The pattern of tube-sharing in Micro deutopus gryllotalpa (Crustacea: Amphipoda)

The amphipod crustacean Microdeutopus gryllotalpa builds tubes on solid substrata. Mature animals usually reside in individual tubes. When more than one individual is present in a tube it is always a single heterogametic pair. Tube-sharing occurs with the greatest frequency 12 h before the female's moult. Following the female's moult, most males leave the tube. The pattern of tube-sharing is the behavioural analoque of precopulation in epibenthic amphipods. It is demonstrated that (1) no more than two individuals are found in a tube because (2) one individual will not permit another individual of the same sex to cohabit the same tube, and (3) the female determines the time of tube-sharing, for most females tube-share only shortly before they moult. It is hypothesized that after the male leaves the female's tube, he cruises from tube to tube until he gains entry into the tube of another receptive female.     

Bayesian mating decisions in an amphipod, Gammarus lawrencianus Bousfield

In G. lawrencianus, females of a given fecundity and proximity to copulation day are assessed by males to be more valuable as mates, the longer the males have amplexed with the females (i.e. the greater their past investment in the act of amplexus). This behaviour is consistent with the hypotheses that (1) males are uncertain of the true reproductive state of a female on first contacting her; (2) the prior distribution of unamplexed females likely to be encountered by the males affects their initial estimates of a female's state; and (3) the time spent amplexing with a female allows information to be gathered which reduces a male's uncertainty of the female's state.     

The effects of mate separation on pair re-formation in the Texas cichlid fish (Cichlasoma cyanoguttatum)

This study tested whether mated pairs of Texas cichlid ($\text{Cichlasoma cyanoquttatum}$) would reform and continue rearing their offspring after being separated for either one, four, or ten days. All pairs successfully re-formed after one day, while only 50% re-formed after 4 days. No pair re-formed after 10 days of separation. In successfully re-formed pairs, the female was always more aggressive than her mate, irrespective of whether the female was the resident or the returning individual. Aggressiveness seemed related to the female's attempt to control the male's parental behavior. For pairs which did not re-form, the resident parent, either the male or female, violently and continously attacked the returned mate.     

Mating behavior ofPhytoecia rufiventris Gautier (Coleoptera: Cerambycidae)

Sexual behavior between males and females, as well as between males, is described and discussed for the cerambycid beetlePhytoecia rufiventris. The beetles' taxis toward plants taller than average height brings the sexes together from a distance. A male may mount another individual (male or female) and attempt copulation without sex discrimination. The male can discern the sex of another individual only when the terminal part of his abdomen touches the ventral surface of the fifth visible sternite of the latter. No evidence of a sex pheromone is found in this species. Within 1.5–5.5 cm the substrateborne vibrations produced by a moving individual may be the important factor which elicits males to approach a moving individual and attempt copulation. If a female is receptive when a male touches her, he can copulate with her without any courtship display. However, if the female runs away and appears unreceptive, the male will perform courtship displays. Copulation is usually terminated by males. Homosexual behavior between males is discussed.     

Infanticide in male laboratory mice: Effects of social status,prior sexual experience,and basis for discrimination between related and unrelated young

Several aspects of infanticide, including effects of social status, prior sexual experience, and the basis for discrimination between related and unrelated young, were examined in Swiss-Webster laboratory mice (Mus musculus). Strange males introduced into the female's cage for 24 h on day 1 postpartum significantly reduced pup survival whereas the introduction of sires did not. Direct observations of infanticide were frequent, and the motor patterns used by males to kill pups are described. Males killed their own offspring when those young were in the nest of a strange female, whereas most males did not kill unrelated young in the nest of a familiar female. Thus, past association with the mother appears to be the single most important factor mediating male discrimination of young. Prior contact with a specific female's urine reduced a male's propensity for subsequently killing her young. The act of copulation itself also reduced infanticide. Infanticidal behaviour was correlated with the male's social status: most dominant males killed unrelated pups, whereas none of the subordinates engaged in infanticide. These results are discussed in terms of the population biology of this species.     

Sexual coercion does not exclude luring behavior in the climbing camel-spider <Emphasis Type="Italic">Oltacola chacoensis</Emphasis> (Arachnida,Solifugae, Ammotrechidae)

Sexual coercion in the form of forced copulation has been used as a typical example to illustrate the conflict of interests between females and males. Among arthropods, forced copulation has been reported for some groups of insects and crustaceans, but not for arachnids. In the present work, we analyse and describe the behavioral patterns of mating behavior of the climbing camel-spider, Oltacola chacoensis, relating it to relevant morphological features, In this species, the male forcefully clasps the female’s genital region with his chelicerae and locks her fourth pair of legs with his pedipalps. In some cases, the cuticle of the female’s abdomen was damaged by this cheliceral clasping. In contrast to other camel-spiders, the female O. chacoensis never remained motionless during mating, but continuously shook her body, opening her chelicerae notably towards the male. Despite this coercive context, males performed copulatory courtship (tapping with pedipalps) and females showed an apparent cooperative behavior (they remained still during a short period of the sperm transfer phase). These results strengthen the idea that sexual coercion (in the form of forced copulation) and luring behavior (in the form of copulatory courtship) are not two mutually-exclusive male’s strategies during a single copulation.     

