Behavioral indicators of ovarian phase in white-faced capuchins (Cebus capucinus)

In many primate species, conspicuous behavioral and/or morphological changes are indicators of the fertile phase of the female cycle. However, several primate species, such as the white-faced capuchin, lack these cues. This is referred to as "concealed ovulation," and is argued to be a reproductive strategy that confuses paternity and lowers the risk of infanticide. We studied 10 adult female white-faced capuchins in Santa Rosa National Park, Costa Rica, from January to June 2002. We determined their ovarian cycling patterns by analyzing fecal ovarian hormones, and compared simultaneously collected behavioral data to determine which, if any, cues females use to signal their fertile phases. We found that four females cycled during the study period but ceased to cycle without becoming pregnant. We considered several explanations for the lack of conception during our study, including reproductive seasonality. We found that female C. capucinus showed only small increases in rates of affiliative/proceptive behaviors directed toward adult males during their periovulatory phases. The best indicator of cycle phase was a significant increase in male affiliative behaviors (e.g., following and grooming bouts) and sexual behaviors (e.g., copulations and courtship displays) directed toward females during the periovulatory phase compared to the nonovulatory phase. Our finding that females exhibit little proceptive behavior, but that copulations and male courtship are nonetheless concentrated in periovulatory phases suggests that even though females do not provide behavioral and morphological cues to ovulation, males are still able to detect it. Infanticide occurs with some frequency in these monkeys, and there is evidence for postconceptive mating as a female strategy to lower risk of infanticide via paternity confusion. However, despite this occurrence of nonconceptive mating and the absence of female cues to ovulation, truly concealed ovulation does not appear to be characteristic of this study population of white-faced capuchins.     

Display of proceptive behaviors in relation to urinary and fecal progestin levels over the ovarian cycle in female tufted capuchin monkeys

In many primates species, female sexual attractivity is influenced by behavioral cues as well as by nonbehavioral cues (i.e., visual-morphological or chemical signals). Both kinds of cues are usually related to the ovarian cycle and female hormonal state. Female tufted capuchins (Cebus apella) lack any external morphological change in relation to the ovarian cycle and evidence of scent-marking behavior has never been reported. In addition, tufted capuchin males do not routinely investigate the female's body or urine. Instead, capuchin females are extremely active in sexually soliciting the male(s) and their courtship toward them involves a rich behavioral repertoire. In the present study we defined female tufted capuchin proceptivity and investigated its relationship with female reproductive state. Ovarian hormones were measured in urine and fecal samples from four captive females in order to (a) assess their reliability for monitoring female ovarian function and (b) provide information on the timing of the component cycle phases and in particular the periovulatory phase. Measurements of urinary and fecal progestin metabolites provided the best indicator of ovarian cyclicity and for timing of the periovulatory phase. Through a multivariate analysis of the behavioral data set we distinguished four behaviors (eyebrow raising with vocalization, touching-and-running, nuzzling and head cocking) which showed a marked cyclicity (21.3 days) that matched that of urinary progesterone (21.9 days). Data showed that each period of proceptive behaviors was 2.7 +/- 0.8 days long and the day of a defined luteal phase rise in urinary progesterone levels was markedly shifted toward the end of this period. Furthermore, the ejaculations observed always occurred within proceptive periods. The results clearly indicate that female behavior is a good indicator of the periovulatory phase and can enhance female attractivity.     

Sociosexual behavioral patterns involving nulliparous female orangutans (Pongo sp.) reflect unique challenges during the adolescent period

The primate adolescent period is characterized by a series of changes in physiology, behavior, and social relationships. Orangutans have the slowest life history and the longest period of dependency of all primates. As members of a semisolitary species with high levels of sexual coercion, adolescent female orangutans face a unique combination of challenges when achieving independence from their mother. This study examined the mating behavior of adolescent female orangutans and compared it with that of adult females to assess whether mating behavior reflects distinct strategies at these different points in the life cycle. Data were collected in Gunung Palung National Park on the island of Borneo over 20 years. Mating events from adolescent (n = 19) and adult females (n = 26) were scored and compared. Adolescent female mating events had significantly higher mating scores (indicating more proceptivity) than those of adult females (β = 1.948, p = .001). Adolescent females also engaged in elaborate sociosexual interactions with different flanged males, behaviors that were never observed during mating events of adult females. These interactions involved characteristic behavior on the part of both the adolescent females and the flanged males. Given these findings and the documentation of similar accounts of adolescent female–flanged male mating from the island of Sumatra, we propose that adolescent female orangutans display distinctive behavioral repertoires throughout the genus Pongo which serves to overcome male ambivalence toward nulliparous females, establish familiarity, and evaluate coercive tendencies in flanged males. We suggest that these behavioral patterns are an integral part of female social development in a female philopatric, but highly dispersed species where consistent social support is absent after ranging independence is achieved.     

