Testing for reproductive interference in the population dynamics of two congeneric species of herbivorous mites

When phylogenetically close, two competing species may reproductively interfere, and thereby affect their population dynamics. We tested for reproductive interference (RI) between two congeneric haplo-diploid spider mites, Tetranychus evansi and Tetranychus urticae, by investigating their interspecific mating and their population dynamics when they competed on the same plants. They are both pests of tomato, but differ in the host plant defences that they suppress or induce. To reduce the effect of plant-mediated interaction, we used a mutant tomato plant lacking jasmonate-mediated anti-herbivore defences in the competition experiment. In addition, to manipulate the effect of RI, we introduced founder females already mated with conspecific males in mild RI treatments or founder, virgin females in strong RI treatments (in either case together with heterospecific and conspecific males). As females show first-male sperm precedence, RI should occur especially in the founder generation under strong RI treatments. We found that T. urticae outcompeted T. evansi in mild, but not in strong RI treatments. Thus, T. evansi interfered reproductively with T. urticae. This result was supported by crossing experiments showing frequent interspecific copulations, strong postmating reproductive isolation and a preference of T. evansi males to mate with T. urticae (instead of conspecific) females, whereas T. urticae males preferred conspecific females. We conclude that interspecific mating comes at a cost due to asymmetric mate preferences of males. Because RI by T. evansi can improve its competitiveness to T. urticae, we propose that RI partly explains why T. evansi became invasive in Europe where T. urticae is endemic.     

Effects of predation risk on mating behavior of the Kanzawa spider mite

In the Kanzawa spider mite, Tetranychus kanzawai, adult males locate pre-reproductive quiescent females and engage in precopulatory mate guarding. We found that re-reproductive quiescent females preferred to be near veins, rather than other leaf parts, and moreover, adult males spent more time along the vein than on other parts. Consequently, T. kanzawai males found more quiescent females along veins than those on other parts. However, the predatory mite Neoseiulus womersleyi also found more quiescent T. kanzawai females along veins than those on other parts. Moreover, N. womersleyi found more guarding males than solitary males of T. kanzawai. Thus, we experimentally examined the effects of predation risk on the mating behavior of T. kanzawai. The presence of N. womersleyi reduced T. kanzawai female preference for vein vicinity as a quiescent site. Although the predation risk of guarding T. kanzawai males was lower than that of solitary males after detection by predators, the presence of N. womersleyi also reduced the proportion of guarding T. kanzawai males. These results suggest that the possible benefits of preferring vein vicinity as quiescent sites by T. kanzawai females is outweighed by predation risk in the presence of predators, and that the risk of detection by predators would be more important for T. kanzawai males than the risk of being preyed upon.     

Development of Neoseiulus womersleyi (Schicha) and Euseius ovalis (Evans) feeding on four tetranychid mites (Acari: Phytoseiidae,Tetranychidae) and pollen

The development of the predatory mites, Neoseiulus womersleyi (Schicha) and Euseius ovalis (Evans), feeding on four tetranychid mites (Tetranychus urticae, Tetranychus kanzawai, Oligonychus mangiferus, Panonychus citri), maize pollen or Chinese loofah pollen was studied at 25 °C. Immature stages of N. womersleyi feeding on T. urticae and T. kanzawai had shorter developmental duration (4.71 and 5.02 days for females, 4.77 and 5.19 days for males, respectively) than those feeding on other food sources. Immature stages of E. ovalis females feeding on O. mangiferus and T. urticae developed in 4.99 and 5.13 days, respectively, the shortest developmental duration measured. Immature stages of E. ovalis males feeding on O. mangiferus and T. urticae developed in 5.12 and 5.37 days, respectively. The longevity of N. womersleyi males (13.31 to 14.51 days) and females (17.67 to 21.81 days) feeding on T. urticae, T. kanzawai or maize pollen was longer than the longevity of N. womersleyi feeding on O. mangiferus, P. citri or loofah pollen. E. ovalis males (12.91 to 16.74 days) and females (16.24 to 23.77 days) feeding on O. mangiferus, T. urticae or maize pollen lived longer than E. ovalis males and females feeding on T. kanzawai, P. citri or loofah pollen.     

