Seasonal, size- and age-related patterns in body-mass and composition of Purple Sandpipers Calidris maritima in Britain

The masses of 3229 Purple Sandpipers Calidris maritima from Britain were analysed for differences related to age, season and size. First-year birds were lighter by 2 g. There was only a slight increase in mass in mid-winter, in contrast to other waders wintering in Britain, suggesting that Purple Sandpipers are less at risk to cold weather and food shortages. Their winter fat reserves were low but their breast muscles were relatively larger than other small waders wintering in Britain. Their plumage was slightly heavier and their breast and belly feathers were longer. There was a large increase in mass in May by first-year and adult birds prior to migration. The 'long-billed' population showed a greater relative increase in May, compared with the 'short-billed' population, perhaps reflecting different migration distances.     

Population trends and habitat use of Harlequin Ducks in Rhode Island

ABSTRACT.   To assess population trends of Harlequin Ducks ( Histrionicus histrionicus ) in Rhode Island (U.S.A.), we analyzed Christmas Bird Counts and other historical surveys and also conducted surveys during the winter of 2005–2006. We estimated sex and age ratios, evaluated the effects of tidal regime and time of day on survey precision, and quantified habitat use. The population in Rhode Island experienced logistic growth from 1976 to 2004, with approximately 150 birds now wintering at three primary sites in the state. We estimated that the current ratio of males to females in the region was 1.6:1 (62% males) and that 13% of males were first-winter birds. Most Harlequin Ducks were observed in rocky habitats within 50 m of the shore or offshore islands. We detected the greatest numbers of birds, with the least amount of variation, during morning surveys at low tide, suggesting that this may be the most appropriate time for population monitoring. Increases in the Rhode Island population and male-biased sex ratios may indicate a local population recovery resulting from a hunting ban initiated in 1990. Although most Harlequin Ducks in eastern North America winter in Maine, the population in Rhode Island represents one of the largest in the southern part of their range.     

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.     

Seasonal variation in the sex and age composition of the woodcock bag in Denmark

The Eurasian woodcock is a highly valued game bird in Western Europe from which c. 2.7 million individuals are harvested annually from an estimated population of 20–26 million birds. The population size and status remains uncertain due to the cryptic behaviour and widespread and solitary occurrence of woodcock, on breeding and wintering areas, making reliable population surveys difficult. Hunting bag records provide age ratios amongst bagged birds, but sex ratios remain poorly known because of the sexually monomorphic nature of this species. We used DNA analysis to determine sex ratios amongst 327 shot woodcocks from two hunting seasons in Denmark (1 October–31 January, 2012/13 and 2013/14). Based on bag totals, age ratios and sex ratios, juvenile females constituted 37%, juvenile males 27%, adult females 16% and adult males 20% of the annual woodcock bag. The female bias was related to a significant deviation from parity in the sex ratio amongst juvenile birds in October, although no such deviation was found at other times or amongst adults. Compared to limited data from other European countries, our data suggest that autumn migration of woodcock involves an initial wave of juvenile females followed by juvenile males and adults, and perhaps that males stay further north in Europe than females during autumn and winter. This migratory pattern would suggest that postponing the opening of the hunting season could reduce the hunting bag on reproductively valuable females in this polygamous species.     

Offspring sex ratio is related to paternal train elaboration and yolk corticosterone in peafowl

Several recent experimental studies have provided strong evidence for the ability of birds to manipulate the sex ratio of their offspring prior to laying. Using a captive population of peafowl (Pavo cristatus), we tested experimentally the effects of paternal attractiveness on offspring sex ratio, and related sex ratio deviations to egg-yolk concentrations of testosterone, 17beta-estradiol and corticosterone. When females were mated to males whose attractiveness had been experimentally reduced by removing prominent eyespot feathers from their trains, they produced significantly more female offspring, had significantly higher yolk corticosterone concentrations and tended to have lower levels of yolk testosterone than when mated to the same males with their full complement of feathers. Concentrations of 17beta-estradiol did not vary consistently with sex ratio biases. These findings add to the small number of studies providing experimental evidence that female birds can control the primary sex ratio of their offspring in response to paternal attractiveness, and highlight the possibility that corticosterone and perhaps testosterone are involved in the sex manipulation process in birds.     

