Overwintering in Drosophila melanogaster: outdoor field cage experiments on clinal and laboratory selected populations help to elucidate traits under selection

Insects can adapt to temperate environments by increasing levels of resistance to cold conditions over winter and/or altering reproductive patterns to focus reproduction in favourable conditions. In temperate areas, Drosophila melanogaster persists over winter at the adult stage. A previous experiment, conducted with flies kept in outdoor population cages in the temperate winter, indicated that temperate populations produced more eggs than did tropical populations following an abrupt increase in reproduction in late winter. In contrast, the tropical populations produced more eggs prior to the increase. Both patterns resulted in a higher net number of surviving offspring for temperate populations. Here we again examine the clinal pattern in reproduction using outdoor cages, this time held under tropical winter conditions. In this environment, surprisingly, egg production was higher and on average earlier in populations originating from temperate areas. However, mortality rates also increased with latitude of origin, and the relationship of lifetime egg production to latitude should therefore be measured. To test the role of altered pattern of egg production per se in the reproductive advantage of temperate populations in the temperate winter, we tested the performance of laboratory lines selected for altered reproductive patterns, under temperate winter conditions. Lines selected for high early fecundity exhibited this characteristic in the field cages and lines selected for late reproduction exhibited a relatively high fecundity in spring. The timing of the abrupt increase in egg production was identical in these sets of lines and occurred at the same time in recently collected populations, suggesting evolutionary conservation of the switch. These findings suggest that changes in early and late reproduction per se determine adaptation to temperate winter conditions, and illustrate how laboratory selection lines can be used to understand traits underlying adaptive shifts in field performance.     

Absence of clinal variation in virgin retention capacity in Australian Drosophila melanogaster

The ability of virgin Drosophila melanogaster adults to retain eggs is thought to be an adaptation to persisting in temperate areas, based on differences in this trait between European and African populations, and based on seasonal changes in this trait in France. By retaining eggs in the absence of males and under conditions of poorer nutrition (conditions common in temperate areas during colder months), females reduce the wastage of resources and increase their probability of surviving spring into summer, enabling them to initiate summer population expansions. To test for variation in virgin egg retention along a climatic gradient, we characterized clinal variation in strains collected from eastern Australia extending from temperate Tasmania to tropical northern Queensland. Despite testing a large number of strains and repeated testing of the cline ends, we did not detect any evidence for clinal variation in virgin egg retention. Therefore although D. melanogaster in temperate Australia overwinter at the adult stage, there is no evidence for selection on virgin retention capacity producing clinal patterns. This contrasts with other evidence for clinal variation in egg production patterns over winter.     

极端生理耐受性导致果蝇Zaprionus indianus 在温带地区广泛分布（英文）

在过去的十年中, Zaprionus indianus这一温带适应性果蝇已经入侵印度次大陆, 并扩大了其在该地区的分布。Z. indianus能成功入侵是由于它具有很强的适应性和对极端生理条件的耐受性。对Z. indianus (温带狭域分布种类) 和黑腹果蝇Drosophila melanogaster (全球广域分布种类)在极端温度下未成熟期和成虫期发育阈值的比较研究表明, 两者的死亡率和发育起点温度存在显著差异。为了检测越冬期间未成熟期和成虫期抗逆性和存活率的变化, 以采自印度温带和热带不同地点的Z. indianus种群进行饲养实验。在温带地区的田间养虫笼中以及恒定的实验室条件下监测这些种群的卵孵化率和成虫存活率, 直至全部成虫死亡。结果表明, 由于温带地区卵孵化率和存活率高, 导致总的孵化率和存活率在不同纬度间存在显著差异。卵至成虫发育实验结果表明, 低温条件下产下的卵在温度适中时成功发育成成虫。由此可见, 这种昆虫在未成熟期具有的气候适应性以及在成虫期具有的抗逆性可为该物种提供季节性保护。考虑到气候变暖情况, 即温度增加0.6℃, 温度的少许改变都可能导致种群存活能力的显著增强和发育历期缩短。这些结果可解释Z. indianus为什么能够轻易突破障碍并适应新的环境。     