Relationship between sexual interactions and the timing of the fertile phase in captive female Japanese macaques (Macaca fuscata)

Japanese macaques live in multi‐male/multi‐female social groups in which competition between males, female mate choice, and alternative male mating strategies are important determinants of mating and reproductive success. However, the extent to which adult males rely on female behavior to make their mating decisions as well as the effect of social rank on mating success are not clear as results are inconclusive, varying from study to study. In this study, we combined behavioral and endocrine data of 14 female Japanese macaques to examine the relationship between ovarian cycle phase and frequency of sexual behaviors, and to investigate how social rank influences sexual behavior in this species. We found that there was no increase in female proceptive behaviors during the fertile phase of the ovarian cycle, suggesting that female behaviors did not clearly signal the probability of conception. In spite of that, the frequencies of ejaculatory copulations were highest during this phase, indicating that the attractivity of females increased significantly during the period with higher probability of conception. Males, and especially the highest ranking male, were able to discriminate females nearing ovulation and to concentrate their mating effort, implying that the timing of ovulation was not concealed from them. The α male seemed able to monopolize most female matings, which is probably due in part to the low number of females simultaneously ovulating and to the limited number of inconspicuous places that the lower ranking males have to mate with females and avoid α male aggression. All together, these results suggest that different males may have access to different signals of ovulation and/or are differentially restrained as to how they can act on that information. The exact nature of the estrogen‐related cues males use to recognize female reproductive status, and to what extent males use them warrants further investigation. Am. J. Primatol. 71:868–879, 2009. © 2009 Wiley‐Liss, Inc.     

Exploring the influence of individual courtship behaviors on male mating success in a blow fly

Male courtship behavior and displays influence female mating decisions, and therefore affect mating success in a diverse range of organisms. While there is substantial evidence confirming that females prefer males who invest more in courtship, less attention has been paid to the relative importance of individual behaviors, and the discrete sequences of courtship that result in mating success. The small hairy maggot blow fly, Chrysomya varipes, performs stereotyped courtship behaviors, involving orienting, tapping, waving, arching, wing vibration and mounting. This study aimed to quantify male investment in specific courtship behaviors, and compare courtship investment and behavioral transitions between males who gained mating success (successful males) and those that did not (unsuccessful males). Our results show that mating success was influenced by the behaviors orienting, tapping, arching and mounting. Behavioral transitions revealed a distinct pattern of behaviors leading to a mating attempt, and some differences were observed between successful and unsuccessful males. Overall, our findings suggest that female mating decisions were based on differences in specific male courtship behaviors. This detailed observational study has quantified multiple courtship behaviors for the first time in C. varipes, and highlights the importance of considering multiple behavioral traits when exploring the influence of male courtship on mating success.     

Ovarian cycle phase and same-sex mating behavior in Japanese macaque females

The relationship of the ovarian cycle phase to same-sex mounting activity in adult female Japanese macaques (Macaca fuscata) was studied during the 1997/1998 mating season. Fecal samples were collected from eight female subjects two to three times per week and analyzed by enzyme immunoassay for fecal hormone levels. Hormone profiles of estrone (E1) and pregnanediol (PdG) were used to separate ovarian cycles into three phases: follicular, periovulatory, and luteal. Patterns of same-sex and heterosexual mounting behavior in the females were analyzed for phase variation during conceptive cycles. Same-sex mounting among female Japanese macaques occurred most frequently during the follicular and periovulatory phases of the cycle, and not at all during the luteal phase, paralleling the pattern found in heterosexual mounting behavior. These findings suggest a link between hormonal fluctuations and patterns of sexual mounting, regardless of whether the partner is of the same or opposite sex.     

