Evidence for a mounting sex pheromone in the brown ear tick Rhipicephalus appendiculatus,Neuman 1901 (Acari: Ixodidae)

The presence of a mounting sex pheromone was demonstrated on the surface of fed female Rhipicephalus appendiculatus. This pheromone, which is present on the female cuticle, allows the male to recognise the female. The pheromone was removed by cleaning the female in hexane, resulting in the loss of male mating behaviour in in vitro experiments. Male mating behaviour was resumed when extract made from fed female cuticle was replaced on cleaned females. When the extract was transferred to innanimate objects typical male mating behaviour was released. Preliminary chemical analyses indicated that the active component of the extract was contained in the sterol ester fraction of the extract.     

松墨天牛是否存在雌性接触信息素?

将松墨天牛MonochamusalternatusHope的身体不同部位切除之后的交配试验显示,分别剪去头部、鞘翅以及前胸部后仍然可以正常交配,用雄虫的腹部代替雌虫的腹部后,仍然能够引起雄虫的交配趋势动作,但不能完成真正交配,因此推测雌性接触信息素如果存在,主要集中在雌虫身体后半部,尤其是中后胸部位。但是冻死后的雌虫不再引起雄虫的交配行为,而且冻死的雌虫与活雌虫同时放入培养皿中,雄虫喜欢与活雌虫交配,不选择冻死的雌虫,因此推测视觉在其松墨天牛的交配过程中扮演着重要的角色,而接触性信息素有可能并不存在。将雄虫的触角分别从第5~6节和第1节剪去的试验结果显示,雄虫仍然可以很快地找到雌虫,发生交配行为。这有2种可能的解释一是雄虫不仅仅靠触角来感受雌虫的接触信息素,头部下颚须和下唇须上或者足上也有可以感受信息素的感受器;二是雌虫不存在接触信息素,雄虫通过视觉就可以找到对方,完成交配过程。浸泡致死的雌虫不再引起雄虫的交配行为,并且将正己烷和乙醚的浓缩液滴加在浸泡致死的雌虫以及玻璃棒和鹅卵石的生测试验结果显示,都没有引起雄虫的交配行为。这进一步说明,松墨天牛可能并不存在体表接触信息素。性成熟后的松墨天牛雌雄成虫是靠衰弱寄主的强烈的挥发性气味聚集到寄主树干上,在树干这个较为狭小的环境里,有可能是通过视觉找到配偶,完成交配过程。     

Observations on the mating behavior of the alfalfa blotch leafminer,Agromyza frontella (Rondani) (Diptera: Agromyzidae), and evidence of a female sex pheromone

The mating behavior of Agromyza frontella was studied under laboratory conditions. Adults were able to mate on the day of emergence, with no evident periodicity throughout the photophase. The host plant was essential for mating to occur, its presence affecting female receptivity rather than male copulatory behavior. Males generally entered a stationary phase once in the proximity of a female, before undertaking a final approach. This stationary behavior frequently resulted in male aggregations around a female, and under such conditions males exhibited a characteristic wing vibrating behavior. As male wing vibration was not an essential behavior for successful mating, and rarely occurred during male encounters in the absence of females or when only one male was near a female, it was considered as being primarily a male-male signal. The majority of females that mated exhibited an ovipositor pumping behavior that stimulated the male approach. However, such behavior was not essential to attract mates, as dead females elicited the entire sequence of male mating behavior. This suggested the presence of a cuticular sex pheromone, as reported for other species of higher Diptera. Whole virgin female (<24- h or 3- day-old) hexane extracts applied to male cadavers increased the time males spent on the plant, the number of contacts with the treated cadavers, the incidence of attempted copulations, and the wing vibrating behavior between males compared with controls using untreated cadavers. The results obtained indicate that females control copulation in A. frontella and that both semiochemical and visual cues are important in eliciting male mating behavior.     

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.     

