Fecundity, lifespan and egg mass in butterflies: effects of male-derived nutrients and female size

1. Effects of larval reserves and nutrients received as adults on fecundity and lifespan in female Danaus plexippus (the Monarch Butterfly) were measured to determine the relative importance of different sources of nutrients for reproduction and somatic maintenance.
2. Egg-laying lifespan was correlated with female size but not with the amount of male-derived nutrients or adult food concentration.
3. Lifetime fecundity was higher when females received a large first spermatophore, but was not affected by female size when lifespan was controlled or by adult food concentration.
4. At the end of their lives, females contained unlaid eggs and retained, on average, 88% of their initial mass. This proportion was unchanged in two years, although mean egg-laying lifespan varied from 22·5 to 28·7 days.
5. Egg mass decreased over the female lifespan, and was correlated with female size.
6. These results suggest that larval reserves are more important for somatic maintenance than adult income, but that the protein-rich nutrients received from males contribute to egg production. This supports theoretical predictions and empirical studies of other Lepidoptera showing that larval reserves are less likely to affect fecundity when the adult income can contribute substantially to egg production.     

Using food for different purposes: female responses to prey in the predator Coccinella septempunctata L. (Coleoptera: Coccinellidae)

Abstract.  1. Although predatory insects often feed on diverse prey, their reproductive activity may be linked most strongly to a more restricted range of prey. The propensity of adult females of the ladybird beetle, Coccinella septempunctata L., to attack two natural prey species, pea aphids [ Acyrthosiphon pisum (Harris)] and alfalfa weevil larvae [ Hypera postica (Gyllenhal)], was compared, and the degree to which ladybird egg production depends on consumption of aphids vs. weevils was assessed.
2. Coccinella septempunctata females more readily attacked aphids than weevil larvae. This was true regardless of whether females had fed previously on aphids or on weevil larvae.
3. When females were provided with few to many aphids daily, or few aphids plus an excess number of weevil larvae, their rates of egg production depended primarily on the number of aphids consumed.
4. Addition of weevil larvae to diets of limited numbers of aphids increased egg production, but only modestly. Thus, consumption of weevil larvae may have served mostly for self-maintenance, thereby enabling females to use for egg production more of the nutrients and energy obtained from limited consumption of aphids.
5. The females' linkage of egg production primarily to aphid rather than weevil consumption may be adaptive, as their offspring are much less able as larvae to survive and mature on a diet of weevils rather than aphids.     

Quantitative food utilization and reproductive allocation by adult milkweed bugs, Oncopeltus fasciatus

ABSTRACT. Age-specific and lifetime dry mass budgets were estimated for mated and virgin adult milkweed bugs, Oncopeltus fasciatus (Dallas) Hemiptera: Lygaeidae), fed air-dried milkweed seeds ( Asclepias syriacd ) in the laboratory at LD 14:10 and 23°C. Relative consumption rate (RCR) of all bugs was high during the first 8 days posteclosion (teneral period) as their fresh weight, dry weight, and fat content increased. Thereafter, the physiological syndrome associated with reproduction in mated females was indicated by their higher RCR, earlier and greater rate of egg production, greater lifetime relative metabolic rate and higher net and gross production efficiencies than virgin females and males. Males tended to live longer than virgin and mated females, which had similar lifespans. Mated females weighing less at eclosion remained lighter in weight on the day of mean peak weight, but food consumption, egg production and lifespan were independent of body-weight over a 25% range. Input of nymphal reserves or male reproductive secretions to egg production is probably minor in comparison with the adult female's food budget. The high proportion of the food budget allocated to egg production by mated females of O.fasciatus is consistent with its migratory, colonizing lifestyle.     

Influence of adult nutrition on the relationship between body size and reproductive parameters in a parasitoid wasp

Abstract.  1. An important constraint upon life-history evolution in parasitoids is the limit imposed by body size on allocation of limited metabolic resources to different fitness-related physiological functions such as reproduction and survival.
2. The influence of adult nutrition on reproductive and maintenance variables was studied in the synovigenic ectoparasitoid Mastrus ridibundus , and it was determined whether resource allocation to these different functions depends on body size.
3. Over the course of adult life there was a positive relationship between body size and the number of mature eggs in adult females both in the presence and absence of food. However, only in the presence of food did egg maturation rates increase significantly with body size. Starved wasps produced significantly smaller eggs than fed ones, which has not been documented before. Moreover, starved wasps produced fewer offspring than fed wasps, and attacked fewer hosts.
4. The availability of food had a major effect on longevity, with fed females living about 10 times longer than starved ones. There was also a positive relationship between body size and longevity. In starved wasps, this relationship was the same both in the presence and absence of hosts, but in fed wasps there was a positive relationship between body size and longevity in the absence of hosts only. Allocation to initial eggs relative to lifetime progeny production did not decline with body size.
5. The data reveal that in M. ridibundus the trade-off between maintenance and reproduction varies with life expectancy.     