Climatic selection on genes and traits after a 100 year-old invasion: a critical look at the temperate-tropical clines in Drosophila melanogaster from eastern Australia

Drosophila melanogaster invaded Australia around 100 years ago, most likely through a northern invasion. The wide range of climatic conditions in eastern Australia across which D. melanogaster is now found provides an opportunity for researchers to identify traits and genes that are associated with climatic adaptation. Allozyme studies indicate clinal patterns for at least four loci including a strong linear cline in Adh and a non-linear cline in alpha-Gpdh. Inversion clines were initially established from cytological studies but have now been validated with larger sample sizes using molecular markers for breakpoints. Recent collections indicate that some genetic markers (Adh and In(3R)Payne) have changed over the last 20 years reflecting continuing evolution. Heritable clines have been established for quantitative traits including wing length/area, thorax length and cold and heat resistance. A cline in egg size independent of body size and a weak cline in competitive ability have also been established. Postulated clinal patterns for resistance to desiccation and starvation have not been supported by extensive sampling. Experiments under laboratory and semi-natural conditions have suggested selective factors generating clinal patterns, particularly for reproductive patterns over winter. Attempts are being made to link clinal variation in traits to specific genes using QTL analysis and the candidate locus approach, and to identify the genetic architecture of trait variation along the cline. This is proving difficult because of inversion polymorphisms that generate disequilibrium among genes. Substantial gaps still remain in linking clines to field selection and understanding the genetic and physiological basis of the adaptive shifts. However D. melanogaster populations in eastern Australia remain an excellent resource for understanding past and future evolutionary responses to climate change.     

Temperature tolerance of the diamondback moth, Plutella xylostella (Lepidoptera: Yponomeutidae) in tropical and temperate regions of Asia

Temperature tolerance was investigated in nine populations of Plutella xylostella Linnaeus from tropical and temperate regions of Asia. At all rearing temperatures between 15 and 35 degrees C, no clear differences were observed in female egg production or larval development between tropical and temperate populations. Thus, tropical populations did not show a high-temperature tolerance superior to that of the temperate populations. In all populations, the net reproductive rate (number of new females born per female) largely depended on the number of eggs laid per female, and egg production significantly decreased with increasing temperature (P < 0.001). Larval developmental rate also showed a significant positive correlation with temperature (P < 0.001). Per cent hatch of eggs and larval survival did not show a significant correlation with temperature: hatching was constant between 15 and 32.5 degrees C, but considerably lower at 35 degrees C. Larval survival was similar between 15 and 30 degrees C, appreciably lower at 32.5 degrees C and declined to 0% at 35 degrees C. Based on these results, environmental conditions under which P. xylostella can maintain a high population density throughout the year in tropical and subtropical regions are discussed.     

Clinal variation in post‐winter male fertility retention; an adaptive overwintering strategy in Drosophila melanogaster

In insects including Drosophila melanogaster, females can overwinter at the adult stage by adopting a shallow reproductive diapause, but almost nothing is known about male reproductive diapause. In this study, we test for the maintenance of fertility in overwintering males from the eastern Australian D. melanogaster cline. Males from southern temperate populations maintained in field cages in temperate Melbourne had a higher fertility in spring compared with males from tropical locations. Temperate males successfully inseminated more females, and there were also more offspring produced per inseminated female. The resulting linear post‐winter fertility clines were unrelated to male body size. In contrast, there was no clinal variation for fertility in nonoverwintering males held in the laboratory. The cline in overwintering male fertility is likely to have evolved as an adaptive response to latitudinal climatic variation over the last 100 years.     

