Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.     

No differential consequences of reproduction according to sex in Juniperus communis var. depressa (Cupressaceae)

In dioecious plant species, males and females are thought to have dissimilar allocation patterns. Females are believed to invest more in reproduction and less in growth and maintenance than males. This differential investment between sexes could result in distinct growth patterns and contrasting survival rates, thereby affecting the sex ratio of a population and the age and size distribution of males and females, possibly leading to habitat segregation according to sex. These effects might become more apparent under particularly limiting conditions, such as in nutrient-deficient soils or in climatically stressed environments. To verify these predictions, growth patterns, microsite characteristics, and age and size distribution of male and female individuals were compared, and population sex ratio was determined in three populations of the dioecious shrub Juniperus communis var. depressa (Cupressaceae, Pinophyta) along a short latitudinal gradient on the eastern coast of Hudson Bay (Northern Québec, Canada). We found that the northernmost population had a male-biased sex ratio, but that the southernmost one had a higher proportion of females. Our results failed to reveal any significant differences in radial growth patterns, mean sensitivity, annual elongation of the main axis, and size and age frequency distribution between males and females in any population. Furthermore, there was no evidence of microhabitat segregation according to sex as indicated by the lack of differences in the physicochemical characteristics of the substrate under males and females. Clearly, the expected ecological consequences of a presumed greater investment of females in reproduction were not apparent even under the very stressful conditions prevailing on subarctic dunes. Many factors could reduce differences in the cost of reproduction between males and females, such as the number and quality of reproductive structures produced annually by individuals of each sex, the possible photosynthetic activity of the immature female cones, and the complexity of the source/sink relationship within individuals. Alternatively, there may be no differences between sexes in their reproductive investment.     

Effects of population structure and density on calf sex ratio in red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>)—implications for management

There is ongoing interest to assess what factors affect offspring sex ratio, especially in ungulates. Wildlife managers might be interested in influencing this sex ratio for two reasons: either in order to limit population growth more effectively by reducing the proportion of females born or to increase revenues by a higher proportion of trophy bearing males in the population. While previous studies mostly focused on how maternal traits affect offspring sex ratio, we included here also male traits in our analysis. We achieved this by investigating data from 30 areas covering entire Lower Austria, collected over the past 12 years from both hunted red deer and those killed in road accidents. We focused our analyses on parameters that can be easily assessed by managers on the population and individual level, i.e. the numbers of animals culled in different age/sex classes and their body mass. We found that the proportion of females among calves increased with population density. Furthermore, we found that calf sex ratio (i.e. the proportion males among calves aged between 2 and 7 months) increased with increasing proportions of adult females and males older than 10 years, independent of the density effect. We conclude that wildlife managers interested in the effective reduction of red deer abundance and/or increasing the proportion of males among offspring should select a culling regime leading to a low population density dominated by adult, prime-aged females and males. This can be achieved by over-proportional removal of young females and warranting that a high number of strong males reach an age of at least 10 years.     

Age‐ and sex‐specific survival of California sea lions (Zalophus californianus) at San Miguel Island,California

We conducted a mark‐recapture study of California sea lions (Zalophus californianus) using pups branded on San Miguel Island, California, from 1987 to 2014, and annual resightings from 1990 to 2015. We used the Burnham model (Burnham 1993), an extension of the Cormack‐Jolly‐Seber mark‐recapture model, which includes recoveries of dead animals, to analyze age, sex, and annual patterns in survival. Generally, females had higher survival than males. For female pups, the average annual survival was 0.600 and for male pups it was 0.574. Yearling survival was 0.758 and 0.757 for females and males, respectively. Peak annual survival was at age 5 and was 0.952 for females and 0.931 for males. Pups with larger mass at branding had higher survival as pups and yearlings, but the effect was relative within each cohort because of large between‐cohort variability in survival. Annual variability in sea surface temperature (SST) affected survival. For each 1°C increase in SST, the odds of survival decreased by nearly 50% for pups and yearlings; negative SST anomalies yielded higher survival. Annual variation in male survival was partly explained by leptospirosis outbreaks. Our study provides a unique view of one demographic parameter that contributed to the successful recovery of the California sea lion population.     

