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1.
Understanding why females mate multiply is a major issue in evolutionary ecology. We investigated the consequences of an asynchronous arrival pattern on male competition and multiple paternity in the apparently monoandrous agile frog ( Rana dalmatina ). The largest frogs arrived first and both males and females lost weight significantly during the spawning period. Asynchronous arrival at breeding sites resulted in a male-biased operational sex ratio (OSR). The OSR was more strongly male-biased at the beginning and at the end of the breeding period when the number of satellite males increased. All females mated only once, but multiple paternity within clutches occurred at the beginning and the end of the breeding period. The influence of asynchronous arrival and biased sex ratio suggests that reduced variance or bet-hedging promoting female fitness had only a reduced role in the evolution of polyandry, and polyandry is likely to be associated with male benefits. Polyandry in frogs can be explained either by forced mating as a result of sexual conflict or by clutch piracy. By modifying intrasexual competition, asynchronous arrival and changes in OSR may have a decisive influence upon the evolution of mating systems and favour both polyandry and stable coexistence of alternative mating behaviour.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 191–200.  相似文献   

2.
Conventional sex roles imply caring females and competitive males. The evolution of sex role divergence is widely attributed to anisogamy initiating a self‐reinforcing process. The initial asymmetry in pre‐mating parental investment (eggs vs. sperm) is assumed to promote even greater divergence in post‐mating parental investment (parental care). But do we really understand the process? Trivers [Sexual Selection and the Descent of Man 1871–1971 (1972), Aldine Press, Chicago] introduced two arguments with a female and male perspective on whether to care for offspring that try to link pre‐mating and post‐mating investment. Here we review their merits and subsequent theoretical developments. The first argument is that females are more committed than males to providing care because they stand to lose a greater initial investment. This, however, commits the ‘Concorde Fallacy’ as optimal decisions should depend on future pay‐offs not past costs. Although the argument can be rephrased in terms of residual reproductive value when past investment affects future pay‐offs, it remains weak. The factors likely to change future pay‐offs seem to work against females providing more care than males. The second argument takes the reasonable premise that anisogamy produces a male‐biased operational sex ratio (OSR) leading to males competing for mates. Male care is then predicted to be less likely to evolve as it consumes resources that could otherwise be used to increase competitiveness. However, given each offspring has precisely two genetic parents (the Fisher condition), a biased OSR generates frequency‐dependent selection, analogous to Fisherian sex ratio selection, that favours increased parental investment by whichever sex faces more intense competition. Sex role divergence is therefore still an evolutionary conundrum. Here we review some possible solutions. Factors that promote conventional sex roles are sexual selection on males (but non‐random variance in male mating success must be high to override the Fisher condition), loss of paternity because of female multiple mating or group spawning and patterns of mortality that generate female‐biased adult sex ratios (ASR). We present an integrative model that shows how these factors interact to generate sex roles. We emphasize the need to distinguish between the ASR and the operational sex ratio (OSR). If mortality is higher when caring than competing this diminishes the likelihood of sex role divergence because this strongly limits the mating success of the earlier deserting sex. We illustrate this in a model where a change in relative mortality rates while caring and competing generates a shift from a mammalian type breeding system (female‐only care, male‐biased OSR and female‐biased ASR) to an avian type system (biparental care and a male‐biased OSR and ASR).  相似文献   