Geographic variation in diapause incidence, life-history traits, and climatic adaptation in Drosophila melanogaster

In Drosophila melanogaster, exposure of females to low temperature and shortened photoperiod can induce the expression of reproductive quiescence or diapause. Diapause expression is highly variable within and among natural populations and has significant effects on life-history profiles, including patterns of longevity, fecundity, and stress resistance. We hypothesized that if diapause expression is associated with overwintering mechanisms and adaptation to temperate environments, the frequency of diapause incidence would exhibit a latitudinal cline among natural populations. Because stress resistance and reproductive traits are also clinal in this species, we also examined how patterns of fecundity and longevity varied with geography and how stress resistance and associated traits differed constitutively between diapause and nondiapause lines. Diapause incidence was shown to vary predictably with latitude, ranging from 35% to 90% among natural populations in the eastern United States Survivorship under starvation stress differed between diapause and nondiapause lines; diapause phenotypes were also distinct for total body triglyceride content and the developmental distribution of oocytes in the ovary following stress exposure. Patterns of longevity, fecundity, and ovariole number also varied with geography. The data suggest that, for North American populations, diapause expression is functionally associated with overwintering mechanisms and may be an integral life-history component in natural populations.     

Adaptation to a seasonally varying environment: a strong latitudinal cline in reproductive diapause combined with high gene flow in Drosophila montana

Adaptation to seasonal changes in the northern hemisphere includes an ability to predict the forthcoming cold season from gradual changes in environmental cues early enough to prepare for the harsh winter conditions. The magnitude and speed of changes in these cues vary between the latitudes, which induces strong selection pressures for local adaptation.We studied adaptation to seasonal changes in Drosophila montana, a northern maltfly, by defining the photoperiodic conditions leading to adult reproductive diapause along a latitudinal cline in Finland and by measuring genetic differentiation and the amount of gene flow between the sampling sites with microsatellites. Our data revealed a clear correlation between the latitude and the critical day length (CDL), in which half of the females of different cline populations enter photoperiodic reproductive diapause. There was no sign of limited gene flow between the cline populations, even though these populations showed isolation by distance. Our results show that local adaptation may occur even in the presence of high gene flow, when selection for locally adaptive life-history traits is strong. A wide range of variation in the CDLs of the fly strains within and between the cline populations may be partly due to gene flow and partly due to the opposing selection pressures for fly reproduction and overwinter survival. This variation in the timing of diapause will enhance populations' survival over the years that differ in the severity of the winter and in the length of the warm period and may also help them respond to long-term changes in environmental conditions.     

Latitudinal compensation in female reproductive rate of a geographically widespread reef fish

Latitudinal variation in fitness-related traits has often been attributed to local adaptation to climates. In poikilotherms including fishes, lower temperatures and shorter reproductive seasons at high latitudes would be expected to cause a reduction in annual reproductive output of an individual. Theories of latitudinal compensation predict that organisms at high latitudes should evolve compensatory responses for these climatic effects. Therefore, latitudinal compensation in female reproductive rate (egg production rate), that individuals from high latitudes produce eggs at higher rates than those from lower latitudes, is likely to occur. I tested this hypothesis with a latitudinally widespread reef fish Pomacentrus coelestis that is a multiple batch spawner, from three different localities, from temperate to subtropical waters, within Japan. I used common-environment experiments at three different temperatures to compare reproductive capacity among local populations. In the experiments, average inter-spawning intervals were the shortest and average size-specific clutch weight was the heaviest in fish from the most northern locality across all temperatures, showing clear latitudinal clines. Thus, the northern fish can achieve higher reproductive output per unit time both by shortening inter-spawning intervals and increasing size-specific clutch weight. Additionally, faster egg production rate of the northern fish did not result from increased food consumption. This finding suggests that gross egg production efficiency was higher in the northern fish and that northern fish had a superior capacity for reproduction within a season. These results support the prediction that latitudinal compensation occurs in the female reproductive rate of P. coelestis. As the reproductive season of this species decreases drastically with increasing latitude, the observed cline in the reproductive rate must be an adaptive response to the local selective regime, i.e., length of the reproductive season. Such latitudinal compensation in female reproductive rates may be a common pattern in latitudinally widespread fishes.     

