Behaviour and fitness of γHCH/dieldrin resistant and susceptible female Anopheles gambiae and An.stephensi mosquitoes in the absence of insecticide

The effects of gamma HCH/dieldrin resistance genes on various fitness components of mosquito larvae and adult females in the absence of insecticide were investigated in backcrossed strains of Anopheles gambiae Giles and An.stephensi Liston. Among larvae, heterozygotes (RS) developed slightly but significantly faster than homozygotes for resistance (RR) or susceptibility (SS). The lifetime fecundity of RR females in population cages was only half to two-thirds that of SS and RS females despite similar longevities; several reasons were identified: RR gravid females were less responsive to oviposition-site stimuli, their spontaneous activity--as measured in an acoustic actograph--was only half that of SS or RS females, and RR females produced fewer eggs per unit bloodmeal. When inseminated females were recorded in LD 12:12, RR were again less active than SS or RS. When the lighting was switched to a regime simulating full-moonlight, the activity pattern of SS and RS changed and they flew for longer periods. In contrast, the activity of RR females was the same in LD 12:12 as in 'moonlight'. In a test simulation of potential predation, RR mosquitoes took to flight least readily. All component tests on adult females therefore point to RR as being the least fit of the three genotypes. The behavioural tests suggest that resistance has raised the response threshold of RR females to diverse stimuli. A possible physiological mechanism underlying RR behaviour is that a change in the cyclodiene receptor on the chloride channels has increased their permeability to chloride ions, causing hyper-inhibition of the nervous system.     

Changes in the circadian flight activity of the mosquito Anopheles stephensi associated with insemination, blood-feeding, oviposition and nocturnal light intensity

Abstract. The circadian activity patterns of Anopheles stephensi Liston were examined before and after mating and throughout the gonotrophic cycle. In LD 12:12h, males and virgin females are active at dusk. After insemination, females are active at dusk and also for short bursts during much of the night. After blood-feeding, inseminated females are inactive for two nights. On becoming gravid, these females become highly active at dusk and, to a lesser extent, are also active later in the night. In contrast, blood-feeding and egg maturation have minimal effects on the activity pattern of virgin females, who continue to fly only during dusk periods, i.e. virgins continue with the activity pattern which would most likely lead to mating encounters. After oviposition, parous females resume the activity pattern characteristic of inseminated nulliparous females. In a light regime which simulates moonlit nights and normal days, inseminated females are more active overall and flight-burst durations are much longer than in LD 12:12h.     

Flight activity of insecticide resistant and susceptible Anopheles stephensi mosquitoes in actograph chambers lined with malathion, γ HCH or dieldrin

The activity and resting behaviour of resistant and susceptible Anopheles stephensi Liston were recorded in acoustic actograph chambers lined with residual deposits of malathion, dieldrin or gamma HCH. In gamma HCH-treated flight chambers, SS and RS mosquitoes became active only after picking up lethal doses of insecticide, which explains why few SS and RS mosquitoes survive release into gamma HCH-treated experimental huts. Similar results were obtained in flight chambers treated with dieldrin; however, more mosquitoes would be expected to survive dieldrin under field conditions because resistance to this insecticide is greater than to gamma HCH. Mosquitoes in contact with malathion showed a three-phase activity pattern: an initial active phase, an inactive phase, and hyper-activity/convulsions. Initial activity or irritability was especially pronounced in SS and RS but absent in RR mosquitoes. Whether or not irritability would protect RS mosquitoes from malathion would probably depend on the ratio of sprayed to unsprayed surfaces in treated huts.     

Assortative mating in mixed swarms of the mosquito Anopheles gambiae s.s. M and S molecular forms,in Burkina Faso,West Africa

The molecular form composition of Anopheles gambiae Giles s.s. (Diptera: Culicidae) mating swarms and the associated mating pairs (copulae) were investigated during two rainy seasons (July to October, 2005 and July to November, 2006) in the villages of Soumousso and Vallée du Kou (VK7). Although the habitats of these villages differ markedly, sympatric populations of M and S molecular forms of An. gambiae s.s. occur in both places periodically. The main aim was to assess the degree to which these molecular forms mate assortatively. In Soumousso, a wooded savannah habitat, the majority of swarm samples consisted of only S‐form males (21/28), although a few M‐form males were found in mixed M‐ and S‐form swarms. In VK7, a rice growing area, the majority of swarm samples consisted of only M‐form males (38/62), until October and November 2006, when there were nearly as many mixed‐form as single‐form swarms. Overall, ～60% of M‐ and S‐form swarms were temporally or spatially segregated; the two forms were effectively prevented from encountering each other. Of the remaining 40% of swarms, however, only about half were single‐form and the rest were mixed‐form. Of the 33 copulae collected from mixed‐form swarms, only four were mixed‐form pairs, significantly fewer than expected by random pairing between forms (χ² = 10.34, d.f. = 2, P < 0.01). Finally, all specimens of inseminated females were of the same form as the sperm contained within their spermatheca (n = 91), even for the four mixed‐form copulae. These findings indicate that assortative mating occurs within mixed‐form swarms, mediated most probably by close‐range mate recognition cues.     

