Polyandry restores female fertility and paternal effects diminished by inbreeding in Hippodamia convergens

1. The fertility restoration hypothesis posits that polyandry can evolve when female fertility is reduced by matings with related males, but restored by matings with unrelated ones. 2. Using a promiscuous ladybird, this hypothesis was tested by mating mature, virgin females twice with sib and non‐sib males in all four permutations and observing female fertility. The development of progeny from the first and 10th clutches was also followed to test for differences in paternal effects. 3. Mating treatment did not affect fecundity, but egg viability was reduced by sib matings, and restored by non‐sib matings, regardless of mating sequence. In addition, negative paternal effects of sib matings on progeny (lower survival) were compensated by non‐sib matings. 4. The survival of offspring in first clutches was sensitive to the paternal mating sequence and was higher if a sibling male was preceded by a non‐sib male, compared with the reverse, consistent with a time lag in response to male epigenetic signals. Offspring survival did not differ between these two treatments in the 10th clutch, suggesting a blending of paternal effects over time. 5. The results are indicative of interactions between the paternal effects contributed by different males, and between paternal and maternal effects, the latter causing faster development in later clutches. 6. Thus, the reproductive benefits of polyandry for H. convergens females are potentially both genetic and epigenetic, as both egg fertility and beneficial paternal effects were diminished by sib matings, but restored by matings with unrelated males.     

Melanism in a Chinese Population of Harmonia axyridis (Coleoptera: Coccinellidae): A Criterion for Male Investment with Pleiotropic Effects on Behavior and Fertility

In Beijing, China, females of Harmonia axyridis are promiscuous but prefer typical (succinea form) males to melanic ones in the spring generation, ostensibly due to the thermal disadvantages of melanism during summer. We used laboratory observations to test whether males invested differentially in females according to their elytral color, and whether male behavior was phenotype-dependent. Video-recording was used to monitor no-choice mating tests between virgin adults in all phenotype combinations and females were isolated post-copula to observe their egg retention times and reproduction over 5 days. Females tended to wait longer before using the sperm of melanic males, and melanic females delayed longer than succinic females. Melanic males spent longer in copula with succinic than melanic females and the latter received fewer bouts of male abdominal shaking that correlate with sperm transfer, regardless of the phenotype of their mate. Although melanic males abandoned melanic females faster than did succinic males, they remained in copula with females of both phenotypes for a longer period after shaking, suggesting a larger investment in mate guarding by the less-preferred male phenotype. Although female fecundity did not vary among phenotype combinations, egg fertility was lower for females mated to melanic males, suggesting a pleiotropic effect of melanism on male fertility in addition to its effects on male mating behavior.     

Single and multiple mating reduces longevity of female dumpling squid (Euprymna tasmanica)

For many species, mating is a necessary yet costly activity. The costs involved can have an important influence on the evolution of life histories and senescence. Females of many species mate multiply, and this behaviour can inflict a longevity cost. Most studies investigating the effects of multiple mating on female survival have been conducted on insects, and the effects in other taxa are largely unknown. We investigate the effects of both a single mating and a second mating on longevity in female dumpling squid (Euprymna tasmanica), a species in which both sexes mate multiply. Through comparing the longevity of virgin, once‐mated and twice‐mated females, we found that a single mating reduced female life span by 15 days on average. A second mating resulted in an additional 8 day (on average) longevity cost, despite no difference in total clutch mass, number of clutches, single egg mass or number of eggs per clutch between once‐mated and twice‐mated females. This demonstrates a cost to multiple mating which may be independent of the cost of egg production. Furthermore, total clutch mass and female life span were positively correlated, whereas female life span decreased with increasing average water temperature. The presence of an additive effect of reproduction on longevity suggests that multiple mating in cephalopods may have benefits that outweigh these costs, or that there is a conflict in optimal mating frequency between males and females.     