Female response to predation risk alters conspecific male behaviour during pre‐copulatory mate guarding

Mating behaviour often increases predation risk, but the vulnerability within mating pairs differs between the sexes. Such a sex difference is expected to lead to differences in responses to predation risk between the sexes. In the two‐spotted spider mite Tetranychus urticae, males engage in pre‐copulatory mate guarding because only the first mating results in fertilisation. We investigated (i) whether pre‐copulatory pairs are more conspicuous to the predatory mite Phytoseiulus persimilis than solitary females, (ii) whether the vulnerability to the predator differs between sexes within the pre‐copulatory pair, (iii) whether each sex of T. urticae responds to predation risk during pre‐copulatory mate guarding and (iv) whether T. urticae's response to predation risk affects predator behaviour. Because T. urticae females are immobile during pre‐copulatory mate guarding, we observed male behaviour to evaluate effects of predation risk. We found that the predators detect more pre‐copulatory pairs than solitary females and that more females than males of the pre‐copulatory pairs are preyed upon by the predators. The preference of spider mite males for pre‐copulatory pairs versus solitary females was affected by whether or not the female had been exposed to predators during development. Male T. urticae exposed to predation risk did not alter their behaviour. These results suggest that only the most vulnerable sex, that is the female, responds to predation risk, which modifies male behaviour. Regardless of T. urticae females’ experience, however, P. persimilis detected more T. urticae pre‐copulatory pairs than solitary females, suggesting that pre‐copulatory mate guarding itself is dangerous for T. urticae females when these predators are present. We discuss our results in the context of sex‐dependent differences in predation risk.     

Effects of density experience on mate guarding behavior by adult male Kanzawa spider mites

In the Kanzawa spider mite, Tetranychus kanzawai (Acari: Tetranychidae), adult males guard pre-reproductive quiescent females. I experimentally examined the effects of density experience during development and/or after adult emergence on precopulatory mate guarding behavior by T. kanzawai males. Mate guarding behavior was modified by density experience after adult emergence. When males had previously experienced high density after adult emergence (n = 71), 73.2% of them engaged in precopulatory mate guarding. In contrast, when males had previously experienced low density after adult emergence (n = 82), 61.0% of them did not guard females. Mate guarding with physical contact occurred more frequently when males had previously experienced a high density of potential rivals than when they had not, but the difference in behavior between the two groups of males was marginally not significant. Nevertheless, these results suggest overall that T. kanzawai males change mate guarding behavior in response to previously experienced density.     

Predation rates of Neoseiulus womersleyi (Schicha) and Euseius ovalis (Evans) feeding on tetranychid mites (Acari: Phytoseiidae,Tetranychidae)

Predation rates of Neoseiulus womersleyi and Euseius ovalis feeding on eggs, larvae, or protonymphs of Tetranychus urticae, Tetranychus kanzawai, or Oligonychus mangiferus were measured in a chamber at 25 °C. N. womersleyi immatures consumed 9.73 T. urticae eggs, 13.53 larvae, or 11.57 protonymphs, while gravid females consumed 12.13 T. urticae eggs, 14.37 larvae, or 12.07 protonymphs daily. Female N. womersleyi consumed a total of 218.12 T. urticae eggs, 260.85 larvae, or 222.33 protonymphs, while male N. womersleyi consumed a total of 96.39 T. urticae eggs, 112.23 larvae, or 99.65 protonymphs. When O. mangiferus larvae or protonymphs were offered to E. ovalis, immatures consumed 18.57 larvae or 17.47 protonymphs. Gravid females consumed 16.83 larvae or 12.83 protonymphs daily, with a total of 330.68 larvae or 252 protonymphs. Adult E. ovalis males consumed fewer O. mangiferus larvae (107.69) or protonymphs (91.51) than females. Conversion rate of predation to reproduce was expressed as “Food–reproduction exchange rate” of N. womersleyi was lower on T. urticae than on T. kanzawai. E. ovalis showed a higher food–reproduction exchange rate on O. mangiferus than on T. urticae. The results suggest that N. womersleyi and E. ovalis feed mainly on larvae and protonymphs rather than on the eggs of T. urticae, T. kanzawai, and O. mangiferus. We recommended using T. urticae eggs are suitable food for mass rearing for both N. womersleyi and E. ovalis.     

