Time constraints and multiple choice criteria in the sampling behaviour and mate choice of the fiddler crab,Uca annulipes

Active female sampling occurs in the fiddler crab Uca annulipes. Females sample the burrows of several males before remaining to mate in the burrow of the chosen partner. Females time larval release to coincide with the following nocturnal spring tide and must therefore leave sufficient time for embryonic development after mating. Here we show how this temporal constraint on search time affects female choosiness. We found that, at the start of the sampling period (when time constraints are minimal), females selectively sample the larger males in the population. Towards the end of the sampling period (when the temporal constraints increase the costs of sampling), females are less selective. Furthermore, we suggest that the number of males sampled (and other indices of ‘‘sampling effort’’) may not be reliable indicators of female choosiness and may not reflect the strength of female mating preferences under certain conditions. Burrow quality also emerged as an important criterion in final mate choice. Burrow structure potentially influences reproductive success, and mate acceptance based on burrow structure appears to involve a relatively invariant threshold criterion. Since there is no relationship between male size and burrow quality, females are using at least two independent criteria when choosing potential mates. We envisage mate choice as a two-stage process. First, females select which males to sample based on male size. They then decide whether or not to mate with a male based on burrow features. This sampling process explains how two unrelated variables can both predict male mating success. Received: 23 March 1995/Accepted after revision: 14 January 1996     

Female reactions to male absence after pairing in the pied flycatcher

Mating with an already mated pied flycatcher (Ficedula hypoleuca) male is costly for a female. Two hypotheses explain why some females still mate with already mated males. The deception hypothesis suggests that some females mate with already mated males since it is difficult to assess perfectly the mating status of a male with separate territories (polyterritoriality). The search cost hypothesis states that females are aware of male mating status but the costs of searching for an unmated male exceed costs associated with the status of secondary female. One potential cue that could disclose a male's mating status is the existence of brief visits to the primary territory by polyterritorial males. To mimic such visits I removed the male from the territory for 60 min soon after female settlement. Only few females abandoned their mates as a consequence of male removal even though they had available unmated males close by. This result is most consistent with the deception hypothesis. Females did not use occasional male absence as a cue to avoid presumably polyterritorial males and in this respect they were not perfect in assessing males.     

Female mate selection in the natterjack toad: active choice or passive atraction?

Summary Variation in seasonal mating success among male natterjack toads (Bufo calamita) was influenced by the number of nights that males spent at the pond and by male body size. Large males produced louder and lower frequency calls than small males, and maintained larger acoustic territories. After arriving at the pond, one half of all observed females mated with the first male they encountered. The remainder visited several males before initiating amplexus, but no criteria could be identified that females might have used when deciding whether to accept or reject a male. Movements between several males seem to be best explained by low female responsiveness to male advertisement calls on cold nights which were nonoptimal for oviposition. Females attempted to reject non-calling males both before and after amplexus, but this may be a mechanism to avoid mismating with males of the common toad (Bufo bufo), an explosive breeder that utilised the same pond. In two-choice playback experiments using synthetic advertisement calls, females showed no preference for calls based on their frequency. Females preferred calls of intermediate pulse repetition rate equivalent to those produced by a male at the same body temperature. Pulse rate is thus potentially subject to stabilishing selection and may be an important character for species-recognition. Females preferred fast rather than slow call rates, but only when the alternative rates were extreme. They also preferred calls which they perceived at the highest sound pressure level, but did not discriminate between absolute sound pressure levels of alternative stimuli at different distances. Since females that delay mating and oviposition may suffer predation, it is suggested that female preference for loud, rapidly repeated calls may be adaptive in the sense of minimizing the costs of locating conspecific males, rather than maximizing the probability of obtaining a high quality mate. Competition between males to maintain large acoustic territories and produce calls that can be easily detected by females would seem to be a sufficient mechanism to explain the evolution of the striking calls produced by male natterjacks.     

Why do female pied flycatchers mate with already mated males: deception or restricted mate sampling?

