The assessment of female reproductive state during courtship and scramble competition in the fiddler crab, Uca paradussumieri

Male fiddler crabs, Uca paradussumieri, mate underground during a 4- to 7-day period each full and new moon. As soon as the tide recedes, males enter the burrows of females that will ovulate the following day ('pre-ovigerous' females). Males copulate with and guard these females until they ovulate. When interrupted by an intruding male, the first male to reach the female is usually able to defend her and successfully mate with her. In fiddler crabs, females mate multiply and there is last male sperm precedence. Before each semi-lunar mating period, male U. paradussumieri were more likely to court females with whom they would later mate than other nearby females with whom they did not mate. This suggests that males collect information on female reproductive state prior to the females becoming ovigerous. In this species, aggression was common between males that courted the same female. When previously courted females were approached by other males, the initial courter attempted to forcefully disrupt the courtship. This behavior may allow males the exclusive use of information on female reproductive condition. It also suggests a type of scramble competition between males over females. Furthermore, it indicates that males are able to locate receptive females prior to their becoming ovigerous. The shorter guarding period observed in this species, as compared with other fiddler crabs, is caused by females rejecting longer guarding periods. Male ability to assess female reproductive status may therefore be advantageous because it increases male mating success within a scramble type of competitive polygyny.     

Mating strategies of a nocturnal,desert rodent (Dipodomys spectabilis)

Summary The mating system of a nocturnal, desert rodent, the banner-tailed kangaroo rat (Dipodomys spectabilis) was studied through direct observation, live-trapping, and radiotelemetry over a 13-month period from August 1986 to August 1987. Mating behavior varied from exclusive matings between male and female neighbors to competitive mate searching and direct male competition. In summer matings and early in a November to May breeding season, males located receptive females and mated exclusively with them without disturbance from other males. As the operational sex ratio changed in favor of males, multiple males converged on an estrous female's territory and competed for access to her. However, an older, experienced male usually monopolized the matings of the same one to two close female neighbors for the entire breeding period, and females mated with the same male neighbor over several estrous cycles. Monopolization of females by neighbor males was facilitated by female relaxation of individual territorial defense. Dominant males spent considerable time in the territories of the females they monopolized before and during mating. This relaxation in territorial defense was seen in dyadic encounters in which females tolerated all males but allowed significantly more contact by neighbor than stranger males. Neighbor recognition, therefore, seems important in coordinating the mating interactions of this solitary rodent.     

Selected polyandry: female choice and inter-sexual conflict in a small nocturnal solitary primate (<Emphasis Type="Italic">Microcebus murinus</Emphasis>)

Sex-specific interests over the maximization of reproductive success lead to an inter-sexual conflict over the optimal mating system in a species. Traditionally, the outcome of this inter-sexual conflict has been studied from the male perspective but it also depends on female mating strategies, such as manipulating the temporal distribution of sexual activity, advertisement, and mate choice. We used a small nocturnal primate, the gray mouse lemur (Microcebus murinus) to determine the relative importance of female mating strategies on the outcome of this conflict in a species where females are solitary during their activity period. We studied their mating behavior over three consecutive annual mating seasons and determined the genetic relationships among more than 300 study animals to quantify individual reproductive success. We found that most females were receptive asynchronously. Females did not exhibit any obvious direct mate choice, probably due to a highly male-biased operational sex ratio and the corresponding costs of choosiness. However, females exercised indirect choice for multiple matings. They mated with 1–7 males up to 11 times during their single night of receptivity. As a result, mixed paternity was common but heavier males sired more offspring, meaning that indirect female choice for superior males cannot be excluded. Females exhibited a mixed mating strategy, avoiding costly direct mate choice but still counteracting male efforts to monopolize mating, successfully increasing genetic variability among offspring. Thus, females had a major influence on the outcome of the inter-sexual conflict despite male monopolization attempts.Communicated by J. Setchell     

Mating effort and female receptivity: how do male guppies decide when to invest in sex?

