Effect of pairing in vitro on the glutathione level of male Schistosoma mansoni

The effect of in vitro incubation on the level of the intracellular nucleophile, glutathione (GSH), in adult Schistosoma mansoni was investigated. The GSH levels of freshly collected adult male and female parasites were 8.5 +/- 2.5 and 2.7 +/- 0.7 nmol/10 worms, respectively, as determined by an enzymatic assay. Twenty-four-hour incubation of unpaired males in RPMI-1640 medium at 37 C resulted in a 1.7-fold increase (P less than 0.001) in GSH level that remained elevated for at least 7 days. The increase was dependent on exogenous L-cystine, suggesting that it was due to biosynthesis of GSH. Biosynthesis in male S. mansoni was confirmed by isolating [3H] GSH from parasites incubated in medium containing L-[3H] cystine or [3H] glycine. In contrast to unpaired males, the GSH level of paired males as well as that of unpaired or paired females did not increase after 24 hr in vitro. When males that had been incubated unpaired for 24 hr were allowed to couple in vitro with freshly collected females, their GSH level fell to that of continuously paired males. These observations provide evidence that in vitro female schistosomes can influence the physiology of the male.     

Male Distribution and Mate Searching in the Yellow Dung Fly Scathophaga stercoraria: Comparison between Paired and Unpaired Males

Movements by individual males were examined in the yellow dung fly, Scathophaga stercoraria. Males were observed from their arrival until they found a female (paired males) or departed (unpaired males). The focal variables were the male mating status, body size and the number of males and females at the site. Paired males, independent of size, spent more time in the best mate-searching area (pat and the first up-wind zone) than unpaired males. Paired males in all size classes moved around and attacked other males more often than unpaired males. Among paired males, males that caught a single female and those that took over a female from another male were very similar in their mate-searching behaviour. The total time spent searching at the pat was positively related to its resource value as indicated by the number of pairs. Time spent in the best mate-searching area was negatively related with male numbers. The causes of differences in movements and aggression between paired and unpaired males are discussed. The male distribution around cow pats can be understood only if the differences in movement patterns by paired and unpaired males are taken into account.     

Male Defense of the Breeding Cavity and Factors Affecting the Persistence of Breeding Pairs in the Stomatopod,Gonodactylus bredini (Manning) (Crustacea: Hoplocarida)

In Caribbean Panama, nonreproductive male and female stomatopods are solitary and defend their own coral-rubble cavities. When breeding pairs form, however, males assume all responsibility for cavity defense. To compare success in cavity defense and defensive tactics among paired and unpaired males, and to examine the tendency for paired stomatopods to exchange their present mates for larger (higher quality) individuals, we introduced same-sized and 15% larger male, and same-sized and 15% larger reproductive female intruders to paired and unpaired male residents in a balanced design. Paired males were more successful at cavity defense than unpaired males, evidently because paired males strike intruders more than unpaired males, and because intruders fight less intensely against paired males than against unpaired males. Paired males occasionally attempted extrapair copulations, but showed little tendency to abandon their mates in favor of larger females. Paired females, however, mated readily with intruder males that evicted resident males. Populationwide female breeding synchrony and prolonged female receptivity before oviposition reduce variance in male mating success and may force males to guard the breeding cavity to assure their paternity. Uncertainty about the reproductive condition of intruder females may prevent males from exchanging mates.     

Pairing success and sperm reserve of male Gammarus pulex infected by Cyathocephalus truncatus (Cestoda: Spathebothriidea)

Manipulative parasites with complex life cycles are known to induce behavioural and physiological changes in their intermediate hosts. Cyathocephalus truncatus is a manipulative parasite which infects Gammarus pulex as intermediate host. G. pulex males display pre-copulatory mate guarding as a response to male-male competition for access to receptive females. In this paper, we tested the influence that C. truncatus-infection might have on male G. pulex sperm number and pairing success. We considered 3 classes of G. pulex males in our experiments: (i) uninfected males found paired in the field, (ii) uninfected males found unpaired in the field, or (iii) infected males found unpaired in the field. Both infected males and uninfected unpaired males paired less with a new female than uninfected paired males did. Furthermore, infected males appear to be at a strong disadvantage when directly competing for females with a healthy rival male, and had fewer sperm in their testes. We discuss the potential effect of male and female mating strategies on such male host mating alteration.     