Analysing the effect of movement on local survival: a new method with an application to a spatially structured population of the arboreal gecko Gehyra variegata

Mortality during movement between habitat patches is the most obvious cost of dispersal, but rarely it has been demonstrated empirically. An approach is presented, which uses capture–mark–recapture data of an arboreal gecko species to determine the effect of individual movement on local survival in a spatially structured population. Because capture–mark–recapture data are widely available for a range of animal species, it should be possible to extend their application to other species. The method is based on the assumption that the tendency to be a territorial animal or to be a floating animal is fixed during the study period. The advantage of our approach is that only one additional parameter has to be estimated for describing movement risks. We further tested the power of our approach to detect an association of movement and mortality with simulated capture histories. The study revealed a strong negative effect of movement on local survival. Hence, animals that moved more often between trees had a lower survival rate. Interestingly, the mean movement rate for males was significantly higher than for females, which should lead to a biased sex ratio towards females in the population. As there was an even sex ratio in the population, we discuss not mutually exclusive explanations for this finding like differences in emigration rates between sexes, differences in survival rates between sexes, or a skewed sex ratio in offspring.     

Population ecology of Trituras cristatus and T. vulgaris (Urodela) in SW Sweden

The population ecology of newts was investigated between 1969 and 1974 in two localities near Gothenburg, SW Sweden: the Gunnebo park pond (about 300 m²) where Triturus cristatus Laurenti and T. vulgaris L. are sympatric, and a rock-pool (6 m²) in Billdal with only T. vulgaris. At Gunnebo, a population of about 350 breeding T. cristatus was found with about as many males as females but males being more aquatic than females. An annual survival of 0.7–0.8 was estimated for the adults. The T. vulgaris population at the same locality was estimated to about 250–300 breeding adults with a sex ratio possibly in favour of males. The survival of adults was found to be extremely low and a very small percent of the adults seems to breed more than once. The Billdal T. vulgaris population was estimated to contain nearly 70 breeding adults with a sex ratio very near to 1:1. The annual survival of adults was found to be 0.5. The different survival of the two T. vulgaris populations are discussed with particular reference to predation. The age at which newts breed for the first time seems to vary; for T. vulgaris five years may well be the normal age. Factors that may limit the sizes of the populations, possible reasons for differences in frequency of occurrence in water, and possible predators are discussed.     

Bigger is better: age class-specific survival rates in long-lived turtles increase with size

Vital rates for small, non-breeding individuals are important components of population dynamics for many species, but often individuals of these sizes are difficult to locate, capture, and track. As such, biologists frequently lack reliable estimates of juvenile survival because sample sizes and recapture rates for this life stage are low. Long-lived animals often take many years to reach sexual maturity and spend much of this time in the smaller size classes, making them sensitive to changes in survival rates. We estimated the survival rates of all size classes for the northern map turtle (Graptemys geographica) using a mark-recapture dataset with >3,500 captures from 2019–2021 and 210 nests from 2018–2021. As turtle size increased, annual survival probability increased regardless of sex. Estimated annual survival probability for turtles >18 cm long (i.e., adult females >15 years) was about 0.95, over 4 times higher than turtles that were 3 cm long (i.e., hatchlings <1 year; 0.22 annual survival probability). Although we did not observe a difference in survival probability between sexes of any size class, adult females are nearly twice the size of adult males, leading to an increased annual survival probability for females of 0.95, compared to 0.80 for males. Changes in adult survival had the greatest influence on population estimates over time, with temporary decreases, such as those due to poaching or an environmental disaster, potentially leading to unrecoverable decreases in the overall population size. Our study provides detailed survival rates for all size classes in a long-lived turtle, which are necessary to assess population stability and can be used to determine the most effective conservation or management practices.     

Sex ratio, survival and territorial behaviour of polygynous Hen Harriers Circus c. cyaneus in Orkney

Polygyny is widespread among Orkney's Hen Harriers Circus cyaneus . From 1975–81, it was associated with a sex ratio estimated from resightings of colour-marked birds up to six years of age to be 29 females:ten males. The sex ratio of fledglings changed significantly from a greater proportion of females in the 1950s and early 1960s to a greater proportion of males subsequently. The mean estimated 'survival' rates (birds colour-marked in Orkney which were seen there in later years) of males and of females 0–2 years old were 14 and 29%. The mean estimated annual survival rates of males and females from 2–6 years old were 72 and 90%. It is suggested that the uneven sex ratio resulted in more frequent intra-sexual encounters and displays by females. The results of temporary removals of two females (immediately replaced) and of two males (not replaced) in spring indicates that there was a shortage of males in the population.     

