of oviposition and produce fewer eggs in smaller egg masses compared with females that emerge and enter diapause later in autumn. To some extent, N. viridula remains maladapted to Osaka environmental conditions. Ecological perspectives on establishment in recently colonized areas are discussed. A review of available data suggests that terrestrial and aquatic Heteroptera species respond to climate change by shifting their distribution ranges, changing abundance, phenology, voltinism, physiology, behaviour, and community structure. Expected responses of Heteroptera to further climate warming are discussed under scenarios of slight (<2°C) and substantial (>2°C) temperature increase.     

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.     

Time of the season: the effect of host photoperiodism on diapause induction in an insect herbivore,Leptinotarsa decemlineata

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Effects of experimentally imposed climate scenarios on flowering phenology and flower production of subarctic bog species

Climate scenarios for high‐latitude areas predict not only increased summer temperatures, but also larger variation in snowfall and winter temperatures. By using open‐top chambers, we experimentally manipulated both summer temperatures and winter and spring snow accumulations and temperatures independently in a blanket bog in subarctic Sweden, yielding six climate scenarios. We studied the effects of these scenarios on flowering phenology and flower production of Andromeda polifolia (woody evergreen) and Rubus chamaemorus (perennial herb) during 2 years. The second year of our study (2002) was characterized by unusually high spring and early summer temperatures. Our winter manipulations led to consistent increases in winter snow cover. As a result, average and minimum air and soil temperatures in the high snow cover treatments were higher than in the winter ambient treatments, whereas temperature fluctuations were smaller. Spring warming resulted in higher average, minimum, and maximum soil temperatures. Summer warming led to higher air and soil temperatures in mid‐summer (June–July), but not in late summer (August–September). The unusually high temperatures in 2002 advanced the median flowering date by 2 weeks for both species in all treatments. Superimposed on this effect, we found that for both Andromeda and Rubus, all our climate treatments (except summer warming for Rubus) advanced flowering by 1–4 days. The total flower production of both species showed a more or less similar response: flower production in the warm year 2002 exceeded that in 2001 by far. However, in both species flower production was only stimulated by the spring‐warming treatments. Our results show that the reproductive ecology of both species is very responsive to climate change but this response is very dependent on specific climate events, especially those that occur in winter and spring. This suggests that high‐latitude climate change experiments should focus more on winter and spring events than has been the case so far.     

Phenology of Eucalyptus weevil, Gonipterus scutellatus Gyllenhal (Coleoptera: Curculionidae), and chrysomelid beetles in Eucalyptus globulus plantations in south-western Australia

1 In south‐western Australia, Eucalyptus globulus plantations are defoliated by a complex of beetle species, yet only scant information exists on these species under such climatic conditions. To improve management of these defoliating beetles in the region, canopy fogging and shoot clipping were conducted in plantations between 1999 and 2002 to identify and document the phenology of the beetle species present. 2 Eucalyptus weevil, Gonipterus scutellatus, was the most common and destructive defoliating beetle. Gonipterus scutellatus undergoes one principal generation each year with a lesser second generation or cohort in some seasons, which contrasts greatly with reports of two to four annual generations for the species in other regions. This limited reproduction by G. scutellatus may be due to the limited availability from summer onwards of new flushing foliage, which is essential for feeding and oviposition. 3 Several species of chrysomelid beetles were collected in plantations, but these were present in much lower numbers than G. scutellatus and were only a minor concern. However, some species, such as Chrysophtharta variicollis, appear to be capable of developing short‐lived outbreaks. 4 A diverse suite of natural enemies was fogged from plantations but they were significantly less abundant than defoliating beetles and are not likely to provide significant control of beetles. 5 In terms of managing these defoliating beetles, monitoring and control should focus on G. scutellatus, and be conducted during spring when most damage occurs.     

Environmental Cues,Endocrine Factors,and Reproductive Diapause in Male Insects

Environmental cues, mostly photoperiod and temperature, mediated by effects on the neuroendocrine system, control reproductive diapause in female insects. Arrest of oocyte development characterizes female reproductive diapause, which has two major adaptive functions: It improves chances of survival during unfavorable season(s), and/or it confines oviposition to that period of the year that is optimal for survival of the eggs and progeny. Although reproductive diapause is less well studied in male insects, there may be no sex-dependent differences in regard to the first of these functions. The second one, however, is not valid for the male; instead, selection pressure directs the male's reproductive strategy toward maximum chances of fertilization of the female's eggs with minimum waste of energy. Therefore, in species with female reproductive diapause, the males may or may not exhibit diapause, but if they do, their diapause must be adapted to that existing in conspecific females. Male reproductive diapause is defined as a reversible state of inability of the male to inseminate receptive females. In relation to reproductive diapause, there are several patterns of coadaptations between male reproductive strategy and timing of female receptivity, (a) In some insects, the females are receptive in the early part of their diapause; mating occurs during this period and there is no diapause in the male. The male dies shortly after copulation and the female stores the sperms to fertilize the eggs that develop after termination of the female's diapause, (b) In some species, as in the grasshopper Anacridium aegyptium, females are receptive during diapause; though oocyte development is arrested, copulation occurs and the stored sperms fertilize the eggs when the female's diapause ends. Males were claimed to have no diapause, but recent studies have revealed the presence of a reproductive diapause in a proportion of the males. This and other cases show that female receptivity during reproductive diapause may or may not be accompanied by male reproductive diapause. If there is a reproductive diapause in the male, it is controlled by the same endocrine mechanism, the corpora allata (CA), as in the females, (c) In many species females are refractory during their diapause. In these cases, males exhibit reproductive diapause, which may be light, as in the beetle Oulema melanopus, or well established, as in certain grasshoppers, butterflies, and beetles. In the latter cases, male diapause is controlled by similar environmental cues (photoperiod, temperature) and by the same intrinsic mechanism (neuroendocrine system, especially CA) as female diapause. Nevertheless, male diapause is less intense; the environmental cues leading to its termination are less complex and/or less extreme, so male diapause terminates before that of the females. Presumably, male diapause is under two antagonistic selection pressures: A male should not waste energy by courting dia-pausing refractory females, but he should be ready to copulate as soon as the females become receptive, otherwise he may lose in the competition between males for females. Some further strategies, which do not seem to fit the above patterns, are also outlined.