The polygyny threshold model suggests that females make an optimal choice between mated and unmated males. However; in birds in which males provide parental care, the fitness of secondary females is often lower than expected from this model. This has been explained by the deception hypothesis, which states that males hide their mating status and deceive females into polygyny. Yet there is no direct evidence that secondary females are unaware of male mating status when they settle. Alternatively, females settle with mated males as a result of mate competition and costs of searching. We used videofilming at nestboxes defended by males to study mate sampling of female pied flycatchers Ficedula hypoleuca. The females visited on average only 2.74 males (range 1–8, n = 43). Most (16 of 19) of the polygynous matings occurred because females had only visited mated males, or the unmated males visited became occupied by competitors during the sampling period. Among females that could choose between both mated and unmated males, the majority (13 of 16) settled with unmated males. These results lend little support to the deception hypothesis but are consistent with the view that females are able to detect male mating status but sometimes settle with mated males because of cost of searching. Prospecting females seemed willing and able to suffer the cost of fighting with aggressive primary females in the males' secondary territory if no alternative mating options were available. In addition to male mating status, females took male quality (plumage colour, age) into account in mate choice but the former appeared to be the more important. Correspondence to: T. Slagsvold     

Mate choice in the brown thornbill (<Emphasis Type="Italic">Acanthiza pusilla</Emphasis>): are settlement decisions,divorce and extrapair mating complementary strategies?

In socially monogamous birds, females may express mate preferences when they first select a breeding partner, through divorce and subsequent breeding dispersal to a new partner and through extrapair mating. We examined settlement patterns, divorce and breeding dispersal in a sedentary Australian passerine, the brown thornbill (Acanthiza pusilla), in relation to two traits known to influence extrapair paternity (male age and male size). Settlement decisions, divorce and territory switching behaviour were all female strategies that reduced their likelihood of breeding with 1-year-old males. Females preferred to settle in territories with 2+ -year-old males, were more likely to divorce 1-year-old males, and only switched territories if they had an opportunity to form a new pair bond with an old male. In contrast, female settlement and divorce decisions were not influenced by male size. Female thornbills obtain a direct benefit from preferring older males as social mates because breeding success improves with male age in brown thornbills. Nevertheless, divorce rates in this species were low (14% of pair bonds were terminated by divorce), and individuals rarely switched territories following the death of a mate. Both of these mating strategies appeared to be primarily constrained by the distance adults moved to initiate a new pair bond (1–2 territories) and by the limited availability of unpaired older males in the immediate neighbourhood.Communicated by M. Webster     

Female tactics to reduce sexual harassment in the Sumatran orangutan (Pongo pygmaeus abelii)

Orangutan males demonstrate intrasexual dimorphism with corresponding alternative mating strategies. Sexual harassment is the predominant feature of the mating strategy that subadult males pursue. This study investigated the countertactics that females employ to reduce sexual harassment by subadult males. I observed 207 copulations during more than 9,000 h of focal follows of wild Sumatran orangutans at the Suaq Balimbing Research Station over a 23-month period. Rates of copulations initiated by subadult males increased during months of high fruit abundance, and most mating attempts were directed toward females with weaned infants. Simultaneous harassment by multiple subadult males increased significantly during months of high fruit abundance, and nearly all adult female-adult male consortships occurred during periods of high fruit abundance. Females who maintained spatial association with adult males, either via consortship or by nonmating temporary parties, received lower rates of harassment, as measured by the success rate of subadult male mating attempts. Adult female-adult male parties did not always result in mating between the associating dyad. Female initiation of protective services by adult males is one social tactic that female orangutans employ to reduce sexual harassment. Females therefore receive direct services from adult males, which may be one factor that influences female mate choice in Sumatran orangutans.     

Do male plumage and song characteristics influence female off-territory forays and paternity in the hooded warbler?