Fluctuation in abundance and age structure of a breeding population of the Japanese brown frog,Rana japonica gunther (Amphibia,anura)

A breeding population of Rana japonica was studied at a marsh on the campus of Hiroshima University in Higashi-Hiroshima during the five years 1995-1999. The mark-recapture study showed that the size of the breeding population varied from year to year, and increased more than twofold in 1999 in comparison with the preceding years. The sex ratio of the breeding population (male/female) was from nearly 1.0 to 1.6. Frogs of both sexes were estimated to breed for the first time at the age of one or two years, and their maximum age was four years according to skeletochronology using phalanges and mark-recapture. Modes of the estimated ages were one year for males during the study years except 1997, but one or two years for females. Two thirds of breeding frogs, irrespective of their sex, were estimated to breed only once throughout their lives.     

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-biased hatching sequences in the cooperatively breeding Noisy Miner

The Noisy Miner Manorina melanocephala (Meliphagidae) is a cooperatively breeding bird species in which sons often remain on their natal home ranges and help one or both of their parents. In a population of Noisy Miners in SE Queensland, Australia, a molecular technique was used to explore adult and offspring sex ratios, and also hatching sequences. Among the adult population, there were 2.31 males for every female, and roughly 99% of helping was performed by males. At hatching and fledging, the population sex ratio was even, with exactly 57 males and 57 females. However, in 17 out of 18 broods the first egg to hatch was male. First-hatched males were significantly larger and heavier than their sisters just prior to fledging. Through their helping behaviour, large healthy sons could clearly enhance the future reproductive success of parents, and benefit the entire group. Sex-biased hatching sequences could potentially provide cooperatively breeding birds with a subtle and precise way of varying investment in the helping sex.     

Adult survival and numbers in a coastal breeding population of Redshank Tringa totanus in northwest England

The results of a long-term capture-mark-recapture ringing programme carried out on a coastal population of breeding Redshanks Tringa totanus between 1974 and 1988 are presented.
Both sexes were equally likely to be recaptured in subsequent years, as were birds captured for the first time compared with those that had been captured previously. Older birds were more frequently recaptured than were young birds. There was no significant difference in male and female adult survival rates, with a mean of 72% of females and 75% of males surviving each year.
The breeding population fluctuated annually with estimated breeding densities of 122–285 pairs per km². Variation in breeding numbers (males), but not survival, was partially attributable to winter air temperature.     

Dispersal of male ortolan buntings away from areas with low female density and a severely male-biased sex ratio

Dispersal is expected to enhance individual fitness, and individuals should thus disperse from areas with poor conditions to areas with more favourable conditions. Few studies have compared conditions before and after dispersal of the same individuals, and in birds little is known about the effects of sex ratio and female density on male dispersal decisions. In this study we examined various fitness-related parameters that adult male ortolan buntings, Emberiza hortulana, might use as cues in their decisions to disperse and settle. The study population has a strongly male-biased sex ratio. Using pairwise comparisons of pre- and post-dispersal conditions, we found that males moved from areas with low female density and a severely male-biased sex ratio to areas with higher female density and a less male-biased sex ratio. Male density and male age structure did not affect male dispersal and settlement. The sex ratio of the pre-dispersal sites was below the population average, but post-dispersal sites were not significantly better than the population average. This suggests that dispersal was triggered by poor conditions, whereas settlement may have been unrelated to the conditions at the new site. In the year of dispersal, males that undertook dispersal were less successful at acquiring a female than males that remained faithful to the site, but in subsequent years there was no difference. We suggest that dispersal might be adaptive under more natural conditions with a more balanced sex ratio. These results show that male ortolan buntings abandon areas with low female density and a severely male-biased sex ratio, and thereby end up in better places, although probably not better than the population average.     