Effects of hexane extract of Lantana camara leaves on reproductive bioactivities of Dysdercus koenigii Fabricius (Heteroptera: Pyrrhocoreidae)

Effects of hexane extract of Lantana camara leaves were investigated on reproductive bioactivities of Dysdercus koenigii by assessing mating behavior, oviposition behavior and fertility of the adults emerged from treated fifth instar nymphs. Leaves of L. camara were extracted in hexane by ‘cold extraction method’. The fifth instar nymphs were treated with the extract by ‘dry film residual method’ for 24 h and the adults emerged from these nymphs were used for the experiments. The results indicated that treated males showed altered courtship behaviour, lesser number of mounting attempts and took relatively more time to mate than the normal males. The treated females, very often, exhibited mating refusal and non-receptive behaviour towards the courting males. This led to decrease in percent successful mating. Also, the mating in the treated insects frequently got disrupted and terminated prior to insemination. The treatment of the females with the extract resulted in alteration of their oviposition behaviour. Consequently, the treated females laid lesser number of egg batches and total number of eggs in their life span. Although the eggs laid by the treated females were fertile, the percent hatchability was lesser than normal. The results signify that hexane leaf extract of L. camara possesses phytochemicals, which adversely impaired the reproductive bioactivities of D. koenigii. Therefore, some of these compounds individually or synergistically can be employed in integrated pest management of D. koenigii by hampering its reproductive potential.     

Cuticular lipids and odors induce sex-specific behaviors in the male cricket Gryllus bimaculatus

Male crickets display sex-specific (e.g., mating and agonistic) behaviors towards conspecific individuals. One of the key signals for these behaviors is the chemical substance on the cricket body surface. In the present study, we analyzed female and male cuticular substances in behavioral assays. Antennal contact stimulation using female forewings elicited a mating behavior in males, while that using male forewings elicited an agonistic behavior in males. Thin-layer-chromatographic and other techniques analysis showed that saturated cuticular lipids were present in both female and male cuticles and that unsaturated lipids were present only in the male cuticle. Filter papers soaked with saturated or unsaturated cuticular lipids were applied to antennae of male crickets. Males showed mating behavior in response to stimulation with saturated lipids from both females and males but showed avoidance behavior in response to stimulation with male unsaturated lipids. Because cuticular lipids did not induce agonistic behavior in males, we collected odors from male crickets and found that these odors induced agonistic behavior in males. Therefore, we concluded that the key signals for mating, avoidance and agonistic behaviors of male crickets are comprised of at least three different components, saturated and unsaturated cuticular lipids and male odors, respectively.     

Visual and Olfactory Female-Borne Cues Evoke Male Courtship in the Aphid Parasitoid Aphidius colemani Viereck (Hymenoptera: Braconidae)

We investigated the courtship and mating behavior of the pan-tropical polyphagous endoparasitoid Aphidius colemani Viereck. The courtship and mating displays, the magnitude of male-male sexual approaches and the role of female-borne cues evoking male courtship behavior were quantified. The sequence of events leading to copulation in this parasitoid did not differ from that found for other braconids. Females refused to copulate more than once. Same-sex courtships were observed among males and their possible role in an adaptive context is discussed. Olfactory female-borne cues played a key role in eliciting the courtship responses of males. Males were attracted by freshly dead females, but not by dead females soaked in hexane, nor by visual cues from females alone. Intense male wing fanning behavior was elicited by crushed abdomens of virgin females, suggesting that the female abdomen is the source of a short-distance pheromone crucial in evoking male courtship. Further studies are required to clarify the exact nature of the chemicals involved.     

Mating behavior and contact pheromones of Batocera horsfieldi (Hope) (Coleoptera: Cerambycidae)

The mating behavior of Batocera horsfieldi (Hope) was observed in the laboratory. The results showed that copulation of this longhorned beetle consisted of three phases: (i) encountering and pair‐bonding; (ii) mating attempt and ejaculation; and (iii) post‐copulatory guarding. Frozen females, with cuticular hydrocarbons stripped by hexane extraction, showed no attraction for males. Reapplying the solvent extract of frozen females to both washed dead males and females caused mating attempts by males, confirming that cuticular hydrocarbons (contact sex pheromones) played an important role in recognition of females by male B. horsfieldi.     