Extra-pair fertilizations are common in many socially monogamous species, and paternity studies have indicated that females may use male vocal performance and plumage ornaments as cues to assess male quality. Female off-territory forays may represent a key component of female choice and male extra-pair mating success, and female foray behaviour is expected to be strongly influenced by indictors of male quality. In this study, we examined how male song and ornamentation affect how often females left their territories, which males they visited and extra-pair paternity in a socially monogamous passerine, the hooded warbler (Wilsonia citrina). We radiotracked 17 females during the fertile period and quantified male vocal performance (song output and rate) and plumage characteristics (size of the black melanin hood and colour of the black hood, yellow cheeks and breast areas). We obtained blood samples and determined paternity at 35 nests including those of 14 females that we radiotracked. Eleven (65%) of the 17 females forayed off-territory, whilst fertile and female foray rate was positively correlated with the number of extra-pair young in the nest. Females that left their territories more frequently were paired with males that sang at a low rate. In addition, extra-pair mates had higher song rates than the social mates they cuckolded (5.3 songs/min vs. 4.4 songs/min). Female off-territory forays or extra-pair paternity were not significantly related to male plumage characteristics. Our results indicate that a high song rate influences both the foray behaviour of a male’s social mate and the likelihood that he will sire extra-pair offspring with neighbouring females.     

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     

Competition for a mate restricts mate search of female pied flycatchers

Summary We studied the mate sampling behaviour of female pied flycatchers, Ficedula hypoleuca, in a 40-ha area containing 10–12 unmated males whose nestboxes were monitored with videocameras. The main results were: (1) The females undertook a restricted mate search. The females that mated in the area during three monitoring periods (n = 20, 12 females released by us and 8 females that arrived naturally) sampled 1–10 males (median 4.5). This was about 40% of the available mating options. (2) Search costs in terms of time and energy were low. The search period was short (median 5.1 h) and only a small proportion of the search period was spent at the nestboxes of males (median 4%). The females visited up to seven different males in 1 h, and the time elapsing between visits to different males was short (median 13 min). The minimum distance travelled during the search was also short (median 1.4 km). (3) There was competition between the females. We recorded seven cases of two females visiting the same male at the same time, including at least one case involving physical fighting. (4) Females that experienced a high level of competition had a more restricted mate search than females that experienced a low level of competition. (5) The search pattern of most of the females did not conform to the best-of-n-males rule nor to the threshold criterion rule, because they made repeated visits to many of the males sampled.     

Polygyny and male parental care in Savannah sparrows: effects on female fitness

Summary To determine the effects of male mating status on female fitness, we compared the reproductive success, survival, and future fecundity of female Savannah sparrows (Passerculus sandwichensis) mated to monogamous vs. polygynous males in a 5-year study on Kent Island, New Brunswick, Canada. The proportion of males with more than one mate varied from 15 to 43% between years and sites. Polygynous and monogamous males fledged young of equal size in every year of the study. Females who shared paternal care with other females laid as many eggs per clutch and clutches per season as monogamously mated females. In most years polygynously mated females showed no delay in laying a second clutch, and they suffered no reduction in fecundity the following year. Recruitment of a female's offspring into the breeding population was generally independent of her mating status. Fitness costs of being mated to a polygynous male were only apparent in one year of the study, during which females mated to polygynous males had higher over-winter mortality than those mated to monogamous males. That same year, young raised by polygynous males were only one-third as likely to survive to reproductive maturity (as inferred by returns) as those raised by monogamous males. A male's mating status had no effect on his own survivorship. A male's mating status did not necessarily reflect his contributions to raising nestlings, which may partially explain why monogamously and polygynously mated females had equal fitness. At 35 nests the proportion of food deliveries brought by individual males varied from 0 to 75%; on average, males brought fewer than 30% of all food deliveries. Yet parental care by polygynous males was no less than that of monogamous males, at least at the nests of their primary females. Secondary females tended to receive less male assistance during the nestling stage, but their reproductive success was indistinguishable from that of primary females. Females feeding young without male assistance made as many food deliveries/h as did pairs in which males brought at least 30% of all food deliveries. Unassisted females did not suffer diminished fledging success or produce smaller fledglings. The benefits of polygyny for male Savannah sparrows are clear: polygynous males recruit more surviving offspring into the breeding population than monogamous males. The fitness of females, on the other hand, appears to be unaffected by whether their mate was monogamous or polygynous except in occasional years. Polygyny may be maintained in this population by the constraints of a female-biased sex ratio, the inability of females to predict a male's paternal care based on his morphology or behavior, the poor correlation between a male's mating status and his assistance at the nest, and inconsistent natural selection against mating with a polygynous male. Correspondence to: N.T. Wheelwright     

