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.     

The role of gene flow asymmetry along an environmental gradient in constraining local adaptation and range expansion

Theoretically, asymmetric gene flow along an environmental gradient can limit species range expansion by keeping peripheral populations from locally adapting. However, few empirical studies have examined this potentially fundamental evolutionary mechanism. We address this possibility in the cricket Allonemobius socius, which exist along a season‐length gradient where the probability of producing a single generation per year (univoltinism) increases with latitude. As the probability of univoltinism increases northwards, populations are expected to hedge their bets by producing a greater proportion of diapause eggs when exposed to a mild diapause cue. However, gene flow from southern populations may disrupt local adaptation in the north by reducing the proportion of diapause eggs (expected to be 100% in pure univoltine environments). This may limit range expansion along the northern periphery where A. socius compete with A. fasciatus, a sister species that exhibits an invariant diapause‐only egg‐laying strategy. To assess the potential for range limitation, we examined diapause incidence (the proportion of diapause eggs produced under diapause conditions), gene flow symmetry and population structure across nine A. socius populations. We found that gene flow was asymmetric and biased northwards towards the periphery. Furthermore, peripheral populations that inhabited pure univoltine environments produced numerous nondiapause eggs (a southern, bivoltine diapause phenotype), which we assume to be a suboptimal phenotype. These patterns suggest that asymmetric gene flow along the gradient constrains adaptation in peripheral populations, potentially constraining species range expansion.     

Variability in egg-to-adult development time in the bee Ptilothrix plumata and its parasitoids

1. Ptilothrix plumata (Hymenoptera) is a neotropical solitary bee that nests in bare, sandy soils. Data on the biology and behaviour of this species are provided. Egg‐to‐adult development time of a neotropical solitary bee and its parasitoids is quantified for the first time. 2. The bee showed huge variability in egg‐to‐adult development time. There are two categories of eggs relative to adult emergence. The same season eggs are those from which adults emerge in the same reproductive season in which the eggs were laid. Adults from next season eggs emerge in the next dry reproductive season. This variability in egg‐to‐adult development time is reached through prepupal diapause. The bee larval parasitoids Leucospis genalis (Hymenoptera) and Anthrax sp. (Diptera) showed similar range in their development times. The data show that diapause is controlled by season. 3. By synchronising adult emergence, this variability in egg‐to‐adult development time facilitates mate finding, and population recruitment is a consequence of the bees avoiding the unfavourable reproductive season.     

Adaptions in the life cycle ofDermacentor variabilis (Say) andIxodes dammini (Spielman,Clifford, Piesman,and Corwin) marginal populations (Acari: Ixodidae)

Both theDermacentor variabilis andIxodes dammini adult diapause seasons are followed by a breeding period. At the edge of the ticks' range, where diapause and breeding occur under adverse climatic conditions, the species still conserves its fixed response for the onset of diapause. As a result, most of the following breeding effort is wasted; the population can only be maintained by an aberrant breeding period during the normal diapause season.When the diapause and breeding periods are suitable for the climate at the edge of the ticks' range, the rate of immature development versus the temperature regime can result in the appearance of the immature stages out of phase with the timing for the optimal life cycle in the marginal climate. The ticks are limited to minor adjustments of their development rates but, if this is all that is required for an optimum life cycle, this response can preadapt the species to the foreign climate.     

Diapause in the seasonal cycle of stink bugs (Heteroptera,Pentatomidae) from the Temperate Zone

