A trade-off between diapause duration and fitness in female parasitoids

Abstract 1. Diapause allows insects to cope with adverse weather conditions but also poses substantial fitness costs, e.g. through reduced survival of the diapausing stage or sublethal effects in the following season.
2. The energetic costs of diapause in females of the parasitoid Asobara tabida Nees (Hymenoptera: Braconidae) were assessed using experimental manipulation of diapause duration.
3. An increase in diapause length not only led to higher mortality among diapausing pupae but also caused a significant decrease in egg load, fat reserves, and dry weight of the emerging adult females.
4. Only larvae with sufficient resources were able to survive the entire diapause period and there was a trade-off between the metabolic costs of diapause and adult fitness components.
5. In contrast, the size (tibia length) of emerging females increased with increasing diapause duration. It is proposed that this may be due to a genetic correlation between larval dry weight and adult size, so that selection for heavier larvae would result in larger adults even when resources are limited.
6. Ultimately, the evolution of diapause duration and its consequences for adult performance will depend on the costs of overwintering and the ecological conditions in the habitat.     

Age-dependent clutch size in a koinobiont parasitoid

Abstract.  1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid.
2. From egg-limitation and time-limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host-deprived individuals.
3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured.
4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host-deprivation time.
5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host-deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size.
6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short.
7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size.     

Components of male fitness in relation to body size in Epirrita autumnata (Lepidoptera, Geometridae)

Abstract. 1. The effect of body size on different components of male fitness was studied for Epirrita autumnata , a geometrid known for its eruptive population dynamics. Body size is the main determinant of female fecundity in this species.
2. Longevity of males was found to have a weak negative correlation with body size at low temperatures. No significant correlation was found at higher temperatures.
3. We found no correlation between male size and female fecundity or egg size which is consistent with the small size of spermatophores in this species.
4. Small and large males were equally successful when allowed to compete for females in laboratory conditions.
5. In one or two field collections, males found mating were larger than males found singly. Large males also had an advantage in finding of virgin females, offered experimentally. No size-assortative mating was recorded.
6. We conclude that size-dependent mate location ability is the factor accounting for most of the variance in male fitness in E.autumnata. The dependence of fitness on body size may well be equally strong in males and females.     

Dispersal and foraging behaviour of Platygaster californica: hosts can't run, but they can hide

Abstract.  1. Host–parasitoid models often identify foraging behaviour and dispersal distance as important for system persistence.
2. Laboratory observations and field trials were used to characterise foraging behaviour and dispersal capability of Platygaster californica Ashmead (Platygasteridae), a parasitoid of the gall midge Rhopalomyia californica Felt (Cecidomyiidae).
3. Although foraging parasitoids meticulously searched plants in laboratory observations, none of the laboratory trials resulted in 100% parasitism, and the proportion of parasitism declined as midge egg density increased.
4. The field trials showed that the distribution of parasitism over distance from a central release point was hump-shaped, as predicted by a simple diffusion model. Mean parasitoid dispersal distance was 4.5 m, considerably farther than the 1.7 m mean midge dispersal found in previous work.
5. Although the parasitoid appears to search thoroughly for midge eggs and to disperse farther than the midge, the results of this study show how this host–parasitoid system may persist due to spatially variable incomplete parasitism.     

Effects of size and fluctuating asymmetry on field fitness of the parasitoid Trichogramma carverae (Hymenoptera: Trichogrammatidae)

1. The effects of body size and asymmetry in morphological traits on field fitness were studied in the parasitoid Trichogramma carverae .
2. Significant fluctuating asymmetry was detected in four bristle counts and two wing measurements made on forewings. There was no evidence for directional asymmetry in any of the traits. An estimate of field fitness was obtained in grapevines by collecting released wasps arriving at oviposition sites consisting of egg rafts of the tortricid Epiphyas postvittana .
3. Comparisons of ovipositing and emergence samples indicated that wasps at oviposition sites were relatively larger and more variable in their size distribution. A non-parametric analysis of the relationship between fitness and size indicated that extremely large wasps had the highest fitness and suggested that small as well as large wasps may have had a fitness advantage.
4. For asymmetry, the only trait showing an association with field fitness was wing length. Wasps with a low length asymmetry were more likely to be collected at oviposition sites, although fitness curves indicated that wasps had a similar fitness once an intermediate length asymmetry was exceeded.
5. Mother–daughter comparisons for wasps from a genetically heterogeneous stock provided no evidence that size measures or asymmetries were heritable when wasps were reared on a factitious host.
6. These findings have implications for improving parasitism rates in inundative releases.     