3.
Even though numerous metrics exist, we still appear to be far from a consensual view on the best way of measuring or predicting the potential strength of sexual selection. One of the earliest and simplest metrics devised was the operational sex ratio (OSR) (i.e. the ratio between sexually active males and females in a population), and even, if heavily criticized, the OSR can still be viewed as a valuable measure of the potential levels of intrasexual competition. Because this ratio is influenced by the time that individuals spend in the mating pool, the OSR depends on our ability to determine who is indeed ready to mate. Moreover, because the proximate effects of OSR on mate monopolization might not be immediately apparent, we should be prepared to account for its association with the conditions making selection favour traits that reduce the time needed to acquire additional mating opportunities. Using the worm pipefish as a working model, we conducted a more stringent calculation of the OSR by eliminating individuals that, although present, do not appear to be able to reproduce, as determined by an analysis of female size classes with immature or spent ovaries and male classes without pregnancy events. Accordingly, the OSR was not only capable of correctly highlighting the potential for intrasexual competition, but also was able to translate the desertion of individuals from the mating pool as the breeding season progressed into meaningful correlations with variables associated with reproductive investment and costs (i.e. gonad investment and energy reserves). As demonstrated, the predictive power of the OSR can be broader than anticipated, depending primarily on our ability to effectively discriminate which individuals are indeed ready to mate. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 477–484.  相似文献   

4.
The reproductive success of the sneaky mating tactic is difficult to determine in fish with internal fertilization. We approached this problem by developing an assay that allows the recovery of sperm DNA from the reproductive tract of females. This assay was used to test whether sperm transfer occurs between sympatric populations of the guppy, Poecilia reticulata , and its putative sister species, Poecilia picta . Six hundred females of both species from two sympatric sites in Trinidad were collected, and then the contents of the females' reproductive tracts were analysed using reciprocally diagnostic microsatellite markers. Conspecific sperm occurred in approximately 86% of females, whereas heterospecific sperm were found in only 4% of females. Because females of either species do not mate willingly with heterospecific males, the results indicate that sneaky mating results in the transfer of sperm. The data are consistent with the idea that sexual conflict can result in differences in the strength of behavioural isolation between the sexes, and they suggest that behavioural isolation is unlikely to have driven speciation between the guppy and P. picta .  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 397–402.  相似文献   

5.
Cooperatively breeding animals commonly avoid incestuous mating through pre-mating dispersal. However, a few group-living organisms, including the social spiders, have low pre-mating dispersal, intra-colony mating, and inbreeding. This results in limited gene flow among colonies and sub-structured populations. The social spiders also exhibit female-biased sex ratios because survival benefits to large colonies favour high group productivity, which selects against 1 : 1 sex ratios. Although propagule dispersal of mated females may occasionally bring about limited gene flow, little is known about the role of male dispersal. We assessed the extent of male movement between colonies in natural populations both experimentally and by studying colony sex ratios over the mating season. We show that males frequently move to neighbouring colonies, whereas only 4% of incipient nests were visited by dispersing males. Neighbouring colonies are genetically similar and movement within colony clusters does not contribute to gene flow. Post-mating sex ratio bias was high early in the mating season due to protandry, and also in colonies at the end of the season, suggesting that males remain in the colony when mated females have dispersed. Thus, male dispersal is unlikely to facilitate gene flow between different matrilineages. This is consistent with models of non-Fisherian group-level selection for the maintenance of female biased sex ratios, which predict the elimination of male dispersal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 227–234.  相似文献   

6.
Between-individual variance in potential reproductive rate theoretically creates a load in reproducing populations by driving sexual selection of male traits for winning competitions, and female traits for resisting the costs of multiple mating. Here, using replicated experimental evolution under divergent operational sex ratios (OSR, 9:1 or 1:6 ♀:♂) we empirically identified the parallel reproductive fitness consequences for females and males in the promiscuous flour beetle Tribolium castaneum. Our results revealed clear evidence that sexual conflict resides within the T. castaneum mating system. After 20 generations of selection, females from female-biased OSRs became vulnerable to multiple mating, and showed a steep decrease in reproductive fitness with an increasing number of control males. In contrast, females from male-biased OSRs showed no change in reproductive fitness, irrespective of male numbers. The divergence in reproductive output was not explained by variation in female mortality. Parallel assays revealed that males also responded to experimental evolution: individuals from male-biased OSRs obtained 27% greater reproductive success across 7-day competition for females with a control male rival, compared to males from the female-biased lines. Subsequent assays suggest that these differences were not due to postcopulatory sperm competitiveness, but to precopulatory/copulatory competitive male mating behavior.  相似文献   