The paper reviews the data on diapause and related phenomena in stink bugs (Heteroptera: Pentatomidae). Using stink bugs as examples, the consecutive stages of the complex dynamic process of diapause (such as diapause preparation, induction, initiation, maintenance, termination, post-diapause quiescence, and resumption of direct development) are described and discussed. Out of 43 pentatomid species studied in relation to diapause in the Temperate Zone up to date, the majority (38 species) overwinter as adults, two species—as eggs, and another two species—as nymphs. Pentatoma rufipes is believed to be able to overwinter at different stages of its life cycle. Less than 5 % of pentatomid species are probably able to overwinter twice. Only five species have obligate diapause, others have the facultative one. Day-length and temperature are the main diapause inducing factors in the majority of species. The role of food in the control of seasonal development is essential in the pentatomid species feeding on plant seeds. In different species, different stages are sensitive to day-length. Some pentatomids retain sensitivity to photoperiod even after diapause, others lose it and become photo-refractory (temporarily or permanently). In Pentatomidae, such seasonal adaptations as photoperiodic control of nymphal growth rates, seasonal body colour change, migrations, and summer diapause (aestivation) are widely represented, whereas wing and/or wing muscle polymorphism has not been reported yet. In the subfamily Podopinae, induction of facultative reproductive winter diapause is under the control of photoperiod and temperature. All species feed on seeds and their seasonal development to a great extent reflects availability of food. However, the same food preferences and pattern of seasonal development are also characteristic to many species from the subfamily Pentatominae. All species of the subfamily Asopinae are predators. Among them, Picromerus bidens and Apateticus cynicus have obligate embryonic winter diapause, which is rear among true bugs. At the same time, A. cynicus and Podisus maculiventris belong to the same tribe but have different types of diapause: obligate embryonic diapause in A. cynicus and facultative adult diapause in P. maculiventris. Other Asopinae species studied up to date have facultative adult diapause controlled by photoperiod and temperature with probably only one exception: in Andrallus spinidens, adult diapause is controlled by temperature, and photoperiod plays only a secondary role. Thus, in spite of the similar habits and feeding types among Asopinae, the species of this subfamily have different types of diapause and the latter is controlled by different factors. In the subfamily Pentatominae, most species overwinter as adults and induction of their diapause is controlled by the long-day type photoperiodic response, in spite of the differences in their feeding preferences (within phytophagy). However, there are some exceptions in this subfamily, too: Palomena prasina, P. angulosa and Menida scotti have obligate diapause, which conditions univoltinism in these pentatomids. In M. scotti, only females have obligate adult diapause, whereas males remain physiologically active through the whole winter, this pattern being unusual for Heteroptera. The univoltine seasonal cycle of this species with summer diapause (aestivation) and apparent migrations is similar to that of shield bugs (Scutelleridae). According to the analysis of seasonal development, the evolution of seasonal adaptations in Pentatomidae does not directly reflect their phylogeny. However, individual genera, small tribes or even subfamilies have similar complexes of seasonal adaptations. At the same time, Pentatominae is a large and apparently collected taxon, but most of species in this subfamily have the same facultative adult diapause.     

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.     

Effects of host-plant species on diapause induction of the Kanzawa spider mite, Tetranychus kanzawai

In many herbivorous arthropods, incidence of diapause, which is considered to reflect the timing of diapause, changes depending on the host plants they utilize. Several theoretical studies suggest that the optimal timing of diapause induction depends on life‐history traits; if the development time of the arthropod is short, fecundity is high, or survival rate remains high throughout the season, the optimal timing of diapause induction would be shifted toward the end of the season. For herbivorous arthropods, these life history traits may change among their host plants. Here we examined whether a population of the Kanzawa spider mite, Tetranychus kanzawai Kishida (Acari: Tetranychidae), shows the predicted pattern of diapause induction on two host plants, the kidney bean [Phaseolus vulgaris (Leguminosae)] and Japanese Orixa [Orixa japonica (Rutaceae)], on which the mites show different performances. Rearing conditions were controlled in two ways. In the first experiment, day length and temperature were kept constant throughout the mite lifetime at either of three conditions from mid October to early November. In the second experiment, the conditions were changed from 20 °C and L11.5:D12.5 at immature stages to 18 °C and L11:D13 at adult stage in order to better approximate field conditions. In the first experiment, diapause incidence on P. vulgaris was lower than on O. japonica. This tendency became stronger in the second experiment, suggesting a difference in the timing of diapause induction among host plants. On the other hand, P. vulgaris was proven to allow high performance, i.e., greater lifetime fecundity and shorter development times, although it had no effect on the survival rate. The relationship between diapause incidence and performance is consistent with the prediction of theoretical studies that a short development time or high fecundity delays the timing of diapause induction.     