Different cost of reproduction for the males and females of the rare dioecious shrub Corema conradii (Empetraceae)

Males and females of dioecious plant species often differ in their reproductive investment. Such differences frequently result in differential demographic costs represented by lower growth, survival, and/or frequency of reproduction, and/or by more variable reproductive effort through time for females. We present the results of a study on Corema conradii, a rare dioecious shrub of the coastal dune heathlands of northeastern North America. We estimated the reproductive investment of both males and females, determined their age structure, and compared their spatial patterns in a population at ?les-de-la-Madeleine, Quebec. We also determined the sex ratio of the four populations known to occur on the islands. Males invested more in reproduction at flowering, but when fruit production was considered, female reproductive investment was higher in terms of biomass, Mg, and Ca, but not in terms of N, P, and K. The age frequency distribution of males and females did not differ significantly from one another. The population dispersion pattern was contagious, with patches of similar-age individuals. There was no spatial segregation between males and females, although the sex ratio varied somewhat spatially. Females did not start reproducing at a later age than males and did not appear to have a shorter longevity. However, the crown and radial growth rates of females were lower than those of males. When estimated by the crown intercept method, the sex ratio of all four populations was male biased. However, because males had a higher crown growth rate, genet sex ratio was in fact balanced. Higher investment in reproduction was associated with a lower growth rate, which represents a differential cost of reproduction according to sex in this species.     

Sex ratio,survival, and recapture rate in a Cuban population of the damselfly <Emphasis Type="BoldItalic">Hypolestes trinitatis</Emphasis> (Odonata: Megapodagrionidae)

Male-biased sex ratios in adult odonate populations have been the subject of vigorous discussion between the students of this order of insects. The debate has centered on whether the observed male bias in many populations is real, perhaps due to unequal survival rates, or whether it is an artifact caused by differences in recapture probabilities. A mark–recapture study to assess the relative contribution of survivorship and recapture rates on male-biased sex ratio was performed in a Cuban population of the damselfly Hypolestes trinitatis. Maximum likelihood theory and Akaike information criterion were used for parameter estimation and model selection, respectively. Females in the sample were outnumbered two to one by males. Estimated recapture and survival rates were 0.188 (females) and 0.638 (males), and 0.933 (females) and 0.944 (males), respectively. Recapture rates only partially explained the bias since the population sex ratio estimated after correcting for differences in this parameter was male biased (1.5). The observed higher survival probabilities in males could have generated the male-biased population sex ratio. Therefore, we concluded that the observed male-biased population sex ratio in H. trinitatis is real.     

Sex ratio from germination through maturity and its reproductive consequences in the liverwort Sphaerocarpos texanus

Summary The dioecious winter ephemeral liverwort, Sphaerocarpos texanus disperses spore tetrads consisting of two males and two females. I examined the subsequent sex ratio of S. texanus at different stages in its life cycle to detect possible mechanisms affecting deviations from a 1:1 sex ratio and the effect of sex ratio on reproductive success. As S. texanus occurs in pure male, pure female and mixed sex clumps, I examined the proportions and sizes of these, the reproductive success of pure female and mixed sex clumps in the field and the sex ratio of germinating plants in a growth chamber. In both the field and the growth chamber the most abundant clump type was pure female followed by mixed sex and pure male clumps. These abundance patterns suggest that males have a lower survival rate than females before emergence and this lower survival rate continues through the gametophytic stage. This disadvantage may be due to the higher susceptibility of males to environmental conditions, to their competitive inferiority to females, and/or to differential resource allocation to the sexes within the spore tetrad. The female biased sex ratio at germination is consistent with predictions from sex ratio theory. Further my field data indicate that males may gain a survival benefit from growing in a mixed sex clump and both males and females benefit reproductively when they occur in mixed sex clumps.     

What causes male‐biased sex ratios in mature damselfly populations?

1. Several hypotheses to account for biased sex ratios in mature insect populations were tested by monitoring two field populations of the damselfly Lestes sponsa and by performing experiments in field cages. The population sex ratios are heavily male biased in this species. 2. The observed sex ratio at emergence was even and both sexes emerged synchronously. Females had longer maturation times but these were insufficient to explain the observed sex ratio shift. 3. Mass increases during maturation were consistently larger in females. In agreement with this, immature females made more flights per unit of time, which should make them more vulnerable to predation, however maturation probabilities were lower in females only in one field cage experiment. This inconsistency may be due to long bad weather conditions. Interestingly, predators reduced mass increase and this reduction was larger in females than in males. 4. Calculations based on the sex specific maturation times show that only slightly lower daily survival probabilities during maturation in females are enough to generate the observed sex ratio shift. 5. Mature survival was higher in males than in females in one field population but not in another, indicating that this cannot be a general mechanism causing the sex ratio. A higher maturation probability in males is therefore the most plausible mechanism causing the sex ratio shift in damselfly populations.     