7.
Operational sex ratio (OSR) has been traditionally thought ofas a force imposing competition for mates rather than also acue used to regulate the intrasexual competition individualsencounter. To assess whether eastern red-spotted newts, Notophthalmusviridescens, could appropriately compare OSRs, we quantifiedfield responses to traps containing four males, a sexually receptivefemale, four males plus a female, or nothing as a control. Earlyin the breeding season, males from two populations chose competitivemating opportunities over no mating opportunity at all, butgenerally preferred less competitive mating prospects. Laterin the breeding season, as the OSR of newt populations becomesmore male biased, males accordingly increased their acceptanceof intrasexual competition. Females avoided groups of four males,and for both sexes, avoidance of male-biased courting groupsincreased their probability of amplexus courtship. We then isolatedan approximately 33-kD protein from male cloacal glands thatwas used by males to compare OSRs. To our knowledge, this proteinrepresents the first isolated and characterized component ofan olfactory cue used to evaluate OSR. These results supporttwo important principles regarding mating systems: (1) OSR cansomewhat paradoxically be both the source imposing competitionfor mates and the source used to reduce it, and (2) analogousto the sex in short supply often being "choosy" selecting mates,the sex in excess (here, males) appears to be choosy about itsacceptance of intrasexual competition.  相似文献   

8.
Males developing relatively large, costly sexually selected traits may be of superior body condition compared to small-ornamented males. Thus, males developing the largest secondary sexual trait in a given environment may also be able to augment their investment into ejaculate quality, and fertilize a larger proportion of a female's eggs. We tested the prediction that the degree of expression of a condition-dependent secondary sexual trait, the male sex comb, in a Cape Tribulation (northeastern Australia) population of Drosophila bipectinata Duda, reveals male ability to fertilize eggs in the absence of sperm competition. This test permitted us also to evaluate whether pre-copulatory sexual selection and fertilization efficiency might act additively to influence male reproductive success because a previous study of the same population demonstrated a positive association between comb size and copulation probability. The results obtained indicate that, although genotypes developing smaller sex combs collectively had a significantly higher rate of insemination failure compared to larger comb genotypes, the hatch rate and the number of eggs laid by females inseminated by the two genotypic categories were not statistically different. The results fail to support the prediction that comb size reveals noncompetitive fertilization efficiency of males in this Australian population.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 406–413.  相似文献   

9.
One of the various male strategies to prevent or impede female remating is the production of a mating plug that covers the female genital opening or remains inside of the female genital tract after mating. Such structures have been described for many species in many animal taxa; however, in most cases, we know little or nothing about their specific adaptive value. Our investigations demonstrate that females of the dwarf spider species Oedothorax retusus (Westring, 1851) (Linyphiidae, Erigoninae) exhibit a substance on one or both of her paired genital openings only after copulation. We performed double-mating trials and forced the second male to mate into the previously used or unused spermathecal duct of the female by amputating one of his paired male gonopods (pedipalps). Furthermore, to investigate whether the duration of the first mating has an effect on the size and efficiency of the mating plug, we interrupted first matings after either 1 or 3 min, categorized plug size and recorded mating behaviour of subsequent males. The amount of secretion transferred was larger in long compared to short copulations. A long first copulation successfully prevented subsequent males from mating into the used ducts, whereas mating success after short first matings was similar to matings into unused copulatory ducts of the females. The present study demonstrates that a male O. retusus can prevent a rival from transferring sperm into the same spermatheca by applying a mating plug, but only if he mates for long enough.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 574–583.  相似文献   