Latitudinal clines in the timing and temperature-sensitivity of photoperiodic reproductive diapause in Drosophila montana

Reproductive diapause is a primary mechanism used by arthropods to synchronize their life cycle with seasonal changes in temperate regions. Our study species, Drosophila montana, represents the northern insect species where flies enter reproductive diapause under short day conditions and where the precise timing of diapause is crucial for both survival and offspring production. We have studied clinal variation in the critical day length for female diapause induction (CDL) and their overall susceptibility to enter diapause (diapause incidence), as well as the temperature sensitivity of these traits. The study was performed using multiple strains from four latitudinal clines of the species – short clines in Finland and Alaska and long clines in the Rocky Mountains and the western coast of North America – and from one population in Kamchatka, Russia. CDL showed strong latitudinal clines on both continents, decreasing by one hour per five degrees decline in latitude, on average. CDL also decreased in all populations along with an increase in fly rearing temperature postponing the diapause to later calendar time, the effects of temperature being stronger in southern than in northern population. Female diapause incidence was close to 100% under short day/low temperature conditions in all populations, but decreased below 50% even under short days in 19°C in the southern North American western coast populations and in 22°C in most populations. Comparing a diversity of climatic data for the studied populations showed that while CDL is under a tight photoperiodic regulation linked with latitude, its length depends also on climatic factors determining the growing season length. Overall, the study deepens our understanding of how spatial and environmental parameters affect the seasonal timing of an important biological event, reproductive diapause and helps to estimate the evolutionary potential of insect populations to survive in changing climatic conditions.     

QTL analysis of the photoperiodic response and clinal distribution of period alleles in Nasonia vitripennis

In seasonal environments, organisms synchronize their life cycle with the annual cycle of environmental factors. In many insect species, this includes a diapause response: a timed dormant stage that allows to survive harsh winter conditions. Previously, we have shown that larval diapause in the parasitic wasp Nasonia vitripennis is induced by the mother upon exposure to a threshold number of short photoperiods (named switch point) and diapause response follows a latitudinal cline in natural populations. Here, we present a QTL analysis using two lines derived from the extremes of this clinal distribution: a northern line from Oulu, Finland and a southern line from Corsica, France. A genomic region on chromosome 1 and one on chromosome 5 were found to be associated with photoperiodic diapause induction. Interestingly, these regions contain the putative clock genes period, cycle (chromosome 1) and cryptochrome (chromosome 5). An analysis of period polymorphisms in seven European populations showed a clinal distribution of two main haplotypes that correlate with the latitudinal cline for diapause induction.     

Effects of photoperiodically induced reproductive diapause and cold hardening on the cold tolerance of Drosophila montana

Coping with seasonal and daily variation in environmental conditions requires that organisms are able to adjust their reproduction and stress tolerance according to environmental conditions. Females of Drosophila montana populations have adapted to survive over the dark and cold winters at high latitudes and altitudes by spending this season in photoperiodically controlled reproductive diapause and reproducing only in spring/summer. The present study showed that flies of a northern population of this species are quite tolerant of low temperatures and show high seasonal and short-term plasticity in this trait. Culturing the flies in short day length (nearly all females in reproductive diapause), as well as allowing the flies to get cold hardened before the cold treatment, increased the cold tolerance of both sexes both in chill coma recovery time test and in mortality assay. Chill coma recovery time test performed for the females of two additional D. montana populations cultured in a day length where about half of the females enter diapause, also showed that diapause can increase female cold tolerance even without a change in day length. Direct linkage between diapause and cold tolerance was found in only two strains representing a high-altitude population of the species, but the phenomenon will certainly be worth of studying in northern and southern populations of the species with larger data sets.     

