Genetic Analysis of Esterase Polymorphism in the Soybean Cyst Nematode, Heterodera glycines

The genetic basis of esterase polymorphism in Heterodera glycines was investigated through controlled matings and analysis of F₁ and F₂ progeny. Three nematode lines, each fixed for a different esterase phenotype, were isolated and purified through repeated directional selection and inbreeding. Each phenotype was characterized by its distinct pair of closely spaced bands of esterase activity. Single-female single-male crosses were conducted according to a modified agar-plate mating technique. F₁ progeny were homogeneous, exhibiting both parental esterase phenotypes (codominant heterozygotes) but no hybrid bands. Approximately 1,500 F₂ progeny segregated in a 1:2:1 ratio for expression of the esterase phenotypes of the female parental line, the heterozygote, and the male parental line. Apparently the three esterase phenotypes correspond to three codominant alleles of a single esterase locus. Reciprocal crosses gave similar results, suggesting no maternal inheritance.     

Selection Against Heterodera glycines Males by Soybean Lines with Genes for Resistance

Soybeans with genes for resistance select against Heterodera glycines with the corresponding genes for avirulence. There may be a differential effect of sex with some specific gene interactions, which would influence the magnitude of gene frequency changes. No effect on H. glycines males was detected with one selected nematode population and the resistant soybean line PI88788. The selective effect of PI89772 against male nematodes was greater with two inbred nematode populations than with one selected (on PI88788) population, presumably due to differences in H. glycines gene frequencies. ''Peking'' also had few males with the one inbred nematode population, whereas Forrest and ''Pickett 71'' had intermediate numbers. Apparently Forrest and Pickett 71 did not get all the Peking genes for resistance that affect male as well as female nematode development. Other H. glycines-soybean genes stop only females, since there were few or no cysts, except on the susceptible soybean Williams. The number of males'' phenotype will help identify specific genes in both organisms.     

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.     

Egg presence, egg loss, and female mate preferences in the sand goby (Pomatoschistus minutus)

We studied the effect of egg presence on female mate choicein a fish with paternal care. Females who were allowed a freechoice between two males mated within a shorter time than femaleswho were randomly assigned to a particular male. When a secondfemale was allowed to choose among the males, she preferredthe same male as the previous female. This result shows thatfemales are concordant in their mate choice. When the initialfemale was randomly assigned to mate with one of two males (forcedchoice), the second female mated randomly with respect to thefirst one. Thus females do not prefer males with eggs. If theinitial female was given a free choice, but the eggs were removedfrom the chosen male, the test female mated randomly. When boththe males initially had mated but one randomly determined male'seggs were removed, the test female preferred the male who wasstill guarding eggs. These experiments show that females avoidspawning in unsuccessful nests. When the females in the freechoice/egg removal experiment mated with the unsuccessful malethere was a considerably bigger size difference in favor ofthis male than when the females mated with the other male. Weconclude that female sand gobies show clear mate preferences,but that they do not prefer males with eggs over males withouteggs. They do, however, avoid mating with males guarding unsuccessfulnests. We therefore suggest that egg loss could be an importantfactor selecting for egg preference.     

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.     

Sperm storage mediated by cryptic female choice for nuptial gifts

Polyandrous females are expected to discriminate among males through postcopulatory cryptic mate choice. Yet, there is surprisingly little unequivocal evidence for female-mediated cryptic sperm choice. In species in which nuptial gifts facilitate mating, females may gain indirect benefits through preferential storage of sperm from gift-giving males if the gift signals male quality. We tested this hypothesis in the spider Pisaura mirabilis by quantifying the number of sperm stored in response to copulation with males with or without a nuptial gift, while experimentally controlling copulation duration. We further assessed the effect of gift presence and copulation duration on egg-hatching success in matings with uninterrupted copulations with gift-giving males. We show that females mated to gift-giving males stored more sperm and experienced 17% higher egg-hatching success, compared with those mated to no-gift males, despite matched copulation durations. Uninterrupted copulations resulted in both increased sperm storage and egg-hatching success. Our study confirms the prediction that the nuptial gift as a male signal is under positive sexual selection by females through cryptic sperm storage. In addition, the gift facilitates longer copulations and increased sperm transfer providing two different types of advantage to gift-giving in males.     

