The function of female accessory reproductive gland secretion and a cost to polyandry in the yellow dung fly

Abstract Female yellow dung flies have large paired accessory reproductive glands, the function of which remains unclear. However, gland contents are secreted during copula and egg laying. Other female flies produce a range of anti-bacterial substances in their accessory reproductive glands that protect them and their eggs from pathogens, and it is possible that gland secretion acts similarly in yellow dung flies. A series of experiments was conducted to test this idea. Because the volume of secretion remaining in the glands is negatively related to copula duration, egg hatchability and longevity of females was compared in groups that copulated experimentally once or three times. A zone inhibition assay was used to see if gland extract inhibited bacterial growth. Egg number was positively associated with female body size, but the proportion of eggs hatching was not. Neither copula number nor duration influenced egg number or hatch number or proportion. In accordance with this, gland extract did not inhibit bacterial growth. However, female longevity was reduced in females that copulated with more males. This suggests that gland secretion does not serve to protect eggs, but as with a number of taxa, copula is costly to female yellow dung flies.     

Variations in fecundity over catchment scales: Implications for caddisfly populations spanning a thermal gradient

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梨小食心虫蛹重对成虫繁殖力和寿命及下一代幼虫发育的影响

【目的】本文旨在明确营养状况不同造成的梨小食心虫Grapholitha molesta(Busck)雌、雄蛹重量差异对其羽化的成虫产卵量、产卵期、寿命及下一代(F1)幼虫发育的影响。【方法】室内条件下,通过不同的饲养方法,获得个体重量不同的梨小食心虫雌、雄蛹,待其羽化交配后,记录其产卵量、产卵时间和成虫寿命;卵孵化前后,分别测量卵和初孵幼虫大小,计算卵孵化率,统计幼虫发育历期。【结果】雌蛹重量对梨小食心虫的成虫产卵量影响显著,其重量与产卵量呈正相关(y=15.505x-59.292);同一条件下,雌蛹与雄蛹重量也呈正相关(y=0.823x-0.538)。同时,雌蛹重量对成虫产卵期影响也较大,蛹重大的个体羽化的雌虫比蛹重小的个体羽化的雌虫产卵高峰期提前1 d;较重、中等和较轻蛹羽化出的雌虫个体每天产卵量高于10粒/雌的时间分别为9~10,7和5~6 d;产卵量高于5粒/雌的时间分别为12~13,9和6~7 d。而雄蛹重量对产卵量、雄成虫寿命影响没有明显影响。较轻的蛹羽化的雌成虫寿命比较重蛹羽化的雌成虫短2~3 d;而雄蛹重量对其羽化的雄成虫寿命影响没有明显规律。雌、雄蛹重量对其羽化成虫的卵孵化率、卵和初孵幼虫的大小影响均不显著,对F_1幼虫发育历期影响也不显著。【结论】梨小食心虫雌蛹重对羽化成虫的产卵量和产卵期等影响显著,田间防治时应注意在不同条件下完成发育的个体,尤其是雌虫,由于营养差异引起的个体大小对随后种群增长的影响。     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

Female reproductive pattern of Callinectes sapidus Rathbun, 1896 (Brachyura: Portunidae) in Iskenderun Bay,Eastern Mediterranean

We studied the female reproductive pattern of Callinectes sapidus, which was introduced to the Mediterranean in the 20^th century. We assessed female size at first maturity, fecundity, and fecundity relationship to size in Iskenderun Bay, eastern Mediterranean, Turkey. Samples were collected between July 2014 and June 2015 using bottom trawling at depths ranging from 1 to 50 m. A total of 322 crabs were caught of which 308 (95.7%) were females including 116 ovigerous ones. The minimum carapace width of the mature females was 39.1 mm and the mean carapace width 123.8 mm. The carapace width of ovigerous females varied between 95.1 and 144.5 mm, with a mean of 120.3 mm. The highest number of ovigerous females was observed in July and August. Mean fecundity was 1.91 million (667,950–4,669,853) eggs per female. A weak positive linear relationship between fecundity and carapace width was noted, as well as a high correlation with total egg weight. In the eastern Mediterranean, maturity sizes of females were smaller than those in the native region of the species.     