The dynamics of egg production, oviposition and resorption in a parasitoid wasp

1. The extent to which parasitoid wasps are limited by their egg supply is very important in understanding their reproductive strategies. Egg reserves are dynamic, with most wasps maturing new eggs throughout their life (synovigeny) and many species resorbing eggs that are not used in oviposition. We investigated the extent to which a parasitoid modulates its egg reserves in the light of its experience in finding hosts.
2. The egg dynamics of the Encyrtid Wasp Leptomastix dactylopii , a solitary parasitoid of mealybugs, were studied in the laboratory. This species is synovigenic and practises egg resorption.
3. We allowed newly emerged wasps to experience one of four environments of increasing value in terms of reproductive opportunities. We proposed that wasps that experienced good quality environments would maintain more mature eggs ready for oviposition. Dissection of wasps subject to different periods of host deprivation after the experimental treatment failed to confirm the hypothesis: egg load was independent of experience.
4. We also proposed that any adjustment of egg supply to make up for eggs oviposited would be effected through a reduction in egg resorption. Instead, we found that the wasp quickly made up for eggs oviposited by increased egg production.     

Relationships between oviposition date, hatch date, and offspring size in the grasshopper Chorthippus brunneus

Abstract 1. The grasshopper Chorthippus brunneus has been shown to increase egg size with maternal age under constant laboratory conditions, such that late-laid eggs are larger than early laid eggs. In this study, an increase in the size of eggs (and hatchlings from those eggs) was recorded with date of oviposition in a field cage population. This suggests that the relationship between maternal age and egg size, observed previously in the laboratory, also occurs in the field. There was, however, some evidence that the behaviour of the maternal females in field cages was modified by the onset of autumnal weather at the end of the breeding season. Only a small number of eggs was laid in the last 3 weeks of the summer but these yielded relatively small hatchlings.
2. The date on which eggs were laid was correlated positively with their date of hatch in the following year. A consequence of this relationship, and that between oviposition date and egg size, was that size at hatch increased with date of hatch through most of the spring but then declined as the last few eggs hatched. Temporal variation in size at hatch parallelled temporal variation in the maternal females' investment in egg size in the previous year.
3. An undisturbed field population was monitored to assess whether the temporal variation in size at hatch resulted in size variation at eclosion in subsequent developmental stages. There were negative correlations between size at eclosion and date of eclosion at the third and fourth instars, suggesting that despite their smaller size at hatch, early hatchers experienced more favourable conditions for juvenile growth than late hatchers. The larger size at hatch of late hatchers may enhance their survival and compensate (albeit incompletely) for their reduced opportunity for growth. Female reproductive behaviour may represent an adaptive response to a predictable seasonal decline in offspring fitness at hatch.     

Survival of colour-polymorphic Eristalis arbustorum hoverflies in semi-field conditions

1. The theory on adaptive phenotypic plasticity assumes different fitness optima in different environments. To demonstrate these fitness differences in the field or under laboratory conditions can be difficult.
2. The size of the coloured patches on the abdomen of Eristalis arbustorum varies with rearing temperature. At low temperatures small patches develop and at higher temperatures larger patches develop. Consequently, the colour patch size varies seasonally.
3. To study the relative fitness of two extreme phenotypes in abdominal coloration in the summer, a total of 591 laboratory-reared flies were released in six large population cages. As a measure of fitness, survival was estimated.
4. In all replicate cages the survival of the pale phenotypes was higher than that of the dark phenotypes, indicating a higher fitness for the pale flies under summer conditions.
5. The fat content of flies reared under identical conditions to those released in the cages was measured. Pale flies had ≈ 0·7 mg more fat than dark flies (± 25% extra fat). However, it is argued that this difference in fat reserves could not have been the only cause of the observed difference in survival.
6. The results are discussed in relation to mimicry and thermoregulation.     

Female size and fitness in the leaf-cutter bee Megachile apicalis

1. The effects of female size on fitness of the leaf-cutter bee Megachile apicalis Spinola (Hymenoptera: Megachilidae) were examined using artificial nesting sites in the field.
2. Although female size had no significant effect on female longevity or sex ratio of progeny, it did have a significant effect on fecundity; larger females attained greater realized fecundity than did smaller females.
3. This significant effect of female size on fecundity occurred because larger females produced cells faster than did smaller females.
4. Female size also had a significant effect on egg size; larger females laid larger eggs than did smaller females.
5. Female size had a significant effect on the size of investment in each progeny. The size of investment estimated by brood cell weight was greater for larger females than for smaller females. This pattern was largely absent, however, when the size of investment was estimated by adult progeny weight.
6. Female size had a significant effect on nest usurpation behaviour; larger females had a higher capacity to usurp nests than did smaller females.     

