Mating tactics and male wing dimorphism in the damselfly,Mnais pruinosa costalis selys (Odonata: Calopterygidae)

Males of the damselfly,Mnais pruinosa costalis, exhibit wing color dimorphism: one form has orange wings, and the other hyaline wings which resemble female wings. The former is usually territorial and the latter uses sneaky mate securing tactics. When orange-winged males failed to establish territory, they became floaters that day. Hyaline-winged males perched around their territories and often, formed in tandem without any apparent courtship behavior when they found females. Their copulation frequency was higher and copulation duration longer than those of territorial males. A few females oviposited without remating. Total oviposition duration of females with which a hyaline-winged male mated was more than 32 min per male on average in a day Females that copulated with hyaline-winged males often remated with orange-winged residents before oviposition. Total duration of oviposition bouts of females after mating with floaters was short (15 min), while that with territorial residents was long (66 min). As a result, total oviposition duration of females with which an orange-winged male mated was about 40 min in a day. The reproductive success of the hyaline-winged males may be similar to that of the orange-winged males.     

Hammering,mauling, and kissing: stereotyped courtship behavior in <Emphasis Type="Italic">Cardiocondyla</Emphasis> ants

Sex appears to be a rather prosaic and casual event in the life of most social Hymenoptera. In contrast, mating in the ant genus Cardiocondyla is regularly preceded by a prolonged and stereotypic courtship display. Pummeling the head of the female with mandibles and / or antennae and vibrations of the gaster, presumably stridulation, are essential parts of male courtship. The overall structure of the mating pattern is conserved throughout species and between winged and wingless, “ergatoid” males, but exhibits species-specific idiosyncrasies. For example, C. elegans males regularly end the interaction with a female with a short mouth-to-mouth contact. Variation in the duration of the precopulatory phase and the copulation itself might reflect different degrees of inter- and intrasexual selection. More information on the dynamics of sperm transfer and the risk and intensity of sperm competition are needed to better understand the evolution of the complex mating behavior in this genus. Received 15 December 2006; revised 25 June 2007; accepted 11 September 2007.     

Agonistic and courtship behaviour patterns in the skink Chalcides viridanus (Fam. Scincidae) from Tenerife

There have been relatively few attempts to quantitatively describe behaviours in scincid lizards. Chalcides viridanus is a small body-sized skink endemic of Tenerife (Canary Islands). We describe and quantify 18 behaviour patterns (both social and agonistic) of this species, some of which have not been described before for other scincids. Video recordings of male–male, female–female, and male–female interactions were made under laboratory conditions, with controlled light–dark cycle and temperature. We describe several agonistic and courtship behaviour patterns. Within the first context, we detected a new agonistic behaviour for a scincid, “Snout to body”, that appeared at the beginning of agonistic sequences; it consisted of each animal placing its snout in contact with the other individual’s lateral side of the body. The amplitude of head movement during “Head bobbing” was lower than that described for many other lizard species. Agonistic behaviours were shown in intrasexual staged encounters both within males and females. The comparison of behaviour patterns of both types of intrasexual encounters showed that females were more active, exhibiting significantly higher frequencies of behaviour than males. Specifically, females showed the “Snout to body” pattern more frequently than males. In male–female encounters we detected courtship and copulation patterns only in April, when males performed “Bites” and “Snout to body” directed at females.     

Reproductive patterns in mantled howler monkeys: Estrus,mate choice and copulation

The present study was undertaken to evaluate non-random mating patterns in two groups of mantled howler monkeys in two tropical dry forest habitats. Sexual dimorphism, female estrus stage, male dominance rank, sexual solicitations and copulations were assessed. Males are significantly larger than females, but female weight varies more than male weight. The length of female estrus cycles is comparable in both habitats, but females in the more strongly seasonal habitat demonstrate greater estrus synchrony relative to their numbers. Males solicit potential mates more frequently than females, a pattern explained by the relatively high rate of sexual solicitation by high-ranking males. Females in “peak” estrus solicit “alpha” males, while females in other stages of estrus solicit males equally by rank. Intersexual aggression occurs rarely, and “forced copulations” are attempted but, apparently, are unsuccessful. Sexual solicitations by “alpha” males and “peak” estrus females are most likely to lead to copulation, and “alpha” males are more likely to copulate than “gamma” males. In general, latencies from first solicitation to copulation are expensive in time, especially for high-ranking males. Estimated annual reproduction success favors high-ranking males, and results indicate that male and female mating behavior is mutually coordinated and controlled.     