Effects of forced egg-retention in Aedes albopictus on adult survival and reproduction following application of DEET as an oviposition deterrent.

The insect repellent DEET (0.1% concentration), used as a mosquito oviposition deterrent in the laboratory, influenced the retention and maintenance of mature eggs by caged gravid female Aedes albopictus Skuse. This egg-retention mechanism could benefit survival because the gravid females were ultimately able to lay maintained eggs upon availability of water, but the length of forced egg-retention time reduced the number of eggs laid per female. Gravid females with retained eggs also laid a higher percentage of eggs that failed to tan, and this percentage increased with time duration of egg-retention. Percent egg hatch was not significantly affected by DEET when used as an oviposition deterrent; however, percent hatch was affected by time duration of egg-retention in both treated (exposed to DEET) and untreated (control) gravid females. The rate of egg hatch was considerably reduced after three weeks of retention; this reduction declined to zero for treated and control females at six and four weeks post-treatment, respectively. The fecundity and fertility of gravid female Ae. albopictus were affected by the time duration of forced egg-retention.     

Mate guarding and male mate choice in the chameleon grasshopper <Emphasis Type="Italic">Kosciuscola tristis</Emphasis> (Orthoptera: Acrididae)

In the wild, male chameleon grasshoppers (Kosciuscola tristis) are frequently observed mounted on the back of females even when not in copula, and will fight off other usurping males. If this behaviour is mate guarding and reflects investment in male mate choice, then we expect males to preferably guard females based on reliable cues of quality. Cues for female quality likely include female size and egg development that together may indicate fecundity. We investigated male mate choice in the field expressed as mate-guarding preference, by comparing size and egg development in guarded and unguarded females. We found no difference between guarded and unguarded females in measures of fecundity or body size. The majority of females sampled did not contain any viable eggs. This finding suggests that male K. tristis indiscriminately guard females in a scramble mating system.     

Factors affecting the duration of nest defense and reproductive lifespan of female sockeye salmon, Oncorhynchus nerka

Reproductive success of female animals is often affected by a combination of fecundity and parental care. In female salmonid fishes, acquisition of nest (redd) sites and prevention of their use by other females are critical to reproductive success. These factors are particularly important for stocks that spawn at high densities. Body size is positively correlated with fecundity and egg size, and has been hypothesized to control the outcome of intrasexual competition and longevity. We tested this hypothesis by evaluating the influences of body size, intrasexual aggression and arrival date on duration and success of redd guarding by female sockeye salmon, Oncorhynchus nerka, in a small Alaskan creek. Contrary to the hypothesis, larger females guarded their redds no longer than smaller females, and did not live as long in the stream. Aggression was not related to body size or overall longevity but was positively correlated with residence period on the redd. Females that entered the creek earlier lived longer, spent longer on their redds, and spent more time guarding their redds after spawning than females that entered the creek later. However, despite their longevity, early-arriving fish were more likely to have their redds reused by another female because they died before all the females had selected redd sites. The small average body size in this stock is consistent with weak selection for large size, and with our evidence that size provided little if any advantage in nest guarding.     

Optimal copula duration in yellow dung flies: effects of female size and egg content

We examine data on copula duration in dung flies, Scatophaga stercoraria, in relation to female phenotype. We use a marginal value theorem approach based on the plausible mechanisms of sperm competition to predict the effect of female variation on optimal copula duration, t *, from the male perspective. Future fertilizations are expected to have a trivial effect on t * with fully gravid females, but an increasing relative effect on t * towards completion of oviposition. t * is expected to be affected by female size because of variation in (1) a female's egg content, which increases the maximum egg gain available from a mating, and (2) the female reproductive tract, which affects the rate at which sperm are displaced. In fully gravid females, t * was not dependent on egg number variation, but showed a positive relation with egg content in females that had laid a varying proportion of their mature egg load at the time of mating, and were therefore not fully gravid. Our models predict that if a male can estimate egg content only by the distension of a female's abdomen, t * should increase in a similar way to that seen with 'take-over' females. We predict t * for fully gravid females by assuming that males can monitor female size. The data showed that sperm displacement rate decreased, and average egg content increased, with female size. Under two models for a sperm displacement mechanism, one (which assumes indirect displacement at a rate proportional to the increase in spermathecal volume) predicts the observed relation between t * and female size almost exactly. Small males copulated for longer than large males (as predicted and reported previously). Copyright 1999 The Association for the Study of Animal Behaviour.     