Adult sex ratios in wild bird populations

Offspring sex ratios in wild bird populations, and the extent to which they vary from the equality expected by random genotypic sex determination, have received much recent attention. Adult sex ratios (ASRs) in wild birds, on the other hand, remain very poorly described, and many of the questions about them posed by Ernst Mayr in 1939 remain unanswered. This review assesses population-level sex ratio patterns in wild bird populations, with an emphasis on the ASR. A quantitative assessment of over 200 published estimates of ASR, covering species from a wide range of taxa, regions and habitats, supported Mayr's assertion that skewed ASRs are common in wild bird populations. On average, males outnumbered females by around 33%, and 65% of published estimates differed significantly from equality. In contrast, population-level estimates of offspring sex ratio in birds did not generally differ from equality, and mean ASR across a range of wild mammal species was strongly female-skewed. ASR distortion in birds was significantly more severe in populations of globally threatened species than in non-threatened species, a previously undescribed pattern that has profound implications for their monitoring and conservation. Higher female mortality, rather than skewed offspring sex ratio, is the main driver of male-skewed ASRs in birds, and the causes and implications of this are reviewed. While estimates of ASR in wild bird populations may be subject to a number of biases, which are discussed, there is currently no quantitative evidence that an ASR of one male to one female represents the norm in birds. A better understanding and reporting of ASRs in wild bird populations could contribute greatly to our understanding of population processes and could contribute much to theoretical and applied research and conservation.     

Turnover and recruitment in a tundra population of Peregrine Falcons Falco peregrinus

The population dynamics of tundra-nesting Peregrine Falcons Falco peregrinus tundrius was studied over 7 years on a 450 km² study area in the Keewatin District of Canada's Northwest Territories. Peregrines showed strong fidelity to nest sites; none of 25 males ringed changed territories, while five of 38 females ringed were recorded changing territories. Such changes usually occurred after nesting failure. Annual turnover of territorial adults was estimated to be 22% (15% for males and 26% for females). Annual mortality of adults was estimated to be 17% (15% for males and 19% for females). If we assumed that territory vacancies, in addition to mate replacements, were indicative of mortality, then maximum annual mortality was estimated at 24% for each sex. Territories were held only by adult Peregrines. The oldest male on territory was at least 7 years old, the youngest was 2. The oldest female on territory was at least 7 years old, the youngest was 3. Territories were held by individuals of each sex for at least 6 years. One pair remained together for at least 4 years. Less than 4% of all young Peregrines produced on the study area in the first 5 years of the study were recruited into the breeding population. More male than female young were recruited despite an even sex ratio among nestlings. Peregrines did poorly in their first breeding attempts. The single young female recruited into the study population dispersed more than three times the median dispersal distance of six recruited males, suggesting that other females probably dispersed beyond the boundaries of the study area.     

Reverse sex-biased philopatry in a cooperative bird: genetic consequences and a social cause

The genetic structure of a group or population of organisms can profoundly influence the potential for inbreeding and, through this, can affect both dispersal strategies and mating systems. We used estimates of genetic relatedness as well as likelihood-based methods to reconstruct social group composition and examine sex biases in dispersal in a Costa Rican population of white-throated magpie-jays ( Calocitta formosa , Swainson 1827), one of the few birds suggested to have female-biased natal philopatry. We found that females within groups were more closely related than males, which is consistent with observational data indicating that males disperse upon maturity, whereas females tend to remain in their natal territories and act as helpers. In addition, males were generally unrelated to one another within groups, suggesting that males do not disperse with or towards relatives. Finally, within social groups, female helpers were less related to male than female breeders, suggesting greater male turnover within groups. This last result indicates that within the natal group, female offspring have more opportunities than males to mate with nonrelatives, which might help to explain the unusual pattern of female-biased philopatry and male-biased dispersal in this system. We suggest that the novel approach adopted here is likely to be particularly useful for short-term studies or those conducted on rare or difficult-to-observe species, as it allows one to establish general patterns of philopatry and genetic structure without the need for long-term monitoring of identifiable individuals.     

Does breeding population trajectory and age of nesting females influence disparate nestling sex ratios in two populations of Cooper's hawks?

Offspring sex ratios at the termination of parental care should theoretically be skewed toward the less expensive sex, which in most avian species would be females, the smaller gender. Among birds, however, raptors offer an unusual dynamic because they exhibit reversed size dimorphism with females being larger than males. And thus theory would predict a preponderance of male offspring. Results for raptors and birds in general have been varied although population‐level estimates of sex ratios in avian offspring are generally at unity. Adaptive adjustment of sex ratios in avian offspring is difficult to predict perhaps in part due to a lack of life‐history details and short‐term investigations that cannot account for precision or repeatability of sex ratios across time. We conducted a novel comparative study of sex ratios in nestling Cooper's hawks (Accipiter cooperii) in two study populations across breeding generations during 11 years in Wisconsin, 2001–2011. One breeding population recently colonized metropolitan Milwaukee and exhibited rapidly increasing population growth, while the ex‐Milwaukee breeding population was stable. Following life‐history trade‐off theory and our prediction regarding this socially monogamous species in which reversed sexual size dimorphism is extreme, first‐time breeding one‐year‐old, second‐year females in both study populations produced a preponderance of the smaller and cheaper sex, males, whereas ASY (after‐second‐year), ≥2‐year‐old females in Milwaukee produced a nestling sex ratio near unity and predictably therefore a greater proportion of females compared to ASY females in ex‐Milwaukee who produced a preponderance of males. Adjustment of sex ratios in both study populations occurred at conception. Life histories and selective pressures related to breeding population trajectory in two age cohorts of nesting female Cooper's hawk likely vary, and it is possible that these differences influenced the sex ratios we documented for two age cohorts of female Cooper's hawks in Wisconsin.     