Function and genetics of long versus short copulations in the cactophilic fruit fly,drosophila mojavensis (diptera: drosophilidae)

Variation in copulation duration of Drosophila mojavensisstrains was influenced by both sexes. Males maintained predominant control, as copulation duration of pairs from different strains was more similar to that of the strain from which the male was derived, but female origin also contributed significantly to the duration of copulation. Variation among strains was controlled by genes acting additively in both sexes. The size of both males and females also affected copulation duration. Small males copulated longer on average than large males, while males paired with large females copulated longer than those paired with small females. The importance of copulation duration to fitness was tested by correlation analyses with male size, female size, female remating latency, and number of eggs laid prior to female remating. Longer copulations stimulated earlier oviposition, possibly by increasing accessory gland secretions that are passed by males during copulation.     

Role of male scent in the mating behavior ofPieris melete Ménétriès (Lepidoptera: Pieridae)

APieris melete male emits species-specific scent from the wings, while a virgin female ofP. melete is frequently observed to take the so-called “mate-refusal posture” in response to courting males. The role of the male scent in the mating behavior ofP. melete was investigated experimentally by using scented and scentless male models. It is suggested that the male wing scent can function primarily as a sex pheromone to “seduce” the virgin females that assume the mate-refusal posture, and can therefore induce a successful copulation. This work was financially supported by the Japan Society for the Promotion of Science.     

Courtship song, male agonistic encounters, and female mate choice in the house cricket,Acheta domesticus (Orthoptera: Gryllidae)

Whether female crickets choose among males based on characteristics of the courtship song is uncertain, but in many species, males not producing courtship song do not mate. In the house cricket,Acheta domesticus, we examined whether a female chose or rejected a male based on his size, latency to chirp, latency to produce courtship song, or rate of the high-frequency pulse of courtship song (“court rate”). We confirmed that females mated only with males that produced courtship song, but we found no evidence that the other factors we measured affected a female’s decision to mate. In addition, we investigated whether the outcome of male agonistic encounters affected the subsequent production of courtship song. In one experiment, we observed courtship and mating behavior when a single female was placed with a pair of males following a 10-min interaction period between the two males. Winners of male agonistic encounters had higher mating success. However, winners and losers of agonistic encounters were not different in their likelihood or latency to produce courtship song or in the number of times they were disrupted by the other male in the pair. In a second experiment, we allowed two males to interact for a 10-min period, but following this interaction period, we placed a female with each male separately and observed courtship and mating behavior. The mating success of winners and losers was not different under these circumstances, and we found no differences between winners and losers in any subsequent courtship or mating behavior examined. We conclude that winning agonistic encounters influences a male’s mating success in ways other than his production of courtship song and this effect is lost when winning and losing males are separated and each is given an opportunity to mate.     

Allocleaning in an intertidal ocypodid crab,Macrophthalmus banzai

The ocypodid crabMacrophthalmus banzai often forages on the carapace or walking legs of other conspecific individuals. This behaviour can be classified into 2 types, long cleaning and short cleaning. They are distinguished from each other by the cleaner's approach (slow or quick), duration of cleaning and scoops per bout. Long cleaning was done by a male or a female against a larger crab of either sex. Seasonal and daily frequencies of long cleaning were more or less constant. Immediately before and after a long cleaning, the cleaner was mostly engaged in substratum-feeding. After the cleaning, the recipient tended to return to its own burrow. Short cleaning was done mostly by males against smaller females. Seasonal and daily frequencies of short cleaning exhibited positive correlation with waving display. The cleaner frequently performed waving immediately before cleaning. In addition, short cleaning occurred immediately before surface copulation and before underground pairing of male entry into the female's burrow. These data suggest that the long cleaning is related to feeding and the short cleaning with male courtship.     