Males vary in the degree to which they invest in mating. Several factors can explain this variation, including differences in males’ individual condition and the fact that males allocate their energy depending on the context they face in each mating attempt. Particularly, female quality affects male reproductive success. Here, we studied whether male guppies (Poecilia reticulata) strategically allocated more mating effort, in terms of mating behaviour and male–male competition, when they were matched with a receptive (R) female than a non-receptive one. In accordance with our prediction, we found that males increased their mating behaviour when they were with a receptive female. Even though male guppies can inseminate non-receptive females, we only found high levels of courtship between males that were with a receptive female rather than a non-receptive one. Although there was little affect of female receptivity on male–male competition, we found that males chased and interrupted courtships more with receptive females than with non-receptive females regardless of odour. Finally, we also studied whether the sexual pheromone produced by receptive female guppies is a cue that males use in order to increase their mating effort. We found that males were more attracted to a female when they perceived the sexual pheromone, but only increased their mating and aggressive behaviours when females showed receptive behaviour. This strategic increase in mating effort could result in higher male reproductive success because mating attempts towards receptive females are likely to be less costly and males could have a greater probability of fertilisation.     

Reproductive biology of a small isopod symbiont living on a large isopod host: from the maternal marsupium to the protective grip of guarding males

Mating systems of many symbiotic crustaceans are characterised by a high degree of mate guarding. A peculiar case of mate guarding has been reported for small symbiotic janirid isopods where males mate with immature females. Field samples of individual hosts and laboratory experiments were conducted to reveal the mating behaviour of the symbiont in a natural environment, that is, on their hosts. Along the coast of the Magellan Strait, Chile, the janirid isopod Iais pubescens was frequently found on the shore-living isopod Exosphaeroma gigas. Symbiont prevalence (percent hosts occupied) was high at eight of the nine sampling sites. Mean symbiont intensity was very low at one site (<<1 individual host^-1), intermediate at two sites (1-10 individuals host^-1) and high at the other sites (10-40 individuals host^-1). The mean sex ratio (males:females) was male biased at most sampling sites (n=7). Females of I. pubescens reached substantially larger sizes (1.5-3.0 mm body length, BL) than males (1.1-1.9 mm BL). The majority of males were carrying small juveniles (66.15%), and males with juveniles were significantly larger than males without juveniles - this suggests that males prefer virgin juveniles to adult females and that they compete for small juveniles. In laboratory observations, males were seen to manipulate the marsupium of adult females that were about to release small juveniles. Males obtained virgin juveniles in this manner. Juveniles were carried for ~7 days, and they moulted shortly before being fertilised and released by males. The high proportion of juveniles carried by males in the field (68.2%) supports previous observations that males initially are not able to distinguish male and female juveniles. It is suggested that the mating system of symbiotic janirid isopods with long-term sperm storage and continuous receptivity in females and male mating with virgin females has evolved in response to highly unpredictable encounter probabilities between the sexes. Mate guarding and manipulation of small virgin juveniles may be favoured on the highly mobile hosts of symbiotic janirid isopods. Furthermore, adult females may gain by leaving their emerging offspring in the protective grip of guarding males, thereby reinforcing the maintenance of this peculiar mating system.     

Biased sperm use by polyandrous queens of the ant<Emphasis Type="Italic"> Proformica longiseta</Emphasis>