Seasonal cycles of assortative mating and reproductive behaviour in polymorphic populations of Harmonia axyridis in China

Abstract 1. We observed native populations of Harmonia axyridis (Pallas) around Beijing, China, over 2 years and performed choice and no‐choice mating tests between melanic and succinic (non‐melanic) beetles in the laboratory. 2. Succinic phenotypes outnumbered melanics by 5:1 in autumn, but melanics became equally abundant in spring, supporting previous inferences that melanism is advantageous in winter, but costly in summer. 3. Female H. axyridis expressed mate preference overtly, by rejecting less‐preferred phenotypes, and cryptically, by retaining their eggs for longer periods after matings with less‐preferred males, ostensibly to replace their sperm. 4. Succinic pairs formed more quickly in the spring generation, and melanic pairs in the autumn, and the time to copula was affected by both male and female phenotype. The strength of mate preference was contingent on female phenotype, suggesting melanic alleles had pleiotropic effects. 5. Whereas pair formation was under female control, the duration of copula was under male control and lasted longer in the autumn generation than in the spring. Copulations in the choice test tended to be shorter between similar phenotypes, suggesting that males invested more in dissimilar females when alternative mates were available. 6. Although spring and autumn generations were raised under identical conditions, significant contrasts were observed in their reproductive behaviour. 7. Two alternative hypotheses are advanced to explain why gender‐specific reproductive behaviours might vary between generations: maternally‐mediated epigenetic factors that influence the expression of genes in progeny as a function of maternal environment, and linkage disequilibria among alleles that cycle in frequency seasonally as a function of assortative mating.     

Limb ablation and regeneration in Harmonia axyridis: costs for regenerators,but benefits for their progeny

Previous work revealed that Harmonia axyridisPallas (Coleoptera: Coccinellidae) in Beijing, China, were capable of regenerating a forelimb amputated in the fourth instar; 75% of surviving individuals fully regenerated the limb during pupation. In this study, we tested a population of H. axyridis invasive in North America and found that virtually 100% of beetles surviving the operation successfully regenerated the limb. Ablated/regenerated beetles spent longer in pupation, and emerging females were smaller than controls. However, reproductive success was unaffected in all pairwise crosses of control/regenerated adults; there were no differences in pre‐oviposition period, the time required to produce 10 clutches, 10‐day fecundity, or the fertility of eggs, whereas ablated/regenerated parents paid a developmental cost, their progeny obtained benefits. Offspring of crosses that included a regenerated parent tended to have faster larval development than the control cross, although not all were significantly different from controls. However, when either or both parents were ablated and regenerated, their daughters were heavier than controls at emergence. Limb regeneration during pupation appears to activate a physiological cascade which increases the magnitude of beneficial parental effects normally conferred to progeny, possibly via pleiotropic effects. The invasive North American H. axyridis population appears to have higher regeneration capacity than the Chinese population tested previously, although how regeneration capacity might be associated with invasiveness remains unclear. Limb regeneration ability may be a side effect of selection on other traits that confer high fitness under either natural or sexual selection, as it seems unlikely to confer fitness benefits directly in this species.     

Paternity following multiple mating in ladybird <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

We investigated factors which may determine realised paternity following multiple mating in a promiscuous ladybird Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), namely the mating order and ability of males of different morphs to fertilize females. In experiments using males of two colour morphs, we found that the first male had a reproductive advantage over the second male (sperm precedence P2 = 0.38). Males of the melanic morph spectabilis had a reproductive advantage over males of the semi-melanic morph axyridis (P2 = 0.35). Analysis of paternity using microsatellites applied to young larvae, which avoids possible differential mortality of progeny, confirmed the precedence of the first male and higher paternity rate of colour morphs that are more frequent in the wild. Hatchability of eggs increased from 65% after the first mating to 85% after the second mating delayed by five days. Average duration of mating was 157 min and did not differ between the first and second mating and between the colour morphs of males.     