Dispersal of diapausing Tetranychus urticae and T. kanzawai

The spider mites Tetranychus urticae Koch and Tetranychus kanzawai Kishida (Acari: Tetranychidae) overwinter mostly as mated adult diapausing females. Their overwintering survival depends in part on their dispersal towards suitable habitats. We investigated the dispersal behaviour of diapausing females of T. urticae and T. kanzawai with respect to factors known to affect the dispersal of non‐diapausing mites: light, population density, gravity, and humidity. In general, diapausing females of T. urticae showed a stronger tendency to disperse than did those of T. kanzawai under all test conditions. High population density promoted the dispersal of diapausing T. urticae, but not of T. kanzawai. Dispersal of diapausing females of both species was not significantly affected by gravity, humidity, or whether feeding damage was caused by conspecifics or heterospecifics. On plants, more T. urticae than T. kanzawai moved downward. We propose that dispersal after the onset of diapause may be an important life‐history strategy in T. urticae, but not in T. kanzawai.     

Mating interaction of the Japanese horned beetle Trypoxylus dichotomus septentrionalis: does male-excluding behavior induce female resistance?

Female resistance to male mating is thought to have developed as a mechanism to avoid multiple matings or to allow the choice of good quality males. Detailed mating behavior of the Japanese horned beetle Trypoxylus dichotomus septentrionalis (Kono) was examined under natural conditions to clarify the function of female resistance to mating. When a male recognized the presence of a female at the same feeding spot, it immediately began to court and mount the female and attempted to insert its genitalia. During the courtship, females performed intense resistance behavior against the mating, regardless of whether the female would eventually accept males or not. In T. dichotomus septentrionalis, it was observed that female resistance to mating did not occur for reasons of precopulatory mate choice or avoidance of multiple matings; however, additional studies on postcopulatory mate choice are needed. After copulation, males usually tried to keep females away from the sap site. Because of males' excluding behavior, females were able to stay at the sap site for feeding only when they were resisting or copulating. Females that showed resistance behavior before copulation stayed at the sap site 1.56 times longer than females that did not show the resistance behavior. Female resistance behavior in T. dichotomus septentrionalis is thus considered a tactic for prolonging their feeding duration, which is reduced by male-excluding behavior after mating.     

Female and Male Moths Display Different Reproductive Behavior when Facing New versus Previous Mates

Multiple mating allows females to obtain material (more sperm and nutrient) and/or genetic benefits. The genetic benefit models require sperm from different males to fertilize eggs competitively or the offspring be fathered by multiple males. To maximize genetic benefits from multiple mating, females have evolved strategies to prefer novel versus previous mates in their subsequent matings. However, the reproductive behavior during mate encounter, mate choice and egg laying in relation to discrimination and preference between sexes has been largely neglected. In the present study, we used novel and previous mate treatments and studied male and female behavior and reproductive output in Spodoptera litura. The results of this study do not support the sperm and nutrient replenishment hypotheses because neither the number of mates nor the number of copulations achieved by females significantly increased female fecundity, fertility and longevity. However, females showed different oviposition patterns when facing new versus previous mates by slowing down oviposition, which allows the last male has opportunities to fertilize her eggs and the female to promote offspring diversity. Moreover, females that have novel males present called earlier and more than females that have their previous mates present, whereas no significant differences were found on male courtship between treatments. These results suggest that S. litura females can distinguish novel from previous mates and prefer the former, whereas males generally remate regardless of whether the female is a previous mate or not. In S. litura, eggs are laid in large clusters and offspring competition, inbreeding and disease transfer risks are thus increased. Therefore, offspring diversity should be valuable for S. litura, and genetic benefits should be the main force behind the evolution of female behavioral strategies found in the present study.     

Do males evaluate female age for precopulatory mate guarding in the two-spotted spider mite?

In the two-spotted spider mite Tetranychus urticae (Acari: Tetranychidae), because only the first mating results in fertilisation, adult males guard quiescent deutonymph females, the stage immediately before adult emergence. Previous studies showed that T. urticae males prefer to guard older rather than younger females. However, as previous experimental designs have included uncertain factors, findings have not always been sufficiently rigorous. Therefore, we reexamined whether T. urticae males discriminate between the females closest to becoming receptive and younger females. One male was introduced onto leaf squares with two differently aged quiescent deutonymph females, with time lags between the females of 3, 6, 12 or 23 h, and whether the older or younger female was guarded was recorded. When the time lags between the females were 3, 6 or 12 h, males showed no preferences. Males preferred older females over younger ones only when the time lag was 23 h. Under non-choice conditions, fewer males guarded the 23-h younger females than other older females, suggesting that the attractiveness of the younger females is weaker than older females. Our results are not fully consistent with previous studies; T. urticae males do not obviously discriminate between females closest to becoming receptive and younger females.     