Genotype-by-environment interactions and adaptation to local temperature affect immunity and fecundity in Drosophila melanogaster

Natural populations of most organisms harbor substantial genetic variation for resistance to infection. The continued existence of such variation is unexpected under simple evolutionary models that either posit direct and continuous natural selection on the immune system or an evolved life history "balance" between immunity and other fitness traits in a constant environment. However, both local adaptation to heterogeneous environments and genotype-by-environment interactions can maintain genetic variation in a species. In this study, we test Drosophila melanogaster genotypes sampled from tropical Africa, temperate northeastern North America, and semi-tropical southeastern North America for resistance to bacterial infection and fecundity at three different environmental temperatures. Environmental temperature had absolute effects on all traits, but there were also marked genotype-by-environment interactions that may limit the global efficiency of natural selection on both traits. African flies performed more poorly than North American flies in both immunity and fecundity at the lowest temperature, but not at the higher temperatures, suggesting that the African population is maladapted to low temperature. In contrast, there was no evidence for clinal variation driven by thermal adaptation within North America for either trait. Resistance to infection and reproductive success were generally uncorrelated across genotypes, so this study finds no evidence for a fitness tradeoff between immunity and fecundity under the conditions tested. Both local adaptation to geographically heterogeneous environments and genotype-by-environment interactions may explain the persistence of genetic variation for resistance to infection in natural populations.     

INTERACTIVE EFFECTS OF OFFSPRING SIZE AND TIMING OF REPRODUCTION ON OFFSPRING REPRODUCTION: EXPERIMENTAL,MATERNAL, AND QUANTITATIVE GENETIC ASPECTS

We demonstrate that egg size in side-blotched lizards is heritable (parent-offspring regressions) and thus will respond to natural selection. Because our estimate of heritability is derived from free-ranging lizards, it is useful for predicting evolutionary response to selection in wild populations. Moreover, our estimate for the heritability of egg size is not likely to be confounded by nongenetic maternal effects that might arise from egg size per se because we estimate a significant parent-offspring correlation for egg size in the face of dramatic experimental manipulation of yolk volume of the egg. Furthermore, we also demonstrate a significant correlation between egg size of the female parent and clutch size of her offspring. Because this correlation is not related to experimentally induced maternal effects, we suggest that it is indicative of a genetic correlation between egg size and clutch size. We synthesize our results from genetic analyses of the trade-off between egg size and clutch size with previously published experiments that document the mechanistic basis of this trade-off. Experimental manipulation of yolk volume has no effect on offspring reproductive traits such as egg size, clutch size, size at maturity, or oviposition date. However, egg size was related to offspring survival during adult phases of the life history. We partitioned survival of offspring during the adult phase of the life history into (1) survival of offspring from winter emergence to the production of the first clutch (i.e., the vitellogenic phase of the first clutch), and (2) survival of the offspring from the production of the first clutch to the end of the reproductive season. Offspring from the first clutch of the reproductive season in the previous year had higher survival during vitellogenesis of their first clutch if these offspring came from small eggs. We did not observe selection during these prelaying phases of adulthood for offspring from later clutches. However, we did find that later clutch offspring from large eggs had the highest survival over the first season of reproduction. The differences in selection on adult survival arising from maternal effects would reinforce previously documented selection that favors the production of small offspring early in the season and large offspring later in the season—a seasonal shift in maternal provisioning. We also report on a significant parent-offspring correlation in lay date and thus significant heritable variation in lay date. We can rule out the possibility of yolk volume as a confounding maternal effect—experimental manipulation of yolk volume has no effect on lay date of offspring. However, we cannot distinguish between genetic effects (i.e., heritable) and nongenetic maternal effects acting on lay date that arise from the maternal trait lay date per se (or other unidentified maternal traits). Nevertheless, we demonstrate how the timing of female reproduction (e.g., date of oviposition and date of hatching) affect reproductive attributes of offspring. Notably, we find that date of hatching has effects on body size at maturity and fecundity of offspring from later clutches. We did not detect comparable effects of lay date on offspring from the first clutch.     