Mating behavior of Aegla platensis (Crustacea, Anomura, Aeglidae) under laboratory conditions

The mating behavior of several decapod crustaceans has been extensively studied; however, this aspect of anomuran biology is still poorly known in some groups. Aeglids are the only anomurans inhabiting freshwaters, and the mating behavior of the species in this family is unknown. We provide the first account of the mating behavior of an aeglid, Aegla platensis, under laboratory conditions. The precopulatory phase was characterized by male agonistic display, male approach, and courtship. Males exhibited the agonistic display toward immature and mature females, but only physiologically mature females allowed males to approach. Male approach led to display of courtship behaviors (body vibration, thrust, body lifting, and abdomen flapping). During the copulatory phase, males and females touched each other with the antennae (antennae touch), and males positioned themselves beneath the females (supine position). Although sperm transfer was not directly observed, a “white mass” was detected among oocytes in the female abdominal chamber shortly after some copulations. Finally, in the postcopulatory phase, males guard females during the process of egg attachment. Despite their morphological similarities with other anomurans, the mating behavior of aeglids seems to be unique, and the freshwater environment appears to have an important role in driving these differences.     

Relatively large males lower reproductive success in female zebrafish

Females can adjust their reproductive effort in relation to their partner’s perceived fitness value. In zebrafish (Danio rerio), large males are typically preferred mating partners. However, females have been observed to reduce their reproductive output with exceptionally large males but it remains unknown whether it is due to sexual harassment or aggressive behavior to establish and maintain dominance. Here, we study the association between relative male size, sexual harassment and dominance behavior, female stress status (stress behaviors and whole-body cortisol concentration), and reproductive success during a 4 day spawning trial. We found female cortisol to correlate negatively with female body size and positively with female dominance behavior. However, male and female behavior as well as female cortisol level were not related to relative male size. Females mating with relatively large males produced more and most of their eggs during the first spawning day, while females with smaller males produced few eggs during the first day but then increased egg production. Despite females produced more eggs when mating with relatively larger males, their eggs had substantially lower fertilization rates compared to females mating with relatively smaller males. Hence, overall, the reproductive fitness was lowest when females mated with a relatively large male. These findings could help to explain the maintenance of male size variation under natural conditions.     

Mate Choice and Spawning Success in the Fighting Fish Betta splendens: the Importance of Body Size, Display Behavior and Nest Size

Courtship displays should be exaggerated enough to attract mates and yet tempered so as not to deter them. We tested this hypothesis in the fighting fish Betta splendens by studying courtship displays and body size and their relationships with male parental quality and female fecundity, as well as the effects of display behavior and body size on mate choice decisions and spawning success. Because of their high degree of parental investment, males are expected to be discriminating in their choice of mates. Males who displayed more frequently built larger nests, a measure of parental quality, but larger males did not. When females were paired with males with high display rates, however, the pair had fewer eggs in their nest, even when accounting for female body mass. In a mate choice test using computer‐generated male stimuli that differed only in display behavior, females showed no preferences for displaying males vs. non‐displaying males, or for males with higher display rates vs. lower display rates. In similar tests in which the computer‐generated males differed only in size, females preferred larger males, but also preferred males that differed with respect to body size (negative assortative mating). Males preferred computer‐generated females that performed courtship displays over non‐displaying females, but showed no preferences for female body size. Neither a female's body size nor her display behavior was a significant predictor of her fecundity as estimated by the number of eggs released during spawning. Thus, our results suggest that female B. splendens must balance male parental quality (nest size) with the risk of potentially disruptive or dangerous behavior during spawning, and that females may minimize these risks through negative size‐assortative mating. Female display behavior, while unrelated to fecundity in our study, may attract males because it indicates reproductive readiness or serves a species‐recognition function.     

Field Observations on the Mating Behavior of Aclees sp. cf. foveatus Voss (Coleoptera: Curculionidae), an Exotic Pest Noxious to Fig Orchards

Aclees sp. cf. foveatus Voss is a new and exotic pest recently reported as responsible for severe damages to Italian fig orchards and nurseries. Investigating its mating behavioral sequence could help to unravel mate assessment and choice dynamics of this pest, adding useful information to build-up behavior-based control strategies. In this study, we observed A. sp. cf. foveatus mating behavior under field conditions. The mating behavior is divided into precopulatory, copulatory and postcopulatory phases. Simultaneous antennal tapping and rostral rubbing acts performed by the male on female’s body raises male’s chances to copulate, while males that performed only antennal tapping on the partner’s body did not achieved mating success. Copulation attempt phase lasted 37.80?±?7.65 s. Copulation lasted 1116.00?±?229.73 s and was followed by a motionless postcopulatory phase (8.40?±?1.21 s). Same-sex sexual interactions were observed among A. sp. cf. foveatus males. Male-male sexual interactions lasted about 140.17?±?19.45 s and were composed of antennal tapping and rostral rubbing followed by one or more mounting attempts. Further research is ongoing to investigate the exact nature of male- and female-borne olfactory and tactile cues routing mate recognition in A. sp. cf. foveatus.     