The relative role of male vs. female mate choice in maintaining assortative pairing among discrete colour morphs

Mate choice has important evolutionary consequences because it influences assortative mating and the level of genetic variation maintained within populations. In species with genetically determined polymorphisms, nonrandom mate choice may affect the evolutionary stability and maintenance (or loss) of alternative phenotypes. We examined the mating pattern in the colour polymorphic Gouldian finch (Erythrura gouldiae), and the role of mate choice, both female and male, in maintaining the three discrete head colours (black, red and yellow). In both large captive and wild populations, Gouldian finches paired assortatively with respect to head colour. In mate choice trials, females showed a strong preference for mates with the most elaborate sexually dimorphic traits (i.e. more chromatic UV/blue plumage and longer pin-tail feathers), but did not discriminate assortatively. Unexpectedly, however, males were particularly choosy, associating and pairing only with females of their own morph-type. Although female mate choice is generally invoked as the major selective force maintaining conspicuous male colouration in sexually dichromatic species, and is typically thought to drive nonrandom mating, these findings suggest that mutual mate choice and male mate choice in particular, are an important yet neglected component of selection.     

Testing the role of coadapted genes versus bet-hedging for mating strategies in colour polymorphic pygmy grasshoppers

Recent investigations of mate choice indicate that the genetic effect of sires on offspring fitness may depend on the interaction between maternal and paternal genotypes and the environmental conditions experienced by the offspring. Alternative colour morphs of the pygmy grasshopper, Tetrix subulata , represent ecological strategies that differ in body size, life history, thermoregulatory behaviour, and habitat selection. The hypothesis that selection promotes behaviours maintaining coadapted gene complexes predicts individuals to mate assortatively with respect to colour morph. On the other hand, the bet-hedging hypothesis predicts that the temporal variability of the environment inhabited by these animals may select for disassortative mating behaviour resulting in heterogeneous offspring. To distinguish between these competing hypotheses, we investigated mating behaviours using dual-choice experiments. Our results were not in agreement with the prediction of assortative mating but suggest instead that matings were random with regard to colour morph. Polyandry was common, and females mated with the second male regardless of whether the first mating was assortative or disassortative. Polyandry also was equally frequent among females in triads in which the two males belonged to different colour morphs as in triads where both males belonged to the same colour morph. A field experiment confirmed that polyandry occurred also among free-ranging individuals, and uncovered variation in mating success among male colour morphs, probably due to indirect effects of coloration on activity or habitat use. The consequences of this random and polyandrous mating strategy for the evolutionary dynamics of the colour polymorphism remain to be explored.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 491–499.     

Size‐Dependent Male Mate Preference and its Association with Size‐Assortative Mating in a Mangrove Snail,Littoraria ardouiniana

In many animals, body size plays a crucial role in mating success in the context of competition and preference for mates. Increasing evidence has shown that male mate preference can be size‐dependent and, therefore, an important driver of size‐assortative mating. To test this theory, mate choice experiments were performed during the three consecutive stages of mating behaviour, namely trail following, shell mounting and copulation, in the dioecious mangrove snail, Littoraria ardouiniana. These experiments identified two possible forms of size‐dependent male mate preference which could contribute to the formation of size‐assortative mating in these snails. Firstly, whereas small males were unselective, large males were selective and preferred to follow mucus trails laid by large females. Alternatively, the results can also be interpreted as all males were selective and adopted a mating strategy of selecting females similar to, or larger than, their own sizes. Both small and large males also copulated for longer with large than with small females, and this was more pronounced in large males. When two males encountered a female, they engaged in physical aggression, with the larger male excluding the smaller male from copulating with the female. This study, therefore, demonstrated that size‐dependent male mate preference may, along with male–male competition, play an important role in driving size‐assortative mating in these mangrove snails, and this may also be the case in other species that exhibit male mate choice.     

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.     