Inbreeding avoidance in a poeciliid fish (Heterandria formosa)

Polyandry is hypothesized to give females an opportunity to avoid inbreeding through postcopulatory selection mechanisms if precopulatory inbreeding avoidance is not possible, for example, because of forced matings. Here, we report a postcopulatory, prefertilization, inbreeding avoidance mechanism in the least killifish, Heterandria formosa, a species in which males can force matings. Females had 50% less sperm in their oviducts and ovarian cavities after mating with a sibling compared to a mating with a nonsibling male. Neither sex showed inbreeding avoidance in a dichotomous precopulatory mate choice test or during mating trials. Females in this species invest substantially in each offspring after fertilization (matrotrophy), whereas males invest little more than sperm. Based on theory, females should therefore be more likely than males to avoid inbreeding in this species. We suggest that females do this by reducing the amount of sibling sperm in their reproductive system. However, the possibility that males invested more sperm to nonsiblings could not be ruled out. In a fertilization success experiment, the first male to mate with a female sired all the offspring in most cases, even if it was a sibling. However, large females were more likely to carry offspring of multiple males. Possibly female sperm storage sites were filled by the first male, and only large females had space for the second male's sperm.     

Active mate choice at cock-of-the-rock leks: tactics of sampling and comparison

Summary The importance of active mate choice as a component of sexual selection remains controversial, despite recent studies identifying traits that correlate with male mating success. We analyzed the sampling sequences of individual female Guianan cock-of-the-rock, providing quantitative documentation of active female choice in this lekbreeding species. We compared observed sampling patterns with null models of random female movement and of movement weighted by the observed distribution of male success. The courts actually visited by individual females were significantly more clumped and centrally located than predicted by computer simulations of either random or weighted movement. Contrary to simplified hypotheses of female choice, hen cock-of-the-rock were not limited either to sequential comparisons or to threshold decision rules when selecting mates. Instead, they repeatedly sampled a subset of potential mates, using the pool-comparison tactic. Within this pool, females preferentially mated with the males ranked highest in terms of total courtship visits received, and rejected low-ranking males. Our results suggest complex interactions between patterns of female sampling and male dispersion in lek mating systems, and also provide insights into the constraints on female sampling in lek species.     

Loss of mating opportunities influences refuge use in the Iberian rock lizard,<Emphasis Type="Italic"> Lacerta monticola</Emphasis>

Because time spent in refuge may be costly if prey lose opportunities to forage, fight, or mate, prey allow predators to approach closer before beginning to flee when opportunity costs are high. Because the same opportunity costs may apply to refuge use as to escape, prey should make similar trade-offs between risk of emerging and cost of remaining in refuge. In the Iberian rock lizard, Lacerta monticola, we studied the effects of sex, reproductive season, speed of predator approach, and potential loss of mating opportunities on time spent in refuge following simulated predatory attacks. Lizards of both sexes adjusted refuge use to the level of risk by spending more time in refuge when approached rapidly than slowly. Females remained in refuge for equal times in the mating and postreproductive seasons, but males emerged sooner during the mating season, suggesting adjustment to a cost of lost opportunity to search for mates during the mating season. When a tethered female was nearby, males emerged from refuge earlier than if no female was present, indicating a trade-off between risk and mating opportunity. Approach speed affected emergence time when females were absent, but not when a female was present. Approach speed did not affect the probability that, after emerging, a male would return to court the female. For males that courted females intensely (bit them) before entering refuge, approach speed did not affect latency to emerge, but males that courted less intensely emerged sooner if approached slowly than rapidly. These findings show that males adjust the length of time spent in refuge to both risk of predation and reproductive cost of refuge use.Communicated by A. Mathis     