Female choice for good genes and sex-biased broods in guppies

In a population of guppies Poecilia reticulata females were found to prefer large males and the offspring of these males had a higher survival rate than those sired by smaller males, suggesting that females were selecting their mates on the basis of their good genes. The possibility that females differentially invested in male or female offspring depending on the size or attractiveness of their mate was also investigated, but no relationship was found between a male's attractiveness or body size and the sex ratio of the offspring he sired. However, a strong female-biased sex ratio was detected in the broods and the proportion of males produced decreased with the amount of time that a female was confined with a male. The possible reasons for this are discussed.     

Sex ratio of North Island kaka (

The sex ratio of the kaka population inhabiting the Waihaha Ecological Area, Pureora Forest Park was estimated between late October 1994 and January 1995. The observed sex ratio estimate was three males to one female compared to a capture rate (using mist nets) of six to one in the same area between January and June 1994. Females appeared to be less susceptible to capture than males. The skewed sex ratio toward male kaka was significant and suggests that female kaka suffer higher mortality (probably due to predation at the nest) than males. The slow breeding rate of kaka combined with a high loss of breeding females is a serious threat to the long term survival of the kaka population within the Waihaha Ecological Area.     

Sex‐related site fidelity of humpback whales (Megaptera novaeangliae) to the Fueguian Archipelago feeding area,Chile

We investigated sex‐related site fidelity by humpback whales to the Fueguian Archipelago, a new feeding area in the eastern South Pacific, by examining the resighting histories of 45 males and 39 females recorded from 2003 to 2012. Results indicated an overall annual return to the feeding area of 74.8%, and annual sex ratio is roughly equal in the population. The probability of an individual being resighted across years and in subsequent years was not significantly different for both males and females, however, the proportion of resighting within a year was significantly higher for individual males compared to females. Potential sources of sex‐related bias were analyzed, but none were found to be significant. Greater intraannual resighting frequency for males may reflect sex‐based differences in spatial occupation and short‐range movements due to potential differences in energy budgets.     

Influential factors for natal dispersal in an avian island metapopulation

Dispersal is a key parameter in evolutionary, demographic and conservation theory, but the factors influencing dispersal between populations are rarely known, and the contribution of immigrants to population stability remains uncertain. Using dispersal data from nine island populations of song sparrows, we show that female and male immigrants responded differently to population structure: in females, immigration varied with adult sex ratio; whereas immigration by males was more influenced by population density. These patterns are consistent with the hypothesis that intra-sexual competition for breeding resources influenced recruitment patterns. Immigrants often constituted a substantial fraction of local population size, and in six cases immigration by females prevented the extirpation of that sex from the island. Breeding vacancies and extirpations may have been more likely in females because their apparent survival was lower than in males. Local recruitment and immigration varied markedly among islands, perhaps as consequence of island size and isolation. Overall, our results suggest that immigration varied with local demography in a sex-specific way, stabilized population numbers and reduced extinction rates in the smallest populations.     

Sex-role reversal of a monogamous pipefish without higher potential reproductive rate in females

In monogamous animals, males are usually the predominant competitors for mates. However, a strictly monogamous pipefish Corythoichthys haematopterus exceptionally exhibits a reversed sex role. To understand why its sex role is reversed, we measured the adult sex ratio and the potential reproductive rate (PRR), two principal factors influencing the operational sex ratio (OSR), in a natural population of southern Japan. The adult sex ratio was biased towards females throughout the breeding season, but the PRR, which increased with water temperature, did not show sexual difference. We found that an alternative index of the OSR (Sf/Sm: sex ratio of 'time in') calculated from the monthly data was consistently biased towards females. The female-biased OSR associated with sex-role reversal has been reported in some polyandrous or promiscuous pipefish, but factors biasing the OSR differed between these pipefish and C. haematopterus. We concluded that the similar PRR between the sexes in C. haematopterus does not confer reproductive benefit of polygamous mating on either sex, resulting in strict monogamous mating, and its female-biased adult sex ratio promotes female-female competition for a mate, resulting in sex-role reversal.     