Plasma steroid hormones in relation to behavioral sex role reversal in the spotted sandpiper, Actitis macularia

Plasma samples collected from spotted sandpipers during the reproductive season were analyzed for testosterone, dihydrotestosterone (DHT), estradiol-17 beta and progesterone. Prior to incubation, plasma testosterone and DHT levels were significantly greater in males than in females. Estradiol levels of paired females were significantly greater than those of paired males. Testosterone and DHT levels of unpaired resident and paired males were significantly greater than those of incubating and brooding males. A 25-fold decline in testosterone occurred in males from the 1- or 2-egg stage to the 3-egg stage, when incubation is initiated. In females, testosterone values were low in unpaired, brooding, and transient birds. Paired females had levels 7-fold greater than unpaired birds. In both sexes, there was a strong correlation between testosterone and DHT levels. Prolactin values were negatively correlated with testosterone and DHT in males. These results indicate that the high level of intrasexual competition for mates among female spotted sandpipers is not based upon a total reversal of the normal male/female levels of androgens and estradiol. Territoriality and intense competition for mates in females may be based upon enhanced receptivity of neural centers to moderate hormone levels. Relative changes in testosterone between unpaired and paired females indicates that this hormone may play a role in mate acquisition and territoriality of these sex role-reversed females.     

Increased vigilance of paired males in sexually dimorphic species: distinguishing between alternative explanations in wintering Eurasian wigeon

In animal pairs, males are often more vigilant than females.This is generally assumed to result from mate guarding (eitheragainst predators or other males). However, when males haveconspicuous secondary sexual characteristics, they could beconstrained to be more vigilant because of a higher predationrisk than females. We attempted to distinguish between the "maleconstraint hypothesis" and two variations of the mate-guardinghypothesis by studying the vigilance behavior of the sexuallydimorphic wigeon during early winter, when some males are inbreeding plumage and some are not and when not all males arepaired. The proportion of time spent vigilant by paired malesin breeding plumage was five times higher than any other categoryof males or females. We found no significant differences betweenthe vigilance levels of unpaired male wigeon in cryptic andin breeding plumage and therefore rejected the male constrainthypothesis. As vigilance levels of paired and unpaired femalesdid not differ either, we rejected the hypothesis that pairedmales invest in vigilance to reduce their mate's need to bevigilant to predation risks. Paired females interacted lessfrequently with other wigeon than unpaired ones, and it is probablyto protect their female from other males that paired male wigeonincrease their vigilance times.     

Copulation patterns in the golden orb-web spider Nephila madagascariensis

A consequence of multiple mating by females can be that the sperm of two or more males directly compete for the fertilisation of ova inside the female reproductive tract. Selection through sperm-competition favours males that protect their sperm against that of rivals and strategically allocate their sperm, e.g., according to the mating status of the female and the morphology of the spermatheca. In the majority of spiders, we encounter the otherwise unusual situation that females possess two independent insemination ducts, both ending in their own sperm storage organ, the spermatheca. Males have paired mating organs, but generally can only fill one spermatheca at a time. We investigated whether males of the African golden orb-web spider Nephila madagascariensis can prevent rival males from mating into the same spermatheca and whether the mating status of the female and/or the spermatheca causes differences in male mating behaviour. There was no significant difference in the duration of copulations into unused spermathecae of virgin and mated females. We found that copulations into previously inseminated spermathecae were generally possible, but shorter than copulations into the unused side of mated females or with virgins. Thus, male N. madagascariensis may have an advantage when they mate with virgins, but cannot prevent future males from mating. However, in rare instances, parts of the male genitals can completely obstruct a female genital opening.     

Postcopulatory female choice increases the fertilization success of novel males in the field cricket, Gryllus vocalis

Although recent studies have demonstrated that female crickets prefer novel males to previous mates, the relative contribution of pre- and postcopulatory behaviors to this advantage remain unknown, as do the reproductive consequences to males. I paired females either with previous or novel mates, and recorded the latency to mating and the time after mating at which the female removed the male's spermatophore, terminating sperm transfer. Females that mated with familiar males removed their spermatophores sooner than females that mated with novel males. Females paired with novel males also mated more quickly than females paired with familiar males, but this difference was not statistically significant. A molecular-based paternity analysis was used to determine whether the postcopulatory preference of females for novel males influences a male's fertilization success. Females were assigned to either mate three times with the same male and then once with a novel male, or four times with four different males. The paternity of the last male was higher when the female previously had mated repeatedly with the same male than when she had mated previously with different males. These results suggest that female spermatophore removal behavior influences male paternity such that novel males receive a fertility benefit.     