Menstrual cycle patterns of hormones and sexual behavior in gorillas

Oppositely sexed pairs of gorillas were tested behaviorally during the menstrual cycle to determine the relationship between hormone concentrations of the female and the frequency of sexual activity by the pair. Five females were tested individually during two cycles with each of two males, but serum samples for hormone assay were obtained from each female only during the first cycle of testing. There was no clear relationship between hormones and behavior for the single cycle in which the serum samples were obtained, with the exception that no copulations occurred after the early luteal phase, when progesterone was greater than 5 ng/ml. Normalized behavioral data from all four test cycles for all pairs suggested that female-solicited copulations were restricted primarily to the periovulatory period. Male sexual initiative (by one of the males) accounted for most copulations temporally dissociated from the periovulatory period. Normalized hormone data for all of the females suggested that (1) attractivity was associated with estradiol concentrations during the follicular phase, (2) proceptivity with estradiol and testosterone at midcycle, whereas (3) receptivity was not associated with hormone patterns or cycle phase. The data suggest that hormones are one of several variables that contribute to the regulation of sexual behavior in gorillas.     

Male chimpanzees' grooming rates vary by female age, parity, and fertility status

Copulation preferences in our closest living relative, the chimpanzee, suggest that males prefer older females who have had previous offspring. However, this finding is counter to some behavioral models, which predict that chimpanzee males, as promiscuous breeders with minimal costs to mating, should show little or no preference when choosing mating partners (e.g. should mate indiscriminately). To determine if the preferences indicated by copulations appear in other contexts as well as how they interact, we examined how male chimpanzees' grooming patterns varied amongst females. We found that males' preferences were based on interactions among females' fertility status, age, and parity. First, grooming increased with increasing female parity. We further found an effect of the estrous cycle on grooming; when females were at the lowest point of their cycle, males preferentially groomed parous females at peak reproductive age, but during maximal tumescence, males preferred the oldest multiparous females. Nulliparous females received relatively little grooming regardless of age or fertility. Thus, male chimpanzees apparently chose grooming partners based on both female's experience and fertility, possibly indicating a two-pronged social investment strategy. Male selectivity seems to have evolved to effectively distribute costly social resources in a pattern which may increase their overall reproductive success.     

Relationship between ovarian cycle phase and sexual behavior in female Japanese macaques (Macaca fuscata)

We conducted behavioral observations simultaneously with fecal sample collection on eight nonlactating females 2-3 times per week, October 1997-March 1998, to examine the relationship between ovarian hormones and the sexual behavior of female Japanese macaques (Macaca fuscata) during the mating season. We analyzed samples by enzyme immunoassay for fecal hormone levels. Hormone profiles of estrone-glucuronide (E1) and pregnanediol-glucuronide (PdG) were used to separate ovarian cycles into three phases (follicular, periovulatory, and luteal). Hormonal profiles indicate average cycle lengths of 27.6 +/- 4.2 days (+/- SD; n = 26). Average lengths of the luteal and follicular phases were 12.3 +/- 3.8 days (+/- SD) and 8.3 +/- 3.4 days (+/- SD), respectively. We observed female Japanese macaques engaging in sexual activity throughout the ovarian cycle, with the highest rates occurring during the follicular and periovulatory phases as compared to the luteal phase. The attractivity of female Japanese macaques increased significantly during the follicular and periovulatory phases of the ovarian cycle, when E1 levels are peaking and PdG levels drop to baseline. In addition, females displayed a significant increase in proceptive behavior during the follicular and periovulatory phases. Grooming bouts, as well as proximity between female and male macaques, also increased significantly during the follicular and periovulatory phases. We conclude that fluctuating levels of ovarian hormones in different phases of the cycle are significantly associated with variable rates of copulatory and pericopulatory behaviors in these Japanese macaque females.     