10.
Recent investigations of mate choice indicate that the genetic effect of sires on offspring fitness may depend on the interaction between maternal and paternal genotypes and the environmental conditions experienced by the offspring. Alternative colour morphs of the pygmy grasshopper, Tetrix subulata , represent ecological strategies that differ in body size, life history, thermoregulatory behaviour, and habitat selection. The hypothesis that selection promotes behaviours maintaining coadapted gene complexes predicts individuals to mate assortatively with respect to colour morph. On the other hand, the bet-hedging hypothesis predicts that the temporal variability of the environment inhabited by these animals may select for disassortative mating behaviour resulting in heterogeneous offspring. To distinguish between these competing hypotheses, we investigated mating behaviours using dual-choice experiments. Our results were not in agreement with the prediction of assortative mating but suggest instead that matings were random with regard to colour morph. Polyandry was common, and females mated with the second male regardless of whether the first mating was assortative or disassortative. Polyandry also was equally frequent among females in triads in which the two males belonged to different colour morphs as in triads where both males belonged to the same colour morph. A field experiment confirmed that polyandry occurred also among free-ranging individuals, and uncovered variation in mating success among male colour morphs, probably due to indirect effects of coloration on activity or habitat use. The consequences of this random and polyandrous mating strategy for the evolutionary dynamics of the colour polymorphism remain to be explored.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 491–499.  相似文献   

11.
Female choice is thought to increase the fitness returns of females. The complementary choice model states that the best mate depends on the particular genotype of a female. Aculeate Hymenoptera represent a special case of complementary female choice because males should be chosen on the basis of their allele at the sex determination locus. The prevalent sex determination mechanism in bees and wasps (single-locus complementary sex determination) requires that, to produce a daughter, diploid offspring are heterozygous at the sex determination locus. Otherwise, infertile diploid males result. Inevitably, the proportion of diploid males increases with the rate of inbreeding. In the European Beewolf, males scent mark territories to attract mates and the composition of the pheromone might provide a basis for female choice. One crucial prerequisite for females to be able to discriminate against brothers and avoid inbreeding is that the male sex pheromone varies with familial affiliation. This hypothesis was tested by analysing the pheromone of male progeny of eight mothers using gas chromatography and mass spectrometry. A significantly higher similarity was found among brothers than among unrelated individuals. Such a genetic component of a male sex pheromone has not yet been described from aculeate Hymenoptera.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 433–442.  相似文献   

12.
In resource-based promiscuous mating systems, synchronous spawning of females affects competition among males and variation in the reproductive success of males. We documented the mating behavior of Japanese charr (Salvelinus leucomaenis japonicus) through an annual breeding season to examine the relationship between female spawning synchrony and male mating behavior. Females spawned highly synchronously in the population studied, i.e., approximately half the spawning was finished within the first three days of the entire spawning season (11 days). The daily operational sex ratio (OSR) was nearly 1:1 through the spawning period. The number of males around a spawning female was very small (1.21+/-0.49 males per female) over the spawning ground and period, suggesting that a competitive male could effectively chase subordinate males away from a spawning female. A few males attempted to sneak near the oviposition site of females (16%; 9 of 57 breeding groups), while some males adopted sneaking tactics in the initial phase of females' spawning (24%). We did not observe any males to succeed in sneak fertilizations. We conclude that in this Japanese charr population, the synchronous spawning of females was related to the unbiased daily OSR, male aggregation around females, and consequently whether and how efficiently males engaged in sneak mating behavior.  相似文献   

13.
Sex allocation theory predicts that in a population with a biased operational sex ratio (OSR), parents will increase their fitness by adjusting the sex ratio of their progeny towards the rarer sex, until OSR has reached a level where the overproduction of either sex no longer increases a parent''s probability of having grandchildren. Furthermore, in a monogamous mating system, a biased OSR is expected to lead to lowered mean fecundity among individuals of the more abundant sex. We studied the influence of OSR on the sex ratio of newborns and on the population birth rate using an extensive data set (n = 14,420 births) from pre-industrial (1775-1850) Finland. The overall effect of current OSR on sex ratio at birth was significant, and in the majority of the 21 parishes included in this study, more sons were produced when males were rarer than females. This suggests that humans adjusted the sex ratio of their offspring in response to the local OSR to maximize the reproductive success of their progeny. Birth rate and, presumably, also population growth rate increased when the sex ratio (males:females) among reproductive age classes approached equality. However, the strength of these patterns varied across the parishes, suggesting that factors other than OSR (e.g. socioeconomic or environmental factors may also have influenced the sex ratio at birth and the birth rate.  相似文献   