The effects of exposure to seaweed on willingness to mate, oviposition, and longevity in seaweed flies

Abstract  1. Large male seaweed flies (Diptera: Coelopidae) are more likely to mate than smaller males. This is due to sexual conflict over mating, by which females physically resist male attempts to copulate. In some species, large males are simply more efficient at overpowering female resistance.
2. Female reluctance to mate is likely to have evolved due to the costs of mating to females. In many dipterans, males manipulate female behaviour through seminal proteins that have evolved through sperm competition. This behavioural manipulation can be costly to females, for example forcing females to oviposit in sub-optimal conditions and increasing their mortality.
3. Previous work has failed to identify any ubiquitous costs of mating to female coelopids. The work reported here was designed to investigate the effects of exposure to oviposition sites ( Fucus algae) on the reproductive behaviour of four species of coelopid. Algae deposition in nature is stochastic and females mate with multiple males in and around oviposition sites. Spermatogenesis is restricted to the pupal stage and there is last-male sperm precedence. It was predicted that males would avoid wasting sperm and would be more willing to mate, and to remain paired with females for longer, when exposed to oviposition material compared with control males. Females were predicted to incur longevity costs of mating if mating increased their rate of oviposition, especially in the presence of algae.
4. The behaviour of males of all four species concurred with the predictions; however mating did not affect female receptivity, oviposition behaviour, or longevity. Exposure to algae induced oviposition and increased female mortality in all species independently of mating and egg production. The evolutionary ecology of potential costs of mating to female coelopids are discussed in the light of these findings.     

Assortative mating by multiple ornaments in northern cardinals (Cardinalis cardinalis)

In positive assortative mating, individuals of similar phenotypemate together more frequently than expected by chance. Assortativemating by a variety of qualities, including ornamentation, iswell documented in birds. Studies of assortative mating by ornamentshave focused on single, highly conspicuous ornaments, but manyspecies of birds possess multiple ornaments in both sexes. Wecompared ornament expressions between mates of northern cardinals(Cardinalis cardinalis) to determine if assortative mating occurredby one or more of the four ornaments displayed by both sexes.All cardinals possess tall head crests and red-orange bills.In addition, males have black face masks and entirely red bodyplumage, whereas females have blackish face masks and red underwingcoverts. We predicted that cardinals mate assortatively by plumagecolor because red plumage expression has been shown to indicatequality in both sexes. We found that cardinals mate assortativelyby plumage and bill color, the two ornaments colored by carotenoidpigments, but not by mask expression or crest length. Whetherthis mating pattern arises by mutual mate choice or intrasexualselection is not known.     

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。     

Polyandry enhances offspring viability with survival costs to mothers only when mating exclusively with virgin males in Drosophila melanogaster

A prominent hypothesis for polyandry says that male–male competitive drivers induce males to coerce already‐mated females to copulate, suggesting that females are more likely to be harassed in the presence of multiple males. This early sociobiological idea of male competitive drive seemed to explain why sperm‐storing females mate multiply. Here, we describe an experiment eliminating all opportunities for male–male behavioral competition, while varying females’ opportunities to mate or not with the same male many times, or with many other males only one time each. We limited each female subject's exposure to no more than one male per day over her entire lifespan starting at the age at which copulations usually commence. We tested a priori predictions about relative lifespan and daily components of RS of female Drosophila melanogaster in experimental social situations producing lifelong virgins, once‐mated females, lifelong monogamous, and lifelong polyandrous females, using a matched‐treatments design. Results included that (1) a single copulation enhanced female survival compared to survival of lifelong virgins, (2) multiple copulations enhanced the number of offspring for both monogamous and polyandrous females, (3) compared to females in lifelong monogamy, polyandrous females paired daily with a novel, age‐matched experienced male produced offspring of enhanced viability, and (4) female survival was unchallenged when monogamous and polyandrous females could re‐mate with age‐ and experienced‐matched males. (5) Polyandrous females daily paired with novel virgin males had significantly reduced lifespans compared to polyandrous females with novel, age‐matched, and experienced males. (6) Polyandrous mating enhanced offspring viability and thereby weakened support for the random mating hypothesis for female multiple mating. Analyzes of nonequivalence of variances revealed opportunities for within‐sex selection among females. Results support the idea that females able to avoid constraints on their behavior from simultaneous exposure to multiple males can affect both RS and survival of females and offspring.     