Body size, host choice and sex allocation in a spider-hunting pompilid wasp

Two important relationships in parasitoid evolutionary ecology are those between adult size and fitness and between host quality and sex ratio. Sexually differential size–fitness relationships underlie predicted sex-ratio relationships. Despite each relationship receiving considerable attention, they have seldom been studied simultaneously or using field data. Here we report the biology of Anoplius viaticus paganu s Dahlbom, a little known parasitoid of spiders, using field and laboratory data. We found that larger foraging females were able to select larger host spiders from the field, thus identifying a relatively novel component of the size–fitness relationship. Larger offspring developed from larger hosts and, in agreement with the prediction of the host quality model of sex allocation, were generally female. Data on the size–fitness relationship for males are lacking and, in common with many prior studies, we could not evaluate sexually differential size–fitness relationships as an explanation for the observed sex-ratio patterns. Nonetheless, A. v. paganu s exhibited one of the strongest relationships between host size and offspring sex ratio yet reported.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 285–296.     

Physiological costs of cold exposure on the parasitoid <Emphasis Type="Italic">Aphidius ervi</Emphasis>, without selection pressure and under constant or fluctuating temperatures

The aim of this study is to evaluate the physiological costs and the consequences on fitness of cold exposure on the parasitoid Aphidius ervi (Hymenoptera: Aphidiinae) under non-lethal conditions. We exposed 1-day-old mummies to different treatments: control at 20°C, 7°C constant, 7°C fluctuating (7°C for 22 h and 20°C for 2 h). Two performances of fitness were particularly affected after two weeks at 7°C constant: female longevity and the sex ratio of the progeny was male biased. In contrast, egg load at emergence, lifetime fecundity, frequency of asymmetric individuals in both sexes, and mating success were not significantly affected under all treatments. The effects of these treatments on trends such as utilisation of fat reserves and fertility of males and females are discussed.     

A comparison of life history traits of sexual and asexual strains of the parasitoid wasp,Lysiphlebus fabarum (Braconidae: Aphidiinae)

1. This study examined biological characteristics of sexual and asexual strains of the parasitoid wasp, Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae). 2. Strains were reared in different instar hosts (the black bean aphid, Aphis fabae Scopoli) under identical environmental conditions (21 °C, 65–75% RH, and LD 16:8 h). 3. Results showed that the second instar of the aphid is the most suitable growth stage for both strains, as the wasps that emerged from the second instar hosts were larger, more fecund, and had larger egg size. Trade‐offs between the fitness components of the parasitoid were clearer when the parasitoids were reared in suboptimal instars. 4. According to the results, sexual females emerged around 1 day earlier and lived around 0.5 day less than asexual females. Also, sexual females emerged with a lower initial egg load, although these wasps tend to have larger eggs than asexual females. Asexual females may enjoy greater longevity and higher developmental plasticity which suggests a higher degree of synchronization with pest population dynamism. 5. The results suggest that sexual wasps, in contrast to asexual wasps, invest more in egg size than in egg load. This study suggests strain‐specific adaptations of L. fabarum to different instars of the black bean aphid by which the allocation of nutritional resources to various functions differs between strains. 6. Furthermore, differences in life history traits between strains can greatly influence the population dynamics of each strain, and hence their effectiveness in suppressing pest populations.     

Maternal effects and their consequences for offspring fitness in the Yellow Dung Fly

1. Maternal adult diet and body size influence the fecundity of a female and possibly the quality and the performance of her offspring via egg size or egg quality. In laboratory experiments, negative effects in the offspring generation have often been obscured by optimal rearing conditions.
2. To estimate these effects in the Yellow Dung Fly, Scathophaga stercoraria , how maternal body size and adult nutritional status affected her fecundity, longevity and egg size were first investigated.
3. Second, it was investigated how female age and adult nutritional experience, mediated through the effects of egg size or egg quality, influenced the performance of offspring at different larval densities.
4. Maternal size was less important than maternal adult feeding in increasing reproductive output. Without food restriction, large females had larger clutch sizes and higher oviposition rates, whereas under food restriction this advantage was reversed in favour of small females.
5. Offspring from mothers reared under nutritional stress experienced reduced fitness in terms of egg mortality and survival to adult emergence. If the offspring from low-quality eggs survived, the transmitted maternal food deficiency only affected adult male body size under stressful larval environments.
6. Smaller egg sizes due to maternal age only slightly affected the performance of the offspring under all larval conditions.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999     