Demonstration of Multiple Mating in Heterodera glycines with Biochemical Markers

Controlled crosses of Heterodera glycines were carried out by placing one o r more virgin females of known esterase phenotype on an agar plate and adding, at various time intervals, one or more males of different esterase phenotypes. Progeny (second-stage juveniles) of such crosses were propagated on soybeans, and 30 days later young females were subjected to electrophoretic analysis to determine their esterase phenotype. Esterase phenotypes that represented the heterozygous state of the maternal and paternal genomes confirmed the hybrid nature of the progeny and identified their male parent. When each of 74 females was given the opportunity to mate successively with two males of different esterase phenotypes, 43 mated with a single male and 31 mated with both males. One female mated with three males, i.e., with a male of its own population (sib mating) and the two males provided for the cross. Inseminated females could mate for a second time soon after, or as late as 24 hours after, their first mating. When single males were given the opportunity to mate with many females, about equal numbers of them inseminated zero, one, two, or three females. In greenhouse tests, 12 females were given the opportunity to mate with many males of three different esterase phenotypes. Two females mated with one and possibly more males of the same phenotype, and 10 females mated with males of two different esterase phenotypes. In conclusion, multiple mating appears to be a common behavior of males and females of H. glycines.     

Age‐specific maternal effects interact with larval food supply to modulate life history in Coleomegilla maculata

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Multiple Mating Increases Fecundity, Fertility and Relative Clutch Mass in the Female Leopard Gecko (Eublepharis macularius)

Females across many taxa may mate with several males or mate more than once with the same male within one reproductive event. Although many researchers have discussed the effects of multiple mating on reproductive success of females, few studies have attempted to disentangle whether the reproductive success of females differs with respect to whether females mate with multiple males or mate more than once with one male. In this study, we hypothesized that female leopard geckos (Eublepharis macularius) increase aspects of their reproductive success, such as fecundity, fertility and relative clutch mass, by mating more than once within one reproductive event, either by mating repeatedly with the same male or multiply mating with different males. We controlled for the potentially confounding variables of mating frequency and mate number by allowing females to mate once with one male, twice with the same male, or twice with two different males. We found that females that mated with more than one male laid more clutches, exhibited increased egg fertility and invested more in clutches relative to females that mated only once with one male, whereas females that mated twice to the same male were intermediate for these variables. Thus, reproductive success is higher among female leopard geckos that mated with more than one male compared to female leopard geckos that mated only once.     

High levels of polyandry,but limited evidence for multiple paternity,in wild populations of the western rock lobster (Panulirus cygnus)

Polyandry, where multiple mating by females results in the temporal and spatial overlap of ejaculates from two or more males, is taxonomically widespread and occurs in varying frequencies within and among species. In decapods (crabs, lobsters, crayfish, and prawns), rates of polyandry are likely to be variable, but the extent to which patterns of multiple paternity reflect multiple mating, and thus are shaped by postmating processes that bias fertilization toward one or a subset of mated males, is unclear. Here, we use microsatellite markers to examine the frequency of multiple mating (the presence of spermatophores from two or more males) and patterns of paternity in wild populations of western rock lobster (Panulirus cygnus). Our data confirm that >45% of females had attached spermatophores arising from at least two males (i.e., confirming polyandry), but we found very limited evidence for multiple paternity; among 24 clutches sampled in this study, only two arose from fertilizations by two or more males. Single inferred paternal genotypes accounted for all remaining progeny genotypes in each clutch, including several instances when the mother had been shown to mate with two or more males. These findings highlight the need for further work to understand whether polyandry is adaptive and to uncover the mechanisms underlying postmating paternity biases in this system.     

Strong male‐biased operational sex ratio in a breeding population of loggerhead turtles (Caretta caretta) inferred by paternal genotype reconstruction analysis

Characterization of a species mating systems is fundamental for understanding the natural history and evolution of that species. Polyandry can result in the multiple paternity of progeny arrays. The only previous study of the loggerhead turtle (Caretta caretta) in the USA showed that within the large peninsular Florida subpopulation, multiple paternity occurs in approximately 30% of clutches. Our study tested clutches from the smaller northern subpopulation for the presence of multiple paternal contributions. We examined mothers and up to 20 offspring from 19.5% of clutches laid across three nesting seasons (2008–2010) on the small nesting beach on Wassaw Island, Georgia, USA. We found that 75% of clutches sampled had multiple fathers with an average of 2.65 fathers per nest (1–7 fathers found). The average number of fathers per clutch varied among years and increased with female size. There was no relationship between number of fathers and hatching success. Finally, we found 195 individual paternal genotypes and determined that each male contributed to no more than a single clutch over the 3‐year sampling period. Together these results suggest that the operational sex ratio is male‐biased at this site.     