Sperm stratification and paternity success in red flour beetles

Abstract. When females are inseminated by multiple males, male paternity success (sperm precedence) is determined by the underlying processes of sperm storage and sperm utilization. Although informative for many questions, two-male sperm competition experiments may offer limited insight into natural mating scenarios when females are likely to mate with several males. In this study, genetic markers in Tribolium castaneum are used to trace paternity for multiple sires, and to determine whether displacement of stored sperm that occurs after a third mating equally affects both previous mates, or if fertilizations are disproportionately lost by the female's most recent mate. For 20 days after triple-matings, first males retain significantly higher paternity success (relative to first male paternity in double-matings) compared with second males. These results demonstrate that when females remate before sperm mixing occurs, sperm stratification results in differential loss of sperm from the most recent mate. This study provides insight into the mechanisms underlying sperm precedence in a promiscuous mating system, and suggests that T. castaneum females could limit paternity success of particular mates by remating with more highly preferred males.     

Switchover to the mated state by spermathecal activation in female Anopheles gambiae mosquitoes

Both Aedes aegypti and Anopheles gambiae mosquitoes undergo physiological and behavioral changes after the females mate, but unlike Ae. aegypti, the mated state in An. gambiae is not attained through the action of male accessory gland substances. Experiments in which the spermathecae of mated females were manipulated suggest that a spermatheca filled with sperm is responsible for triggering oviposition behavior. An estimated 48% of the previously mated An. gambiae females mated again when the interval between encounters with males was less than 24h. Unlike male Ae. aegypti that can inseminate up to seven females after their testes are removed, An. gambiae do not store sperm in the vas deferens and without testes would be unable to inseminate any females.     

Costs of Mate Guarding and Opportunistic Mating Among Wild Male Japanese Macaques

A trade-off relationship between mating and feeding effort is important when considering reproductive strategies of long-lived species. I compared the influence of male sexual activities, female mate-choice behaviors and the daily activity budget on male mating success among males in a group of wild Japanese macaques (Macaca fuscata yakui) on Yakushima Island. The 1st-ranking male, which had immigrated into the troop at this rank, more frequently approached peri-ovulatory females, spent more time grooming peri-ovulatory females and in mounting series and spent less time feeding than subordinate males did. The 1st-ranking male attained the highest mating success as a result of his high expenditure of time and energy in sexual behaviors directed toward peri-ovulatory females. Mating success of subordinate males did not relate to the amount of sexual effort, but instead to the frequency of female approaches, female rush toward males and the number of peri-ovulatory females within the group. The pattern of intermale competition shifted from nearly contest competition to scramble competition as the number of peri-ovulatory females in the group increased. Feeding time of subordinate males did not vary between the days when they copulated and the days when they did not. The findings demonstrate that mate guarding in the 1st-ranking male is a high-cost mating tactic, while opportunistic mating in subordinate males is a low-cost mating tactic. The differences in male mating tactics are probably related to male life history and to the formation of groups with a high socionomic sex ratio.     

Assortative Mating and Lack of Temporality Between Corn and Rice Strains of Spodoptera frugiperda (Lepidoptera,Noctuidae) from Central Colombia

Spodoptera frugiperda is a Neotropical moth that has diverged into “corn” and “rice” strains. In the US, prezygotic isolation studies have shown that both populations mate assortatively. In addition, recent studies have demonstrated that mating by the same strain individuals is enhanced by an allochronic shift in mating activity and male pheromones are important during courtship. In Colombia, studies on mate choice have never been performed previously, although earlier analyses made in populations from Central Colombia showed no significant differences in time of first mating and copula duration between the strains. Here, we performed multiple choice experiments using a tetrad design composed of individuals of the corn and the rice strains. We found that corn strain females rarely mate with rice strain males, but rice strain females mate with both strains of males. In addition, no temporal isolation was found. A ML (maximum likelihood) approach was used to discriminate between mating propensity and mate choice behaviors in S. frugiperda strains. This approach showed that mating propensity of corn strain males is three times greater than rice strain males. In contrast, in females, the propensity of mating was slightly higher for the corn strain. Finally, the isolation index between the corn and the rice strains from Colombia produced a value of I?=?0.33. Our results suggest that prezygotic isolation at the behavioral and temporal levels differ between the US and Colombia. Moreover, in the US temporal isolation appears to have an important role in behavioral isolation but in Colombia temporality is not necessary to reduce encounters between S. frugiperda strains.     