Developmental characteristics of two closely related species of Allonemobius and their hybrids

Summary Two species of ground crickets, Allonemobius fasciatus (from Corvallis, 44.3 N) and A. socius (from Gainesville, 30.3 N), and their hybrids were compared for nymphal development, wing form, and embryonic diapause. A. fasciatus matured more rapidly at a short photoperiod than at a long photoperiod and also than A. socius which showed no developmental response to photoperiod. The proportion of long-winged forms varied with nymphal photoperiod greatly in A. fasciatus but slightly in A. socius. A. fasciatus laid only diapause eggs, while A. socius responded to a long parental photoperiod and a high incubation temperature by averting diapause. A. fasciatus showed temperature-dependent shift of the diapause stage butA. socius entered diapause only at the early stage. These differences are related to their voltinism and distribution.Reciprocal crosses between the two species produced viable hybrids. In development rate, female hybrids were intermediate between their parents while male hybrids were closer to their mothers, suggesting that the development rate is controlled primarily by the X chromosome. Overdominance occurred in wing form; reciprocal crosses produced either higher or lower proportions of long-winged forms than their parents. Diapause of F1 hybrid embryos was determined only by the female parent and the paternal effect appeared in F2 embryos.     

Reproductive seasonality in tropical satyrine butterflies: strategies for the dry season

Abstract.

• 1 In tropical savanna environments rainfall is often very seasonal, so that much of the year is characterized by a long and unpredictable dry season. Because the timing and availability of rain exerts a major influence on plant growth and production, many species during the dry period exhibit dramatic reduction in leaf quality. Accordingly, and kind of behaviour shown by phytophagous insects that synchronizes larval feeding with food availability will be adaptive.
• 2 The reproductive status of three Mycalesis butterflies was monitored over a 2-year period (1989–90) in north-eastern Queensland, Australia, at a lowland site (Cardwell, 18°16's, 146°02′E) which experiences a pronounced dry season. Females of these species and of five other satyrines (Ypthima, Hypocysta spp.) were also examined less intensively during the dry season in areas throughout northern and central Queensland, north of the tropic of Capricorn.
• 3 These relatively sedentary butterflies exhibit three different strategies for dealing with the unpredictable dry period and associated deterioration of larval food plants (grass). First, five species appear to breed continuously, though for most reproductive activity (mature egg number) declines markedly in the late dry season. Two of these (Hypocysta irius, H.metirius) are restricted to less seasonal and more favourable (wetter) areas but the three others (Ypthima arctous, H.adiante, H.pseudirius) occur widely in the relatively dry savanna, where they may specialize on grass in moister microenvironments. Second, two species (M.terminus, M.sirius) live in predictably moist habitats which are buffered from climatic extremes; they breed for much of the season but reproductive activity declines as the dry season progresses and may cease late in the season. Third, one species (M.perseus) is more opportunistic, breeding for only a limited interval during the favourable (wet) periods; during the long dry season adults contract to moist refugia and remain in reproductive diapause.
• 4 Spending the late dry season as an adult, either in diapause or with mature eggs, may improve the capacity to utilize new growth of grasses at the start of the favourable season, thereby enhancing population growth during good times. It may also provide additional flexibility to counter the temporal uncertainty of the dry season.
• 5 The strategy of residing in more equitable habitats or specializing on predictable foods may be the most restrictive in terms of distribution.

