Fighting for females in the toad Bufo calamita is affected by the operational sex ratio

Fighting between male natterjack toads, Bufo calamita, for access to females was studied in relation to both the influence of male and female density each night and the operational sex ratio. Fighting males were larger than other males and were present in the breeding area on more nights but their average mating success was the same. Fighting between males was associated with not strongly male-biased operational sex ratios. The density of breeding individuals, however, was irrelevant. Take-over rate was very low with an insignificant effect on mating success.     

The effect of brood cannibalism on the operational sex ratio in parental teleost fishes

The operational sex ratio (OSR), an important determinant of the intensity of sexual selection, can vary with the population sex ratio and the potential reproductive rates of males and females. In parental teleosts, different forms of brood cannibalism, filial and non-kin, could affect the potential reproductive rates of males and females, and thereby the OSR. The direction of the effect of brood cannibalism on the OSR would depend on the sex of the cannibal and the parent. No evidence has been found that brood cannibalism can affect the OSR by altering the sex ratio.     

Urea synthesis and osmotic stress in the terrestrial anurans Bufo woodhousei and Hyla cadaverina

• 1.1. Bufa woodhousei. a terrestrial toad, and Hyla cadarerina, a terrestrial treefrog, accumulate and tolerate high concentrations of urea in the body fluids during dehydration or reduced water turnover (up to 280 mM urea in B. woodhousei, 140 mM urea in H. cadaverina).
• 2.2. In contrast to Rana cancrivora. Xenopus laevis and Scaphiopus couchi (considered to be unusual in their ability to handle osmotic stress), B. woodhousei and H. cadaverina do not increase the rate of urea synthesis and therefore the rate of urea accumulation in the body fluids in response to dehydration.
• 3.3. Hepatic activity of CPS-I. the first enzyme of the urea-ornithine cycle, is correlated with a greater proportion of N waste being formed as urea rather than as NH₃ during times of reduced water turnover or osmotic stress. This results in preferential accumulation of urea rather than of NH₃ in the body fluids.
• 4.4. CPS-I activity is also correlated with the increased N waste production that follows from increased dietary N intake.

     

Do females adjust the sex of their offspring in relation to the breeding sex ratio?

When the adult sex ratio differs between years in local populations, but still is predictable between adjacent years, it has been proposed that the best strategy would be to bias the offspring sex ratio in favour of the rare sex. We tested this hypothesis using a data set of great reed warbler offspring, sexed by molecular techniques, that were collected over 11 breeding seasons at two adjacent reed marshes. Three important assumptions for this hypothesis are fulfilled in the studied great reed warbler population. First, a substantial proportion of great reed warblers are living in small local populations where sex ratio distortions would be sufficiently large and common. Second, breeding adults and their offspring return to breed in the local population to a high degree. Third, females have a possibility to assess the breeding sex ratio before laying their eggs. At our study site, the breeding sex ratio was positively correlated between successive years. However, contrary to our prediction, female great reed warblers seemed not to adjust their offspring sex ratio in relation to the local breeding sex ratio.     

Adaptation to experimental alterations of the operational sex ratio in populations of Drosophila melanogaster

Theory predicts that males adapt to sperm competition by increasing their investment in testis mass to transfer larger ejaculates. Experimental and comparative data support this prediction. Nevertheless, the relative importance of sperm competition in testis size evolution remains elusive, because experiments vary only sperm competition whereas comparative approaches confound it with other variables, in particular male mating rate. We addressed the relative importance of sperm competition and male mating rate by taking an experimental evolution approach. We subjected populations of Drosophila melanogaster to sex ratios of 1:1, 4:1, and 10:1 (female:male). Female bias decreased sperm competition but increased male mating rate and sperm depletion. After 28 generations of evolution, males from the 10:1 treatment had larger testes than males from other treatments. Thus, testis size evolved in response to mating rate and sperm depletion, not sperm competition. Furthermore, our experiment demonstrated that drift associated with sex ratio distortion limits adaptation; testis size only evolved in populations in which the effect of sex ratio bias on the effective population size had been compensated by increasing the numerical size. We discuss these results with respect to reproductive evolution, genetic drift in natural and experimental populations, and consequences of natural sex ratio distortion.     

Asynchronous arrival pattern, operational sex ratio and occurrence of multiple paternities in a territorial breeding anuran, Rana dalmatina

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.     

Opposing effects of mortality factors on progeny operational sex ratio may thwart adaptive manipulation of primary sex ratio

Despite extensive research on mechanisms generating biases in sex ratios, the capacity of natural enemies to shift or further skew operational sex ratios following sex allocation and parental care remains largely unstudied in natural populations. Male cocoons of the sawfly Neodiprion abietis (Hymenoptera: Diprionidae) are consistently smaller than those of females, with very little overlap, and thus, we were able to use cocoon size to sex cocoons. We studied three consecutive cohorts of N. abietis in six forest stands to detect cocoon volume‐associated biases in the attack of predators, pathogens, and parasitoids and examine how the combined effect of natural enemies shapes the realized operational sex ratio. Neodiprion abietis mortality during the cocoon stage was sex‐biased, being 1.6 times greater for males than females. Greater net mortality in males occurred because male‐biased mortality caused by a pteromalid parasitic wasp and a baculovirus was greater and more skewed than female‐biased mortality caused by ichneumonid parasitic wasps. Variation in the susceptibility of each sex to each family of parasitoids was associated with differences in size and life histories of male and female hosts. A simulation based on the data indicated that shifts in the nature of differential mortality have different effects on the sex ratio and fitness of survivors. Because previous work has indicated that reduced host plant foliage quality induces female‐biased mortality in this species, bottom‐up and top‐down factors acting on populations can affect operational sex ratios in similar or opposite ways. Shifts in ecological conditions therefore have the potential to alter progeny fitness and produce extreme sex ratio skews, even in the absence of unbalanced sex allocation. This would limit the capacity of females to anticipate the operational sex ratio and reliably predict the reproductive success of each gender at sex allocation.     