Development of the solitary endoparasitoid Microplitis demolitor: host quality does not increase with host age and size

Abstract.  1. Many studies examining the relationship between host size, an index of host quality, and parasitoid fitness use development time and/or adult parasitoid size as currencies of fitness, while ignoring pre-adult mortality. Because the physiological suitability of the host may vary in different stages, sizes, or ages of hosts, a misleading picture of host quality may therefore be obtained in cases where fitness is based on only one or two developmental traits.
2. The development of the solitary koinobiont endoparasitoid Microplitis demolitor is examined in different larval age-classes of its host the soybean looper Pseudoplusia includens . Hosts were parasitised on days 1–8 after hatching from the egg, and development time, adult body size, and mortality of the parasitoid were compared.
3. A comparison of larval growth trajectories (using dry body mass) of M. demolitor revealed that parasitoid larvae attained over twice as much body mass in old hosts than in younger hosts. Similarly, adult parasitoid size at eclosion generally increased with host size, although parasitoids developing in smaller hosts lost a much lower proportion of mass between pupation and eclosion.
4. Overall egg-to-adult development was most rapid in intermediate-aged hosts, and longer in hosts at opposite ends of the age continuum. Moreover, parasitoid mortality varied non-linearly with host stage, and was generally higher in very young and older hosts.
5. Based on these results and other empirical data for koinobionts, it is argued that fitness functions in this group of parasitoids are not simply a positive function of host size or age, but instead may be distinctly dome-shaped, both patterns reflecting the degree of physiological and nutritional compatibility between the two organisms.     

Preference and performance of the hyperparasitoid Syrphophagus aphidivorus (Hymenoptera: Encyrtidae): fitness consequences of selecting hosts in live aphids or aphid mummies

Abstract.  1. Theoretical models predict that ovipositional decisions of parasitoid females should lead to the selection of the most profitable host for parasitoid development. Most parasitoid species have evolved specific adaptations to exploit a single host stage. However, females of the aphid hyperparasitoid Syrphophagous aphidivorus (Mayr) (Hymenoptera: Encyrtidae) display a unique and atypical oviposition behaviour by attacking either primary parasitoid larvae in live aphids, or parasitoid pupae in dead, mummified aphids.
2. In the laboratory, the correlation between host suitability and host preference of S. aphidivorus on the host Aphidius nigripes Ashmead parasitising the aphid Macrosiphum euphorbiae (Thomas) was investigated.
3. The relative suitability of the two host stages was determined by measuring hyperparasitoid fitness parameters (survival, development time, fecundity, sex ratio, and adult size of progeny), and calculating the intrinsic rate of population increase ( r _m). Host preference by S. aphidivorus females and the influence of aphid defence behaviour on host selection was also examined.
4. Hyperparasitoid offspring performance was highest when developing from hosts in aphid mummies and females consistently preferred this host to hosts in parasitised aphids. Although aphid defensive behaviour may influence host selection, it was not a determining factor. Ecological and evolutionary processes that might have led to dual oviposition behaviour in S. aphidivorus are discussed.     

Spatial dynamics in a host–multiparasitoid community

1. The harlequin bug, a herbivore on bladderpod, is attacked by two specialist egg parasitoids Trissolcus murgantiae and Ooencyrtus johnsonii . Ooencyrtus can out-compete Trissolcus in the laboratory, but coexistence is the norm in field populations. Despite the heavy mortality inflicted by the two parasitoids, the host–parasitoid interaction is persistent in all sites that have been studied in southern California.
2. I manipulated inter-patch distances in a field experiment to determine whether spatial processes drive parasitoid coexistence and/or host–parasitoid dynamics. I first tested the hypothesis that the parasitoids coexist via a dispersal–competition trade-off. Both parasitoid species took significantly longer to colonize isolated patches than well-connected patches, suggesting that they have comparable dispersal abilities. Ooencyrtus did not exclude Trissolcus even when inter-patch distances were reduced to 25–30% of those observed in natural populations. These data suggest that parasitoid coexistence can occur in the absence of a dispersal advantage to the inferior competitor.
3. Since the treatments with isolated vs. well-connected patches did not differ in parasitoid composition, I next asked whether isolation would destabilize, or drive extinct, the host–multiparasitoid interaction. No local extinctions of bugs or parasitoids were observed during the 18-month study period. Bug populations in the isolated patches were no more variable than those in the well-connected patches. In fact, temporal variability in the experimentally isolated patches was comparable to that observed in highly isolated natural populations.
4. These data argue against a strong effect of spatial processes on host–parasitoid dynamics. Local processes may mediate both parasitoid coexistence as well as the host–parasitoid interaction.     