Facultative and non‐facultative sex ratio adjustments in a dimorphic bird species

If parental allocation to each offspring sex has the same cost/benefit ratio, Fisher's hypothesis predicts a sex ratio biased towards the cheaper sex. However, in dimorphic birds there is little evidence for this, especially at hatching. We investigated the pre‐fledgling 1) sex ratio, 2) body condition and 3) sex‐differential mortality in a population of the glossy ibis Plegadis falcinellus, in southern Spain between 2001 and 2011. We defined two age groups for the period between hatching and fledging. We also compared pre‐fledgling with the autumn sex ratio. Metabolic rates were estimated by the doubly labeled water (DLW) technique to establish that sons (the bigger sex) were 18% more energy demanding than daughters, and to compute the predicted Fisher's sex ratio (0.465). As population size increased between years, body condition decreased in both sexes, and mortality increased more for daughters than sons prior to fledging. At the same time, the proportion of males among chicks close to fledging increased (average sex ratio: 0.606) while the proportion close to hatching decreased (average sex ratio: 0.434, in line with Fisher's prediction). Furthermore, the proportions of males at fledging and the following autumn were negatively correlated across years. We suggest that, as population density increased and conditions worsened the larger sex had relatively higher survival. These differences in survival produce a shift from a facultative female‐biased sex ratio at hatching into a non‐facultative male‐biased sex ratio of fledglings. Additionally, the excess of males at fledging was counterbalanced by sex‐related dispersal during the autumn. Overall, glossy ibis sex ratio is a product of a combination of facultative and non‐facultative adjustments triggered by environmental conditions, driven by rapid population growth, and mediated by highly interrelated life‐history traits such as body condition, mortality, and dispersal.     

Modelling the population consequences of age- and sex-related differences in winter mortality in the oystercatcher, Haematopus ostralegus

A modelling approach is used to explore the effect of age and sex differences in oystercatcher ( Haematopus ostralegus ) winter mortality on population size, population structure and the population response to habitat loss or change. Increasing the mortality of first and second year birds reduced population size, but had very little effect on the proportion of the population that were adults. Increasing female mortality reduced population size and resulted in a male-biased population. A sex bias amongst birds of breeding age meant that there were fewer potential breeding pairs for a given population size, reducing the size of the breeding population and the breeding output. Increasing the mortality of one sex relative to the other reduced population size, even when mean adult mortality rates remained unchanged. Increasing the strength of density-dependent mortality in young birds caused a greater reduction in population size as habitat was lost. Increasing the strength of female density-dependent mortality had the same effect, even though male density-dependent mortality had been correspondingly reduced. Increasing density-independent or density-dependent winter mortality in one sex relative to another also exaggerated the disproportional effect of winter habitat loss on separate breeding subpopulations using the same overwintering area. These results suggest that any study of population dynamics should be aware of both age and sex differences in mortality. Conservationists should be particularly aware of any age or sex differences in diet or habitat use that may result in a differential response to environmental change.     

High progesterone during avian meiosis biases sex ratios toward females

Evidence of altered primary sex ratios in birds shows that mothers can manipulate the sex of their offspring before oviposition. In birds, females are the heterogametic sex (ZW) and males are homogametic (ZZ). Sex is determined in the first meiotic division, when one sex chromosome is retained in the oocyte and the other segregates to the polar body. Altered primary sex ratios suggest that birds may be capable of biasing the segregation of sex chromosomes during meiosis I. During the time of meiosis I, follicular steroid production is limited primarily to progesterone (P4). We experimentally manipulated the levels of P4 in female domestic chickens during the approximate time of meiosis I. We advanced the ovulation of the first egg of a sequence (or clutch) with a subcutaneous injection of P4. We found a significant effect of P4 dose on the sex of the resulting egg. The high progesterone group produced 25% males whereas the low progesterone group produced 61% males and the control group produced 63% males in the first ovulation of the sequence. We propose that variation in maternal progesterone during the critical time for genetic sex determination is the mechanism for primary sex ratio manipulation in birds.     