Differences in biological and behavioural characteristics of Harmonia axyridis (Coleoptera: Coccinellidae) according to colour patterns of elytra

Samples of overwintering populations of Harmonia axyridis were classified into five colour patterns, succinea 1, succinea 2, conspicua, spectabilis and axyridis. Generally, the succinea 1 colour pattern was dominant at six collecting sites in Daejon and Chungnam provinces of Korea and their occurrence proportion was over 50%. Non‐melanic forms such as succinea 1 and 2 were noted in about 90% of collected individuals. On the other hand, conspicua and spectabilis as melanic forms were dominant in the laboratory. One hypothesis is that the difference between field and laboratory populations is related to non‐random mating and environmental conditions. Practically, in mate‐choice experiments, most colour pattern ladybirds seemed to mate with the melanic form, especially the conspicua form. The body size (length) of H. axyridis in colour patterns was slightly different, but in females, there was no significant difference among colour patterns. However, both lengths of males were significantly different among colour patterns. In reproductive ability (fecundity), there was no difference among colour patterns.     

Remating, sperm transfer, and sperm displacement in the cactophilic species Drosophila buzzatii Patterson & Wheeler (Diptera: Drosophilidae)

The mating system of Drosophila buzzatii is characterized by short copulation duration, frequent remating in both males and females, and male ejaculate partitioning. Additional features of the system are strong sperm displacement and a high frequency of sterile matings. Remating frequencies and the effects of remating on various mating parameters were studied. In order to characterize variation, five isofemale lines from geographically distant localities in Australia (three localities), Brazil and the Canary Islands were used. Mating parameters studied were: premating time, copulation duration, interval between successive matings, and progeny number as a measure of sperm transfer. Variation for sperm displacement was studied in crosses between laboratory stocks and a number of isofemale lines from Australia. There were significant between‐line differences in female remating frequencies, premating time, copulation duration, interval between successive matings, and progeny numbers, indicating genetic variation for these traits. Females from the five lines mated on average 1.6 to 3.1 times in 4 h, with a maximum of eight matings for one female. The males were given a maximum of ten virgin females in sequence and more than one‐third of the males mated all ten females in the 2 h observation period. Copulation duration decreased and interval between matings increased with copulation number in multiply mated males. Mean copulation duration was c. 2 min. Sperm transfer, measured as the average number of progeny from a single mating, was low (c. 25) and multiply mated females gave more progeny than single mated females, although with much lower progeny numbers than observed in wild‐caught non‐virgin females. A surprisingly high proportion of observed matings gave no progeny, i.e. they were sterile matings. Sperm displacement was strong in most crosses and remained strong in multiply mated females. The results are discussed in relation to the evolution of mating patterns in Drosophila.     

Small spermatophore size and reduced female fitness in an isolated butterfly population

1. The Glanville fritillary butterfly (Melitaea cinxia L.) has a small population (N_e ～ 100) on the small island of Pikku Tytärsaari (PT) in the Gulf of Finland. The population has remained completely isolated for ～100 generations, which has resulted in greatly reduced genetic variation and high genetic load (low fitness). In particular, females lay small egg clutches with a low egg‐hatching rate in comparison with a large reference population in the Åland Islands (ÅL). 2. In the present study, to what extent egg clutch size and egg‐hatching rate are influenced by male population and spermatophore size was analysed. 3. Spermatophore size increases with male body size, is smaller after the first mating, and is smaller in the small PT population. In the ÅL population but not in the PT population, the egg‐hatching rate increases with spermatophore size. The egg‐hatching rate of PT females is higher when mated with ÅL males than when mated with PT males (heterosis), but there is no such effect on clutch size. The clutch size of ÅL females is, however, reduced when mated with PT males. 4. These results indicate that both male and female traits contribute to reduced reproductive fitness in the small isolated population.     