Effects of mating rates on oviposition,sex ratio and longevity in a predatory mite <Emphasis Type="Italic">Neoseiulus cucumeris</Emphasis> (Acari: Phytoseiidae)

Two aspects of mating effects on the fecundity, sex ratio and longevity of Neoseiulus cucumeris (Acari: Phytoseiidae) were examined in laboratory experiments: (1) females mated by one, two or three different males (unmated and 3 days old) at 5-day intervals, and (2) females mated by males with different age/mating status (number of females mated previously by the male). Females allowed to mate with a second or third male at 5-day intervals produced 39 eggs on average, but those mated with a single male produced 28 eggs on average. Matings with additional males 5 or 10 days after the first male increased the duration of the oviposition period of these females by 5–7 days and at the same time reduced the post-oviposition period by about 10 days. Overall, females with additional matings by one or two different males at 5-day intervals survived a few days shorter than females without additional males. Mating with a different female each day, a male of N. cucumeris could mate with 5–8 females, which produced a total of 85–116 eggs: females mated with a male during days 1 and 2 in its adulthood and with a male of the last 2 days of life (days 7 and 8) produced about half as many eggs as females mated with a male during 3–6 days of its adulthood. Females mated with males that are too young or too old had a shorter oviposition period and a longer post-oviposition period and longevity than females mated with middle-aged males. In both experiments, rates of oviposition remained similar in females with high or low fecundity. This indicates that in both cases, the increased fecundity is due to the extension of the oviposition period through additional sperm supplied by the second male and or third male (in experiment 1) or more sperm by males not too young nor too old (experiment 2).     

Evidence for Inbreeding Depression and Pre-Copulatory,but Not Post Copulatory Inbreeding Avoidance in the Cabbage Beetle Colaphellus bowringi

Inbreeding is known to have adverse effects on fitness-related traits in a range of insect species. A series of theoretical and experimental studies have suggested that polyandrous insects could avoid the cost of inbreeding via pre-copulatory mate choice and/or post-copulatory mechanisms. We looked for evidence of pre-copulatory inbreeding avoidance using female mate preference trials, in which females were given the choice of mating with either of two males, a sibling and a non-sibling. We also tested for evidence of post-copulatory inbreeding avoidance by conducting double mating experiments, in which four sibling females were mated with two males sequentially, either two siblings, two non-siblings or a sibling and a non-sibling in either order. We identified substantial inbreeding depression: offspring of females mated to full siblings had lower hatching success, slower development time from egg to adult, lower survival of larval and pupal stages, and lower adult body mass than the offspring of females mated to non-sibling males. We also found evidence of pre-copulatory inbreeding avoidance, as females preferred to mate with non-sibling males. However, we did not find any evidence of post-copulatory inbreeding avoidance: egg hatching success of females mating to both sibling and non-sibling males were consistent with sperm being used without bias in relation to mate relatedness. Our results suggest that this cabbage beetle has evolved a pre-copulatory mechanism to avoid matings between close relative, but that polyandry is apparently not an inbreeding avoidance mechanism in C. bowringi.     

Cost of reproduction in male medflies: The primacy of sexual courting in extreme longevity reduction

In polygynous insect species, male reproductive success is directly related to lifetime mating success. However, the costs for males of sexual activities such as courting, signaling, and mating are largely unknown. We studied the cost of sexual activities in male Mediterranean fruit flies, Ceratitis capitata (Tephritidae), a polygynous lekking species, by keeping cohorts of individual male flies under relaxed crowding conditions in the laboratory. We used 5 cohorts among which individuals differed in their opportunities to interact with con-specifics and recorded life span, and in one treatment, mating rate. We found that males kept singly lived more than twice as long as males that interacted intensively with mature virgin females, while male-male interactions caused a smaller reduction in longevity. Because longevity of males that could court but not mate was not significantly different from those that could court and mate, we conclude that courting (not mating) was responsible for the observed longevity reduction. Moreover, we detected high variability in male mating success, when 5 virgin females were offered daily. In contrast to the cohort level, individual males that mated at a high rate lived relatively long, thus indicating heterogeneity in quality or sexual strategy among males.     