Juggling priorities: female mating tactics in Phayre's leaf monkeys

Extended sexual receptivity in primates is thought to facilitate paternity confusion, thus decreasing the risk of infanticide. However, females might also provide some indication of ovulation to attract preferred males during fertile periods. We examined female mate preferences across defined receptive periods (N = 59) in a group of wild Phayre's leaf monkeys (Trachypithecus phayrei crepusculus) at Phu Khieo Wildlife Sanctuary (February-September 2006; 2,603 contact hours). The group contained seven cycling adult females and three reproductively active males (one adult and two adolescents). We predicted that females would prefer the adult male during periovulatory (POP) receptive periods, but the adolescent males during nonperiovulatory (NPOP) and postconceptive (PC) periods. We collected focal and ad libitum data on sexual and agonistic behaviors to determine female preferences and male awareness of female fertility. We also determined the degree of mating overlap to assess if males were capable of monopolizing females. Our results indicate that females were more frequently proceptive and receptive toward the adult male during POP. By contrast, females were more proceptive and receptive toward one of the adolescent males during PC periods, but rarely interacted with the other adolescent. Patterns of attractivity and agonism across receptive periods suggested that the adult male could detect fertility, while the preferred adolescent could not. Finally, we found a high degree of overlap in total receptive period days, but a low degree of overlap in POP receptive days, suggesting that the adult male might have monopolized females, especially since he seemed to be aware of female fertility. Although these results suggest that females provide some information on ovulation, they also suggest that females attempt to confuse paternity, perhaps capitalizing on male differences in the ability to detect fertility.     

Seasonality of adult male Japanese macaques (Macaca fuscata): androgens and behavior in a confined troop

Seasonal variations in levels of serum testosterone, dihydrotestosterone (DHT), reproductive behavior, and social behavior were investigated in 12 adult males (5 to 20+ years of age) of the Oregon troop of Japanese macaques (Macaca fuscata). Blood samples were collected at 2- to 4-month intervals, and behaviors were monitored twice weekly over a 15-month period. Significant seasonal variations in levels of testosterone and DHT, and in frequencies of mount series, ejaculations, number of female partners, displays, courtship, and aggression were observed. Seasonal variations in reproductive and social behaviors did not correlate with seasonal variations in androgen levels because seasonal increases in these behaviors followed seasonal increases in the androgens with a 1- to 2-month delay. However, significant correlations between increased androgen levels and the onset of mating activity occur when mean monthly frequencies of mount series are shifted 1 to 2 months earlier to coincide with the rise in serum androgen levels. The frequency of adult male play and male-male mounting increased significantly when androgen levels were low. We suggest that photoperiod changes may function as a proximate cue in male Japanese macaques which induces a state of biological readiness for mating, and the behavioral consequences (i.e., mating) are then dependent upon the presence of receptive females.     

Female Mate Preferences Among Pan troglodytes schweinfurthii of Kanyawara, Kibale National Park, Uganda

Studies of reproduction among chimpanzees traditionally have focused on the mating strategies of males. However, less is known about the mating strategies of female chimpanzees and whether they demonstrate mate choice. I investigated sexual behavior and female mate preference in the chimpanzees of the Kanyawara community. To estimate mate preferences, I analyzed female proceptivity and resistance rates of 6 estrous females toward a total of 13 males as well as male solicitation and aggression rates toward females. Males solicited some females more often than others for mating and preferred them throughout estrus, not only during the periovulatory period (POP), when conception was most likely. In contrast, though females had strong mate preferences in both non-POP and POP, their mate preferences were not consistent between the 2 phases. The shift in mate preferences is evidence of a promiscuous yet tactical mating strategy to confuse paternity. Further, females were more proceptive and generally less resistant toward eschewed males in non-POP and more proceptive and less resistant toward preferred males in POP. Hence, the results indicate that females attempted to mate selectively during the fertile phase. Kanyawara female chimpanzees appear to change their mating strategies and selectivity during estrus and thus may pursue a mixed reproductive strategy. The tactic may allow females to deceive males, indicating that promiscuity among chimpanzee females may be more strategic than previously thought.     

Mating Tactics in Response to Costs Incurred by Mating with Multiple Males in Wild Female Japanese Macaques

We investigated the costs of mating with multiple males in terms of feeding time, traveling distances, sexual proceptivity, and male aggression, for wild female (Macaca fuscata yakui) on Yakushima Island, Japan. We analyzed all-day focal sampling data from 7 females during the mating season (Sept.-Nov. 1996). On days when estrous females copulated with multiple males, they decreased their feeding time to half that of anestrous days, traveled longer distances, showed more proceptive sexual behaviors and received more aggression from subordinate males than on days when they copulated with only the 1st-ranking male. On days when females copulated with only the 1st-ranking male, they showed no difference in feeding time with that of anestrous days, and expended less effort than the above mating pattern because of short traveling distances, diminished sexual proceptivity and a lower frequency of aggression received. The results suggest that the costs of estrous vary according to female sexual proceptivity and the number and social status of mating partners. Female Japanese macaques exhibit a mixed mating strategy over prolonged estrous periods, which may provide females with opportunities to maximize the benefits of copulating with multiple males and to minimize the costs of estrus by mating with only the 1st-ranking male. During an estrous cycle, females may be adjusting efforts for reproduction and survival; i.e., mating vs. feeding.     