Pape in Panorpa scorpionflies and a general rape hypothesis

The adaptive significance of heterosexual rape is difficult to demonstrate because (1) female coyness is difficult to distinguish from apparent rape, and (2) male fitness must be enhanced by rape. Male Panorpa scorpionflies inseminate unwilling females by securing the female's wings in an abdominal clamp. Panorpa rapists gain fitness through avoidance of risks of predation by web-building spiders associated with the more typical male behaviours of feeding females a dead insect or salivary secretion during copulation. Males should be most strongly selected to rape in species in which males provide resources important for female reproduction. In such species, the strength of selection on males to rape should exceed the strength of selection on females to prevent rape.     

Group mating among Norway rats II. The social dynamics of copulation: Competition,cooperation, and mate choice

The social interactions within groups of Norway rats (Rattus norvegicus) had a strong impact on the individual pattern of copulation which, in turn, affects sperm precedence and the probability of implantation in this species. Males alternated uninterrupted ejaculatory series, augmenting each others' copulatory investment. Females took turns mating after receiving an intromission, collectively potentiating the males' copulatory behaviour; increasing the number of oestrous females increased the number of intromissions and ejaculations achieved by each male but did not affect the amount of copulation experienced by each female. These turn-taking patterns within each sex provided the opportunity to change partners and permitted the emergence of different sex-typical patterns of copulation. Furthermore, the dominant male contributed more intromissions and tended to give each female more ejaculations than the subordinates did. Dominant males were also more likely to inhibit the subordinates' sperm transport. Females competed among themselves for the opportunity to mate with a male as he approached ejaculation and were likely to protect more of the dominant male's sperm transport than the subordinate male's.     

Post-copulatory mate guarding by males of the demselfly Hetaerina vulnerata Selys (Odonata: Calopterygidae)

Males of the calopterygid damselfly Hetaerina vulnerata remain with their mates after copulating with them. The species exhibits two unusual features of post-copulatory mate guarding. First, a male will often leave his territory to accompany a female in tandem on a search for oviposition sites elsewhere. Second, a male will perch near his ovipositing female even though she completely submerges when egg-laying and cannot be captured and mated by another male while she is underwater. These activities carry two potential costs: (1) a male may miss other receptive females while guarding one mate and (2) he may lose his territory to an interloper while he is absent. These costs were low, however, because territorial males secured only one mating per 3.6 days on average. Moreover, 23 times out of 26, territorial males reclaimed their plots quickly after being away for 30–60 min. The gain from postcopulatory guarding came from being present to recapture a female should she fly up from the water after rejecting an oviposition site. There was a 40% chance that a female would leave one site to search for another during an oviposition bout. If the male were not present, his mate would be captured and mated by another individual (no female ever selected an oviposition site without being carried to it by a male). Her new partner would fertilize the remaining eggs in the female's clutch (if sperm precedence occurs in this species). The total number of eggs fertilized by a male will be affected by how well he prevents any one mate from copulating again before she lays her entire clutch and the total number of receptive females he captures. The variation in the degree of mate guarding by male odonates seems to be the evolutionary outcome of differences in fitness gains derived from these two competing activities in different ecological settings.     

The Mating Behaviour of the Velvet Ant, <Emphasis Type="Italic">Nemka viduata</Emphasis> (Hymenoptera: Mutillidae)

In the present study, the mating behaviour of the velvet ant Nemka viduata (Pallas) (Mutillidae) is described both from field and laboratory observations. The whole pairing interaction, lasting around two hours, includes several behavioural phases. During pre-copula, the male seizes the female’s neck with his mandibles, and then starts to rhythmically stroke the prothorax of the female with his forelegs (this behaviour is also resumed after copulation) before curving his abdomen in order to couple the genital parts, including genital armatures (the male parameres remaining outside the female body); just prior to copulation, the female extrudes the sting, and immediately after copulation begins, she stridulates for 7–10 s, this behaviour is repeated when the pair separates. During copulation (lasting around two minutes), the male moves his antennae rhythmically, hitting the back of the female’s head with the scape. Generally, recently-mated males become aggressive towards females, but more tolerant after a few days. During the whole pairing act, females are held by the males’ mandibles, and in the field they are carried off in flight or by walking to a safe place to copulate. This would suggest that larger males, which can lift a wider range of female sizes, have a reproductive advantage, as indicated by data obtained on their load-lifting capacity with respect to the size distribution of females. A review of mating behaviour in mutillid wasps and comparisons with other lineages of aculeate and non-aculeate Hymenoptera are also given.     