Effects of delayed mating on the fecundity,fertility and longevity of females of diamondback moth,Plutella xylostella

Abstract The effects of delayed mating on the copulation duration, female fertility, fecundity, egg fertility, longevity and the number days alive after mating of females of diamondback moth (DBM), Plutella xylostella, were studied. When male mating was delayed, the female fertility, fecundity, egg fertility, longevity and number days alive after mating of DBM decreased, and there was a negative correlation between the age of the moth with those variables except copulation duration. When female mating was delayed, the female fertility, fecundity, percent egg fertility and number days alive after mating of DBM also decreased, but the longevity increased, which also showed a negative relationship between the age of the moth with the variables except copulation duration and longevity. When both males and females delayed mating, the female fertility and fecundity decreased; egg fertility was affected marginally, and the longevity of females increased. The moth age was negatively correlated with those variables.     

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.     

Control of copula duration and sperm storage by female Queensland fruit flies

Copula duration and sperm storage patterns can directly or indirectly affect fitness of male and female insects. Although both sexes have an interest in the outcome, research has tended to focus on males. To investigate female influences, we compared copula duration and sperm storage of Queensland fruit fly females that were intact, or had been incapacitated through decapitation or abdomen isolation. We found that copulations were far longer when females had been incapacitated, indicating that constraints imposed on copula duration by intact females had been relaxed. Repeatability of copula duration for males was very low regardless of female treatment, and this is also consistent with strong female influence. Number of sperm in the spermathecae was not influenced by female treatment, suggesting that female abdominal ganglia control the transport of sperm to these long-term storage organs. However, more sperm were found in the ventral receptacles of incapacitated females compared to intact females. Overall, results implicate cephalic ganglia in regulation of copula duration and short-term sperm storage in the ventral receptacle and abdominal ganglia in regulation of long-term sperm storage in the spermathecae.     

Optimal numbers of matings: the conditional balance between benefits and costs of mating for females of a nuptial gift‐giving spider

In species where females gain a nutritious nuptial gift during mating, the balance between benefits and costs of mating may depend on access to food. This means that there is not one optimal number of matings for the female but a range of optimal mating numbers. With increasing food availability, the optimal number of matings for a female should vary from the number necessary only for fertilization of her eggs to the number needed also for producing these eggs. In three experimental series, the average number of matings for females of the nuptial gift‐giving spider Pisaura mirabilis before egg sac construction varied from 2 to 16 with food‐limited females generally accepting more matings than well‐fed females. Minimal level of optimal mating number for females at satiation feeding conditions was predicted to be 2–3; in an experimental test, the median number was 2 (range 0–4). Multiple mating gave benefits in terms of increased fecundity and increased egg hatching success up to the third mating, and it had costs in terms of reduced fecundity, reduced egg hatching success after the third mating, and lower offspring size. The level of polyandry seems to vary with the female optimum, regulated by a satiation‐dependent resistance to mating, potentially leaving satiated females in lifelong virginity.     

Genetic variation in male effects on female reproduction and the genetic covariance between the sexes

Abstract.— Males of many insect species increase the fecundity and/or egg size of their mates through the amount or composition of their nuptial gifts or ejaculate. The genetic bases of such male effects on fecundity or egg size are generally unknown, and thus their ability to evolve remains speculative. Likewise, the genetic relationship between male and female investment into reproduction in dioecious species, which is expected to be positive if effects on fecundity are controlled by at least some of the same genes in males and females, is also unknown. Males of the seed beetle Stator limbatus contribute large ejaculates to females during mating, and the amount of donated ejaculate is positively correlated with male body mass. Females mated to large males lay more eggs in their lifetime than females mated to small males. We describe an experiment in which we quantify genetic variation in the number of eggs sired by males (mated to a single female) and found that a significant proportion of the phenotypic variance in the number of eggs sired by males was explained by their genotype. Additionally, the number of eggs sired by a male was highly positively genetically correlated with his body mass. The between-sex genetic correlation, that is, the genetic correlation between the number of eggs sired by males and the number of eggs laid by females, was highly positive when eggs were laid on Acacia greggii seeds. This indicates that males that sire many eggs have sisters that lay many eggs. Thus, some of the genes that control male ejaculate size (or some other fecundity-enhancing factor) when expressed in males appear to control fecundity when expressed in females. We found no significant interaction between male and female genotype on fecundity.     