Female pied flycatchers trade between male quality and mating status in mate choice

According to mate choice theory, females should consider both male quality and mating status when choosing a mate. In birds, strong experimental evidence indicates that females prefer males with elaborate traits. No comparable evidence exists to determine whether females take male mating status into account or how they may trade between male quality and male mating status. We studied mate choice of female pied flycatchers (Ficedula hypoleuca) in outdoor aviaries where the effect of territory quality could be eliminated and where we could control which males were mated and which were unmated. We used male plumage colour as our measure of male quality. In the aviaries, focal females could easily compare males and assess their plumage colour and mating status, and resident females were prevented from attacking prospecting females because of separation in different compartments. The study provided evidence for a trade-off in mate choice. Females may compromise by choosing an already mated male if he is more brightly coloured and, presumably, of higher quality than available unmated males. The study did not support the idea that polygyny is based on male deception of females, but the results were consistent with the female aggression hypothesis.     

Male Drosophila melanogaster show adaptive mating bias in response to female infection status

Given the non-trivial cost of reproduction for males and substantial variation in female quality, males have been predicted to show mating bias as an evolved strategy. Using a large outbred population of Drosophila melanogaster, we test this prediction and show that males may adaptively bias their mating effort in response to the infection status of females. Given a simultaneous choice between females infected with pathogenic bacteria and sham infected females, males preferentially mated with the latter, who had a higher reproductive output compared to infected females. This may provide evidence for pre-copulatory male mate choice. Assessment of the reproductive behaviour ensured that the observed pattern of mating bias was not due to differences in receptivity between females infected with pathogenic bacteria and sham infected females. Further, there was no evidence for post-copulatory male mate choice measured in terms of copulation duration.     

FEMALES RECEIVE A LIFE-SPAN BENEFIT FROM MALE EJACULATES IN A FIELD CRICKET

Abstract.— Mating has been found to be costly for females of some species because of toxic products that males transfer to females in their seminal fluid. Such mating costs seem paradoxical, particularly for species in which females mate more frequently than is necessary to fertilize their eggs. Indeed, some studies suggest that females may benefit from mating more frequently. The effect of male ejaculates on female life span and lifetime fecundity was experimentally tested in the variable field cricket, Gryllus lineaticeps. In field crickets, females will mate repeatedly with a given male and mate with multiple males. Females that were experimentally mated either repeatedly or multiply lived more than 32% longer than singly mated females. In addition, multiply mated females produced 98% more eggs than singly mated females. Because females received only sperm and seminal fluid from males in the experimental matings, these life‐span and fecundity benefits may result from beneficial seminal fluid products that males transfer to females during mating. Mating benefits rather than mating costs may be common in many animals, particularly in species where female mate choice has a larger effect on male reproductive success than does the outcome of sperm competition.     

Mating Status Effects on Sexual Response of Males and Females in the Parasitoid Wasp <Emphasis Type="Italic">Urolepis rufipes</Emphasis>

Although mate preferences are most commonly examined in females, they are often found in both sexes. In the parasitoid wasp Urolepis rufipes, both female and male mating status affected certain aspects of sexual interactions. Female mating status mattered only in the later stages of mating. Males did not discriminate between virgin and mated females in terms of which they contacted or mounted first. However, once mounted, most virgin females were receptive to copulation, whereas very few mated females were. Whether a male’s mating status affected his own sexual response depended on the female’s ability to respond and the stage of mating. Examining male behavior toward dead females allowed elimination of the role of female behavior in how males responded. Virgin and mated males are both attracted to dead females as evidenced by their fanning their wings at such females. However, mated males were quicker than virgin males to contact and to mount in an experiment with dead females, whereas there was no such differential response in an experiment with live females. This difference is consistent with greater female sexual responsiveness to virgin males. Male mating status also affected female receptivity to copulate. Once mounted, live virgin females were less likely to become receptive to copulation by mated males than to virgin males, but only in a choice experiment, not in a no-choice experiment.     