Female loggerhead sea turtles (Caretta caretta L.) rarely remate during nesting season

The goal of this study was to assess the consequences of single versus multiple paternity by identifying paternity of clutches per female to identify whether there were detectable costs or benefits. Multiple mating can occur when the benefits of mating outweigh the costs, but if costs and benefits are equal, no pattern is expected. Previous research on loggerhead sea turtle (Caretta caretta) populations found male‐biased breeding sex ratios and multiple mating by many females nesting in southwestern Florida. A sample of nesting loggerhead females who laid more than one nest over the course of the season and a subset of their hatchlings were examined from 36 clutches in 2016 on Sanibel Island, Florida. Males that fathered hatchlings in the first clutch sampled were identified in subsequent clutches. Interestingly, 75% of the females analyzed had mated singly. No male was represented in more than one female's clutches. The results suggest that females likely mate at the beginning of the season and use stored sperm for multiple clutches. Evidence for mating between laying events was limited. There was no consistent pattern across the subsequent multiple paternity clutches, suggesting benefits to loggerhead females likely equal their costs and subsequent mating is likely determined by female preference.     

Effects of multiple matings on reproductive fitness of male and female Diaeretiella rapae

Mating frequency and the amount of sperm transferred during mating have important consequences on progeny sex ratio and fitness of haplodiploid insects. Production of female offspring may be limited by the availability of sperm for fertilizing eggs. This study examined multiple mating and its effect on fitness of the cabbage aphid parasitoid Diaeretiella rapae McIntosh (Hymenoptera: Aphidiidae). Female D. rapae mated once, whereas males mated with on average more than three females in a single day. The minimum time lag between two consecutive matings by a male was 3 min, and the maximum number of matings a male achieved in a day was eight. Sperm depletion occurred as a consequence of multiple mating in D. rapae. The number of daughters produced by females that mated with multiple‐mated males was negatively correlated with the number of matings achieved by these males. Similarly, the proportion of female progeny decreased in females that mated with males that had already mated three times. Although the proportion of female progeny resulting from multiple mating decreased, the decrease was quicker when the mating occurred on the same day than when the matings occurred once per day over several days. Mating success of males initially increased after the first mating, but then males became ‘exhausted’ in later matings; their mating success decreased with the number of prior matings. The fertility of females was affected by mating with multiple‐mated males. The study suggests that male mating history affects the fitness of male and female D. rapae.     

Effects of parental age at mating on the fecundity and progeny sex ratio of <Emphasis Type="Italic">Campoletis chlorideae</Emphasis>, an endolarval parasitoid of the pod borer, <Emphasis Type="Italic">Helicoverpa armigera</Emphasis>

Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), commonly called gram pod borer, is a serious pest of pulse crops in India and damages chickpea (Cicer arietinum L). On average a 30% crop loss is reported. One of the potential natural enemies reported for its biological control is Campoletis chlorideae Uchida (Hymenoptera: Ichneumonidae), an idiobiont parasitic wasp that attacks second instar larvae of the pod borer. Male-biased sex ratios hinder efforts to mass release parasitic Hymenoptera for biological control by making the production of females costly. Parental age at time of mating is known to affect the progeny sex ratio in some Braconidae. In this view, we examined the reproduction and survival of the parasitoid C. chlorideae in the laboratory [22 ± 4°C, 70 ± 10% R.H. and 10:14 h light:dark photoperiod]. All the nine (i.e. 3 × 3) combinations of possible mating were made for males and females, being each of different age groups (0–12, 48–60 and 96–108 h old). Second instar larvae of the pod borer were reared on chickpea and exposed to mated females of different mating age group combinations. Results revealed that (a) newly emerged females (0–12 h old) mated to aged males yielded more progeny contrasted to the older females inseminated by newly emerged males (0–12 h old). This implies that increased maternal age at mating caused greater reduction in the progeny yield than that of paternal age; (b) the progeny sex ratio was lowest (0.344 ± 0.057) (mean ± SD) when parents mated just after emergence (0–12 h old). It became more and more male biased as parents were deprived of mate and was highest (0.666 ± 0.701) for 96–108 h old parents. The analysis of variance of the data shows that mate deprived males caused greater contribution in the production of sons, resulting a male-biased progeny sex ratio. Our conclusion indicate that newly emerged parasitoids (0–12 h old) are most fecund and should be used in rearing practices in order to enhance the progeny yield, particularly with respect to daughters. Handling Editor: Torsten Meiners     