The occurrence and genetic effects of polyandry were studied in the ant Proformica longiseta using three microsatellite markers. The average queen mating frequency (QMF) estimated from the sperm dissected from the spermathecae of 61 queens was 2.4 with 69% of the queens being multiply mated. QMF estimated from worker offspring in a subsample of eight monogynous colonies was 3.5, but the effective paternity (m_e,p) was only 1.23. The difference between these values reflected unequal sperm use by the queens. Most colonies of P. longiseta were polygynous and the average relatedness among workers was 0.35. Polyandry thus added only marginally to the genetic diversity of colonies, and our results gave little support to the genetic-variability hypothesis for explaining polyandry. Diploid male load was low, as only 1% of males were diploid. A large majority (92%) of nests produced one sex only, with males produced in colonies that had higher than average worker relatedness. This contradicted the predictions derived from worker control of sex ratios. Males produced enough sperm to fill the spermathecae of several queens. Thus, the results indicated that diploid male load, sperm limitation and sex ratio conflict are also unlikely explanations of polyandry. Plausible hypotheses for polyandry include mating by convenience, as the sex ratio is male biased and the mating costs to a female can be low because the females are wingless and have no mating flight. The observed unequal sperm use furthermore points to sperm choice and sperm competition as important factors in the evolution of polyandry.     

Tests of the mate-guarding hypothesis for social monogamy: does population density, sex ratio, or female synchrony affect behavior of male snapping shrimp (Alpheus angulatus)?

There are at least two mechanisms by which social monogamy in the absence of biparental care may evolve: as a form of territorial cooperation, in which one or both sexes benefits by sharing a territory with a partner, and as a form of extended mate guarding, in which males guard females through entire, and perhaps multiple, reproductive cycles. I examined the effects of population variables (density, sex ratio, female synchrony) on male pairing behavior in the snapping shrimp, Alpheus angulatus, to test the hypothesis that social monogamy in this genus has evolved as a result of selection on males for long-term mate guarding of females. There was no evidence that pairing behavior changes with differences in population density; in a natural population, there was a 1:1 relationship between the number in pairs and local population density. In a laboratory experiment, males altered their pairing behavior in response to manipulated differences in sex ratio. Males in female-biased sex ratios were significantly more likely to abandon recently mated females than males in equivalent sex ratios, though there was no significant difference in the duration of pairing or the number of times males switched females. Observations of shrimp maintained for an extended period in the laboratory revealed no evidence that females molt and become sexually receptive synchronously, which would reduce the likelihood that a searching male would encounter additional receptive females. These data suggest that sex ratio may have contributed to the evolution of social monogamy in snapping shrimp, but provide no evidence that population density or female synchronous receptivity have contributed to the evolution of social monogamy.     

Bi-directional sex change: testing the growth-rate advantage model

Bi-directional sex change in coral-dwelling fishes (genera Gobiodon and Paragobiodon) has been attributed to a growth-rate advantage for females during the non-breeding season. This model predicts that the smallest individual in a newly formed pair should always be female. To determine if a growth-rate advantage exists for female Gobiodon histrio, I monitored the growth of males and females in natural pairs during the breeding and non-breeding season. I then used a manipulative field experiment to test four predictions of the growth-rate advantage model: (1) the larger individual should change sex to male in new pairs containing two females; (2) the smaller individual should change sex to female in new pairs containing two males; (3) neither individual should change sex in heterosexual pairs where the male is larger than the female; and (4) both individuals should change sex in heterosexual pairs where the female is larger than the male. A growth-rate advantage was detected for female G. histrio during the non-breeding season; however, only the first three of the predicted outcomes were observed in the manipulative experiment. Sex change did not occur in heterosexual pairs where the female was larger than the male. Furthermore, growth did not differ between sex-changed and non-sex-changed fish; therefore, the absence of sex change in these pairs is not due to a growth cost to sex change. I propose that the risk of moving among spatially isolated habitat patches and the low probability of finding a mate have been more important than sex-specific differences in growth rates to the evolution of bi-directional sex change in coral-dwelling gobies.     