Cultural influences on female mate choice: an experimental test in cowbirds, Molothrus ater

Previous studies of brown-headed cowbirds Molothrus ater, have shown that social learning and cultural transmission can influence courtship and mating patterns. These earlier studies did not test whether cultural background influenced mate choice in females and therefore whether culture could potentially play a role in sexual selection in this species, as has been suggested by recent theory. Here, we tested whether culture influences female mate choice in brown-headed cowbirds. Female cowbirds from a South Dakota population were housed with adult cowbirds from the same South Dakota population or with adult cowbirds from a behaviourally distinct population from Indiana. We tested the mating preferences of females of the South Dakota culture and females of the Indiana culture in sequential mate-choice trials with males, controlling for intrasexual interactions. The males were South Dakota cowbirds that had also been housed either in the South Dakota culture or in the Indiana culture. Females showed mating preferences for males from their own culture. These results suggest that mate choice in female cowbirds can be influenced by cultural background. We briefly discuss the effect that culture may have on sexual selection and on the evolution of female mating preferences. Copyright 1999 The Association for the Study of Animal Behaviour.     

Analysis of the mating behavior and some possible causes of male copulatory success in Dryas iulia alcionea (Lepidoptera, Nymphalidae, Heliconiinae)

In this study we examined the mating behavior of Dryas iulia and the acceptance and rejection mechanisms of females during courtship activity. An ethogram of mating behavior was organized on the basis of 100 h of observation in an insectarium. Several different behaviors were catalogued and separated into two behavioral repertoires (pre-coupling, post-coupling). The behavioral sequence of mating behavior was also analyzed using a total of 53 pairs of D. iulia. The courtship activities involved interactions between the sexes in three sequential phases—the aerial, air-ground, and ground phases. In 49% of observations the courtship activities led to copulation. The mean time to the occurrence of the first interaction between males and females did not differ between interactions with and without copulation. The behavior of females and males in the two groups were analyzed and differences were found between behavioral activities of both sexes. The results indicate that males’ insistence on courtship does not affect their copulatory success, and that females play a decisive role in copulation occurrence. Analysis of behavioral transitions showed there are many alternative behavioral routines in interactions with and without copulation. The number of behavioral transitions recorded was smaller in the group in which copulation occurred, indicating that males with copulatory success modified their behavior less frequently. Successful males recorded more transitions with a probability of occurrence greater than 0.4, and their behavioral activity was also less reticulated. Analysis using the stereotyping index showed that situations in which copulations occurred were more stereotyped than those in which it did not.     

Seasonally of adult male japanese macaques (Macaca fuscata): Androgens and behavior in a confined troop

Seasonal variations in levels of serum testosterone, dihydrotestosterone (DHT), reproductive behavior, and social behavior were investigated in 12 adult males (5 to 20 + years of age) of the Oregon troop of Japanese macaques (Macaca fuscata). Blood samples were collected at 2- to 4-month intervals, and behaviors were monitored twice weekly over a 15-month period. Significant seasonal variations in levels of testosterone and DHT, and in frequencies of mount series, ejaculations, number of female partners, displays, courtship, and aggression were observed. Seasonal variations in reproductive and social behaviors did not correlate with seasonal variations in androgen levels because seasonal increases in these behaviors followed seasonal increases in the androgens with a 1- to 2-month delay. However, significant correlations between increased androgen levels and the onset of mating activity occur when mean monthly frequencies of mount series are shifted 1 to 2 months earlier to coincide with the rise in serum androgen levels. The frequency of adult male play and male-male mounting increased significantly when androgen levels were low. We suggest that photoperiod changes may function as a proximate cue in male Japanese macaques which induces a state of biological readiness for mating, and the behavioral consequences (i.e., mating) are then dependent upon the presence of receptive females.     