Effects of spermatophores on male and female monarch butterfly reproductive success

Summary I present the results of experiments designed to measure the effects of spermatophores produced by male monarch butterflies on male and female reproductive success. There was wide variation in the number of matings by captive males, suggesting the potential for strong sexual selection on males. Male lifespan was not affected by total number of matings, nor did it differ between males that were allowed to mate and those not exposed to females. Two effects of spermatophores on female behavior or fecundity are reported; (1) Females that received large spermatophores delayed remating longer than those receiving small ones. (2) Females allowed to mate several times laid more eggs than singly-mated females. The relative importance of these effects is discussed in relation to monarch mating patterns.     

Black Grouse leks on ice: Female mate sampling by incitation of male competition?

Male-male competition is assumed to limit female choice of mates, but it may also help females to choose the most vigorous males. We studied the mate sampling behaviour of female black grouse (Tetrao tetrix) at spatially unstable leks on ice-covered lakes. In the absence of territories and site-dependence in outcomes of fights, the male dominance hierarchy is very evident on ice. When being courted by dominant males, females frequently tried to approach other males. This was frequently prevented because (1) the courting male and the approached male were involved in physical fight, or (2) the dominant male followed the female and the approached male escaped and avoided contact with him. These behaviours express dominance relationships, and the female behaviour could be considered as incitive. Rank in dominance hierarchy was a significant predictor of male mating success. In this case competition between males and female choice worked in parallel favouring male traits correlated with dominance.     

Sex-specific reproductive behaviours and paternity in free-ranging Barbary macaques (Macaca sylvanus)

In a wide variety of species, male reproductive success is determined by contest for access to females. Among multi-male primate groups, however, factors in addition to male competitive ability may also influence paternity outcome, although their exact nature and force is still largely unclear. Here, we have investigated in a group of free-ranging Barbary macaques whether paternity is determined on the pre- or postcopulatory level and how male competitive ability and female direct mate choice during the female fertile phase are related to male reproductive success. Behavioural observations were combined with faecal hormone analysis for timing of the fertile phase (13 cycles, 8 females) and genetic paternity analysis (n = 12). During the fertile phase, complete monopolisation of females did not occur. Females were consorted for only 49% of observation time, and all females had ejaculatory copulations with several males. Thus, in all cases, paternity was determined on the postcopulatory level. More than 80% of infants were sired by high-ranking males, and this reproductive skew was related to both, male competitive ability and female direct mate choice as high-ranking males spent more time in consort with females than low-ranking males, and females solicited copulations mainly from dominant males. As most ejaculatory copulations were female-initiated, female direct mate choice appeared to have the highest impact on male reproductive success. However, female preference was not directly translated into paternity, as fathers were not preferred over non-fathers in terms of solicitation, consortship and mating behaviour. Collectively, our data show that in the Barbary macaque, both sexes significantly influence male mating success, but that sperm of several males generally compete within the female reproductive tract and that therefore paternity is determined by mechanisms operating at the postcopulatory level.     