Uganda kob mating success does not increase on larger leks

One explanation for the movement of sexually receptive females to clusters of male territories in lek-breeding species is that larger clusters provide females with higher-quality mating partners, as would be the case if males were distributed between leks in an ideal free distribution for unequal competitors. This ideal free model of lek evolution predicts that male competitive ability and mating success will be greater on larger leks than smaller ones. I tested these predictions by comparing the mean number of males on 19 different Uganda kob leks with the sex ratio, the availability of oestrous females, mating rates, male fighting rates and male turnover rates. Contrary to the predictions of the model, numbers of females, receptive females and fights increased proportionally with lek size, but were no greater per male on larger leks. Multivariate analyses of male and female numbers on leks showed that male numbers were associated with female numbers, female numbers in the past, and a variety of habitat variables which may have related to the costs of holding a lek territory, but female numbers varied only with male numbers and female density in the area. These data do not provide evidence that females gain access to superior males by mating on larger leks, though they do support the possibility that lekking may be promoted by a tendency for larger leks to retain females longer.     

Evidence for seasonal mate limitation in populations of an intertidal amphipod, Corophium volutator (Pallas)

Potential rates of reproduction (PRR) differ between the sexes of many animal species. Adult sex ratios together with PRR are expected to determine the operational sex ratio (OSR) defined as the ratio of fertilizable females to sexually active males at any given time. OSR is expected to determine the degree to which one sex competes for another—the limiting sex. We explored the potential for mate limitation in an intertidal amphipod, Corophium volutator (Pallas). Males have higher PRR than females, but males may be limiting because of extreme female-biased sex ratios observed in this species. Consistent with this idea, late season females were less likely to be ovigerous and had smaller size-specific clutches, both of which were associated with seasonal declines in availability of males of reproductive size. Seasonal changes in ovigery could not be explained by seasonal changes across sites in other factors (e.g., female body size or phenology of breeding). Smaller females were less likely to become ovigerous later in the season at three of four sites. Seasonal reductions in clutch size also occurred among small females expected to be reproducing for their first time. In complimentary laboratory experiments, reduced likelihood of ovigery and reduced fecundity occurred when the number of receptive females was increased relative to availability of a reproductively active male. Our results suggest male mate limitation can occur seasonally in this species and that male limitation is regionally widespread and may affect recruitment.     

Gonad morphology, sexual development, and colony composition in the obligate coral-dwelling damselfish Dascyllus aruanus

Gonad morphology and colony composition support the existing supposition that the obligate coral-dwelling damselfish Dascyllus aruanus has a protogynous hermaphroditic sexual pattern. Adults had either an active ovary containing vitellogenic oocytes, an ovotestis, or a spermiated testis and were classified as adult female, hermaphrodite, or adult male, respectively. Among individuals having male function, the testis (or testis portion of the ovotestis) takes the form of an unrestricted spermatogonial lobular testis. Among hermaphrodites having an ovotestis, a small proportion of individuals had a gonad in which both the ovarian and testicular portions were inactive (inactive hermaphrodites), whereas the majority had a predominantly testicular ovotestis that contained spermatozoa (male-active hermaphrodites). The size range of individuals within gonadal classes indicates that all D. aruanus first develop an ovariform gonad. Some individuals then undergo ovarian maturation to become adult females while others develop testicular tissue to form an ovotestis and become male-active hermaphrodites. Subsequently, progressive loss of ovarian tissue results in the development of a secondary testis from an ovotestis with the retention of a residual, afunctional lumen among adult males. The wide size range of individuals having an ovotestis suggests that some hermaphrodites function as adult females before developing testicular tissue while other individuals do not pass through an adult female stage. If this is the case, D. aruanus exhibits a diandric protogynous hermaphroditic sexual pattern. The apparent prolonged retention of an ovotestis with both healthy oocytes and an ovarian-type lumen in a spermiated ovotestis, as well as a functional sex ratio of 1:1 for adult females:adult males plus male-active hermaphrodites also raises the possibility that D. aruanus may be capable of bidirectional sex change during the hermaphroditic stage. Such a capability would be highly adaptive for a species having limited mobility and unpredictable recruitment of new colony members resulting in unpredictable mating opportunities.     