State-dependent pairing behaviour in male Gammarus pulex (L.) (Crustacea, Amphipoda): effects of time left to moult and prior pairing status

Because mating can be costly in terms of time and energy, an individual's propensity to engage in courtship and mating activities might be modulated by its physiological state. However, so far, state-dependent mate choice has received little attention The present study examined the effect of both prior pairing status and time left to the moult on the ability of male Gammarus pulex (Crustacea, Amphipoda) to enter in precopula with receptive females. In the lab, males that were freshly collected in precopula pairs in the field had a higher probability of re-pairing and were quicker to enter in precopula with receptive females compared to males of similar size that were freshly collected unpaired. In addition, unpaired males found in the field were closer to their moult than paired males. Considered together, our results strongly suggest that time left to the moult and prior mating status directly influence male propensity to pair in G. pulex.     

Female mating status affects male mating tactic expression in the wolf spider Rabidosa punctulata

Males and females have conflicting interests on the frequency and outcomes of mating interactions. Males maximize their fitness by mating with as many females as possible, whereas choosy females often reduce receptivity following copulation. Alternative male mating tactics can be adaptive in their expression to a variety of mating contexts, including interactions with a relatively unreceptive mated female. Male Rabidosa punctulata wolf spiders can adopt distinctive mating tactics when interacting with a female, a complex courtship display, and/or a more coercive direct mount tactic that often involves grappling with females for copulation. In this study, we set up female mating treatments with initial trials and then paired mated and unmated females with males to observe both female remating frequencies and the male mating tactics used during the interactions. Males adopted different mating tactics depending on the mating status of the female they were paired with. Males were more likely to adopt a direct mount tactic with already-mated females and courtship with unmated females. Already-mated females were considerably less receptive to males during experimental trials, although they did remate 34% of the time, the majority of which were with males using a direct mount tactic. Whereas males adjusting to these contextual cues were able to gain more copulations, the observation of multiple mating in female R. punctulata introduces the potential for sperm competition. We discuss this sexual conflict in terms of the fitness consequences of these mating outcomes for both males and females.     

Variable Social Mating System in the Sedge Warbler,Acrocephalus schoenobaenus

We studied the mating system in a local population of colour-ringed sedge warblers in south Central Sweden in 1990–92. Of 58 territorial males, 59% were socially monogamous, 14% socially polygynous and 27% unpaired. Socially polygynous males in general paired with two females: the only exception was one male that formed pair bonds with three females. Among the males that formed a pair bond, 38% resumed singing after their first female had started egg-laying or incubation and 50% of the paired males that resumed singing succeeded in attracting a second female. Hence, despite a consistently male-biased sex ratio in the population a large proportion of the males tried to become polygynous and they were often successful. The frequency of extra-pair matings did not differ between monogamous and polygynous males. Of 47 breeding females, 6.4% were sequentially socially polyandrous. In two out of three cases, the females fed the young of their first broods until independence before initiating the second brood. In the third case the female deserted her newly fledged young and these were instead cared for by a neighbouring male. DNA fingerprinting revealed that this male had not sired any of these young. Each of the sequentially polyandrous females successfully raised both their broods, and their annual reproductive success was slightly higher than the average for the polygynous males. When the sequentially socially polyandrous females initiated their second brood, their primary male (in all cases polygynous males), cared for young in their secondary female's nest. In all cases, the sequentially polyandrous females formed second pair bonds with unpaired males that were close neighbors. This suggests that females switched pair male for their second brood to obtain a mate that was more likely to provide them with direct benefits (e.g. parental care).     