Sexual Cannibalism in the Brown Widow Spider (Latrodectus geometricus)

Sexual cannibalism may represent an extreme form of male monogamy. According to this view, males gain reproductive success by sacrificing themselves to females. We studied the occurrence and timing of sexual cannibalism in the brown widow spider Latrodectus geometricus and compared male courtship and mating behavior with virgin and with previously mated females. We found that events of sexual cannibalism are frequent, that they occur during copulation and that males initiate cannibalism by placing the abdomen in front of the female’s mouth‐parts during copulation (somersault behavior). Both the somersaults and mating occurred more frequently with virgins than with previously mated females. Our results support the hypothesis that sexual cannibalism is a male strategy in this species. The somersault behavior was previously known only from the redback spider, Latrodectus hasselti. It is as yet unknown whether self‐sacrifice has evolved more than once in this genus.     

Sexual Dimorphisms in the Dermal Denticles of the Lesser-Spotted Catshark,Scyliorhinus canicula (Linnaeus, 1758)

The dermal layers of several elasmobranch species have been shown to be sexually dimorphic. Generally, when this occurs the females have thicker dermal layers compared to those of males. This sexual dimorphism has been suggested to occur as a response to male biting during mating. Although male biting as a copulatory behaviour in Scyliorhinus canicula has been widely speculated to occur, only relatively recently has this behaviour been observed. Male S. canicula use their mouths to bite the female’s pectoral and caudal fins as part of their pre-copulatory behaviour and to grasp females during copulation. Previous work has shown that female S. canicula have a thicker epidermis compared to that of males. The structure of the dermal denticles in females may also differ from that of males in order to protect against male biting or to provide a greater degree of friction in order to allow the male more purchase. This study reveals that the length, width and density of the dermal denticles of mature male and female S. canicula are sexually dimorphic across the integument in areas where males have been observed to bite and wrap themselves around females (pectoral fin, area posterior to the pectoral fin, caudal fin, and pelvic girdle). No significant differences in the dermal denticle dimensions were found in other body areas examined (head, dorsal skin and caudal peduncle). Sexually dimorphic dermal denticles in mature S. canicula could be a response to male biting/wrapping as part of the copulatory process.     

The effect of differences in body size on the male territorial system of the fly Dryomyza anilis

Drymyza anilis nales defend carcasses, the oviposition sites for females. On carcasses less than 100 g in weight, a single male establishes a teriitory. Ther sex ratio at carcasses is male-biased. Territorial males are larger than other males on average, and move and attack other males more frequently than non-territorial males do. Large carcasses attract more males than smaller ones, but the female, but the advantage of territorial behaviour decreases with increasing density of males. Copulating males are significantly larger than average males, but smaller than territorial males, suggesting than some non-territorial males have access to females. Males seem to be albe to assess a female's egg load, and to adjust the duration of copulation accordingly.     

Sexual attractivity,proceptivity, and receptivity in female mammals

Special definitions are proposed for three concepts representing characteristics of female mammals when they are in estrus. Attractivity refers to the female's stimulus value in evoking sexual responses by the male. Proceptivity connotes various reactions by the female toward the male which constitute her assumption of initiative in establishing or maintaining sexual interaction. Receptivity is defined in terms of female responses necessary and sufficient for the male's success in achieving intravaginal ejaculation. Attempts are made to measure each variable in the S-R paradigm and to identify the causal agents determining each aspect of the estrous female's behavioral characteristics.     

Receptivity of female remating and sperm number in the sperm storage organ in the bean bug,Riptortus clavatus (Heteroptera: Alydidae)

This study examines the relationship between the number of sperm in the seminal receptacle (spermatheca) and the receptivity of female remating in the bean bugRiptortus clavatus Thunberg. On the 21 st day after the first mating when receptivity to remating was > 70%, females receptive to remating had significantly fewer sperm ( < 40 on average) in the spermathecae than females reluctant to do (about 150 on average). However, averages of the number of eggs laid by receptive and reluctant females within 21 days were almost same. The proportion of fertilized eggs for receptive females at 15–21 days after copulation was significantly lower than that for reluctant females. Spermatozoa transferred from a male to a female’s spermatheca were detected 5 min after copulation and then increased continuously to about 500 with the first hour. When copulation durations were manipulated artificially, the shorter the copulation period (=females had less sperm in their spermathecae), the higher the remating rate became. Females may perceive the number of sperm in their seminal receptacles and then determine whether they copulate or not. These results support the hypothesis that females mate multiply in order to replenish inadequate sperm supplies to fertilize all eggs produced.     