Operational sex ratios and behavioural sex differences in a pipefish population

In the pipefish Syngnathus typhle, only males brood embryos in specially developed brood pouches, supplying oxygen and nutrients. Laboratory studies have shown that this elaborate paternal care has led to sex-role reversal in this species: males limit female reproductive rate, females are the primary competitors for mates and males exercise greater selectivity in accepting mates. In the first field study of this pipefish, we describe mating behaviour in the wild and test the hypothesis that temporal variations in the operational sex ratio (OSR) determine sex differences in mating behaviour. Our study comprised two reproductive seasons of two sequential mating periods each, the latter separated by a lengthy interval of male brooding. During mating periods, females displayed to all males without wandering and males moved about searching for females, without reacting to all females. The OSR was least female-biased (or even male-biased) at the onset of the breeding season, when most pipefish were simultaneously available to mate, but became strikingly female-biased as males' pouches were filled. The OSR remained substantially female-biased during the second mating period, because few males became available to remate at any one time. As hypothesised, female-biased OSRs resulted in more female-female meetings. As well, females were above the eelgrass more often than brooding males, thus exposing themselves to conspecifics and/ or predators. In the second year, males arrived earlier than females on the breeding site and male pregnancies were shorter, because of higher water temperatures, so rematings occurred earlier. Males met more often during that year than the previous one, but male competitive interactions were still not observed. The field results support laboratory studies and demonstrate that behaviours associated with female-female competition are more prominent when the OSR is more female-biased. Correspondence to: A. Vincent     

Parental investment, adult sex ratios, and sexual selection in a socially monogamous seabird

Although most birds are monogamous, theory predicts that greater female parental investment and female-biased adult sex ratios will lower the polygyny threshold. This should result in polygynous mating, unless obligate biparental care or the spatial and temporal distribution of fertilizable females constrains a male’s ability to take advantage of a lowered polygyny threshold. Here we present data on the extent of male sexually dimorphic plumage, adult sex ratios and breeding season synchrony in three populations of a socially monogamous seabird, the brown booby Sula leucogaster. For one of these populations, San Pedro Mártir Island, we also present data on differences in male and female parental investment, mortality and probability of pairing. The extent of plumage dimorphism varied among populations. Sex ratios were female biased in all populations. On San Pedro Mártir Island, parental investment was female biased, females failed more often than males to find a mate, but there was no polygyny. We suggest that on San Pedro Mártir: (1) a period of obligate biparental care coupled with a relatively synchronous breeding season constrained the ability of males to take advantage of a high environmental polygamy potential and (2) the resulting socially monogamous mating system, in combination with the female-biased adult sex ratio, caused females to be limited by the availability of males despite their greater parental investment. Received: 18 November 1999 / Accepted: 24 January 2000     

Mating patterns of woolly spider monkeys,Brachyteles arachnoides: implications for female choice

Summary Woolly spider monkeys show a promiscuous, polygynous mating system in which a receptive female mates with several males, often in rapid succession. Copulation is prolonged with an average duration of 4.1±1.5 (SD) min. Coitus consists of a stationary phase with the male in intromission, followed by stercotypic behaviours of the female which appear to cue and/or accompany male climax and ejaculation. Subadult and adult males show different association patterns with individual females. Subadult males form long-term consortships with particular females while adult males appear strongly attracted to a particular female only when the is receptive. These different behavior patterns of males are viewed as age-specific mating tactics. Males in a mating aggregation show little intermale aggression for sexual access to a receptive female. Large testis size in this species suggests that much intermale competition for reproductive success may be carried out at a postcopulatory level, perhaps by sperm competition. The copulatory pattern of woolly spider monkeys may function primarily as a mechanism of female choice, aiding a female in assessing the quality of males in hei mating aggregation while helping to ensure that maximal high quality spermatozoa will be available at the proper time for fertilization.     