Sexual reproduction in Daphnia pulex (Crustacea: Cladocera): observations on male mating behaviour and avoidance of inbreeding

1.  Mating behaviour in Daphnia appears to rely on random contact between males and sexual females rather than diffusible pheromones. Males may be able to discriminate sexually receptive females from females in other developmental stages and increase their mating efficiency. Males may also use chemical signals to avoid mating with females from the same clone and avoid the severe inbreeding depression that has been documented for intraclonal mating. The present study used experiments to test for the avoidance of intraclonal mating and assess male mating efficiency in D. pulex .
2.  Three clones were examined for the avoidance of intraclonal mating by providing males with an opportunity to mate with females of the same or two different clones. The proportion of intraclonal matings did not differ from the proportion of interclonal matings, suggesting that D. pulex males do not use kin discrimination to avoid mating with females from the same clone.
3.  The proportion of mated females decreased with increasing numbers and density of sexual females when exposed to a single male. This observation suggests that a male spends more time pursuing and copulating with sexually receptive females than non-receptive females and there is insufficient time to mate with all sexual females. The decrease in proportion of females mated could also be the result of sperm depletion in the male. Sperm depletion is unlikely to occur in nature because sexually receptive females are much rarer than in the experimental conditions.     

Silk‐ and volatile‐based male mate choice in the genital plug‐producing spider

Female mating history can have a strong effect on male fertilization success. Although males often prefer to mate with virgin females, they often also engage with mated females. As the intensity of sperm competition can differ among mated females, males are expected to evolve means to identify their status. In spiders, males often use female silk to gather information about female quality. Males of many spider species deposit mating plugs into female genitalia to hinder further copulations. We tested whether males of the foliage‐dwelling, plug‐producing spider Philodromus cespitum, which is an important natural enemy of pests, discriminate between females of different mating status and whether they can determine the extent of genital plugging in mated females solely on the basis of cues gained from deposited female silk. We presented males with draglines of females that varied in either mating status (virgin vs. mated), the extent of plugging (small vs. big plug), or the age of the plug (fresh vs. old plug) and examined their mate preferences. Additionally, we tested whether males were attracted to volatile cues produced by female bodies. Our experiments revealed that males preferred draglines of virgin females to those of mated females, and mated females with small plugs to those with large plugs. They were also attracted to female volatile cues. This study suggests that males are able to extract fine‐scale information on mating status from female draglines.     

The role of mating history and male size in determining mating behaviours and sexual conflict in a water strider

We conducted three experiments to test the effects of mating history of both sexes and of male body size on mating behaviours in the water strider, Gerris buenoi. Our manipulations influenced the interests of both sexes and, thus, the degree of conflict over mating behaviours. Mating history was a dichotomous variable (deprived/mated), depending on holding conditions in the laboratory. Experiment 1 considered and found independent effects of male and female mating history on latency to copulation and copulation duration. In experiment 2, we manipulated only female mating history, using unsuccessful struggle rates as evidence for female reluctance and conflict over mating. Finally, we investigated the relation between male body size and mating history on copulation duration. We predicted that intersexual conflict over mating would be lowest when females were deprived, because female interests under these conditions should more closely match those of males. Deprived females began mating in half the time of mated females and were twice as likely to mate because of reduced reluctance. Furthermore, copulation duration for deprived males was about one and a half times longer than that for mated males. Although previous studies examining nonrandom mating patterns by size predicted longer copulations for small males, we found that small males prolonged copulation when deprived more than large males. We conclude that females primarily influence copulation frequency, but males primarily influence copulation duration. Our results favour the hypothesis that reduced mating opportunity for small males accounts for their extended copulation duration. Finally, our findings provide evidence for strong effects of male body size on selection mechanisms in water striders, and support the hypothesis of conflicting pre- and postcopulatory selection mechanisms in this group. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.     