The role of genetic constraints and social environment in explaining female extra-pair mating

Why do females of socially monogamous species engage in extra-pair copulations? This long-standing question remains a puzzle, because the benefits of female promiscuous behavior often do not seem to outweigh the costs. Genetic constraint models offer an answer by proposing that female promiscuity emerges through selection favoring alleles that are either beneficial for male reproductive success (intersexual pleiotropy hypothesis) or beneficial for female fecundity (intrasexual pleiotropy hypothesis). A previous quantitative genetic study on captive zebra finches, Taeniopygia guttata, reported support for the first, but not for the second hypothesis. Here, we re-examine both hypotheses based on data from lines selected for high and low male courtship rate. In contrast to previous conclusions, our new analyses clearly reject the hypothesis that male and female promiscuity are genetically homologous traits. We find some support for a positive genetic correlation between female promiscuity and fecundity. This study also shows that the behavioral outcome of extra-pair courtships primarily depends on individual-specific female preferences and not on the “attractiveness” of the social mate. In contrast, patterns of paternity are strongly influenced by the social partner and the pair bond, presumably reflecting variation in copulation behavior, fertility, or sperm competitiveness.     

EVIDENCE FOR WIDESPREAD COURTSHIP DURING COPULATION IN 131 SPECIES OF INSECTS AND SPIDERS,AND IMPLICATIONS FOR CRYPTIC FEMALE CHOICE

Male courtship behavior is generally thought to function prior to copulation, as an inducement to the female to allow the male to copulate with her; this study indicates however, that male courtship during and following copulation (“copulatory courtship”) is common in insects and spiders (81% of 131 species in 102 genera and 49 families, mostly Coleoptera, Hemiptera, Diptera, and Araneioidea). Copulatory courtship is apparently evolutionarily labile, as expected if it is under sexual selection; intrageneric variation occurred in all 17 genera in which more than one species was observed. In 81% of 94 species with copulatory courtship, the male abandoned the female soon after copulation ended; thus, copulatory courtship appears not to function generally to induce acceptance of further copulatory attempts. The most likely explanation for copulatory courtship is that it represents attempts by males to influence cryptic female choice. This suggests that an aspect of sexual selection by female choice not considered by Darwin may be more important than previously appreciated and that the common practice in evolutionary studies of measuring male reproductive success by counting numbers of copulations may sometimes be misleading because of cryptic female choice during and after copulation.     

Behavior of triatomines (Hemiptera: Reduviidae) vectors of Chagas' disease. I. Courtship and copulation of Panstrongylus megistus (Burm., 1835) in the laboratory

A study of the courtship and copulation behavior of Panstrongylus megistus was carried out in the laboratory. Fifty-five newly-fed virgin couples were used. Experiments were performed during the day (9:00 to 12:00 a.m.) and at night (7:00 to 10:00 p.m.). Behavior was recorded by direct observation and was found to consist of the following sequence of behavioral patterns: the male approached the female and jumped on her or mounted her; he took on a dorsolateral position and immobilized the female dorsally and ventrally with his three pairs of legs; the male genital was placed below those of the female; the paramers of the male immobilized the female's genitals; copulation started. The couple joined by the iniciative of the male. The female could be receptive and accept copulation, or nonreceptive and reject the male. Copulation occurred more often on the occasion of the first attempt by the male. Duration of copulation was X = 29.3 +/- 9.3 min (CV = 83%). No behavioral differences were observed between couples tested during the day or at night.     

Variation in the number of sperm transferred during mating among males of the Colorado potato beetle (Coleoptera: Chrysomelidae)

We suggest an explanation for the bimodal distribution in the number of sperm transferred into the female during mating by males of the Colorado potato beetle (Leptinotarsa decemlineata). While in most matings females receive 20,000 sperm, in others they receive less than 1000. A variety of potential proximate causes for this bimodal distribution including experimental procedures and several physiological and behavioral causes were examined: male body size, female body size, male mating history, socio-sexual environments, duration and number of mounts, and duration and number of copulations. Surprisingly, none of these explained the observed distribution. However, we did find that the total time spent in copulation was significantly less variable among males that transferred more than 20,000 sperm than among males that passed less than 1000 sperm. In addition, males transferring large amounts of sperm resulted in proportionally more sperm inside the female’s spermatheca within a short period of time than males passing less than 1000 sperm. The lack of pattern in male sperm ejaculate suggests definite differences in sperm delivery tactics and may be related to the female condition rather than the male.