Search behavior of pheromone-stimulated potato tuber moth males (Lepidoptera: Gelechiidae)

The processes of female searching by male potato tuber moths,Phthorimaea operculella, were analyzed. The behavioral components to copulation were antennal cleaning, quiescence, walking, wing fanning, contact with female, hair brush display, copulation attempt, and copulation. Males did not always succeed in mating on their first attempt. Searching behavior of males changes to “area-restricted searching” after contact with a female. Males could, therefore, find females efficiently and copulate.     

Male Competition over Access to Females in a Spider with Last-Male Sperm Precedence

Agonistic behaviour between male cellar spiders (Pholcus phalangioides) was investigated to test whether (1) size difference determines which male achieves access to the female, (2) males are able to monopolize access to the female until egg laying and whether (3) female resource value increases before egg laying because of last‐male sperm precedence. We further investigated whether (4) there is variation in time and energy spent on courtship and copulation depending on the degree of sperm competition, i.e. with or without rival present. In three experimental settings we introduced two males of either different or similar sizes, or a single male to a female. The mating units were constantly video‐observed until the females produced their first egg sac. Experience, ownership and female resource value in terms of body size was controlled. Our results show that larger males achieve almost exclusive access to females. Size symmetrical settings resulted in increased fighting activity and duration but dominance did not influence mating success. If copulations were disturbed by the rival male, copulations were terminated earlier in symmetrical settings compared with asymmetrical settings. In 94.8% of trials only one copulation took place, suggesting that the copulating male successfully monopolized access to the female. Males confronted with a rival copulated longer but courted significantly shorter than lone males. Although the last male to copulate sires 88% of the offspring in P. phalangioides, neither fighting nor courtship activity increased before the female laid a batch of eggs. This suggests that males have no indication of the timing of oviposition.     

Do bonobos copulate more frequently and promiscuously than chimpanzees?

The copulatory activities of bonobos (Pan paniscus) of Wamba, Zaire, were compared with those of chimpanzees (P. troglodytes schweinfurthii) of Mahale, Tanzania. The copulation rates of adult male bonobos were equal to or lower than those of adult male chimpanzees. The copulation rates of adult female bonobos were approximately equal to those of adult female chimpanzees who were in maximal genital swelling, but it should be much higher than those of the adult female chimpanzees throughout the birth interval. The copulation rates of adolescent male bonobos were lower than those of adolescent male chimpanzees, whereas the copulation rates of adolescent female bonobos were much higher than those of adolescent female chimpanzees. It was suggested that the bonobos of Wamba did not copulate more promiscuously than did the chimpanzees of Mahale. The female bonobos may show “receptivity”, whereas female chimpanzees may show rather “proceptivity”.     

Mating patterns,mate choice,and birth season heterosexual relationships in free-ranging rhesus macaques

