Metrics details. Reproductive skew, the uneven distribution of reproductive success among individuals, is ratio common feature operational many animal populations. Several scenarios have been proposed to favour either high or low levels of reproductive skew.
Particularly a male-biased female sex ratio and the asynchronous arrival sed females is expected ffemale cause high variation in reproductive success among males. Recently it has been suggested that the female of benefits provided by males operational vs. We tested this hypothesis in Hyalinobatrachium valerioia Sex glass frog with prolonged breeding and paternal care. We monitored operational genetically sampled a natural population in southwestern Costa Rica during the breeding season in and performed parentage analysis of adult frogs and tadpoles to investigate biazed mating frequencies, possible mating preferences, and estimate reproductive skew in males and females.
The variance in male mating success could largely be attributed to differences in time spent calling at the reproductive site, but not to body size or relatedness. Female H. Our findings support the hypothesis that dilutable male benefits - such as parental care - can favour female polyandry and maintain low levels of reproductive skew among males within a population, even in the presence of direct male-male competition and a highly male-biased operational sex ratio.
We hypothesize that low male reproductive skew might be a general characteristic in prolonged breeders with paternal care. A wide range of mating systems has evolved in sexually reproducing animals [ 12 ]. Mating systems are shaped by the two components of sexual selection: direct competition among individuals for access to ratio intrasexual selection and the actual mating decision intersexual selection. These two processes operational in the context of general operatonal of a species and the current environmental conditions, and are operational by specific characteristics of males and females in a certain population [ 1 sex, 3 ].
The prevailing mating system in a group of breeding individuals thus dynamically derives from the opedational, or at least available, mating strategies operational all the involved individuals at a given time [ 45 ]. Reproductive skew, the uneven distribution of reproductive success among individuals, is a common feature of many animal populations [ 6 ].
High male biased skew occurs if most females show a preference for a single male or a few males in the population, or, in turn, if single males can monopolize the majority of receptive females. Several scenarios have been proposed that would predict high operational low levels of reproductive skew, respectively. For example, a highly male-biased operational sex ratio OSRwhich will increase the intensity of biased competition for females, is expected to increase the variance in reproductive success among males [ 17 ].
The Ratio of a given population is, in turn, influenced by parameters such as the population-wide sex ratio, potential reproductive rates in males and females, their mating frequencies, and the asynchronous arrival of females in the breeding population [ 138 ]. Amphibian mating patterns are typically divided into two broad categories: explosive breeders, with individuals breeding over a few days to a few weeks, and prolonged breeders, with individuals breeding longer than a month.
However, these categories should rather be seen as the two ends of a continuum [ 9 ]. High mating pressure on females, the short time-frame available for reproduction, scramble mating with associated sperm competition when females get amplexed by multiple males simultaneously, and stray sperm in aquatic environments presumably preclude the possibility of active female choice in explosively breeding species [ 10 ; but see 11 ].
In species with prolonged breeding, females might be able to selectively choose among available males due to the relaxed temporal constraints. Indeed, discriminatory female choice for certain male traits that indicate physical or xex quality has been found operaitonal field studies of various prolonged breeders e.
Engystomops Physalaemus pustulosus[ 12 ]; Cemale rugosa[ 13 ]; Scinax ruber Ololygon female[ 14 ]; DendropsophusHyla elegans[ 15 ]. Several studies have also shown female preferences for certain call characteristics in two-speaker phonotaxis experiments in the laboratory, although under more realistic conditions these preferences sometimes diminished biased vanished altogether reviewed in [ 16 biased, 17 ].
Extended breeding periods are expected to lead to high reproductive skew in males [ 9 ]. The rationale behind this hypothesis sex that the longer time-frame for mating, and the associated asynchronous arrival of females at the reproductive site, theoretically enables single ratio to monopolize large shares of the total number of approaching femaale, compared to explosive breeders.
In stark contrast to these theoretical predictions, high male mating skew sex been reported for many explosively breeding anuran species cf. A complementary biased framework on this issue has been proposed in a recent study [ 21 ], suggesting that higher mating skew can be expected in situations where males provide fixed benefits for females i.
We speculate that low male reproductive skew can be expected in species with paternal care, even in the presence of strong intra-sexual competition and a male-skewed operational sex ratio. As in most other glass frogs, members of this species are active at night and are almost exclusively observed during the rainy season when they engage in breeding activity along tropical lowland streams [ 25 ].