Males mate with females even after sperm depletion in the two-spotted spider mite

Generally, males increase their reproductive success by mating with as many females as possible, whereas females increase their reproductive success by choosing males who provide more direct and indirect benefits. The difference in reproductive strategy between the sexes creates intense competition among males for access to females, therefore males spend much energy and time for competition with rival males for their reproduction. However, if they do not need to engage themselves into male competition and females are in no short supply, how many females can a male mate with and fertilize? We address this question in the two-spotted spider mite, Tetranychus urticae Koch. In this study, we investigated how many females a young, virgin male mated in 3 h, and checked whether the mated females were fertilized. We found that on average males mated with 12–13 females (range: 5–25). As latency to next mating did not change with the number of matings, the males are predicted to engage in even more matings if the mating trial were continued beyond 3 h. Copulation durations decreased with the number of matings and typically after 11 copulations with females any further copulations did not lead to fertilization, suggesting that males continued to mate with females even after sperm depletion. We discuss why spider mite males continue to display mating and copulation behaviour even after their sperm is depleted.

     

Effects of host-plant quality on male two-spotted spider mite (Acari: Tetranychidae) mate location and guarding behavior

Effects of host-plant quality on two-spotted spider mite,Tetranychus urticae Koch, mate location and guarding behaviors were described using a no-choice bioassay. Males and quiescent deutonymphs were collected from lima bean leaves of one of two host qualities. High-chlorophyll (HC) leaves had been infested with spider mites for 6–10 days, while low-chlorophyll (LC) leaves had been infested for>21 days. Three parameters of maleT. urticae guarding behavior were quantified: approach arrestment, and arrestment duration. HC males approached quiescent deutonymphs more often than did LC males, even though host quality of females had no effect on male approach frequency. HC males were arrested more frequently by HC quiescent deutonymphs than were LC males, while LC males were arrested more often by LC females than were HC males. However, a different pattern was observed for arrestment duration. HC males were arrested for twice as long by LC quiescent deutonymphs than by HC females, while the LC-male arrestment durations elicited by HC and LC females did not differ. These results show that host-plant quality affectsT. urticae intersexual communication, in terms of both the female signal and the male response. Whether the differing male responses observed in this study indicate alternativeT. urticae mating strategies or are incidental by-products of host-induced physiological changes remains to be determined.     

Mate choice promotes inbreeding avoidance in the two-spotted spider mite

Since inbreeding in Tetranychus urticae can reduce offspring fitness, sexual selection may favour disassortative mate choice with respect to relatedness of the mating partners. We tested whether T. urticae shows this preference for mating with unrelated partners. We chose an experimental set-up with high potential for female choosiness, since females only mate once and are therefore expected to be the choosier gender. An adult virgin female was placed together with two adult males from the same population. One male was unrelated and the other male was related—a brother with whom she had grown up. Significantly more copulations (64%) took place with the unrelated male. Time to mating did not depend on the female-to-male relatedness. The remaining (non-copulating) male tried to interfere with the ongoing mating in the majority of cases, but this interference did not depend on the female-to-male relatedness. These results imply that T. urticae (a) can recognize kin (via genetic and/or environmental similarity) and (b) has the potential to avoid inbreeding through mate choice.     

Possible ecological consequences of heterospecific mating behavior in two tetranychid mites

Interspecific mating between the two-spotted spider mite,Tetranychus urticae Koch, and the Banks grass mite,Oligonychus pratensis (Banks), was documented using laboratory populations. The incidence of mating betweenT. urticae males andO. pratensis females was 26.0%, while that for the reciprocal mating was 18.8%. The incidence of mating was affected by both male and female species. Such matings may have several important ecological consequences. Interspecific matings resulted in all-male progenies. Thus, progeny sex ratios may be distorted by misdirected mating behavior. In addition, heterospecific mating resulted in lower fecundity than conspecific matings in the two-spotted spider mite, although not in the Banks grass mite. Aerial dispersal behavior of the two-spotted spider mite was also affected. Under crowded conditions and deteriorating resource quality, female mites exhibit an aerial dispersal posture that helps them to become airborne, and allows them to disperse long distances. Forty-two percent ofT. urticae females that mated with conspecific males exhibited this dispersal behavior, compared to only 3.6% for virgin females. The incidence of aerial dispersal behavior for females that mated with heterospecific males was intermediate (27.3%). The effects of these behavioral alterations on male and female fitness may depend on the population structure and resource distribution.     