     

Why Callosobruchus rhodesianus causes limited damage during storage of cowpea seeds in a tropical humid zone in Togo

The sympatric bruchids Callosobruchus rhodesianus (Pic) and Callosobruchus maculatus (F.) (Coleoptera: Bruchidae) colonize cultures of cowpea, Vigna unguiculata (Walp.) (Papilionaceae), in Togo at the end of the rainy season. Seeds containing larvae of the two bruchid species were introduced into the storage systems. Callosobruchus rhodesianus adults emerged from the seeds at the beginning of storage, but this species disappeared quickly and C. maculatus became dominant. Analysis of the reproduction of females collected at the beginning of storage made it possible to distinguish between four groups of females differing in their reproductive activity. Three groups were sexually active but they differed in their fecundity and the duration of the latency period before the first oviposition phase. The fourth group of females was in reproductive diapause. In interspecific competition, the presence of C. maculatus larvae reduced the survival chances of C. rhodesianus and therefore is disadvantageous for this species. The main factors explaining the rapid decrease of the C. rhodesianus populations under storage conditions were the variability of female reproductive activity with the emergence of insects in reproductive diapause and the significant larval mortality in interspecific competition with C. maculatus.     

Diapause in copepods (Crustacea) from ephemeral habitats with different hydroperiods in Everglades National Park (Florida,U.S.A.)

Water management practices in the Everglades have severely stressed the natural system, particularly by reducing the hydroperiods of much of the region. During the dry season of 1999, we investigated the influence of hydroperiod on the species composition and dormancy patterns of freshwater copepod communities in seasonal wetlands of Everglades National Park, Florida, U.S.A. The habitats were characterized by an annual dry season, from December through June. We sampled at two locations: the Long Pine Key area of the Rocky Glades region (short hydroperiod, ca. 4–5 months), and western Taylor Slough (intermediate hydroperiod, ca. 8–10 months). Both areas have experienced a reduction in natural hydroperiods and an increase in the frequency of dry-down. We collected weekly plankton samples from Rocky Glades solution holes to assess the potential species pool of copepods. To document the taxa capable of surviving dry-down by resting, we performed three immersion trials in which we rehydrated, in laboratory aquaria, sediment patches from solution holes and surface soils from all stations. Only a subset of the planktonic species collected emerged from the dried sediments. The cyclopoids Microcyclops rubellus and Paracyclops poppei were dominant. This is the first record of diapause for P. poppei. Species distributions from the different hydroperiod soil patches indicated that more diapausing species occurred at the sites that dried for shorter periods. Emerging individuals of M. rubellus and P. poppei were mainly ovigerous females, demonstrating a resting strategy seldom before recorded. The cyclopoid Diacyclops nearcticushad not been previously reported to diapause, but they emerged from the dried sediments in our trials. Our collections included six new records for Florida: Diacyclops nearcticus, Megacyclops latipes, Orthocyclops modestus, Elaphoidella marjoryae, Bryocamptus sp. and Bryocamptuscf. newyorkensis. Paracyclops poppei, Macrocyclops fuscus and Arctodiaptomus floridanus are new records for Everglades National Park. Clearly, diapause is an important strategy for the persistence of copepods in short-hydroperiod wetlands. The duration of the dry period appears to be inversely related to the number of species that emerge from diapause.     

Cloning and characterization of Methoprene-tolerant(Met)and Kruppel homolog 1(Kr-h1)genes in the wheat blossom midge,Sitodiplosis mosellana

Juvenile hormone(JH),a growth regulator,inhibits ecdysteroid-induced meta-morphosis and controls insect development and diapause.Methoprene-tolerant(Met)and Krippel homolog I(Kr-h1)are two proteins involved in JH action.To gain some insight into their function in development of Sitodiplosis mosellana,an insect pest undergoing obligatory larval diapause at the mature 3rd instar stage,we cloned full-length complemen-tary DNAs of Met and Kr-h1 from this specics.SmMet encoded a putative protein,which contained three domains typical of the bHLH-PAS family and eight conserved amino acid residues important for JH binding.SmKr-h1 encoded a protein showing high sequence homology to its counterparts in other specics,and contained all eight highly conserved Zn-finger motifs for DNA-binding.Expression patterns of SmMet and SmKr hl were de-velopmentally regulated and JH III responsive as well.Their mRNA abundance increased as larvae entered carly 3rd instar,pre-diapause and maintenance stages,and peaked during post-diapause quiescence,a pattern correlated with JH titers in this species.Different from reduced expression of SmMer,SmKr-h1 mRNA increased at mid-to-late period of post-diapause development.Topical application of JH II on diapausing larvac also induced the two genes in a dose-dependent manner.Expression of SmuMer and SmKr-h1 clearly declined in the pre-pupal phase,and was significantly higher in female adults than male adults.These results suggest that JH-responsive SmMet and SmKr-h1 might play key roles in diapause induction and maintenance as well as in post-diapause quiescence and adult reproduction,whereas metamorphosis from larvae to pupac might be correlated with their reduced expression.     