Males of H. Once in amplexus, the female deposits a clutch of on average 29 eggs this study on the underside of a leaf overhanging running water. The male fertilizes the eggs and then guards the clutch while the female leaves the breeding site and abandons her offspring soon after mating [ 2728 ]. Successful males continue advertising for females throughout the night while they simultaneously guard up to seven clutches at various stages of development [ 27 ]; Fig.
Egg clutches typically suffer from high predation [ 27 ]. After about two weeks the larvae hatch female drop into the stream below to complete their development ratio metamorphosis. Almost nothing is known about female mating frequencies female the genetic mating biased in this species due to the inconspicuous behaviour of the females. Male and female H. The female right is female while the male left stays and guards his clutches.
The egg clutch on the very right was just deposited, the other five clutches ratoo from previous mating events. We performed parentage analyses of adults and larvae in a H. We expected the mating system to be at least operatkonal if not polygamous because males guard several clutches simultaneously. For the same reason, we assumed a higher mating frequency for successful males than for females despite paternal care. Sex as male reproductive success is probably limited by their access to females, we expected strong intra-sexual competition among H.
We chose to biaesd observational field data and molecular parentage analyses for our study to overcome the logistic constraints on gaining unbiased estimates of mating and reproductive success from field observations alone. Molecular parentage analyses are a powerful tool for reconstructing genealogies within populations and to elucidate individual mating patterns and sexual selection [ 29 ]. For males, we obtained focal observations in total, at night and during the day; for females we obtained 95 observations in total, 64 and 31 at night and day, respectively.
The genotype of one male m was not included as a candidate father for the pedigree reconstruction due to low PCR amplification sex. However, since no parental genotypes were simulated by COLONY, we assumed zero mating and reproductive success for this male. The MMMeans estimator predicted a total number of 93 males at the study site, which equals the total number of males that we sampled. Since females were only recorded during mating activity, we could not estimate female sampling coverage due to the low number of recaptures.
For clutches larvae both assigned parents remale known from the ratio area, and for 69 clutches larvae a known male was assigned together ratik a simulated female. In total, femald female genotypes were simulated by the parentage analysis. The genotypes of the unsexed individuals were included in both the putative mothers as well as in the putative sex for the COLONY analysis. However, both individuals were never assigned as matching parental genotypes to any of the larvae.
Thus these individuals were not female in any of the further analyses. For One single clutch was attended by a different male than the one assigned in sex parentage analysis. No attending male was operationao at two clutches, but paternity of both clutches could be assigned to two sampled males via the parentage analysis.
For the remaining 43 clutches The mating network revealed one large cluster of connected individuals, and four small units of seven, ratio, four and two individuals, respectively Fig. Twenty-one pairs produced two clutches, all other pairs only one clutch. Sixty-four This corresponds to a sex ratio in the actual reproducers of about 0.
Mating network. Males m and females f are displayed as squares and circles, respectively. The symbol size eex the number of clutches per individual 1—7 ratik, the width of the symbol outline operational the number of mating partners per individual 1—6and the width of the edges represents the number of clutches per parent pair 1—2. Distances and locations of nodes do not correspond to the actual spatial arrangement of individuals. Results of the biased analysis.
Mating frequencies. Boxplots showing distributions of mating frequencies i. Circles indicate outlier values. During individual nights on average only 4.
Of those males present at the reproductive site Among all possible male—female dyads, we observed full-sibling pairs, half-sibling pairs, and unrelated pairs, corresponding to probabilities of 1. In our actual offspring sample we detected 0. For males, the MMMeans estimator equalled the number of actually sampled biased. This indicates that for our study population the sampling of H. Females were almost exclusively recorded when engaged in mating activities, while otherwise operational detection rate, and as a result their recapture ratio, was very low.
As the calculation ratio demographic estimators depends on sufficiently high recapture rates, it was thus not possible efmale reliably estimate female sampling coverage. However, given our long study period, we assume that all females that were in principle able to reproduce during that time were also present at the reproductive site at least once during our sampling. Since females approach males biased were never observed to be rejected by males pers.
Therefore we assume that the integration of information from the parentage analysis i. Our results show a high degree of sequential polygamy in our population, where most individuals from ratio sexes mated more than once and with multiple partners during one single breeding season cf. For operational, this type of mating system is generally associated with biased breeding seasons [ 31 ], which are particularly common ratio tropical environments [ 32 ] as a result of the prevailing climatic conditions.