Egg maturation dynamics of the parasitoid Microplitis rufiventris: starvation speeds maturation in early life

The number of mature eggs carried by a female parasitoid at any given moment (egg load) is a fitness‐related parameter affecting reproductive potential and impacting upon host population dynamics. Microplitis rufiventris Kokujev (Hymenoptera: Braconidae) is a solitary koinobiont endoparasitoid wasp of several noctuid pests, including Spodoptera littoralis. The number of mature eggs carried by females at emergence is approximately 50. The rate of egg maturation is strongly affected both by feeding status and access to host larvae. In early adult life, egg maturation rates are lower for 6–72 h in fed wasps compared with food‐deprived wasps. When given access to hosts, honey‐fed wasps live for approximately 9 days with high lifetime fecundity (226 eggs). By contrast to early adult life, the total realized fecundity is positively affected by feeding status, where water‐fed and starved females have 140 and 107 eggs, respectively. Egg resorption is most pronounced in the later life of females. The results suggest, in addition to confirming the effect of honey‐feeding on total fecundity, that fecundity of starved wasps includes rapid egg maturation early in life, which potentially could improve the performance of the parasitoid as a biological control agent.     

Survivorship characteristics of the mosquito Aedes caspius adults from southern France under laboratory conditions

The survivorship characteristics of two populations of Aedes caspius (Pallas) (Diptera: Culicidae) were compared in the laboratory. One population was sourced from Mourgues, where larvicides have been used continuously for approximately 40 years, and the other from Pont de Gau, where there has been no consistent mosquito control. The aims of the study were to ascertain the basic life history profiles of adults and to determine whether continuous larviciding affects inherent adult survivorship. Life tables were constructed to calculate the following life expectancy parameters: mean lifetime (tau(ad)); maximum lifetime (tau(max)), and daily survival rate (p(ad)). All three parameters were higher for females than for males (paired t-test, P < or = 0.001); male mean lifetime, maximum lifetime and daily survival rate were 4.95 +/- 0.94 days, 20.50 +/- 6.66 days and 0.79 +/- 0.05, respectively; female values were 14.74 +/- 3.68 days, 49.69 +/- 16.55 days and 0.93 +/- 0.02, respectively. No differences were found between the two populations, and no correlations were found between initial adult densities and their respective survival rates. The survivorship curves for Ae. caspius were type IV for males (mortality rates higher for young adults) and type III for females (mortality rates constant).     

Does reproductive investment change with age in tsetse flies, Glossina morsitans morsitans (Diptera: Glossinidae)?

1. Viviparous insects such as tsetse ( Glossina spp.) provide unusual opportunities to compare age-related changes in the proportion of maternal resources transferred to offspring.
2. In laboratory populations of Glossina morsitans morsitans the survival of females was high for the first 60 days of adult life but declined rapidly thereafter.
3. Average longevity did not differ significantly between mated and unmated females (93·6 and 90 days, respectively).
4. Nutritional state in terms of fat content and residual dry mass did not decline with adult female age.
5. The fecundity of mated females was constant for the first 60 days of adult life and declined only slightly thereafter.
6. Offspring size did not change towards the end of the adult female lifespan and there was no evidence of an increase in the allocation of resources to reproduction in older females.
7. Results contrast with those obtained recently for vertebrates and may indicate that age-related changes in offspring size in Glossina are not adaptive, or that so few females reach old age under natural conditions that there is no selection for a strategy of terminal investment.     

Lifetime mating success in the damselfly Coenagrion puella

Lifetime mating success of male azure damselflies (Coenagrion puella) was measured in a natural population. The major determinant of mating success is the number of days a male spends at the breeding site, which is mostly determined by a male's adult lifespan. Long-lived males have a higher mating rate than short-lived males, and daily mating rate increases with age up to 6 days, then falls. Large males live longer, but have a lower daily mating rate than small males. These effects of size are very weak, accounting for no more than 2% of the daily variance in mating success. The only overall effect of size on lifetime mating success is that males at both extremes of the size distribution are more likely to fail to mate. Chance differences in the number of females encountered are sufficient to account for the remaining variance in mating success. The weather is also shown to have a major effect on mating success. We draw attention to the ways in which it may be misleading to draw conclusions about the action of sexual selection from studies of daily, rather than lifetime, reproductive success. We provide evidence to support the view that variance in male reproductive success is neither evidence for sexual selection, nor a measure of its intensity.     