Adult female hamsters avoid interspecific mating after exposure to heterospecific males

When females mate with a heterospecific male, they do not usually produce viable offspring. Thus, there is a selective pressure for females to avoid interspecific mating. In many species, females innately avoid heterospecific males; females can also imprint on their parents to avoid later sexual interactions with heterospecific males. However, it was previously unknown whether adult females can learn to discriminate against heterospecific males. We tested the hypothesis that adult females previously unable to avoid interspecific mating learn to avoid such mating after being exposed to heterospecific males. Syrian hamster (Mesocricetus auratus) females not previously exposed to Turkish hamster (Mesocricetus brandti) males can discriminate between odors of conspecific and heterospecific males, but they mate with either type of male. However, when we exposed adult females to both a conspecific male and a heterospecific male through wire-mesh barriers for 8 days, and then paired them sequentially with the two males, females were more receptive to conspecific males and more aggressive to heterospecific males. When females were paired with the heterospecific male first and the conspecific male second, no female was receptive and all were aggressive to heterospecific males. When females were paired with the conspecific male first, only 43% of females were then aggressive toward the heterospecific male. That is, interactions with conspecific males may decrease a female’s ability to properly avoid heterospecific males. Our study clearly shows for the first time that females can learn during adulthood to avoid interspecific mating just by being exposed to stimuli from heterospecific males.     

Attraction and learning in mate-finding by solitary bees,Lasioglossum (Dialictus) figueresi Wcislo and Nomia triangulifera Vachal (Hymenoptera : Halictidae)

Summary Males of the solitary sweat bee, Nomia triangulifera, patrol over large areas where thousands of females emerge, searching for receptive females. The daily operational sex ratio is strongly male-biased. Males contact dead, frozen (untreated) females more frequently than they contact females which were washed in hexane, showing that olfactory cues are utilized in mate-finding. A major source of female sex pheromone is in the head. Male pouncing on females is temporally non-random, indicative of group stimulation. Bioassays show that newly emerged females are more attractive to males than are older pollen-collecting females. Female odors are individually distinctive, based on male responses, and there is much variation among females in their attractive properties. Male responses to female odors suggest that learning is important for mating in natural populations. In contrast, the following hypotheses are unlikely to account for the observed behavior: (1) dissipation of female odors; (2) site learning and avoidance behavior by males; (3) decay of male motivation; or (4) male-produced repellents effective against other males. Laboratory and field studies show that female Lasioglossum figueresi produce individually-distinctive odors, which are attractive to males. There is considerable inter-individual variation among females in their attractiveness to males among sexually immature females. Male responses to female odors decay over the course of the presentation, suggesting the importance of learning in natural populations, although several alternatives could not be tested.     

Swarming and mating behavior of a mayfly Baetis bicaudatus suggest stabilizing selection for male body size

Large size often confers a fitness advantage to female insects because fecundity increases with body size. However, the fitness benefits of large size for male insects are less clear. We investigated the mating behavior of the mayfly Baetis bicaudatus to determine whether the probability of male mating success increased with body size. Males formed mating aggregations (swarms) ranging from a few to hundreds of individuals, 1-4 m above the ground for about 1.5-2 h in the early morning. Females that flew near swarms were grabbed by males, pairs dropped to the vegetation where they mated and then flew off individually. Some marked males returned to swarms 1, 2 or 3 days after marking. Larger males swarmed near spruce trees at the edges of meadows, but the probability of copulating was not a function of male body size (no large male advantage). Furthermore, the potential fitness advantage of mating with larger, more fecund females was not greater for large males (no size-assortative mating). However, the sizes of copulating males were significantly less variable than those of non-mating males collected at random in swarms. Intermediate male size may be optimal during mating because of trade-offs between flight agility and longevity or competitive ability. Results of this study are consistent with the hypotheses that there is stabilizing selection on adult male body size during mating, and that male body size in this species may be influenced more by selection pressures acting on larvae than on adults.     