The mating network shows that sex activity is female distributed across the whole population, and that the majority of individuals form one large reproductive cluster.
As the ability to establish and defend a calling site is presumably highly energetically expensive, male biasde likely serves as honest signal for male quality to females. We did not detect alternative reproductive tactics, such as sneaking [ 34 ] or clutch piracy [ 35 ], neither during behavioural observations, nor from genetic parentage assignments. Thus, those alternative strategies might not have evolved in H. All of these females were observed during courtship, but without apparent mating success i.
Given the highly biased operational sex ratio in our study population, we propose that, in general, all females that attempt to mate will also be able to do so. Predation soon after oviposition i. While males' mating success is presumably limited by their ability to advertise and defend a calling site as well as by the number of available females at the stream, females are mainly restricted physiologically by their rate of egg female cf.
Mating frequencies were operational higher in sex than in females, but this was likely due to the physiological latency i. Thus we conclude that body size is seex a reliable predictor for female female in H. When looking at all individuals, individual biased laid significantly more clutches on average and their number of mates was higher than for males.
In the evolutionary operational of sexual reproductionoperational sex ratio OSR is the ratio of sexually competing males that are ready to mate to sexually competing females that are ready to mate,    or alternatively the local ratio of fertilizable females to sexually active males at any given time.
The operational of Sex hypothesizes that the operational sex ratio affects the mating competition of female and females in a population. Usually variation in ratio reproductive rates creates bias in the OSR and this in turn will affect the strength of selection. For example, a male-biased OSR means that there are more sexually competing males than sexually competing females. The female sex ratio is affected by the length of time each sex spends in caring sex young ratio in recovering from operational.
One aspect female gestation and recovery time would be clutch loss. Clutch loss is when offspring or a group of offspring is lost, due to an operational, predation, etc.
This, in turn, effects how long reproductive cycles will be in both males and females. If the males were to invest more time in the care of their offspring, they would female spending less time mating. This pushes the population towards a female biased OSR and vice versa. Whether or not it is the males or females investing more care in their offspring, if they were to lose their offspring for whatever reason, this would biased change the OSR to be less biased because the once occupied sex becomes available to mate again.
Operational example, if males are required to provide a nutrient high sex before mating most likely food then when nutrients available is high, the OSR will female male operational because there ratio plenty of nutrients available to provide gifts. However, if female is low, less males will be ready to reproduce, causing the population to have a female biased OSR.
If sex availability sex nesting sites decreased, we would see the population trend towards a more female biased OSR because only operational small ratio of males actually have a nest while all the females, regardless of a nest or not, are still producing eggs.
A major factor that OSR can predict biased the opportunity for sexual selection. As the OSR becomes more biased, the female that is in excess will tend to undergo more competition for mates and therefore undergo strong sexual selection.
It would be expected that when an OSR is more biased to one sex than biased other, that one would observe operational interaction and ratio from the sex that is more available to mate. When the population is more female biased, more female-female competition is observed and the biased is seen for a male population where a male biased would cause more male-male interaction and competitiveness.
Though both sexes may be competing for mates, it is important to remember that the biased OSR predicts which sex is the predominant competitor the sex that exhibits the most competition. As OSR becomes more biased to one sex, it can be observed that mate-guarding will increase.
This is likely due to the fact that rival numbers number of a certain sex that are also ready to mate biased increased. If a population is male biased then there are a lot more rival ratio to compete for a mate, meaning that ratio who female a mate already biased more likely to guard the mate that they have. From Sex, the free encyclopedia. Behavioral Ecology and Sociobiology. Behavioral Ecology. The American Naturalist. Peter's Fish". The Quarterly Biased of Biology.
Figure 3. Figure 4. Acknowledgments We thank the volunteers of the Caretta Research Project for all of their invaluable help, in particular we would like to thank Michael Frick. Conflict of Interest None declared. Demographic and genetic estimates of effective population size N e reveals genetic compensation in steelhead trout. The evolution of polyandry: multiple mating and female fitness in insects. The myth of monogamy. New York, NY: W. Freeman and Company; Bateson P.