A female preference for experienced males in the almond moth, Cadra cautella

Male mating status can affect female reproductive output if male ejaculate investment declines over consecutive matings. Accordingly, females are predicted to mate preferentially with virgin males. In mildly polyandrous lepidopterans, female fitness is less affected by reduced male investment than in more polyandrous species, and so the predictions for female mating preferences are less clear. We examined female mating preferences in the mildly polyandrous almond moth, Cadra cautella, in which ejaculate size does not affect female reproductive output. First, we allowed females to mate with virgin or once-mated males, in which the males were presented individually or simultaneously. We recorded the latency to mating and, in the case of the simultaneously presented trials, the identity of the successful, copulating male. We found that females mated more frequently with mated males (when simultaneously presented with both males), yet females did not differ in the time taken to initiate copulation with any male. We further examined if this mated male advantage was due to differential mate detection or locomotory behaviour of the male treatments. We tested the ability of virgin and mated males to locate a receptive female within a wind tunnel using long-distance pheromone cues and recorded their activity budget. We found no difference in the ability of mated or virgin males to locate or approach a receptive female, or in their activity levels. These data suggest a female preference for mated males in this species, a preference that may minimise other potential costs of mating.     

Female choice in sage grouse: the roles of attraction and active comparison

Previous studies of female choice in sage grouse Centrocercus urophasianus have implicated both the acoustic quality and repetition rate of the stereotyped strut display as putative cues for female choice. Stages in the choice process at which specific components of male courtship display influence female decisions were investigated using field observations of female pre-mating behavior. Females visited a subset of territorial males and then actively chose one of these as a mate. The order in which males were visited suggested that females searched until an acceptable mate was found, rather than employing a “best-of-n” tactic. Numbers of females visiting a male were related to differences in an acoustical component of display (inter-pop interval) whereas the probability that a visiting female mated was related to display rate (Table 3), indicating that initial attraction and active choice are influenced by different components of display. In addition, inter-pop interval and display rate tended to covary inversely (Fig. 1), suggesting that attraction and active choice may impose conflicting selection pressures on display performance. Received: 11 November 1995/Accepted after revision: 16 March 1996     

Large male advantage for access to females: evidence of male-male competition and female discrimination in a territorial salamander

Summary In the natural habitat visited in this study, adult male red-backed salamanders (Plethodon cinereus) found near females were significantly larger than adult males found alone; there was no evidence for size assortative associations. Previous research has indicated that females associate preferentially with males that occupy high quality territories and that larger individuals are more successful at obtaining higher quality areas. In laboratory experiments, when resource quality was held constant and males were restrained so that male-male interactions were prohibited, females discriminated behaviorally between large and small males by spending more time visually and chemically assessing larger males and more time apparently attempting to leave the chamber when near smaller males. Females were found near the larger males at the end of the trials significantly more often than predicted by chance. In a separate experiment in which only females were restrained, larger males spent significantly more time in aggressive postures when paired with smaller males than when alone with the female. Smaller males spent significantly more time in submissive postures when paired with larger males and more time near the female when alone. Therefore, large body size of males may positively affect both intra- and intersexual interactions and, ultimately, mating success of male P. cinereus.     

Experimental evidence for female-driven monandry in the wolf spider, <Emphasis Type="Italic">Pardosa astrigera</Emphasis>

Females of many species mate multiply, yet some taxon females mate with only one male, also known as monandry. Although the underlying mechanism behind female monandry is poorly understood relative to female polyandry, there are two contrasting hypotheses, male control and female control, for the maintenance of monandry. Since females generally benefit from multiple mating for material and/or genetic benefits, cases of monandry may reflect male manipulation on female remating at the expense of female fitness (male control). Alternatively, monandry may be favored by females, if females maximize their fitness by mating once (female control). Here, we tested two hypotheses by manipulating the number of mating (repeated mating and polyandry) on female fitness in a largely monandrous wolf spider, Pardosa astrigera. We allowed females to be inseminated once, twice with the same males (repeated mating) or with two males (polyandry) and determined female fitness consequences. The number of female mating, regardless of a single mating, repeated mating, or polyandry, had no significant effects on female fecundity, fertility, and survival and size of their spiderlings. However, the fitness cost of female multiple mating may to some extent be underestimated under laboratory conditions. In addition, female survival was adversely affected by induced multiple mating. Therefore, our results suggest that monandry of the wolf spider (P. astrigera) may be under the control of females, rather than under the control of males.