Male death resulting from hybridization between subspecies of the gypsy moth,Lymantria dispar

We explored the origin of all-female broods resulting from male death in a Hokkaido population of Lymantria dispar through genetic crosses based on the earlier experiments done by Goldschmidt and by testing for the presence of endosymbionts that are known to cause male killing in some insect species. The mitochondrial DNA haplotypes of the all-female broods in Hokkaido were different from those of normal Hokkaido females and were the same as those widely distributed in Asia, including Tokyo (TK). Goldschmidt obtained all-female broods through backcrossing, that is, F1 females obtained by a cross between TK females (L. dispar japonica) and Hokkaido males (L. dispar praeterea) mated with Hokkaido males. He also obtained all-male broods by mating Hokkaido females with TK males. Goldschmidt inferred that female- and male-determining factors were weakest in the Hokkaido subspecies and stronger in the Honshu (TK) subspecies. According to his theory, the females of all-female broods mated with Honshu males should produce normal sex-ratio broods, whereas weaker Hokkaido sexes would be expected to disappear in F1 or F2 generations after crossing with the Honshu subspecies. We confirmed both of Goldschmidt''s results: in the case of all-female broods mated with Honshu males, normal sex-ratio broods were produced, but we obtained only all-female broods in the Goldschmidt backcross and obtained an all-male brood in the F1 generation of a Hokkaido female crossed with a TK male. We found no endosymbionts in all-female broods by 4,′6-diamidino-2-phenylindole (DAPI) staining. Therefore, the all-female broods observed in L. dispar are caused by some incompatibilities between Honshu and Hokkaido subspecies.     

Is egg carrying attractive? Mate choice in the golden egg bug (Coreidae,Heteroptera)

In several species of fish, females select males that are already guarding eggs in their nests. It is a matter of debate as to whether a female selects a good nest site for her offspring (natural selection) or a male for his attractiveness (sexual selection). The golden egg bug, Phyllomorpha laciniata Vill, resembles fish in the sense that mating males carry more eggs than single males, but in the bugs, female mate choice is decoupled from egg site choice. The sexual selection hypothesis predicts that if females select males using male egg load as a cue for male quality, they should not mate with a male when eggs are removed, regardless of his mating attempts. When individual females were enclosed with an egg-loaded male and an unloaded male, they mated equally often with both males, although the loaded males courted more. In addition, when only successful males were used, females mated equally often with the loaded male and the unloaded male irrespective of sex ratio. Male choice rather than female choice affected mating frequency when sex ratio was equal. Therefore, females do not select the male by the eggs he carries, but successful males may receive many eggs due to egg dumping by alien females while they mate or as a consequence of mate guarding.     

Males of the orb-web spider Argiope bruennichi sacrifice themselves to unrelated females

Costs of inbreeding can lead to total reproductive failure and inbreeding avoidance is, therefore, common. In classical sex roles with no paternal care, the selective pressure to avoid inbreeding is mostly on the female, which carries the higher costs. In some orb-web spiders, this situation is very different because females are polyandrous and males are monogynous or at most bigynous. Additionally, females of many entelegyne orb weavers are thought to bias paternity post-copulatorily towards a desired mate. This increases the selective pressure on males to adjust their investment in a mating with regard to the compatibility to a female.Here, we examine whether genetic relatedness influences mating behaviour in the orb-web spider Argiope bruennichi. We mated either a sibling or a non-sibling male to a female in single copulation trials and compared copulation duration, cannibalism rate and female fecundity.Our experiment revealed that males prolonged their copulation duration and were cannibalized more frequently when mating with a non-sibling female. Males mating with a sibling female were more likely to escape cannibalism by copulating briefly, thus presumably increasing their chances of re-mating with a more compatible female. This suggests that males can adaptively adjust their investment relating to the compatibility of a female.     

Chemosensory cues allow male Tenebrio molitor beetles to assess the reproductive status of potential mates

Males of many insect species, including beetles, choose their mates according to their reproductive status. However, the ways in which male beetles evaluate female reproductive status have received little attention. We tested the existence of male mate choice in the mealworm beetle, Tenebrio molitor, by observing mating and courtship behaviour of males given simultaneous access to pairs of females differing in their reproductive status: (1) mature versus immature; (2) virgin versus previously mated; (3) familiar (mated with the experimental male) versus unfamiliar (mated with a different male). Males courted and mated preferentially with mature and virgin females. To determine whether chemical cues played a role in these discriminations, we exposed males to filter paper squares bearing chemical cues from different types of females: (1) virgin versus mated; (2) mature versus immature. Males were significantly more attracted to those squares bearing chemical cues from virgin and mature females, suggesting that males can assess female reproductive status on the basis of chemical cues alone.