Effects of permethrin at different temperatures on pyrethroid-resistant and susceptible strains of Anopheles

The influence of temperature (16, 22, 28, 37 degrees C) on effects of permethrin was investigated for susceptible and pyrethroid-resistant strains of the mosquitoes Anopheles gambiae and An. stephensi (Diptera: Culicidae). Young unfed female adult mosquitoes were exposed to 0.25% permethrin test papers or to polyester netting treated with permethrin 500mg a.i./m2. The time to 50% knockdown (KT50) declined as temperature increased, i.e. there was a positive temperature coefficient of this effect of the pyrethroid. Resistance ratios (comparing KT50 values) between resistant and susceptible An. stephensi ranged between 2.5 and 4.4 at the different temperatures. Comparative tests of pyrethroid tolerance of different strains would be valid over the 22-28 degrees C range but, when using a discriminating dose to detect resistance, more precise temperature control is desirable. Mortality 24h after exposure to 0.25% permethrin of both susceptible and resistant strains of An. stephensi showed a negative correlation with temperature between 16 and 22 degrees C and a positive correlation at higher temperatures. In An. gambiae, however, the correlation was positive over the whole range. Irritancy of permethrin-treated netting to Anopheles females (measured as time lapse until first flight take-off, and the number of take-offs during 7.5 min exposure) was positively correlated with temperature in all four strains and was much greater for the susceptible than the resistant strains.     

Males,but not females,perform strategic mate searching movements between host plants in a leaf beetle with scramble competition polygyny

Mate searching is assumed to be performed mostly by males, but when females benefit from multiple mating or are under risk of failing to mate, they may also perform mate searching. This is especially important in scramble competition polygynies, in which mate searching is the main mechanism of mate competition. Typically, more mobile individuals are expected to achieve higher mating success because mobility increases their probability of finding mates. If we assume individual movements are mainly explained by mate searching in scramble competition polygynies, we can investigate searching strategies by asking when individuals should leave their location and where they should go. We hypothesize that individuals will leave their locations when mating opportunities are scarce and will seek spatially close sites with better mating opportunities. We tested these hypotheses for males and females of Leptinotarsa undecimlineata, a leaf beetle with scramble competition polygyny in which both sexes are promiscuous. Individuals mate and feed exclusively on Solanum plants, and thus, individual movements can be described as switches between plants. Females were less likely than males to leave isolated plants, and both males and females moved preferentially to neighboring plants. Males were more likely to leave when the local number of females was low, and the number of males was high. They moved to plants with more females, a behavior consistent with a mate searching strategy. Females were more likely to move to plants with fewer males and many females, a behavior consistent with male harassment avoidance. Strategic movement is widely considered in foraging context, but seldom in a mate searching context. Considering that selection to minimize searching costs, maximize mating success, and minimize harassment may be ubiquitous in nature, we argue that strategic movements by mate searching individuals are likely to occur in many species.     

The dynamics of sexually antagonistic coevolution and the complex influences of mating system and genetic correlation

Sexual conflict has been proposed to be a mediator of speciation but recent theoretical work, as well as empirical studies, suggests that sexual conflict may also be able to prevent speciation and to preserve genetic polymorphism within a species. Here, we develop a population genetic model and study the effects of sexual conflict in a polymorphic population. The morphs mate assortatively based on different sexually antagonistic traits and females are assumed to suffer a cost when the proportion of matching males is high. We consider the model in two different mating systems; promiscuity and polygyny. Our results show that genetic polymorphism may be maintained through negative frequency dependent selection established by assortative mating and female conflict costs. However, the outcome significantly differs between mating systems. Furthermore, we show that indirect selection may have profound effects on the evolutionary dynamics of a sexual conflict.     

A survival and reproduction trade-off is resolved in accordance with resource availability by virgin female mosquitoes

The first 2-4 days after an Anopheles gambiae female mosquito emerges are critical to her survival and reproductive success. Yet, the order of behavioural events (mating, sugar feeding, blood feeding) during this time has received little attention. We discovered that among female cohorts sampled from emergence, sugar feeding had a higher probability than blood feeding of occurring first, and mating rarely occurred before a meal was taken. The night after emergence, 48% of females fed on sugar in mesocosms, and 25% fed on human blood; in the absence of sugar, 49% of females fed on human blood. After 5 days, 39% of the sugar-supplied females had blood fed and mated, and were fructose negative, whereas only 8% of the sugar-denied females had both blood fed and mated by this time. The model that best explained the transitions suggests that females made use of two distinct behavioural pathways, the most common one being to sugar-feed, then mate, and then seek blood. Other females sought blood first, then mated, and forwent a sugar meal. Lipid levels were higher in females with access to sugar than in females without access to sugar, particularly for those in later gonotrophic stages, while glycogen levels in the sugar-supplied group were higher throughout. In single-night experiments with females having had access to sucrose since emergence, those given a blood meal 1 day before spending a night with males had higher insemination rates than those not receiving the blood meal. These results indicate that the trade-off between survival and immediate reproduction is resolved by young adult females in accordance with availability of resources and gonotrophic state.