Revisiting telegony: offspring inherit an acquired characteristic of their mother's previous mate

Newly discovered non‐genetic mechanisms break the link between genes and inheritance, thereby also raising the possibility that previous mating partners could influence traits in offspring sired by subsequent males that mate with the same female (‘telegony’). In the fly Telostylinus angusticollis, males transmit their environmentally acquired condition via paternal effects on offspring body size. We manipulated male condition, and mated females to two males in high or low condition in a fully crossed design. Although the second male sired a large majority of offspring, offspring body size was influenced by the condition of the first male. This effect was not observed when females were exposed to the first male without mating, implicating semen‐mediated effects rather than female differential allocation based on pre‐mating assessment of male quality. Our results reveal a novel type of transgenerational effect with potential implications for the evolution of reproductive strategies.     

Effects of male genetic contribution and paternal investment to egg and hatchling size in the cricket,Gryllus firmus

Egg provisions represent the complete energy supply of oviparous organisms from fertilization until hatching, and egg size is generally correlated with initial offspring size and a suite of other early fitness related traits. Since egg size is determined by the mother, little attention has been given to potential sources of paternal effects on either egg size or initial offspring size. This study considers two processes by which the sire can affect the egg size and/or initial body size of his offspring, in the multiply mated cricket, Gryllus firmus. The first is a paternal genetic effect, whereby differences in offspring genotype result in differences in the efficiency of metabolising available resources. The second is a paternal environmental effect, whereby the quality or size of paternal investment varies among male phenotypes and this is correlated with the size of eggs the females subsequently produce. Using a one-locus two-allele recessive mutation for pale eye colour as a marker, a mating experiment was designed which enabled the discrimination between eggs fertilized by two males mated simultaneously to a single female. The results of this experiment suggest that sire effects on egg and initial body size occur through both processes. Eggs fertilised by the two males were significantly different at day ten of development, suggesting that the genetic contribution of the sire is affecting embryonic body size. Further, a negative correlation was found between the head size of the pale eyed male and the size of all the eggs that the female laid, suggesting an effect of male size of the amount of nutrients the female receives from her mates. The results of this study suggest that paternal effects may be both more common and more profound than previously thought, and that studies examining early fitness traits that are correlated to egg size or initial body size, in oviparous animals, should consider the possibility and importance of the paternal contribution.     

Pupation site and emergence time influence the mating success of bagworm females,Oiketicus kirbyi

In commercial oil palm plantations in Costa Rica, we tested the hypotheses that pupation site and emergence time affect the mating success of protogynous female bagworms,Oiketicus kirbyi (Guilding) (Lepidoptera: Psychidae). Greater proportions of female than male pupae on upper leaves of oil palms and greater proportions of mated females in the upper rather than lower crown strata support the hypothesis that selection of pupation site by female larvae influences the mating success of adults. Increasing captures of males with increasing trap height further suggest that enhanced mating success of females in tree tops may be attributed either to most effective dissemination of sex pheromone on higher sites, or to males foraging predominantly in the upper strata of oil palms. As the majority of females pupated in the middle rather than upper crown of oil palms, selection of pupation site by females may be affected by additional as yet unknown factors. Emergence of females significntly preceded emergence of males. Increasing proportions of mated females throughout the emergence seasons probably resulted from an increased ‘availability’ of males. In tropical rainforests with local variations inO. kirbyi developmental time and stage, protogyny may represent an evolutionary strategy that furthers outbreeding.