The function of multiple mating in oviposition and egg maturation in the seed beetle Callosobruchus maculatus

Abstract.  To comprehend the significance of multiple mating in female Callosobruchus maculatus , the material-benefits hypothesis is tested, as well as the potential function and mechanism of additional mating for females. The results show that longevity and lifetime fecundity are significantly higher in doubly-mated females, whereas females subject to an interrupted second copulation (which transfers no ejaculate) show resemblance with singly-mated females in lifetime fecundity and daily fecundity, supporting the material-benefits hypothesis. Female lifetime fecundity increases with ejaculate size in doubly-, but not singly-mated females. Doubly-mated females exhibit an immediate increase in fecundity on the day of remating. Moreover, dissection of ovaries after remating shows that the number of unlaid eggs is lower in doubly-mated females in comparison with singly-mated females, whereas the total number of unlaid and laid eggs shows no significant difference. This suggests that the second mating increases the oviposition rate, but not the egg maturation rate in female C. maculatus . Thus, ejaculate serves as a stimulus for oviposition, at least on the remating day. However, because doubly-mated females lay more eggs in total than singly-mated females, doubly-mated females appear to mature more eggs than the singly-mated counterparts. This is attributed to the 'indirect-driven' hypothesis, which states that the reduction in the number of mature eggs in the oviducts brought about by oviposition stimulates the maturation of oocytes in ovarioles to replenish the number of mature eggs.     

Biology and field performance of gryon clavigrallae (Hymenoptera: Scelionidae), an egg parasitoid of Clavigralla spp. (Hemiptera: Coreidae) in India

The biology and impact of Gryon clavigrallae Mineo, an egg parasitoid of Clavigralla scutellaris Spinola and C. gibbosa (Westwood), was investigated. The calculated developmental threshold temperatures for females and males were 15.6 degrees C and 15.8 degrees C, respectively. Emergence exceeded 94% at temperatures between 22 and 30 degrees C. Adult females lived on average 28-96 days when fed with honey. Without food, adults lived < 6 days. Mean fecundity was 56.4 eggs per female. A significant trend of lower fecundity after longer periods of host deprivation was observed. Gryon clavigrallae females successfully oviposited in host eggs of all ages though eggs < 4 days old were preferred. Total host handling times were significantly longer on C. gibbosa eggs (23.5 min) than on C. scutellaris eggs (12.0 min). Females readily distinguished parasitized from non-parasitized host eggs. Superparasitism was observed when few or no unparasitized eggs were available. Eggs of the two Clavigralla species can be separated by surface structure and condition after eclosion. Clavigralla scutellaris laid significantly larger egg clusters than C. gibbosa (19.9 versus 10.5 eggs per cluster). Gryon clavigrallae was present as soon as the first Clavigralla spp. egg clusters were found on pigeonpea. The percentage of egg clusters parasitized increased early in the season with egg cluster density and remained high (up to 83%) despite fluctuations in host density. Overall, G. clavigrallae parasitized 40 and 58% of C. gibbosa and C. scutellaris eggs. The percentage of egg clusters parasitized and the number of eggs parasitized per cluster increased significantly with egg cluster size. The overall parasitoid sex ratio was highly female biased but varied with the number of eggs parasitized per cluster.     

Maternal and interannual comparison of the ovulatory periodicity, egg production and egg quality of the batch-spawning yellowtail flounder

Estimates of the ovulatory periodicity of yellowtail flounder indicate that a 1-day interval, which predominated over all other intervals, may characterize regular ovulation patterns. Females produced a mean number of 14–22 batches in 1994 and 1995, respectively. Batch fecundities usually remained within a range of 10 000–60 000 eggs. Mean egg production increased from 549 756 eggs per female in 1994 to 1 186 881 eggs in 1995. Mean fertilization rates rose interannually from 38 to 57%, while hatching rates, tested in 1994, had a mean of 63%. Maternal variation in egg production and egg quality was large and independent of size differences among females. Some females had disrupted ovulation patterns which affected the realization of potential fecundity contained within the prespawning ovary. High interbatch variation in egg quality was not related to progressive decreases in egg diameter and dry weight over time. Batches with high survival rates appeared at random within a female's duration of ovulation.     