Sexual dimorphism,survival, and parental investment in relation to offspring sex in a precocial bird

Differential growth rate between males and females, owing to a sexual size dimorphism, has been proposed as a mechanism driving sex‐biased survival. How parents respond to this selection pressure through sex ratio manipulation and sex‐biased parental investment can have a dramatic influence on fitness. We determined how differential growth rates during early life resulting from sexual size dimorphism affected survival of young and how parents may respond in a precocial bird, the black brant Branta bernicla nigricans. We hypothesized that more rapidly growing male goslings would suffer greater mortality than females during brood rearing and that parents would respond to this by manipulating their primary sex ratio and parental investment. Male brant goslings suffered a 19.5% reduction in survival relative to female goslings and, based on simulation, we determined that a female biased population sex ratio at fledging was never overcome even though previous work demonstrated a slight male‐biased post‐fledging survival rate. Contrary to the Fisherian sex ratio adjustment hypothesis we found that individual adult female brant did not manipulate their primary sex ratio (50.39% male, n = 645), in response to the sex‐biased population level sex ratio. However, female condition at the start of the parental care period was a good predictor of their primary sex ratio. Finally, we examined how females changed their behavior in response to primary sex ratio of their broods. We hypothesized that parents would take male biased broods to areas with increased growth rates. Parents with male biased primary sex ratios took broods to areas with higher growth rates. These factors together suggest that sex‐biased growth rates during early life can dramatically affect population dynamics through sex‐biased survival and recruitment which in turn affects decisions parents make about sex allocation and sex‐biased parental investment in offspring to maximize fitness.     

Assessment of costs of reproduction in a pelagic seabird using multistate mark-recapture models

We used a long‐term population band‐resight survey database, a parallel reproduction database, and multistate mark–recapture analysis to assess the costs of reproduction, a keystone concept of life‐history evolution, in Nazca boobies (Sula granti) from Punta Cevallos, Isla Española, Galápagos, Ecuador. We used eight years of resight and breeding data to compare models that included sex‐ and state‐specific survival probabilities and probabilities of transition between reproductive states using multistate mark–recapture models. Models that included state‐specific effects were compared with models lacking such effects to evaluate costs of reproduction. The top model, optimizing the trade‐off of model simplicity and fit to the data using the Akaike Information Criterion (AIC), showed evidence of a temporally varying survival cost of reproduction: nonbreeders showed higher annual survival than breeders did in some years. Because increasing investment among breeders showed no negative association with survival and subsequent breeding success, this evidence indicates a cost to both males and females of initiating, but not of continuing, a reproductive attempt. In some cases, breeders reaching the highest reproductive state (fledging an offspring) showed higher survival or subsequent breeding success than did failed breeders, consistent with differences in overall quality that promote both survival and reproduction. Although a male‐biased adult sex ratio was observed in this population of Nazca boobies, models of state‐ and sex‐specific survival and transition probabilities were not supported, indicating that males and females do not incur different costs of reproduction, and that the observed sex ratio bias is not due to sex‐specific adult mortality.     

Hatching sex ratio and sex specific chick mortality in common terns Sterna hirundo

Bias in sex ratios at hatching and sex specific post hatching mortality in size dimorphic species has been frequently detected, and is usually skewed towards the production and survival of the smaller sex. Since common terns Sterna hirundo show a limited sexual size dimorphism, with males being only about 1–6% larger than females in a few measurements, we would expect to find small or no differences in production and survival of sons and daughters. To test this prediction, we carried out a 2-year observational study on sex ratio variation in common terns at hatching and on sex specific post hatching mortality. Sons and daughters hatched from eggs of similar volume. Post hatching mortality was heavily influenced by hatching sequence. In addition, we detected a sex specific mortality bias towards sons. Overall, hatching sex ratio and sex specific mortality resulted in fledging sex ratios 8% biased towards females. Thus, other reasons than body size may be influencing the costs of rearing sons. Son mortality was not homogeneous between brood sizes, but greater for two-chick broods. Since adults rearing two-chick broods were younger, lighter and bred consistently later than those rearing three-chick broods, it is suggested that lower capacity of two-chick brood parents adversely affected offspring survival of sons. Though not significantly, two-chick broods tended to be female biased at hatching, perhaps to counteract the greater male-biased nestling mortality. Thus, population bias in secondary sex ratio is not limited to strongly size dimorphic species, but species with a slight sexual size dimorphism can also show sex ratio bias through a combination of differential production and mortality of sons and daughters.