Wolbachia-Host Interactions: Host Mating Patterns Affect Wolbachia Density Dynamics

Wolbachia are maternally inherited intracellular bacteria that infect a wide range of arthropods and cause an array of effects on host reproduction, fitness and mating behavior. Although our understanding of the Wolbachia-associated effects on hosts is rapidly expanding, our knowledge of the host factors that mediate Wolbachia dynamics is rudimentary. Here, we explore the interactions between Wolbachia and its host, the two-spotted spider mite Tetranychus urticae Koch. Our results indicate that Wolbachia induces strong cytoplasmic incompatibility (CI), increases host fecundity, but has no effects on the longevity of females and the mating competitiveness of males in T. urticae. Most importantly, host mating pattern was found to affect Wolbachia density dynamics during host aging. Mating of an uninfected mite of either sex with an infected mite attenuates the Wolbachia density in the infected mite. According to the results of Wolbachia localization, this finding may be associated with the tropism of Wolbachia for the reproductive tissue in adult spider mites. Our findings describe a new interaction between Wolbachia and their hosts.     

酪蝇的羽化节律和交配行为

【目的】为了明确酪蝇Piophila casei的羽化节律和交配行为,为深入研究酪蝇的性信息素打下基础。【方法】在室内条件[温度(27±1)℃,光周期L︰D=14︰10]下观察了酪蝇的羽化节律和交配行为。【结果】结果表明,化蛹后第8天成虫羽化最多,羽化高峰期出现在由暗期进入光期的前后,雄虫略早于雌虫羽化。观察结果表明,当雄虫接近蛹壳时会表现出"触摸"、"经过"、"扑"、"抱握"等动作,而雌虫仅有"触摸"、"抱握"的动作。当遇到同性成虫或者已交配雌虫时,雄虫会"追逐"对方,当接触到对方时雄虫会表现出"抱握"、"触摸"动作以进行识别,被识别雄虫会"翘起腹部末端"使对方离开,已交配雌虫则猛烈"摆动身体"使对方离开。当遇到处女雌虫时,雄虫跳跃到雌虫背上迅速进行交配。雌虫羽化后1 min即可交配,雄虫羽化后30 min可交配,交配平均持续时间10 min。雄虫间会有交配竞争行为发生,后来者会跳跃到先来者背上,向下弯曲腹部试图越过先来者与雌虫交配。【结论】酪蝇的羽化高峰出现在进入光期前后。成虫对蛹壳有显著趋性,雄虫主动追逐识别异性,较雌虫表现出更强烈的交配欲望。     

Indirect evidence that guarded quiescent deutonymph females invest energy to attract conspecific males in the Kanzawa spider mite (Acari: Tetranychidae)?

Because only the first mating results in fertilization in Tetranychus kanzawai (Acari: Tetranychidae), adult males guard quiescent deutonymph females (i.e., precopulatory mate guarding). A previous study reported that quiescent deutonymph females guarded by a male attract more conspecific males than solitary females and then hypothesized that guarded females release more chemical signals than solitary ones to attract males. Quiescent deutonymph females do not feed. If the hypothesis is appropriate, guarded females should invest energy in attracting males at the expense of investment in other activities, such as egg production. Therefore, we compared oviposition rates immediately after adult emergence between guarded females and solitary females. On the first day, the oviposition rate of guarded females was lower than that of solitary females. On the second day, however, there was no significant difference between female groups. These results suggest that guarded females invest energy in activities other than egg production before adult emergence and that the energetic cost is easily recoverable. We believe that our finding indirectly supports the hypothesis that guarded females release more chemical signals than solitary females to attract conspecific 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.     

Reproductive Biology and Behavior of Rhabditis pellio, (Schneider) (Rhabditida: Rhabditidae)

Laboratory studies were conducted on the mating behavior of Rhabditis pellio males and females, which were maintained on a culture of Flavobacterium sp. bacteria isolated from earthworms. The mean time that elapsed between first contact of the sexes and their ultimate separation was 23.2 min. However, only 5.0 min were required for copulation (the interval during which male spicules were inserted into the female vagina). Three-day-old females that were permitted to mate once on their first day of adult life produced only one-third as many larvae as did females that were permitted unlimited mating. However, the longevity of females was found to decrease with an increase in the number of matings. Both males and females that were permitted to mate daily produced the greatest number of offspring when they were 4 days old. When the initiation of mating was delayed beyond their third day of life, the number of larvae produced by females decreased. In approximately one-half of the copulations, males failed to inseminate their female partners.