Low mate encounter rate increases male risk taking in a sexually cannibalistic praying mantis

Male praying mantises are forced into the ultimate trade-off of mating versus complete loss of future reproduction if they fall prey to a female. The balance of this trade-off will depend both on (1) the level of predatory risk imposed by females and (2) the frequency of mating opportunities for males. We report the results of a set of experiments that examine the effects of these two variables on male risk-taking behavior and the frequency of sexual cannibalism in the praying mantis Tenodera sinensis. We experimentally altered the rate at which males encountered females and measured male approach and courtship behavior under conditions of high and low risk of being attacked by females. We show that male risk taking depends on prior access to females. Males with restricted access to females showed greater risk-taking behavior. When males were given daily female encounters, they responded to greater female-imposed risk by slowing their rate of approach and remained a greater distance from a potential mate. In contrast, males without recent access to mates were greater risk-takers; they approached females more rapidly and to closer proximity, regardless of risk. In a second experiment, we altered male encounter rate with females and measured rates of sexual cannibalism when paired with hungry or well-fed females. Greater risk-taking behavior by males with low mate encounter rates resulted in high rates of sexual cannibalism when these males were paired with hungry females.     

Repeatability of Male Mate Choice in the Mosquitofish, Gambusia holbrooki

Male mate choice has evolved in many species in which female fecundity increases with body size. In these species, males are thought to have been selected to favour mating with large females over smaller ones, thereby potentially increasing their reproductive success. While male mate choice is known to occur, it is less well studied than female mate choice and little is known about variation in mating preference among individual males. Here, we presented individual male eastern mosquitofish ( Gambusia holbrooki ) with paired females that differed in body size, and we quantified their mate preference on two consecutive days, allowing us to assess repeatability of preferences expressed. When males were allowed to view paired stimulus females, but not to acquire chemical or tactile cues from them, they exhibited a strong preference for large females over smaller ones. However, individual males were not consistent in the strength of their preference and repeatability was not significant. When individual males were allowed to fully interact with pairs of females, the males again exhibited a preference for large females over smaller ones, as revealed by a greater number of attempted copulations with large females than with smaller ones. In the latter social context, individual male preference was significantly repeatable. These results indicate that male eastern mosquitofish from our Florida study population possess, on average, a mating preference for larger females and that this preference is repeatable when males socially interact freely with females. The significant repeatability for mating preference, based on female body size, obtained for male mosquitofish in the current study is consistent with the presence of additive genetic variation for such preferences in our study population and thus with the opportunity for the further evolution of large body size in female mosquitofish through male mate choice.     

Do Male Veiled Chameleons,Chamaeleo calyptratus,Adjust their Courtship Displays in Response to Female Reproductive Status?

Variation in male courtship behavior may be due to inherent differences among males or may arise from males adjusting their courtship displays according to female responsiveness. Female veiled chameleons, Chamaeleo calyptratus , exhibit two distinctive suites of body coloration and behavior patterns that vary according to receptive and non-receptive stages of their reproductive cycle. We presented male chameleons with both receptive and non-receptive females, and recorded differences in their mating frequency, courtship intensity and courtship behavior patterns. As expected, males were more likely to court and attempt mating with receptive females. Although fewer males courted non-receptive females, their courtship displays were significantly longer than those directed towards receptive females. Males also adjusted the contents of their displays according to female reproductive condition. Certain behavior patterns were unique to courtship displays directed towards each class of females. Males exhibited the behavior pattern `head roll' only when paired with receptive females, and `chin rub' was displayed only during courtship of non-receptive females. We hypothesize that these differences in male courtship frequency, intensity and content reflect differences in female reproductive value. Although males may benefit from mating with both receptive and non-receptive females, the costs associated with courtship may depend on female responsiveness. Thus, males adjust their courtship tactics accordingly.     

Male Age Affects Female Mate Preference and Reproductive Performance in the Cabbage Beetle, <Emphasis Type="Italic">Colaphellus bowringi</Emphasis>

The influence of male age on female mate preference and reproductive performance in the cabbage beetle, Colaphellus bowringi was examined, using male and female adults of varying ages (young, middle-aged and old) after a single mating. In a simultaneous choice test, females of all age class preferentially mated with middle-aged males. Mating duration was positively related to male age. Longevity of females was not significantly affected by male age. Young females paired to middle-aged males had significantly higher egg production than those paired to old males, and the eggs of females paired to middle-aged males exhibited significantly higher hatching success than the eggs of females mated to young or old males. These results suggest that middle-aged males are more advantageous for female fitness than young and old males.     