The influence of gene flow on latitudinal clines of photoperiodic adult diapause in the Drosophila auraria species-complex

Temperate species belonging to the Drosophila auraria species complex, D. auraria Peng, D. biauraria Bock & Wheeler, D. triauraria Bock & Wheeler and D. subauraria Kimura, enter reproductive diapause to pass the winter in response to short daylengths. These species from Japan showed latitudinal clines in critical daylength which is longer in populations from higher latitudes. The slopes of these clines coincided well with that of the cline which is approximately predicted from climatic data, suggesting that these clines result from adaptation of the species to the latitudinal gradient of climatic conditions. Between the mainlands and the surrounding islands of Japan, the slopes of clines did not differ significantly, but the deviation from the regression line was usually smaller in mainland populations. It is assumed that gene flow reduces the genetic variation among mainland populations and results in the development of smooth clines. In the plain of east China, D. triauraria did not show clinal variation in critical daylength, although the development of the cline is expected from climatic data. Extensive gene flow among Chinese populations is considered to prevent the development of a cline.     

Variable maternal control of facultative egg diapause in the bushcricket Ephippiger ephippiger

1. Diapause is a term used to describe an arrest in the development of insects as adults, eggs, or embryos, and allows survival during adverse environmental conditions. 2. The influence of maternal age on the proportion of eggs that entered an initial facultative diapause and on fecundity and egg mortality was investigated in the bushcricket Ephippiger ephippiger. 3. In the absence of variable abiotic cues such as temperature and photoperiod, most E. ephippiger females increase the proportion of their eggs that enter diapause as they age, however there are large differences among females, with a minority showing different trends. Fecundity and egg survival decline with female age and also differ among females. 4. The influence of age on the incidence of diapause is likely to be an adaptive reproductive strategy, as those eggs laid later in the season are unlikely to complete egg development before the onset of winter. Females are probably hedging their bets by diversifying the rate of development among their offspring and effectively spreading development over several seasons. The low predictability of future environmental conditions might explain the heterogeneity of female reproductive strategies.     

Life-history strategies of Australian lizards: a comparison between the tropics and the temperate zone

Summary Previous studies have suggested that tropical and temperate-zone lizards may differ fundamentally in life histories. We tested the applicability of this idea to Australian species by comparing temperate-zone species of agamid and scincid lizards with their congeners from the seasonal tropics. Data were derived from dissection of 1,941 specimens and from published information. Clutch size and egg size were positively correlated with mean maternal body size in most lizard species from both climatic zones. Mean body size of the lizards studies did not differ between the tropics and the temperate zone, nor did egg or hatchling size. However, tropical skinks showed considerably (approximately 20%) lower clutch size and relative clutch mass than did temperate-zone skinks. This difference was partly due to the higher incidence of species with low, invariant clutch size in the tropical lizard fauna (as seen in other continents as well), but primarily due to a trend for lineages (especially genera) with relatively high fecundity to be more common in the temperate zone than in the tropics. In contrast to studies on African lizards, our data suggested that modification of clutch size between areas has not occurred within genera: congeneric species from the tropics and temperate zone did not differ in clutch size. Production of more than one clutch per annum by individual females was common in both climatic zones. Tropical lizards may differ from temperate-zone species in showing higher reproductive frequencies, more rapid growth and earlier maturation. However, most of these effects may be due to phenotypic responses to environmental conditions (especially longer annual activity season), rather than to genetically based lifehistory adaptations.     