Estimating preharvest density,adult sex ratio,and fecundity of white‐tailed deer using noninvasive sampling techniques

Adult sex ratio and fecundity (juveniles per female) are key population parameters in sustainable wildlife management, but inferring these requires abundance estimates of at least three age/sex classes of the population (male and female adults and juveniles). Prior to harvest, we used an array of 36 wildlife camera traps during 2 and 3 weeks in the early autumn of 2016 and 2017, respectively. We recorded white‐tailed deer adult males, adult females, and fawns from the pictures. Simultaneously, we collected fecal DNA (fDNA) from 92 20 m × 20 m plots placed in 23 clusters of four plots between the camera traps. We identified individuals from fDNA samples with microsatellite markers and estimated the total sex ratio and population density using spatial capture–recapture (SCR). The fDNA‐SCR analysis concluded equal sex ratio in the first year and female bias in the second year, and no difference in space use between sexes (fawns and adults combined). Camera information was analyzed in a spatial capture (SC) framework assuming an informative prior for animals’ space use, either (a) as estimated by fDNA‐SCR (same for all age/sex classes), (b) as assumed from the literature (space use of adult males larger than adult females and fawns), or (c) by inferring adult male space use from individually identified males from the camera pictures. These various SC approaches produced plausible inferences on fecundity, but also inferred total density to be lower than the estimate provided by fDNA‐SCR in one of the study years. SC approaches where adult male and female were allowed to differ in their space use suggested the population had a female‐biased adult sex ratio. In conclusion, SC approaches allowed estimating the preharvest population parameters of interest and provided conservative density estimates.     

Extreme bias in sex allocation in Eclectus parrots

We investigated extraordinary patterns of sex allocation in captive eclectus parrots (Eclectus roratus). These birds are extremely unusual as they show reverse sexual dichromatism, they are the only cooperatively breeding parrot, and they are one of the few birds with nestlings that are easily sexed. They lay two eggs per clutch, but often only fledge one young, and the sex ratio of 209 fledglings did not differ significantly from parity. However, when two young are fledged together they are very likely to be of the same sex, and some females produce long unbroken runs of one sex (the maximum was 20 males) before switching to the other sex. Monte-Carlo simulations show that these runs of same-sex clutches defy expectation if we assume that the sex of chicks within each clutch is independent of the previous clutch. We use further simulations to show that the sex bias must occur at fertilization (i.e. the primary sex ratio), although the female may make further adjustments via infanticide. Control over sex allocation in eclectus parrots is one of the most extreme reported from birds.     

Egg sex ratio and paternal traits: using within-individual comparisons

Empirical studies of sex ratios in birds have been limited dueto difficulties in determining offspring sex. Since molecularsexing techniques removed this constraint, the last 5 yearshas seen a great increase in studies of clutch sex ratio manipulationby female birds. Typically these studies investigate variationin clutch sex ratios across individuals in relation to environmentalcharacteristics or parental traits, and often they find no relationships. In this study we also found that clutch sex ratiosdid not vary in relation to a number of biological and environmentalfactors for 238 great tit Parus major nests. However, interestingsex ratio biases were revealed when variation in clutch sexratios was analyzed within individual females breeding in successiveyears. There was a significant positive relationship betweenthe change in sex ratio of a female's clutch from one yearto the next and the relative body condition of her partner.Females mating with males of higher body condition in yearx + 1 produced relatively male-biased sex ratios, and the oppositewas true for females mated with lower condition males. Within-individualanalysis also allowed investigations of sex ratio in relationto partner change. There was no change in sex ratios of femalespairing with the same male; however, females pairing with anew male produced clutches significantly more female biased. Comparisons of clutch sex ratios within individuals may be apowerful method for detecting sex ratio variation, and perhapsfemale birds may indeed manipulate egg sex but require personalcontextual experience for such decisions.