Birth season adult heterosexual nonkin relationships of 50 free-ranging female rhesus macaques (Macaca mulatta) in two social groups at Cayo Santiago, Puerto Rico were examined using focal follow (289 hr) and ad lib data. Eighty-eight percent of subjects had at least one relationship characterized by particularly high frequencies of spatial proximity, grooming, or both. These were designated “friendships.” Males intervened in aggressive interactions more frequently on behalf of Friends than non-Friends. Female aggressive support of males was extremely rare. Higher-ranking males experienced more friendships than lower-ranking males. High-ranking females had higher-ranking Friends than low-ranking females. Older females had higher-ranking Friends than younger females. Females groomed high-ranking Friends more than they were groomed by them, whereas they groomed low-ranking Friends less than they were groomed by them. In one social group, high-ranking females were more likely than low-ranking females to groom their Friends more than they were groomed by them. Males were more responsible than females for spatial proximity maintenance in 9 of 14 Friend dyads for which sufficient data were available. Neither male nor female dominance rank affected responsibility for proximity maintenance in Friend dyads. Eight of 24 females had friendships with males with whom they had completed copulations during their conception peri-ovulatory period of the preceding mating season. Two of 19 females completed peri-ovulatory copulations with Friends during the following mating season. Friendship was not correlated with either of two demonstrated female mate choice indicators: (1) proximity maintenance during estrus; or (2) cooperation with male “hip-grasp” courtship attempts. Males directed “muzzle-up” courtship signals at lower rates toward Friends than toward non-Friends. These and other investigators' results indicate that (1) protection from aggression is the primary benefit to female rhesus macaques of birth season heterosexual relationships; (2) the most effective protectors are in greatest demand as Friends; and (3) friendship has no effect or an inhibitory effect on mate choice in this species. Benefits to males of friendships were not apparent from this study but may include coalitional support against lower-ranking males.     

Female sexual receptivity and male copula guarding during prolonged copulations in the damselflyIschnura senegalensis (Odonata:Coenagrionidae)

Laboratory experiments were conducted to clarify the relationship between female sexual receptivity and male copula guarding inI. senegalensis, a species that copulates for several hours. In insectaries, most copulations were initiated early in the morning, and terminated relatively synchronously between 11 00 and 13 00. Females refused males with wing-flutter display and oviposited alone in the afternoon regardless of copulation events of that morning. Females could sexually receive males only in the morning. Males copulated for several hours until 12 00 after which females could oviposit. To determine whether copulations that last for hours function as male copula guarding or only of sperm displacement, emerged males were kept at various densities and permitted to copulate with virgin and mated females in insectaries. Both with virgin and mated females, “social” (not solitary; 2–4 males / insectary) males initiated copulations early in the morning and always terminated at around 12 00. However, both with virgin and mated females, solitary (one male / insectary) males terminated copulations in the morning. In both cases, duration of copulations did not significantly differ for virgin females and mated females. Therefore, long (several hour) copulation is more likely to function as male copula guarding than as sperm displacement, and duration of copulations is predicted to be shortened when male density is very low.     

Absence of social facilitation of courtship in the wolf spider, Schizocosa ocreata (Hentz) (Araneae: Lycosidae)

Males of many animal species are reproductively limited by the difficulty and time costs of finding mates. Males of such species should be selected to take advantage of any cues that might reveal the location of prospective mates. Cues to female location are not restricted to those produced by females, but might also include the highly apparent courtship displays of males that have already found a female. By “eavesdropping” on these courting rivals, initiating sexual displays when courting rivals are detected (i.e., social facilitation of displays); males might effectively exploit the mate-searching efforts of their rivals. We tested the possibility that male Schizocosa ocreata wolf spiders exhibit social facilitation of courtship behaviors using a combination of live behavioral trials and video playback with single stimulus presentations. When exposed to visual cues from another male, male S. ocreata can discern the presence of another individual whether that individual is courting or not. However, we found no evidence of social facilitation of courtship or chemoexploratory behaviors in response to seismic or visual cues presented in isolation or combined. While complex, multimodal, male courtship signals are important in mate choice by female S. ocreata, males do not appear to use these cues to socially facilitate their own courtship.     

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.     