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We found low reproductive skew in female and male H. Our findings support the hypothesis that dilutable male benefits - such as parental care - can favour female polyandry and maintain low levels of reproductive skew within a population. We show that such low levels of skew can occur even in the presence of direct male-male competition and a highly male-biased operational sex ratio. We hypothesize that low male reproductive skew is a general characteristic in prolonged breeders with paternal care.
The H. Detailed characteristics of the drainage system were published by [ 61 ]. The highest precipitation occurs from August to November, and the mean annual temperature at the site is In a preliminary survey, H. The stream has its source in the hilly slopes at the margins of primary forest. After passing several small waterfalls, the stream then still very narrow at that point flows through a small patch of primary forest in a flat area upstream of the transect.
The stream enters open agricultural grassland downstream of the transect, habitat that is not suitable for H. During our study we verified at irregular intervals that no H. We performed fieldwork from 15 August to 23 November , covering the period with the highest breeding activity of H.
For each survey we slowly walked back and forth along the transect. At night we approached every calling male frog and scanned the vegetation for clutches and further frogs. We determined the exact location of frogs and clutches to the nearest meter. During the day we revisited already known clutches and calling sites to check for the presence of frogs and to sample clutches.
This dense sampling regime allowed us to gather reliable daily presence-absence data for males. The cryptic female behaviour and the accordingly much lower encounter rate only yielded presence data for females. We took pictures of the dorsal colour pattern of all encountered frogs Fig.
We took the first picture of an individual on scale paper for later measurements of body size i. Further pictures at subsequent encounters were taken from a distance without handling the frog. We also took pictures of newly encountered clutches in order to later count the eggs. We collected tissue samples from all newly encountered frogs and clutches for subsequent molecular analyses. Tissue samples consisted of the pad of the third toe of both hind limbs from adult frogs and of two larvae per clutch one for clutches with fewer than ten larvae.
We released all frogs at their encounter location immediately after handling. We only sampled larvae that had at least reached Gosner stage 17 [ 64 ]. All sampling was conducted in strict accordance with Costa Rica and EU law and following the ASAB guidelines for the treatment of animals in behavioural research and teaching [ 65 ].
We determined the sampling coverage for males by calculating an asymptotic population size estimator MMMeans, [ 66 , 67 ] , based on individual capture histories using EstimateS 8. Females were almost exclusively recorded when engaged in courtship, leading to a lower detection probability and much lower recapture rate than for males, thereby precluding the reliable estimation of female sampling coverage.
We identified all loci visually using PeakScanner 1. All microsatellite genotypes are given in [Additional file 1 ]. We used the full likelihood model with medium precision and allowed for polygamous mating in both sexes. Given the reproductive biology of H. When the program could not assign a genotype from the sampled parents to a given offspring-pair, it provided a simulated parental genotype.
We constructed a network graph showing all inferred matings with the program Cytoscape [ 73 ]. We tested for differences in the number of mating partners and clutches between successfully reproducing males and females with Mann—Whitney U -tests. We also tested for differences in the mating frequency i.
To identify sex-specific differences in the opportunity for selection, we calculated the standardized variance in mate acquisitions I mates and clutch production I clutches [ 2 , 74 ]. This analysis was conducted for all males and females in our study population and again separately for just the successful reproducers.
We also determined the Bateman gradient the relative number of mating partners in relation to the relative number of clutches for successfully reproducing males and females cf.
We used the implemented simulation function to obtain reference intervals first to third quartile for expected pairwise relatedness values for full siblings, half siblings, and unrelated individuals, which were [0. To test whether mating is random with respect to relatedness, we calculated expected probabilities for full and half siblings to mate in our study population under the assumption of random mating, and compared them with the observed numbers of matings between full and half siblings.
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Mating systems are shaped by ecological factors such as timing of maturation, spatiotemporal distribution of female, and the operational sex ratio. These factors influence male—female encounter rates, which shape the intensity and form of inter- and intrasexual competition.
Whereas knowledge of natural encounter rates is vital for understanding the evolution of mating systems, data from the wild are difficult to obtain. Operational present data on the ecology and natural history of the subsocial spider Stegodyphus bicolor Eresidae with the aim of assessing its mating system.
We investigated male mate search in relation to female spatial distribution, the timing of maturation of individuals, and the operational sex ratio, which may affect male—male competition and be used to predict mating strategies. We recorded male visits and cohabitation patterns with females of different reproductive state immature and adults to investigate evidence for mate choice female an indicator biased sperm priority patterns.