Environmental heterogeneity and the maintenance of genetic variation for reproductive diapause in Drosophila melanogaster

Drosophila melanogaster has colonized temperate habitats on multiple continents over a historical time period, and many traits vary predictably with latitude. Despite considerable attention paid to clinal variation in Drosophila, the mechanisms generating such patterns in nature remain largely unidentified. In D. melanogaster, the expression of reproductive diapause can be induced by exposure to low temperatures and shortened photoperiods. Both diapause expression itself and the underlying genetic variance for diapause expression have widespread impacts on organismal fitness, and diapause incidence exhibits a 60% cline in frequency in the eastern United States. The major aim of this study was to evaluate whether the relative fitness of diapause and nondiapause genotypes varies predictably with environment. In experimental population cages in the laboratory, the frequency of genotypes that express diapause increased over time when flies were exposed to environmental stress, whereas the frequency of nondiapause genotypes increased when flies were cultured under benign control conditions. Other fitness traits correlated with the genetic variance for diapause expression (longevity, mortality rates, stress resistance, lipid content, preadult viability, fecundity profiles, and development time) also diverged between experimental treatments. Similarly, sampling of isofemale lines from natural populations revealed that the frequency of diapause incidence cycled over time in seasonal habitats: diapause expression was at high frequency following the winter season and subsequently declined throughout the summer months. In contrast, diapause expression was low and temporally homogeneous in isofemale line collections from human-associated urban habitats. These data suggest that genetic variation underlying the diapause-nondiapause dichotomy may be actively maintained by selection pressures that vary spatially and temporally in natural populations.     

Temperature,size, reproductive allocation,and life‐history evolution in a gregarious caterpillar

The present study aimed to investigate the relationship between growth rate, final mass, and larval development, as well as how this relationship influences reproductive trade‐offs, in the context of a gregarious life‐style and the need to keep an optimal group size. We use as a model two sympatric populations of the pine processionary moth Thaumetopoea pityocampa, which occur in different seasons and thus experience different climatic conditions. Thaumetopoea pityocampa is a strictly gregarious caterpillar throughout the larval period, which occurs during winter in countries all over the Mediterranean Basin. However, in 1997, a population in which larval development occurs during the summer was discovered in Portugal, namely the summer population (SP), as opposed to the normal winter population (WP), which coexists in the same forest feeding on the same host during the winter. Both populations were monitored over 3 years, with an assessment of the length of the larval period and its relationship with different climatic variables, final mass and adult size, egg size and number, colony size, and mortality at different life stages. The SP larval period was reduced as a result of development in the warmer part of the year, although it reached the same final mass and adult size as the WP. Despite an equal size at maturity, a trade‐off between egg size and number was found between the two populations: SP produced less but bigger eggs than WP. This contrasts with the findings obtained in other Lepidoptera species, where development in colder environments leads to larger eggs at the expense of fecundity, but corroborates the trend found at a macro‐geographical scale for T. pityocampa, with females from northern latitudes and a colder environment producing more (and smaller) eggs. The results demonstrate the importance of the number of eggs in cold environments as a result of an advantage of large colonies when gregarious caterpillars develop in such environments, and these findings are discussed in accordance with the major theories regarding size in animals. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 340–349.     

No patterns in thermal plasticity along a latitudinal gradient in Drosophila simulans from eastern Australia

Phenotypic plasticity may be an important initial mechanism to counter environmental change, yet we know relatively little about the evolution of plasticity in nature. Species with widespread distributions are expected to have evolved higher levels of plasticity compared with those with more restricted, tropical distributions. At the intraspecific level, temperate populations are expected to have evolved higher levels of plasticity than their tropical counterparts. However, empirical support for these expectations is limited. In addition, no studies have comprehensively examined the evolution of thermal plasticity across life stages. Using populations of Drosophila simulans collected from a latitudinal cline spanning the entire east coast of Australia, we assessed thermal plasticity, measured as hardening capacity (the difference between basal and hardened thermal tolerance) for multiple measures of heat and cold tolerance across both adult and larval stages of development. This allowed us to explicitly ask whether the evolution of thermal plasticity is favoured in more variable, temperate environments. We found no relationship between thermal plasticity and latitude, providing little support for the hypothesis that temperate populations have evolved higher levels of thermal plasticity than their tropical counterparts. With the exception of adult heat survival, we also found no association between plasticity and ten climatic variables, indicating that the evolution of thermal plasticity is not easily predicted by the type of environment that a particular population occupies. We discuss these results in the context of the role of plasticity in a warming climate.     