Post‐mating enhancement of fecundity in female Lygus hesperus

Although mated females of the western tarnished plant bug Lygus hesperus Knight are known to produce more eggs than virgins, the nature of the inducing stimuli and the specific changes occurring in the female require additional elucidation. Compared with virgin females isolated from males, those exposed to male precopulatory behaviours produce similar numbers of eggs, whereas inseminated females produce 50% more during the observation period. Although the quantity of seminal fluids received by a female does not influence egg number, mating twice within a 10‐day span causes a 16% increase in fecundity, on average. Females mating more than twice during the same period do not exhibit additional increases in egg number. Because virgin females contain more chorionated eggs than are laid, mating appears to enhance the rate of oviposition. However, to achieve a sustained increase in fecundity, an augmented rate of oocyte maturation would also be required. Male‐derived spermatophores lack substantive quantities of nutrients that might otherwise have enhanced female fecundity. The total amounts of carbohydrate, protein and lipid, as well as eight essential minerals transferred by the male, are insufficient for producing even a single egg, and the female has already produced a large number of chorionated oocytes before she mates. Collectively, the data suggest that seminal fluid contains one or more activational molecules, such as a peptide, which triggers an increase in egg deposition. A prolonged increase in oviposition rate may be achieved through multiple matings to ensure a supply of sperm or to offset the degradation of the putative activational factor.     

Female fecundity traits in wild populations of African annual fish: the role of the aridity gradient

The evolution of life history is shaped by life expectancy. Life‐history traits coevolve, and optimal states for particular traits are constrained by trade‐offs with other life‐history traits. Life histories contrast among species, but may also diverge intraspecifically, at the level of populations. We studied the evolution of female reproductive allocation strategy, using natural populations of two sympatric species of African annual fishes, Nothobranchius furzeri and Nothobranchius orthonotus. These species inhabit pools in the Mozambican savanna that are formed in the rainy season and persist for only 2–10 months. Using 207 female N. furzeri from 11 populations and 243 female N. orthonotus from 14 populations, we tested the effects of genetic background (intraspecific lineage) and life expectancy (position on the aridity gradient determining maximum duration of their temporary habitat) on female fecundity traits. First, we found that variation in female body mass was small within populations, but varied considerably among populations. Second, we found that fecundity was largely defined by female body mass and that females spawned most of their eggs in the morning. Third, we found that the trade‐off between egg size and egg number varied among lineages of N. furzeri and this outcome has been confirmed by data from two separate years. Overall, we demonstrate that local conditions were important determinants for Nothobranchius growth and fecundity and that eggs size in arid region was less limited by female fecundity than in humid region.     

The role of male age, sperm age and mating history on fecundity and fertilization success in the hide beetle

Models of age-related mate choice predict female preference for older males as they have proven survival ability. However, these models rarely address differences in sperm age and male mating history when evaluating the potential benefits to females from older partners. We used a novel experimental design to assess simultaneously the relative importance of these three parameters in the hide beetle, Dermestes maculatus. In a two-part experiment we first explored age-related male mating success and subsequently examined the consequences of male age, sperm age and male mating history on female fecundity and fertilization success. In a competitive mating environment, intermediate-age males gained significantly higher mating success than younger or older males. To test the consequences for females of aged-related male mating success, a second set of females were mated to males varying in age (young, intermediate-age and old), in numbers of matings and in timing of the most recent mating. We found that male age had a significant impact on female fecundity and fertilization success. Females mated to intermediate-age males laid more eggs and attained consistently higher levels of fertilization success than females with young and old mates. A male's previous mating history determined his current reproductive effort; virgin males spent longer in copula than males with prior mating opportunities. However, differences in copulation duration did not translate into increased fecundity or fertilization success. There was also little evidence to suggest that fertilization success was dependent on the age of a male's sperm. The experiment highlights the potential direct benefits accrued by females through mating with particular aged males. Such benefits are largely ignored by traditional viability models of age-related male mating success.