Mating Behavior of Cephalonomia tarsalis (Ashmead) (Hymenoptera: Bethylidae) and the Effect of Female Mating Frequency on Offspring Production

The courtship behavior of Cephalonomia tarsalis, a solitary semiectoparasitoid of Oryzaephilus surinamensis, was investigated in the laboratory. Courtship behavior includes a series of stereotypic movements. Males play the most active role, executing the majority of courtship action, and females respond with relatively limited observable behaviors. Males typically keep antennae still during encounters with females prior to mounting, which may be correlated with recognition of the female's sexual status. After mounting, males display a series of movements on females, such as antennae touching female's antennae, antennae or mouth touching female's head or thorax, and walking around on female, which may serve to stimulate females towards increased receptivity. Females signal receptivity by assuming a stereotypical posture of remaining stationary, with head down, and antennae still in front of the body. The male then inserts his aedeagus and the pair copulates. After an average of 40.4 s of copulation, females signal the end of copulation by waving the antennae and moving away from the copulation site. Males continue copulating for a short time after females start moving but dismount soon thereafter. After dismounting, the two wasps move away from each other immediately, and they typically begin grooming. Neither males nor females exhibit mating preference based on mate's mating status in both choice and no-choice tests. The male is polygynous and the mated female can mate multiple times within the first 3 days after starting oviposition. However, female mating frequency does not affect the production of female progeny.     

Effects of male group size,parity, and cycle stage on female chimpanzee copulation rates at Ngogo,Kibale National Park,Uganda

Chimpanzees have complex and variable mating strategies, but most copulations occur when females with full sexual swellings are in parties with multiple males and mate with most or all of those males. Daily copulation rates for fully swollen females vary at different times of a female’s cycle, among females, and across communities and populations. Variation in female age, parity, and cycle stage underlie some of this variation, but possible demographic effects on copulation rates have not been systematically investigated. Demographic variation can affect many aspects of behavior and ecology, including the frequency and success of different mating tactics. Analysis of data from the unusually large chimpanzee community at Ngogo produces two results that are consistent with the hypothesis that demographic variation affects female copulation rates. Copulation rates were high compared with those reported from other research sites, where females had fewer potential mates available. Daily copulation rates of fully swollen females were also positively related to the number of males with whom they associated. Ngogo data also re-confirm results from other studies, of both wild and captive populations, showing that female copulation rates increase during periovulatory periods. This is consistent with the hypothesis that sexual swellings and extended receptivity and proceptivity help to protect females against infanticide by helping to ensure they mate with all potential sires. As at some other sites, parous females at Ngogo copulated at higher rates than nulliparous females. Possible effects of demography on sexual behavior should be considered in assessments of differences between chimpanzees and bonobos and of variation across chimpanzee populations.     

Role of vocalizations in the reproductive cycle of ring doves (Streptopelia risoria): effects of hypoglossal nerve section on the reproductive behavior and physiology of the female

The role of vocalizations in the reproductive cycle of female ring doves was investigated. Two-stage bilateral hypoglossal nerve section (HNS) was performed on adult females to alter their cooing and they were observed for courtship behavior with the same stimulus males used in preoperation behavioral tests. The HNS females showed a reduction in both the vocalization and wing flipping components of nest-coos but no changes in other female courtship behaviors. In addition, the HNS females failed to show the typical male courtship-induced follicular growth observed in sham-operated females. The behavior of stimulus males did not differ between groups and therefore could not account for the failure of follicular growth in the HNS females. These data suggest that the female's performance of nest-coos, or factors associated with it, may stimulate her own follicular growth.     

Mating speed and the interplay between female and male courtship responses inDrosophila melanogaster (Diptera: Drosophilidae)

The objective of this study was to compare measures of general activity and sexual behavior for various genotypes within a strain of Drosophila melanogaster, which had known differences in mating speed. Three inbred lines of D. melanogaster differed significantly in mating speed when tested in female-choice and in single-pair experiments. Analyses of locomotor activity and sexual activity of females and males revealed no significant differences between the inbred lines. An analysis of the interplay between female and male courtship behaviors enabled the examination of signal-response differences between the inbred lines. The inbred lines with intermediate and slow mean mating speed showed a decreased number of significant transitions between female and male behavioral responses. This decrease was more severe in the slow mating line. Further, the intermediate- and slow-mating females and males displayed courtship responses toward signals of the opposite sex that were different from those of the fastmating line. Models of the relationship between behavioral activity and mating speed in Drosophila are discussed and a different explanation for variation in mating speed among the three inbred lines is considered.