Female choice of sites versus mates in a coral reef fish,Thalassoma bifasciatum

To understand the dynamics of sexual selection, one needs to identify the basis of female choice in mating. Daily censuses showed that most spawns of individual female bluehead wrasses, Thalassoma bifasciatum, occurred at one of several available mating sites, either in a group-spawning aggregation of small males or with a large territorial male. Most females were not located near their principal mating site during the non-spawning period of the day. Females were thus exposed to other males and sites and should have had the opportunity for choice. When a resident territorial male disappeared or was removed experimentally from a mating site, females did not alter their fidelity to that site. Furthermore, manipulations showed that when a territorial male changed the location of his mating activity to an adjacent site, females did not follow him but continued to mate at their customary site. Thus mating sites, rather than the males occupying them, appeared to be the objects of female choice. The bright coloration and courtship behaviour of larger males may demonstrate the safety of a particular site rather than indicating some aspect of male quality.     

Sexual Competition,Coercive Mating and Mate Assessment in the One‐Sided Livebearer,Jenynsia multidentata: Are They Predictive of Sexual Dimorphism?

We investigated the mechanisms of sexual selection operating on body size in the one‐sided livebearer (Jenynsia multidentata), a small fish characterized by male dwarfism. Mating in the one‐sided livebearer is coercive: males approach females from behind and try to thrust their copulatory organ at the female genital pore. Females counter males' mating attempts by either swimming away or attacking them. We tested the hypothesis that the components of sexual selection favouring small size in males (sexual coercion) were more effective than those favouring a large size (male competition and mate choice). When alone, small males had a significantly higher success in their mating attempts than large males. The proportion of successful attempts was also positively correlated with female size. When two males competed for the same female, the large male had a significant mating advantage over the small one. With a 1 : 1 sex ratio, the large‐male mating advantage vanished because each male tended to follow a different female. Large males, however, preferentially defended large females, thus compelling small males to engage with smaller, less fecund females. Males did not discriminate between gravid and non‐gravid females, but preferred mating with larger females. This preference disappeared when males were much smaller than the female, probably in relation to the risk for the male of being eaten or injured by the female. In a choice chamber, male‐deprived females that had their sperm storage depleted remained close to males and showed a preference for large individuals, a behaviour not observed in non‐deprived females. Nonetheless, when placed with males in the same aquarium, all females showed avoidance and aggression. Struggling may represent a way by which the female assesses the skill and endurance of males.     

Mate choice and sexual conflict in <Emphasis Type="Italic">Brachionus plicatilis</Emphasis> (Rotifera)

Male mate choice in Brachionus plicatilis is based on information from coronal receptors and contact chemoreception of a glycoprotein signal on the body surface of females. Male mating behavior is modulated by mating signal quality and strength, which determines male mating persistence and circling intensity. We probe the sensory abilities of males by better resolving their discrimination of female age. Males preferentially initiate mating with young females, but not too young. Males circle 0.2 h old females just hatched from resting eggs only about 1/2 as frequently as 12 h old females. Males are even more discriminating of females with whom they copulate, preferring 3 h old females significantly more than 0.2 h or 6 h old females. In contrast, males cannot distinguish between virgin females and those who have already copulated. Similarly, males circled and copulated with females hatching from resting eggs with equal frequency as those hatching from amictic eggs. The counterpoint to male mate choice is female resistance to mating. When males contact females, the females respond with one of four behaviors: no response, acceleration, foot flipping, or coronal retraction. In 65% of male–female encounters, there was no initial response by females. However, when males began circling females, females accelerated 11.1 times more often than when males were absent. The second type of evasive female behavior was foot-flipping, which tends to knock off males attempting to circle the female. In the presence of circling males, females performed foot-flipping behavior three times more often than in the absence of males. Coronal retraction, where they stop swimming and withdraw their corona, was observed less frequently than acceleration or foot-flipping, and there was no difference in the presence or absence of males. These data are interpreted in the context of sexual conflict, where the behaviors that optimize male and female fitness differ. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

Do female Nicrophorus vespilloides reduce direct costs by choosing males that mate less frequently?