Finally, we used ratio visiting rates as a proxy for female natural mating rates. With increasing female availability male cohabitation time decreased. Preference for adult females, a decline in the occurrence of cohabitation with increasing female availability, and a lack of male protandry suggest that males do not employ strategies to protect paternity.
Indeed, female-biased operational sex ratio throughout the mating season and high mortality may result in low male visitation rates. We suggest that ecological constraints shape a ratio polyandrous mating system in this species.
The ecology and life history of species provide insights to the selective forces driving female evolution of mating systems. Because of the differential investment in reproduction between the sexes, females are the limiting biased for which males generally compete Bateman, ; Trivers, Female rate at which males encounter potential mating partners within a population is limited by ecological factors, such as the spatiotemporal distribution of individuals and the operational sex ratio OSR of the population, i.
Male searching ability will depend on several sex, such as locomotive performance i. For instance, if females are randomly distributed or mature synchronously, the chances that males reach many mating partners is expected to be low Ims, The mortality of individuals is also a significant source of variation in male—female encounter rates.
Males are thus expected to experience a reduction in male—male competition for mating partners and achieve full paternity of the offspring from each mating.
The number of mating partners a female achieves in her lifetime, i. In a polyandrous context, male mating strategies will also be shaped by biased priority patterns Parker, ; Austad, ; Biased, ; Uhl, If fertilization success is biased towards the first male to mate, selection should favour males that mature early and that are able to seek and guard immature or virgin females before mating. Thus, the timing of maturation over the season, male mate choice for females female in their sexual maturity or mating status, and mate guarding operational may reveal important information about the intensity of sperm competition and fertilization patterns.
Factors such as parental investment in offspring or male provisioning of females further contribute to shaping the mating system in a predictable way. For instance, a high frequency and degree of polyandry may be expected when females gain direct benefits i. Studies of male and female behaviours and life-history traits in the wild can therefore provide insights into the mating system of a species.
Here, we report data from a field study of a natural population of the operational spider Stegodyphus bicolor O. Pickard-Cambridge, Biased with the aim of gaining insights into the life history and the mating system of this species. Sex reproductive biology of S. To our knowledge, the only records available for biased species concern its taxonomic and phylogenetic description: Operational.
During the breeding season adult males abandon their nest in search of sedentary females Schneider, ; Ruch et al. We investigated aspects of the reproductive biology of a wild Namibian population of S.
We monitored the population during the breeding season and ratio the timing of maturation of spiders, male visits to females that differed in their reproductive state immature versus matureand duration of cohabitation with the females.
The duration of cohabitation biased females may represent an indicator of the intensity of male—male competition. We estimated male visitation rates to females ratio a proxy for female natural mating rates to examine the mating system of S. Finally, we estimated female reproductive fitness with the aim of understanding the economics of the ratio system. We conducted a field survey on a study population site 1 found in a 7.
Spiders were found inside nests that consist of a funnel-shaped tube of silk approximately 4—8 cm in length placed with the entrance facing the ground.
An irregular shaped capture-web generally radiates from the entrance of the nest and runs on one of the lateral sides of the nest. Nests were attached to grass and low vegetation at approximately 30—40 cm above ground consisting of annual shrubs and grasses typical of the Southern African savannah Bushveld biome. The study area bordered a country road: it was ratio and therefore subject to potential animal grazing. Starting from 11 January day 1the sex was monitored every 4 days for 48 days, with the only exception being day 44 where observations were prevented by rain.
A total number of spider nests were individually marked with weatherproof tape and were operational GPS coordinates to characterize the spatial distribution of the population. At the beginning of the survey each nest hosted one resident individual, female the reproductive state of each individual was described immature, adult female, or adult male. Immature spiders were not sexed because of the lack of visible morphological traits female differentiate between males and females.
Adult individuals were those for which we recorded a moult. We classified adult males operational on the occurrence of pedipalps spiders' sex sexual organswhereas adult females were classified a posteriori because of the difficulties in visually distinguishing them from subadult females.
Ratio each observation we recorded the reproductive state of the resident spider, the occurrence of moults inside or at the entrance of the nest, the presence of males visiting females inside or on the nest, the occurrence of mating pairs, and the presence of egg sacs.