Superparasitism and sex ratio adjustment in a wasp parasitoid: results at variance with Local Mate Competition?

Anaphes nitens is a solitary parasitoid of the egg capsules of the Eucalyptus snout beetle, Gonipterus scutellatus. Some traits of its natural history suggest that Local Mate Competition (LMC) could account for sex ratio adjustment in this species. We tested whether males emerged early, a prerequisite for fully local mating, and investigated the occurrence and effect of superparasitism on adult size and pre-emergence mortality, factors that might influence sex ratio adjustment. We found in field-collected egg capsules that males emerged first. To investigate the effects of superparasitism on adult size, we compared the sizes of parasitoids that emerged early and late from egg capsules collected in the field, and from egg capsules parasitized and superparasitized in the laboratory. Superparasitism reduced parasitoid size, affecting females more strongly than males, and increased pre-emergence mortality. We estimated A. nitens sex ratio and parasitism rate in the field during 2 years in five localities and during 4 years in a sixth. Following LMC we expected an increase in sex ratio (proportion of males) with increasing parasitism rate (assumed to reflect parasitoid density). We found that sex ratio decreased from 0.38 when the parasitism rate was low (0-20%) to 0.21 when parasitism was high (80-100%). In contrast with field results, a laboratory experiment showed that: (1) at a low parasitism level sex ratio was clearly female biased (0.28+/-0.04), (2) at a high parasitism level sex ratio increased (0.40+/-0.07), (3) male larval survivorship was not lower than female survivorship, and (4) low-quality hosts (i.e. superparasitized) were allocated more males. We conclude that LMC cannot explain the sex ratio adjustment observed in the field, even at low parasitism rates, and alternative explications for highly female-biased sex ratios must be found. One such alternative is female-biased dispersal.     

Reproduction in an Irish lake population of the crayfish Austropotamobius pallipes (Lerehoullet)

SUMMARY. 1. Mature crayfish were collected in 1984 from an alkaline Irish lake before and after the mating season.
2. There was some correlation between frequency of mating and decrease in vasa deferentia weight in captive males, but in the field vasa deferentia weight losses were highly variable. The data indicate that one third of adult males may not have mated in the wild.
3. All females appear to have spawned, whether or not mated, with an average 85% reduction in ovary weights over the season.
4. Pleopodal egg counts directly after laboratory spawning were on average 81% of ovarian counts; in the field, an estimated 50 days after spawning, they were only 61% of estimated ovarian egg numbers.
5. Average ovarian egg size was positively correlated with female size. Also, larger crayfish had a wider range of egg sizes than smaller crayfish.
6. It is suggested that aggression and competition for shelter among brooding females may result in a progressive loss of eggs.     

Effects of temperature and season on egg size, hatchling size and adult size in Notiophilus biguttatus

1. Temperature-induced egg size variation and its effects on successive life stages in the carabid beetle Notiophilus biguttatus were examined.
2. In the laboratory, across temperature regimes, egg size and number were inversely related; number of eggs and total egg mass were higher, but egg size was smaller at high temperatures.
3. Food intake rate was shown not to be involved in the temperature effect on egg size.
4. Within the higher temperature regimes, among females, egg size was negatively correlated with number of eggs and with total egg mass.
5. Data on egg mortality and egg development time did not explain why at low temperature eggs were larger than at high temperature.
6. Larvae hatching from eggs produced at a low temperature were heavier than larvae from eggs produced at a high temperature, irrespective of temperature during development.
7. In a prolonged outdoor experiment (January – July), encompassing the main breeding period of N. biguttatus , egg size decreased and both egg production rate and total egg mass increased in the course of the experiment.
8. In the field, an effect of seasonal change in temperature on adult body size was found; teneral beetles that had their juvenile period early in the season were larger than those that had their juvenile period later in the season.
9. The results of the study suggest a mechanistic explanation in which the egg size response to temperature follows from a difference in temperature sensitivity between two processes in oogenesis; adaptiveness of the temperature response was not clarified.