Sex-specific fitness correlates of dispersal in a house sparrow metapopulation

1.  Dispersal affects many important ecological and evolutionary processes. Still, little is known about the fitness of dispersing individuals.
2.  Here, we use data from a long-term study of a house sparrow Passer domesticus metapopulation to compare lifetime reproductive success (LRS) of resident and immigrant individuals, all with known origin.
3.  Lifetime production of recruits by immigrant males was much lower than for resident males, because of shorter life span and lower annual mating success. In contrast, lifetime production of recruits did not differ significantly between immigrant and resident females.
4.  Over their lifetime, dispersers contributed fewer recruits to the local population than residents. This shows that immigrant house sparrows have different, sex specific, demographic effects on the population dynamics than residents.     

The effect of exposure to imidacloprid on Asian longhorned beetle (Coleoptera: Cerambycidae) survival and reproduction

The effect of imidacloprid delivery method and application rate on survival of adult Asian longhorned beetles, Anoplophora glabripennis (Motschulsky), was studied, along with the effect of repeated daily ingestion of imidacloprid on the survival and reproductive capacity of adult females. Beetles exposed repeatedly to 50 ppm imidacloprid died in < 2-3 wk, whether dosed orally each day, or through contact exposure. Beetles given 1 microl of 50 ppm imidacloprid daily for two, three, four, or five consecutive days died sooner with increasing consecutive days: the beetles treated for 5 d all died within 15 d, while 80% of beetles treated for only 2 d lived > 8 wk. For females given 1 microl daily, across a range of doses from 2 to 50 ppm imidacloprid, the total number of viable eggs laid was reduced with increasing dosage, but percentage egg viability was not affected. Survival of females at dosages of 10 or 30 ppm/d was not significantly reduced compared with controls but these females laid 23-38% fewer viable eggs, suggesting a sublethal effect of imidacloprid. Female beetles given 1 microl/d of 40 or 50 ppm imidacloprid died more quickly than controls and viable egg production was reduced 82-93%, because of a combination of lethal and sublethal effects of intoxication.     

Influence of adult and egg predation on reproductive success of Epirrita autumnata (Lepidoptera: Geometridae)

We studied the effects of predation and oviposition activity on reproductive success of a late-season moth, Epirrita autumnata by exposing adult females and eggs to predation in their natural habitat in two successive years. Daily survival rates of adult females ranged from 0.4 to 0.8, average being 0.7. Most predation occurred during nights and was caused by harvestmen and other invertebrate predators. Avian predation did not have an effect on adult survival rates, most likely because of the lateness of E. autumnata flight season. Eggs were also preyed upon by invertebrate predators, although a notable proportion of egg mortality was attributable to causes other than predation. Daily survival rates of eggs were more than 0.99. Using modeling based on empirical data on eclosion of female adults, their oviposition behavior and survival rates of adults and eggs, the daily survival rates were translated into population level consequences. Adult predation was estimated to decrease reproductive success of non-outbreaking E. autumnata by 60–85 percent and egg mortality by 20–40 percent. Predation on adult lepidopterans is a mortality factor potentially as relevant as predation in any other life history stage and thus, should not be ignored in studies of population regulation.     

Influence of host age and host deprivation on longevity, egg load dynamics and oviposition rate in Microplitis rufiventris

Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third （preferred hosts） or fourth （non-preferred hosts） instar larvae of Spodoptera littoralis （Boisd.） were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female＇ s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female＇ s life span, oviposition rate and host size from which the female was derived. These results suggest that： （i） M. rufiventris wasp is a weak synovigenic species; （ii） the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; （iii） the egg load of the newly emerged female is significantly less than the realized fecundity; and （iv） because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.     