Diapause affects cuticular hydrocarbon composition and mating behavior of both sexes in Drosophila montana

Envipnmental cues,mainly photoperiod and temperature,are known to control female adult reproductive diapause in several insect species.Diapause enhances female survival during adverse conditions and postpones progeny production to the favorable season.Male diapause(a reversible inability to inseminate receptive females)has been studied much less than female diapause.However,if the males maximized their chances to fertilize females while minimizing their energy expenditure,they would be expected to be in diapause at the same time as females.We investigated Drosophila montana male mating bchavior under short-day conditions that induce diapause in females and found the males to be reproductively inactive.We also found that males reared under long-day conditions(reproducing individuals)court reproducing postdiapause fermales,but not diapausing ones.The diapausing fies of both sexes had more long-chain and less short-chain hydrocarbons on their cuticle than the reproducing ones,which presumably increase their survival under stressful conditions,but at the same time decrease their attractiveness.Our study shows that the mating behavior of females and males is well coordinated during and afier overwintering and it also gives support to the dual role of insect cuticular hydrocarbons in adaptation and mate choice.     

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.     

Reproduction and egg diapause of the oriental chinch bug,Cavelerius saccharivorus okajima (Heteroptera: Lygaeidae), in the subtropical winter season in relation to its wing polymorphism

Reproduction and egg diapause of the oriental chinch bug, Cavelerius saccharivorus, in the subtropical winter season were investigated in relation to its wing polymorphism. Macropterous females collected from the southern part of Okinawa Is. in the autumn season delayed their oviposition and were less fecund early in the adult life period, but survived much longer than brachypterous females collected from the same locality. The total fecundity was not significantly different between wing morphs. The diapause of eggs laid by brachypters tended to be terminated more easily at a high temperature than that of eggs laid by macropters. This indicated that the eggs laid by macropters were more intense in their diapause than those laid by brachypters. However, irrespective of the parental wing form, diapause showed considerable variation in its intensity within and between clutches.     

ADAPTATIONS TO TEMPERATE CLIMATES AND EVOLUTION OF OVERWINTERING STRATEGIES IN THE DROSOPHILA MELANOGASTER SPECIES GROUP

The Drosophila melanogaster species group is considered to have originated in the tropics and only recently invaded temperate habitats. The temperate species of this group that were studied here may be subdivided into the warm-temperate species (D. lutescens and D. rufa) and the cool-temperate species (species of the auraria complex). The warm-temperate species were more cold-hardy than were their tropical relatives (D. takahashii or D. melanogaster) at the larval and imaginal stages, and the cool-temperate species were more cold-hardy than the warm-temperate species, although only at the imaginal stage. However, these species showed little or no intraspecific variation in cold-hardiness, in spite of great variation in winter temperature within the species' ranges. It is assumed that cold-hardiness is one of the main factors restricting their distributions at high latitudes and that it is the key for evolution of the warm- and cool-temperate species from their subtropical or warm-temperate ancestors. Both warm- and cool-temperate species had photoperiodically controlled reproductive diapause. In the cool-temperate species, the development of cold-hardiness was affected by diapause, but diapause had little or no effect on cold-hardiness in the warm-temperate species. Critical daylengths and the diapause rates varied from species to species according to variation in their overwintering plans and also varied geographically in consequence of their adaptation to local climates. Species showed different responses to temperature in preimaginal and ovarian development. These differences are considered to reflect adaptation to different environmental temperatures.