Behavioural and genetic assessment of reproductive success in a spawning aggregation of the Australian giant cuttlefish, Sepia apama

The only known spawning aggregation of cuttlefish occurs in winter in southern Australia. The operational sex ratio in this aggregation is highly skewed towards males (range 11:1 to 4:1). Using SCUBA, we videotaped females as they mated with up to six males/h. Females rejected and accepted mates of any status or size. We obtained individual eggs and tissue samples from females and their mates and determined paternity by microsatellite DNA analysis. Thirty-nine eggs were sampled during 17 observational sequences (trials); 22 eggs were sired by a sampled male. One-third of the females mated with multiple males and 67% of females' eggs had multiple sires (N=9). Given our restricted sampling, these results probably underestimate actual levels of multiple paternity. Males competed to guard and copulate with females, and spent most of the copulation time flushing water into the female's buccal area. Thus, we predicted a high probability of fertilization of the next-laid egg(s) by the last male to mate, but this prediction was not statistically supported by our data. Fertilization success did not differ significantly between paired consorts (36%) and unpaired males (31%), or between large (33%) and small males (29%). Females used sperm from paired, unpaired and sneaker males as well as from previous matings to fertilize eggs. Our results indicate that this mating system has a high level of multiple mating and multiple paternity and that males of any size or status can obtain successful fertilizations.     

Adaptive significance of winter pair bond in male pintail, Anas acuta

To test the female-advantage hypothesis that has been proposed to explain the adaptive significance of winter pair bonds in ducks, we examined the feeding and social behaviors of the northern pintail, Anas acuta. The female-advantage hypothesis assumes that male attendance offers paired females the benefits of increased social status and access to food, as well as less harassment from conspecifics, allowing them to spend more time feeding. Paired females dominated unpaired females, but neither time budgets of feeding nor frequency of feeding was significantly different between unpaired and paired females. The female-advantage hypothesis predicts that paired males spend less time feeding because they must closely guard their partners from harassment by male conspecifics. Paired males defended their mates by chasing and pecking the unpaired males. However, both time budgets of feeding and frequency of feeding were significantly higher in paired males than in unpaired males. Unpaired males frequently approached females while swimming. They performed courtship displays, mostly toward unpaired females. Paired males spent more time feeding by saving time and energy in courtship. We consider that the advantage of winter pairing for males comes from having a mate plus having an increase in feeding frequency. Received: February 20, 2000 / Accepted: May 22, 2000     

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

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

Mate Switching in a Non-monogamous Species? The Case of the Common Quail (Coturnix coturnix)

We investigated the occutrence of mate switching in the common quail, a non-monogamous species with a temporal pair bond but without either male parental care or territoriality. The study was carried out throughout the breeding seasons of 1993–95 in Mas Esplugues (Catalonia, Spain) by monitoring 28 radio-tagged couples and 17 radio-tagged unpaired males. Agonistic interactions between males inside funnel traps containing a female were also recorded. Mate switching within the laying period occurred in 72% of the females, and new partners always had a higher body condition index than old partners. Agonistic interactions inside the funnel traps also showed that successful males had a better body condition index than losers. These results, along with the observed mate-guarding behaviour of females by their partners throughout the laying period, a highly male-biased sex ratio (five males per female), the lack of territoriality of males and the expected difficulties which unmated males experience in finding pairs, suggest that mate switching is not induced by paired males. Moreover, the constant inflow of new males observed throughout the fertile period of the female and the low costs stemming from mate change strongly support the idea that it is paired females who induce mate switching, in order to improve their fitness by mating with the best quality male available at every moment of their fertile period.     

Non-random Mating in the Dance Fly Empis borealis: The Importance of Male Choice

In the dance-fly Empis borealis (Diptera, Empididae), females form swarms to which males, carrying a nuptial gift, come for mating. We examined whether males or females were choosy and/or competed for mates. First, measurements of the size relationships between copulating males and females, nuptial gifts and the swarming females from different swarms were assessed. Second, male visiting time in differently sized female swarms was recorded. Larger (wing-length) females participated disproportionately in copulations in each swarm, but not for the population at large. Female mating status (virgin/non-virgin) or proximity to oviposition (egg size) did not influence the likelihood of copulation. No assortative mating pattern was found: male size and size of nuptial gift did not correlate with size of the mating female. The time spent by males in swarms increased with the number of females present and it took longer when males left a swarm without copulation than when doing so. Male visiting time per female was negatively correlated with number of females in swarms. Males more often left smaller than larger swarms without mating. We conclude that E. borealis males discriminate among females but find no evidence for male competition or for female choice. It is still a question to what degree females compete for males.