Effects of time limitation and egg limitation on lifetime reproductive success of a parasitoid in the field

We used field observations of freely foraging Aphytis aonidiae parasitoids in conjunction with results of laboratory studies of A. aonidiae and other Aphytis species to simulate lifetime patterns of behavior and reproduction. Field observations provided estimates of encounter rates with three classes of hosts, the mortality rate from predation on adult parasitoids, and host-handling times for oviposition and host feeding by adult wasps. A series of physiological parameters, including the egg maturation rate and the value of host-feeding meals, were estimated from previously published studies. Plasticity in parasitoid behavior was incorporated in two ways. For one set of simulations we used a behavioral rule derived empirically from observations of parasitoids made in the field, and for another we used a dynamic state-variable model to generate a set of behavioral rules that maximize lifetime reproductive success. As was expected, the empirically derived rule led to better matches with field observations than did simulations using the output of the dynamic model. Projections of lifetime reproductive success in the field ranged between three and 37 eggs within the 95% confidence intervals of the mortality rate and host encounter rate and depending on which behavioral rule was used. Lifetime reproductive success from the simulation with central estimates of the mortality and host encounter rates that incorporated the empirical rule was 6.25 eggs. Using the empirical versus the theoretical rule in the simulations led to a 10%-30% decline in projections of lifetime reproductive success, depending on mortality and host encounter rates. Regardless of the behavioral rule, the simulations underscored the observation that the host encounter rate was greater than the egg maturation rate. The overall oviposition rate was sufficiently high to lead to daily episodes of temporary egg limitation during which parasitoids must mature an egg before being able to oviposit.     

Gallery construction and oviposition of the bark beetle Ips typographus (Coleoptera: Scolytidae) at different breeding densities

Abstract. 1. The bole of a Norway spruce, Picea abies (L.) Karst., was cut in 25 cm sections and infested in the laboratory with 2 or 8 females/dm² of Ips typographus (L.) and an equal number of males.
2. One log of each beetle density was dissected every second day, beginning 2 days after female introduction.
3. The oviposition rate was highest at the beginning of the breeding period for both densities and then declined, especially at the high density. Residence time is a useful approximation for the number of laid eggs in the beginning of the oviposition period or when the density is low.
4. Final gallery length and egg number per gallery were on average significantly smaller at density 8 than at density 2.
5. Linear relationships between total gallery length and number of egg niches were found to be similar for the two densities.
6. An estimate of the egg to adult mortality at different densities was found by using data from a field study that related gallery length to breeding density in attacked living trees, together with published relationships between density and egg niches or emergence of new adults.
7. It was concluded that both decreased oviposition rate and earlier re-emergence at higher densities contribute to a declining oviposition with increasing density. The final offspring production is futher reduced by lower survival due to larval competition as density increases.     

Effects of the corpora allata on sexual receptivity and completion of oocyte development in females of the cricket, Gryllus bimaculatus

Abstract Allatectomy of young penultimate nymphs of Gryllus bimaculatus De Geer (Gryllidae) resulted in prothetelic creatures which exhibited reproductive competence. The same operation performed on young last instar nymphs resulted in moulting to morphologically normal adults. Allatectomized morphologically normal adult females, as well as prothetelic ones, showed the same level of sexual receptivity as untreated control females. Allatectomized morphologically normal and prothetelic females laid viable eggs, but rate of egg laying and number of eggs produced by these females were much reduced in comparison with the controls. Administration of methoprene (a Juvenile Hormone analogue) to allatectomized females restored egg production to a more or less normal rate. Removal of the spermatophore within 10 min of copulation had no effect on subsequent sexual receptivity of the females, nor on the reduced rate of egg laying by the allatectomized females, but did affect the rate of egg laying by control females.
It is suggested that the corpora allata (CA) and the Juvenile Hormone (JH) play no major role in controlling basic sexual receptivity of G.bimaculatus females, and do not have an all-encompassing control on egg production, though they do exert a marked quantitative effect on the rate of egg production.     

Time limitation, egg limitation, the cost of oviposition, and lifetime reproduction by an insect in nature

For more than 80 years, ecologists have debated whether reproduction by female insect herbivores and parasitoids is constrained by the time needed to find hosts (time limitation) or by the finite supply of mature eggs (egg limitation). Here we present the first direct measures of permanent time limitation and egg limitation and their influences on the cost of oviposition and lifetime reproduction for an insect in nature. We studied the gall midge Rhopalomyia californica, which neither matures nor resorbs eggs during the adult stage. By sampling females soon after their death and correcting for predation effects, we demonstrate that females lay a large proportion of their total complement of eggs (multiyear mean: 82.9%). The egg supplies of 17.1% of females were completely exhausted, with the remaining 82.9% of females being time limited. As predicted by theory, we estimate that even though egg limitation is a minority condition within the population, egg costs make a substantial contribution (57% of the total) to the cost of oviposition. We conclude that insect life histories evolve to produce a balanced risk of time and egg limitation and, therefore, that both of these constraining factors have important influences on insect oviposition behavior and population dynamics.     