Intrasexual selection in Mirza coquereli : evidence for scramble competition polygyny in a solitary primate

The primates of Madagascar (Lemuriformes) deviate from fundamental predictions of sexual selection theory in that polygynous species lack sexual dimorphism, have even adult sex ratios and often live in female-dominated societies. It has been hypothesized that intrasexual selection in these species is either reduced or primarily focused on traits related to scramble competition. The goal of this study was to examine these hypotheses by studying the mating system of a solitary nocturnal species, Mirzacoquereli. During a 4-year field study in western Madagascar, I captured and followed 88 individually marked animals. I found that adult males were significantly larger than females, providing the first evidence for sexual size dimorphism in lemurs. In addition, the adult sex ratio was biased in favour of females in 3 out of 4 years. There was no significant sex difference in canine size, however. Males showed pronounced seasonal variation in testis size with a 5-fold increase before and during the short annual mating season. During the mating season, males had more injuries than females and more than quadrupled their home ranges, overlapping with those of more than ten females, but also with about the same number of rivals. Only about one social interaction per 10 h of observation was recorded, but none of them were matings. Together, these results indicate that these solitary lemurs are clearly subject to intrasexual selection and that male-male competition is primarily, but not exclusively, of the scramble type. In addition, they suggest that the above-mentioned idiosyncracies may be limited to group-living lemurs, that social systems of solitary primates are more diverse than previously thought, and that the temporal distribution of receptive females is responsible for this particular male mating strategy. Received: 11 January 1997 / Accepted after revision: 18 April 1997     

Mating group composition influences somatic costs and activity in rutting dominant male reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>)

In polygynous species, males devote considerable effort to reproduction during the rut. Both the number of females in the mating group and the ratio of sexually mature males to sexually mature females [adult sex ratio (ASR)] are expected to affect the amount of effort a male devotes to reproductive activities. We predicted the reproductive effort of dominant male reindeer, measured as relative mass loss, proportions of active reproductive behaviors, and frequencies of agonistic behaviors would (1) increase with an increasing number of females in the mating group and eventually level off, and (2) exhibit a dome shape with respect to ASR in the mating group. We tested these predictions using 12 years of data collected from semi-domesticated reindeer in northern Finland. We found a positive relationship between relative mass loss and the mean number of females in the mating group for mature, but not young males. The relationship between the proportion of active reproductive behaviors performed by mature males and the mean number of females in the group was quadratic while agonistic behaviors of mature males increased with the increasing female group size. We also found that active reproductive behaviors decreased with a rising mating group ASR for mature males; whereas, young males performed more agonistic behaviors as group ASR increased. Our results point to age-specific patterns of mass loss and activity during the mating season. They also indicate that both the number of females and ASR in the mating group are important factors in determining the level of reproductive effort of dominant male reindeer.     

Frequency-dependent male mate harassment and intra-specific variation in its avoidance by females of the damselfly Ischnura elegans

We focused on male harassment on different female color morphs of the damselfly Ischnura elegans and on variation in morph-specific mating avoidance tactics by females. In I. elegans, one of the female morphs is colored like the conspecific male (andromorphs) while the other morphs are not (gynomorphs). Our first goal was to quantify morph-specific male mating attempts, hence male harassment, in populations with manipulated population parameters (densities, sex ratios, and proportion of andromorphs). Second, we examined the female's perspective by looking for potential differences in morph-specific mating avoidance tactics and success of those tactics in a natural population. Differences in population conditions did influence the number of male mating attempts per morph. The less frequent female morph was always subject to fewer mating attempts, which contradicts earlier hypotheses on mimicry, but supports those that assume that males learn to recognize female morphs. Gynomorphs occupy less open habitat and often fly away when a male approaches, while andromorphs use more open habitat, do not fly large distances and directly face approaching males. Female morphs did not differ in the proportion of successful mating-avoidance attempts. Our results suggest that the maintenance of the color polymorphism is most probably the result of interactive selective forces depending on variation in all population conditions, instead of solely density- or frequency-dependent selection within populations.