Response of sex ratio to timing of breeding in the small cyprinid Gnathopogon caerulescens

The influence of hatching date on the sex ratio of wild Gnathopogon caerulescens was examined. Cohorts reared from eggs collected in the early and middle parts of the spawning season showed almost balanced sex ratios, with female bias in some cohorts. Cohorts born later in the season mostly displayed male bias, and the mean proportion of males later in the season was significantly higher than in early‐ and mid‐season cohorts. These results indicate that the sex ratio of G. caerulescens changes with the time of breeding, increasing along with the ambient water temperature of the lake.     

The influence of operational sex ratio on the intensity of competition for mates

The evolution and maintenance of secondary sexual characteristics and behavior are heavily influenced by the variance in mating success among individuals in a population. The operational sex ratio (OSR) is often used as a predictor of the intensity of competition for mates, as it describes the relative number of males and females who are ready to mate. We investigate changes in aggression, courtship, mate guarding, and sperm release as a function of changes in the OSR using meta-analytic techniques. As the OSR becomes increasingly biased, aggression increases as competitors attempt to defend mates, but this aggression begins to decrease at an OSR of 1.99, presumably due to the increased costs of competition as rivals become more numerous. Sperm release follows a similar but not significant trend. By contrast, courtship rate decreases as the OSR becomes increasingly biased, whereas mate guarding and copulation duration increase. Overall, predictable behavioral changes occur in response to OSR, although the nature of the change is dependent on the type of mating behavior. These results suggest considerable flexibility of mating system structure within species, which can be predicted by OSR and likely results in variation in the strength of sexual selection.     

Mate choice and the operational sex ratio: an experimental test with robotic crabs

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.     

Effects of angiotensin II and bladder condition on hydration behavior and water uptake in the toad,Bufo woodhousei

• 1.1. Water absorption response (WR) behavior and water weight gain were examined in hydrated toads, Bufo woodhousei, treated with angiotensin II (All) or with a control Ringer's solution. The effects of urinary bladder condition (ad lib. bladder urine or empty bladder) were examined concurrently.
• 2.2. Toads treated with All (100μg/100g body weight), spent more time in WR posture and absorbed more water than Ringer's-injected toads.
• 3.3. Toads with empty bladders maintained WR posture for longer periods of time and gained more weight than toads whose bladders were not emptied.
• 4.4. The effects of All and bladder urine on water absorption by B. woodhousei appear to be separate and additive.

     

Effect of 6-methoxybenzoxazolinone on sex ratio and breeding performance in Microtus montanus

The plant derivative, 6-mathoxybenzoxazolinone (6-MBOA), which has been demonstrated to initiate reproduction in field populations of the montane meadow vole (Microtus montanus), was administered via feeding or Silastic capsule implants to mated pairs of laboratory bred M. montanus. The animals remained paired for 120 days, and the number, size, and sex ratios of the resulting litters were recorded. Both the size and frequency of litters were significantly greater in 6-MBOA-treated pairs than in controls. By using implants, it was possible to treat one or both sexes in a mated pair. The positive effects of this compound on litter size and number of litters occurred when the females received implants, which indicates that the male has no influence on these parameters. The most unusual result of these experiments was that 6-MBOA has a significant effect on the sex ratio of the litters. Animals receiving 6-MBOA produced significantly more females than did control pairs. This result occurred regardless of the method of administration, and in the case of the implant studies, regardless of which sex received the active implants. These findings are discussed in relation to the ecology and life history strategy of Microtus montanus.     

Quantitative measure of sexual selection with respect to the operational sex ratio: a comparison of selection indices

Despite numerous indices proposed to predict the evolution of mating systems, a unified measure of sexual selection has remained elusive. Three previous studies have compared indices of sexual selection under laboratory conditions. Here, we use a genetic study to compare the most widely used measures of sexual selection in natural populations. We explored the mating and reproductive successes of male and female bank voles, Clethrionomys glareolus, across manipulated operational sex ratios (OSRs) by genotyping all adult and pup bank voles on 13 islands using six microsatellite loci. We used Bateman's principles (Is and I and Bateman gradients) and selection coefficients (s' and beta') to evaluate, for the first time, the genetic mating system of bank voles and compared these measures with alternative indices of sexual selection (index of monopolization and Morisita's index) across the OSRs. We found that all the sexual selection indices show significant positive intercorrelations for both males and females, suggesting that Bateman's principles are an accurate and a valid measure of the mating system. The Bateman gradient, in particular, provides information over and above that of other sexual selection indices. Male bank voles show a greater potential for sexual selection than females, and Bateman gradients indicate a polygynandrous mating system. Selection coefficients reveal strong selection gradients on male bank vole plasma testosterone level rather than body size.     

Mating distribution and its temporal dynamics affect operational sex ratio: a simulation study

The present study investigated how variation in mating distribution in time and among males influences the operational sex ratio (OSR) with a simulation inspired by paternally caring fish. Varying (1) the potential reproductive rate of each sex, (2) the mating distribution among males, and (3) the length of male mating phase, we created different mating patterns. In each case, we searched for the adult sex ratio that resulted in an OSR of 50% (where sex-roles switch). This approach enabled a comparison with a previous model. We found that the OSR was influenced by the distribution of matings in time and among males when the male mating phase was limited by a parental phase. Furthermore, the mating dynamics were shaped by the fact that the numbers of males and females and their capacities for collateral investment affected OSR immediately from the start of the reproductive season, whereas their times-out had a delayed effect on OSR.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 551–559.