14.
During the spring emergence of red‐sided garter snakes (Thamnophis sirtalis parietalis) in Manitoba, Canada, the operational sex ratio is strongly skewed towards males, who scramble to locate and court newly emerged females. A high frequency of multiple paternity litters suggests that the females are promiscuous; the gelatinous copulatory plugs (CPs) deposited by males may confer fitness benefits via passive mate guarding. Because precopulatory female choice is limited in large mating aggregations, sexual conflict may place a premium on preventing females from ejecting male sperm. In snakes, sperm are produced in the testes and delivered through the ductus deferens, and the CP is thought to be produced by the renal sexual segment and conveyed through the ureter. We manipulated the delivery of the two fluids separately by surgically ligating the ducts. Ureter‐ligated males did not produce a CP, causing their sperm to leak out of the female's cloaca immediately after copulation. Contrary to previous suggestions, histology revealed sperm distributed throughout the CP. Thus, the CP may function as a spermatophore: the protein matrix contains the sperm, which are liberated gradually as the plug dissolves. The likelihood of a male depositing a CP fell significantly after his second mating, perhaps limiting his reproductive success. These results challenge the hypothesis that passive mate guarding is the primary function of the CP in T. sirtalis parietalis. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 893–907.  相似文献   

15.
In animal species where females mate with multiple males, female mating success might be expected to covary with aspects of female morphology, such as size or shape. Spiders are especially interesting in this regard, as the females of several spider groups weave intricate webs that often accommodate multiple male spiders, all of whom are potential mates. Because web design is likely to be dependent on female size/shape, we use multivariate methods to assess the relationships among female morphology, web design, and reproductive ecology over a range of body sizes in the orb-weaving spider Nephila clavipes . Of the measured variables, only abdomen size explained a significant amount of the variation in number of males on a web, and this relationship holds even after statistically accounting for body size. Because abdomen size is an indicator of body condition in spiders, we suggest that condition is likely to be an important factor relating to potential mating success in female spiders. We found no evidence for an association between web design and number of males on a web, although our data indicate that larger females build webs that are both larger and further from the ground than those of smaller females.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87 , 95−102.  相似文献   

16.
The amount of time taken to copulate varies enormously among mammals. Because copulation likely exposes animals to an increased risk of predation, and uses time and energy that could be spent on foraging, smaller mammals (which are vulnerable to more predators and have a shorter time-to-starvation than larger mammals) should spend less time copulating than do larger mammals. Furthermore, if extended copulation reflects competition among males, then the duration of mating (after correction for body size) should be greater in mammals in which females mate with more than one male. We tested these predictions using comparative data from 113 mammalian species in 85 genera, 40 families, and 14 orders, while controlling for the effect of phylogeny. We found: (1) the relationship between duration of copulation and body size to be negative, not positive; (2) no relationship with inferred multiple mating by females (based upon relative testes mass). We suggest that small mammals may find the sustained maneuvering and body positioning of copulation easier than do large mammals. This hypothesis is supported by an apparently isometric relationship between duration of copulation and ratio of power to mass.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 185–193.  相似文献   