Temperature dependent larval resource allocation shaping adult body size in Drosophila melanogaster

Geographical variation in Drosophila melanogaster body size is a long-standing problem of life-history evolution. Adaptation to a cold climate invariably produces large individuals, whereas evolution in tropical regions result in small individuals. The proximate mechanism was suggested to involve thermal evolution of resource processing by the developing larvae. In this study an attempt is made to merge proximate explanations, featuring temperature sensitivity of larval resource processing, and ultimate approaches focusing on adult and pre-adult life-history traits. To address the issue of temperature dependent resource allocation to adult size vs. larval survival, feeding was stopped at several stages during the larval development. Under these conditions of food deprivation, two temperate and two tropical populations reared at high and low temperatures produced different adult body sizes coinciding with different probabilities to reach the adult stage. In all cases a phenotypic trade-off between larval survival and adult size was observed. However, the underlying pattern of larval resource allocation differed between the geographical populations. In the temperate populations larval age but not weight predicted survival. Temperate larvae did not invest accumulated resources in survival, instead they preserved larval biomass to benefit adult weight. In other words, larvae from temperate populations failed to re-allocate accumulated resources to facilitate their survival. A low percentage of the larvae survived to adulthood but produced relatively large flies. Conversely, in tropical populations larval weight but not age determined the probability to reach adulthood. Tropical larvae did not invest in adult size, but facilitated their own survival. Most larvae succeeded in pupating but then produced small adults. The underlying physiological mechanism seemed to be an evolved difference in the accessibility of glycogen reserves as a result of thermal adaptation. At low rearing temperatures and in the temperate populations, glycogen levels tended to correlate positively with adult size but negatively with pupation probability. The data presented here offer an explanation of geographical variation in body size by showing that thermal evolution of resource allocation, specifically the ability to access glycogen storage, is the proximate mechanism responsible for the life-history trade-off between larval survival and adult size.     

Intraspecific variability in egg maturation patterns and associated life‐history trade‐offs in a polyembryonic parasitoid wasp

1. Life‐history theory predicts a trade‐off between the resources allocated to reproduction and those allocated to survival. Early maturation of eggs (pro‐ovigeny) is correlated with small body size and low adult longevity in interspecific comparisons among parasitoids, demonstrating this trade‐off. The handful of studies that have tested for similar correlations within species produced conflicting results. 2. Egg maturation patterns and related life‐history traits were studied in the polyembryonic parasitoid wasp, Copidosoma koehleri (Hymenoptera: Encyrtidae). Although the genus Copidosoma was previously reported to be fully pro‐ovigenic, mean egg loads of host‐deprived females almost doubled within their first 6 days of adulthood. 3. The initial egg‐loads of newly emerged females were determined and age‐specific realised fecundity curves were constructed for their clone‐mate twins. The females' initial egg loads increased with body size, but neither body size nor initial egg load was correlated with longevity and fecundity. 4. The variation in initial egg loads was lowest among clone‐mates, intermediate among non‐clone sisters and highest among non‐sister females. The within‐clone variability indicates environmental influences on egg maturation, while the between‐clone variation may be genetically based. 5. Ovaries of host‐deprived females contained fewer eggs at death (at ～29 days) than on day 6. Their egg loads at death were negatively correlated with life span, consistent with reduced egg production and/or egg resorption. Host deprivation prolonged the wasps' life span, suggesting a survival cost to egg maturation and oviposition. 6. It is concluded that adult fecundity and longevity were not traded off with pre‐adult egg maturation.