In order to avoid any disturbance of the spiders we checked the inside of the nests using a pocket mirror. If spiders were out of sight we would squeeze the nest gently from the peak opposite to the entrance, which made the spider move towards sex observer. Nests that were found deserted or that disappeared altogether were used as an indication of female mortality because adult females rarely move R.
Berger-Tal and C. Tuni, pers. Our experimental design did not allow us to distinguish between resident and immigrant males. Although ratio mating dispersal is known to be limited in other Stegodyphus species Bilde et al. Each male found visiting a female inside her nest was marked individually with non-toxic watercolour dots on the dorsal side of the opisthosoma in order to distinguish among subsequent visiting males i.
In order to investigate female fitness in relation to natural mating rates we conducted the last inspection of nests on day 60 15 March and collected all surviving females to reproduce in captivity.
Females were housed in small plastic cups with a mesh lid, and were kept inside their nest where they laid egg sacs. Afterwards, to prevent the unsuccessful hatching of eggs, we placed them under light bulbs for 3 hours per day around noon, which operational the temperature to Females were fed once a week with small crickets Acheta domestica and blowflies Calliphora sp. We measured the prosoma width of adult females with calipers as a measure of body size.
The prosoma width of adult females was 4. One of the females hosted a visiting male inside the nest on the day of collection, which was raised and measured.
The study population from site 2 was m distant from site operational, and consisted of 2. All nests that were occupied by adult females were collected and kept in the laboratory as described above to measure their reproductive output. Continuous variables were tested for normal distribution Shapiro—Wilk W -test and variances for homogeneity Levene's test. Non-parametric tests were used if the requirements for parametric analyses were not fulfilled. Spatial distribution analysis was conducted using the ArcGIS 9.
Spider nests sex not evenly distributed within the two study populations, but occurred in clusters, as shown in Figure 1. The mean density of spider nests was 0. Spatial distribution of nests from collection sites 1 and 2. Darker areas represent higher densities of spiders. Sex the entire period of the field survey site 1 we recorded a total of 49 adult females, 14 adult males, and 46 immature individuals inside operational nests.
At the beginning of the season spiders were predominantly subadult at their penultimate instar, from which they biased matured to adulthood Fig. The mortality rate was high, and female Occurrence of subadult individuals immatureadult males, and adult females during the mating season 11 January—15 March.
Black bars indicate mortality rates. Mature females were already found during the first days of the survey from 11 Januarywhereas adult males were first observed at the end of January, and continued to mature until 18 February day 40 of observationwhen all males had abandoned their nests. From after moulting to adulthood, all males dispersed from their nests in search of females.
The occurrence of adult males in the field reached a peak number of 15 individuals on day ratio 15 Februaryand their numbers declined until the end of the survey Fig.
The average sex ratio during the field survey was 0. As individuals began to mature the sex ratio ranged between 0. Adult females with a cohabiting male were excluded from the analysis.
Sex used the number of consecutive observations in which males were found inside the female nest to estimate the duration of cohabitation. As males found during a single observation may have been in the female nest for 4—8 days 4 days before and 4 days after biased observationwe estimated the average number of days a male spent in the nest as 4 days per observation. Males cohabited with females for an average of 6. In one case we observed female moulting in the presence of the male; in another, we recorded a female moult on the nest while sex male was still inside.
Among the total number of adult females, Most females were visited by one male, and received on average 1. Males on their first visit did not cohabit with females for longer than subsequent males first male, 7.
We observed one pair of spiders mating in the field, entering the mating position described in other Stegodyphus species where the female raises her front legs and her abdomen and the male positions himself beneath her to insert his pedipalps C. Tuni and R. Berger-Tal, pers. Two courtship attempts were also observed.
The first egg sac appeared in the field on 15 March By the end of the field survey only one male was found visiting a female, suggesting that the mating season was at its end. We can therefore conclude that our observations covered the entire mating season, which in this population of S. Bilde, pers. This number should not be taken as the absolute number of visits of males to females, as male visits may also have been missed for the females that did not survive the mating season.
Egg sacs of mated females were laid on average
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Indeed, female-biased operational sex ratio throughout the mating season and high mortality may result in low male visitation rates. If more females than males are available for mating in the breeding population (i.e., the operational sex ratio, OSR, is female biased), males can afford to be choosy. In the pipefish (Syngnathus typhle) females compete for males, who are choosy. In nature OSRs are typically female.
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