17.
In anurans, female polyandry under male harassment is distributed across taxa because of external aquatic fertilization. According to the sexual selection theory, male–male competition for access to females is affected by the operational sex ratio (OSR) and population density. The Japanese common toad, Bufo japonicus, is widespread in mainland Japan, and like the European common toad, B. bufo, it engages in explosive breeding. In this study, we observed the breeding behaviour of B. japonicus in isolated local populations for over four years in two breeding ponds with different population sizes and densities: large‐low (L‐pond) and small‐high (S‐pond). We analysed the relative polyandry ratio in egg clutches laid by females and estimated the size‐assortative mating pattern to be an indicator of male–male competition in the two ponds. Both ponds tended to exhibit a size‐assortative mating pattern; however, the frequency of polyandry was different in the two ponds (L‐pond = 20% and S‐pond = 90%). Our results showed that polyandry could occur without multiple amplexus with a high population density, i.e. eggs were often fertilized by free‐swimming sperm in the small shallow pond. We propose that high female polyandry ratios without continuous male harassment are generated because of a male‐biased OSR and a high population density in the small pond. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 236–242.  相似文献   

18.
The operational sex ratio (OSR: sexually active males: receptive females) predicts the intensity of competition for mates. It is less clear, however, under what circumstances, the OSR predicts the strength of sexual selection – that is, the extent to which variation in mating success is attributable to traits that increase the bearer's attractiveness and/or fighting ability. To establish causality, experiments that manipulate the OSR are required. Furthermore, if it is possible to control for any OSR‐dependent changes in the chosen sex (e.g. changes in male courtship), we can directly test whether the OSR affects the behaviour of the choosing sex (e.g. female choice decisions). We conducted female mate choice experiments in the field using robotic models of male fiddler crabs (Uca mjoebergi). We used a novel design with two females tested sequentially per trial. As in nature, the choice of the first female to mate therefore affected the mates available to the next female. In general, we detected significant sexual selection due to female choice for ‘males’ with larger claws. Importantly, the strength of sexual selection did not vary across five different OSR/density treatments. However, as the OSR decreased (hence the number of available males declined), females chose the ‘males’ with the largest claws available significantly more often than expected by chance. Possible reasons for this mismatch between the expected and observed effects of the OSR on the strength of sexual selection are discussed.  相似文献   

19.
Sexual conflict over mating rate is both pervasive and evolutionarily costly. For females, the lifetime reproductive fitness costs that arise through interactions with potential mates will be influenced by the frequency of such interactions, and the fitness cost of each interaction. Both of these factors are likely to be influenced by variation in operational sex ratio (OSR) and population density. Variation in OSR‐ and density‐dependent male alternative reproductive tactics (ARTs) may be particularly important if the fitness costs that females experience vary with the reproductive tactics that males express. Using a simple model, we consider several examples of OSR‐ and/or density‐dependent variation in male ARTs and the frequency of male–female interactions, and find that variation in the expression of male ARTs has the potential to augment or diminish the costs of frequent male interactions for females. Accurately documenting variation in the expression of male ARTs and associated female fitness costs will benefit future work in this area.  相似文献   

20.
The intensity of mating competition and the potential benefits for female of mating with certain males can be influenced by several extrinsic factors, such that behavioral decisions can be highly context-dependent. Short-lived species with a single reproductive season are a unique model to study context-sensitive mating decisions. Through exhaustive sampling in the field and simultaneous choice tests in the laboratory, we evaluated operational sex ratio (OSR) and female mate choice at the beginning and end of the reproductive season in the annual killifish Austrolebias reicherti. We found seasonal change in both OSR and female mate choice. At the start of the reproductive season the OSR did not deviate from parity, and females preferred larger males. Later in the reproductive season, while the proportion of males in the ponds decreased, females became unselective with respect to male size. The particular biological cycle of annual killifish, where both life expectancy and mating opportunities decline sharply over a short timescale, could account for the seasonal change in female choice. Reduction in choosiness could arise from diminished reproductive prospects due to a decline in male availability. Moreover, as the end of the season approaches, any benefits of choosiness are presumably reduced: a female’s fitness will be higher if she mates with any male than if she forgoes reproduction and dies. Future work will disentangle the mechanisms underlying seasonal changes in mating preferences, notably direct responses to demographic factors, environmental cues, or intrinsic changes during development.  相似文献   

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