Sexual conflict occurs when selection to maximize fitness in one sex does so at the expense of the other sex. In the burying beetle Nicrophorus vespilloides, repeated mating provides assurance of paternity at a direct cost to female reproductive productivity. To reduce this cost, females could choose males with low repeated mating rates or smaller, servile males. We tested this by offering females a dichotomous choice between males from lines selected for high or low mating rate. Each female was then allocated her preferred or non-preferred male to breed. Females showed no preference for males based on whether they came from lines selected for high or low mating rates. Pairs containing males from high mating rate lines copulated more often than those with low line males but there was a negative relationship between female size and number of times she mated with a non-preferred male. When females bred with their preferred male the number of offspring reared increased with female size but there was no such increase when breeding with non-preferred males. Females thus benefited from being choosy, but this was not directly attributable to avoidance of costly male repeated mating.     

Density affects mating mode and large male mating advantage in a fiddler crab

Fiddler crabs show two different mating modes: either females search and crabs mate underground in male burrows, or males search and crabs mate on the surface near female burrows. We explored the relationship between crab density, body size, the searching behavior of both sexes, and the occurrence of both mating modes in the fiddler crab Uca uruguayensis. We found that crabs change their mating mode depending on their size and crab density. Crabs mated mostly on the surface at low densities, and underground at high densities. The proportion of wandering receptive females but not courting males accounted for the variation in mating modes. This suggests that whether crabs mate underground (or on the surface) is determined by the presence (or absence) of searching females. We found that the change in the mating mode affected the level of assortative mating; males mating underground were bigger than those mating on the surface, suggesting active female choice. Given that fiddler crabs experience multiple reproductive cycles, they are prone to showing behavioral plasticity in their mating strategy whenever the payoffs of using different mating modes differ between reproductive events. Our results suggest that the incorporation of different levels of environmental variability may be important in theoretical models aimed at improving our understanding of the evolution of alternative mating tactics and strategies.     

Sperm competition and the significance of female multiple mating in the predatory miteParasitus fimetorum

Females of the predatory miteParasitus fimetorum (Gamasida; Parasitina) inhabiting animal manure indiscriminately copulate with many mates. The sperm competition between the males was estimated by electrophoresis of allozymes and the effects of multiple mating on female reproduction were investigated. When females were forced to mate only once, their fecundity decreased drastically compared to the case of multiple mating (but longevity was unaffected). When one female mated with two males, the outcome of sperm competition depended greatly upon the mating interval. When the second mating occurred immediately after the first, the female fecundity decreased as in the case of single mating and the second male fertilized only a few eggs. However, when there was an interval of 1 day between the two matings, the females achieved normal fecundity and the second male fertilized approximately half the eggs. This suggests that the spermatophore deposited by the first male may act as a short-term copulatory ‘plug’ in the female's genital opening. When one female mated with several males with 1 day intervals, three or more males shared fertilization of the eggs. This study suggests that the multiple mating of females is a necessary stimulus to continue oogenesis and some physiological factors for this stimulation may exist in spermatophores.     

Effect of Drosophila melanogaster Female Size on Male Mating Success

In this study, we examined the influence of female size on mating success in Drosophila melanogaster. The results that were obtained from experiments performed in mating chambers allowed us to confirm the results of previous studies, demonstrating higher mating success of larger D. melanogaster males, and to conclude that female size also affects mating success, either when considering a single male or two competing males. We observed that the advantage for larger males depends on their size relative to that of the female, demonstrating a previously unknown role for female size in mating behavior studies. This effect of female size on mating success depends on various factors: males take longer to initiate courtship toward larger females, large females receive more wing vibrations from males prior to mating, and large females tend to keep moving for longer periods during male courtship. The importance of this finding is discussed in the context of recent reports on sexual conflict in D. melanogaster, in which males were observed to depress fitness in females as a result of intercourse.