Control of reproduction and a survival cost to mating in female Bicyclus anynana butterflies

Abstract.  1. Given that sexual conflict is all pervasive, investigating potential costs to mating and the control of female reproduction are important issues.
2. Here, female reproductive output and longevity are investigated in relation to mating status (virgin vs. once-mated females) and host-plant availability in the tropical butterfly Bicyclus anynana .
3. Both factors significantly affected realised fecundity in a manner demonstrating that female reproduction is not entirely under control of the female herself, but is rather subject to additive contributions of the female, her male partner, and intrinsic physiological processes. As evident from the deposition of significant egg numbers by virgin, host-deprived females, B. anynana is effectively unable to completely inhibit oogenesis.
4. Mated females suffered a reduction in adult life span, which cannot be explained as a side-effect of variation in egg size, lifetime, or early fecundity.
5. Such detrimental effects of mating per se are indicative of the cooperation–conflict balance between sexes being shifted towards conflict in B. anynana .     

Effect of temperature on egg production in the common housefly

Musca domestica L. (Diptera: Muscidae), better known as the common housefly, is increasingly considered to be a new, alternative protein source for animal nutrition. By transferring low-value organic side streams into high-value protein products, its commercial production contributes to a circular economy. Next to technical innovations for scaling-up the production capacity, efficient egg production has been identified as one of the bottlenecks of housefly production systems. We investigated egg production in two strains, one originally from Spain (SPA) and one from The Netherlands (GK), at 25 and 32 °C. At 25 °C, duration of preoviposition period, laying phase, and adult longevity was longer than at 32 °C. Lifetime egg production was lower at 32 °C, but the number of clutches laid per female was unaffected by temperature. Daily egg production at 32 °C was higher during the first 7 days, revealing a trade-off between higher early-in-life reproductive effort and adult longevity. The combination of shorter sexual maturation period and higher daily egg-laying rate resulted in reaching 50% of total egg production only 6 days after emergence at 32 °C for both strains, compared to 13 and 14 days at 25 °C for SPA and GK, respectively. We conclude that, in the absence of a need for high adult survival rates, houseflies have favourable production performances at higher temperature, and that efficacy and yield of the production process could be maximized by increasing the rearing temperature to 32 °C.     

Small body size in an insect shifts development, prior to adult eclosion, towards early reproduction

Life-history theory has suggested that individual body size can strongly affect the allocation of resources to reproduction and away from other traits such as survival. In many insects, adults eclose with a proportion of their potential lifetime egg production that is already mature (the ovigeny index). We establish for the solitary parasitoid wasp Aphaereta genevensis that the ovigeny index decreases with adult body size, despite both initial egg load and potential lifetime fecundity increasing with body size. This outcome is predicted by adaptive models and is the first unequivocal intraspecific demonstration. Evidence suggests that a high ovigeny index carries a cost of reduced longevity in insects. Our results therefore contribute to the emerging evidence that small body size can favour a developmental shift in juveniles that favours early reproduction, but which has adverse late-life consequences. These findings are likely to have important implications for developmental biologists and population biologists.     

Pre- and post-maturation survival in adults of the damselfly Pyrrhosoma nymphula (Zygoptera: Coenagrionidae)

The technique of mark-release-recapture was used to study survival before and after sexual maturity in adults of the damselfly Pyrrhosoma nymphula (Sulzer). Fewer females were recaptured upon return to water to breed despite no differences in dispersal or daily survival rate between the sexes over the immature period. Because females took longer to mature than males, their poorer recapture rate was attributed to greater overall mortality during their longer maturation phase. Survivorship curves for tenerals marked at emergence suggested that overall survival of immature adults was similar to, if not better than, that of mature adults. The reasons for this are discussed.
Jolly's model was used to estimate daily survival rates for mature adults. The assumptions of the model were tested rigorously. Estimates for females were statistically less reliable than those for males. Mean reproductive spans for males and females were 6.8 and 6.6 days, respectively, giving mean total adult lifespans of 19.4 days and 21.6 days for individuals surviving the maturation period.
Because neither sex visited the breeding site every day, sampling exclusively at water resulted in underestimation of mean reproductive spans for both sexes. Female reproductive spans were underestimated to a greater extent; because females remain away from water longer between visits, there is a greater chance that they will die before being recaptured.
Mean reproductive spans were also underestimated when only a sub-section of the habitat was sampled. Females were significantly more mobile than males and this increased the likelihood that they would move out of the study area, resulting in more severe underestimation. The importance of obtaining accurate estimates of mature lifespan for females is discussed.