Sexual behavior of adult male chimpanzees of the Mahale Mountains national park,Tanzania

This study, based on 687 hr of focal observations, aims to describe overall patterns of the sexual behavior of the adult male chimpanzees of the Mahale Mountains, to compare the results with previous reports, and to explain the variations between studies. Genital inspection of cycling females by adult males was eight times as frequent as that of lactating females, and twice as frequent as that of pregnant females. Inspection of the genitals of cycling females increased dramatically 7–10 days before the onset of maximal swelling and gradually decreased as the day of ovulation approached. Adult males likely obtained information on the attractivity of females by inspecting their genitals. Mating was usually initiated by male courtship and followed by pelvic thrusts in a dorsoventral posture, performed on, rather than above, the ground, which continued for 7 s. on average, and was typically followed by female squeaking and darting from the male, or by the male grooming the female. Higher-ranking males mated with females in the peri-ovulatory period more frequently than did lower-ranking males. In particular, two alpha males mated with such females more often than did any other adult males. A male who interfered with a mating pair was dominant over the mating male in other agonistic contexts. The duration of intromission was correlated with neither dominance rank nor age. However, when an adult male declined in rank from alpha in 1991 to third in 1992, he showed a significantly shorter duration of intromission. This indicates that for a particular male, the alpha rank guaranteed longer duration of intromission. Allies of alpha males tended to mate with peri-ovulatory females more frequently than expected from their low dominance ranks. The number of mating partners was not correlated with male dominance rank, but was sometimes negatively correlated with male age. Females were significantly more likely to emit a copulatory squeak when mating with younger, rather than older, adult males. Male dominance rank and the rate of female copulatory squeaking were not correlated. Weaning infants regularly interfered with their mothers' mating. Occasionally, unrelated adolescent males and rarely females pushed themselves in between copulating adults. Female choice was indicated when they performed a “penis erection check” or took the initiative in courtship, or on the other hand showed strong reluctance to mate with particular males. Young adult males more often received erection checks than did prime males, while none of the three old adult males did. Courtship initiated by estrous females was not directed to two of the oldest males, the exception of which was the alpha male. The oldest males, except for the alpha, were consistently avoided by many estrous females, both young and old. In response to female reluctance, males behaved violently, however, this was not effective, because other more dominant males came to rescue the female. Neither courtship nor mating was seen between mature sons and their mothers, nor between brothers and sisters.     

Sperm Storage is not Subject to Cephalic Control in the Caribbean Fruit Fly, Anastrepha suspensa

Male Anastrepha suspensa exhibit a repertoire of “copulatory” behaviors involving the female head. This study examined the effect of female decapitation on copulation duration, and quantity and distribution of sperm in the four storage organs. Copulation duration and sperm storage were highly variable (16 to 160 min and 0 to 2,643 sperm, respectively). Sperm quantity variation was due primarily to storage patterns among three spermathecae. There was no significant correlation (Pearson, P > 0.05) between body size and copulation duration, or sperm stored in relation to both variables. Although mean copulation duration was significantly shorter for non-decapitated females (t-test, t = 8.48, P < 0.001), sperm quantity and storage patterns were not significantly different. These data suggest the near autonomy of the abdominal ganglion’s role in any female mediated effects on sperm storage.     

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.     

Male Performance and Body Size Affect Female Re-Mating Occurrence in the Orb-Web Spider Leucauge mariana (Araneae, Tetragnathidae)

Females can affect male probabilities of paternity success through behavioural, morphological and/or physiological processes occurring during or after copulation. These processes under female-control include the acceptance or rejection of mating attempts by subsequent males. Leucauge mariana is an orb weaving spider that shows male mate guarding of penultimate females, male–male competition on female webs and copulatory plugs, suggesting a polyandric mating system. The aim of the present study was to ascertain whether male behaviour during courtship and copulation in L. mariana relate with female re-mating decisions. Forty-three virgin females were exposed to up to three males until they mated. In 24 cases, the copulatory plug was absent after mating and females were exposed the next day to up to three other males. Eighteen females accepted a second mating. Relatively larger females were more receptive to second matings and were more likely to copulate if the second male was smaller. Longer duration of female tapping and abdominal bobbing during courtship, and first copulations with less short insertions and more flubs, were associated with increased female acceptance to second matings. The results indicate cryptic female choice on male courtship and copulatory performance and suggest female-control over the determination of male mating success in this spider species.