Movements and foraging effort of Steller's Eiders and Harlequin Ducks wintering near Dutch Harbor, Alaska

ABSTRACT.   We studied the movements and foraging effort of radio-marked Steller's Eiders ( Polysticta stelleri ) and Harlequin Ducks ( Histrionicus histrionicus ) to evaluate habitat quality in an area impacted by industrial activity near Dutch Harbor, Alaska. Foraging effort was relatively low, with Steller's Eiders foraging only 2.7 ± 0.6 (SE) hours per day and Harlequin Ducks 4.1 ± 0.5 hours per day. Low-foraging effort during periods of high-energetic demand generally suggests high food availability, and high food availability frequently corresponds with reductions in home range size. However, the winter ranges of Harlequin Ducks did not appear to be smaller than usual, with the mean range size in our study (5.5 ± 1.1 km²) similar to that reported by previous investigators. The mean size of the winter ranges of Steller's Eiders was similar (5.1 ± 1.3 km²), but no comparable estimates are available. Eutrophication of the waters near Dutch Harbor caused by seafood processing and municipal sewage effluent may have increased populations of the invertebrate prey of these sea ducks and contributed to their low-foraging effort. The threat of predation by Bald Eagles ( Haliaeetus leucocephalus ) that winter near Dutch Harbor may cause Steller's Eiders and Harlequin Ducks to move further offshore when not foraging, contributing to an increase in range sizes. Thus, the movement patterns and foraging behavior of these ducks likely represent a balance between the cost and benefits of wintering in a human-influenced environment.     

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.     

Survival of captive and free-ranging Harlequin Ducks (Histrionicus histrionicus) following surgical liver biopsy

We measured intra- and postoperative mortality rates of captive and free-ranging Harlequin Ducks (Histrionicus histrionicus) undergoing surgical liver biopsy sampling for determination of the induction of cytochrome P4501A, a biomarker of oil exposure. Liver biopsies were taken from and radio transmitters were implanted into 157 free-ranging Harlequin Ducks over three winters (55 in 2000, 55 in 2001, and 47 in 2002). No birds died during surgery, but seven (4.5%) died during recovery from anesthesia (three in 2001 and four in 2002). None of the deaths could be attributed directly to the liver biopsy. Four of the 150 (2.7%) birds that were released died in the 2 wk period after surgery. All post-release deaths occurred in 2001; no birds died after release in 2000 or 2002. No mortalities of 36 captive birds occurred during surgery or recovery or in the 2 wk period following surgery. Hemorrhage was a minor problem with one captive bird. Surgical liver biopsies appear to be a safe procedure, but anesthetic complications may occur with overwintering ducks.     

Regional, seasonal and annual variations in the structure of Purple Sandpiper Calidris maritime populations in Britain

Samples of Purple Sandpipers were captured around the coasts of Britain. Analysis of their bill-length distributions enabled the sex ratios and percentages of 'long-billed' and 'short-billed' birds at each locality to be estimated. The sex ratio for the 'long-billed' population was estimated to be one female to 2–11 males, and one female to 1 -34 males for the 'short-billed' population. During winter, proportionately more 'long-billed' birds occurred in northern and western Scotland, Wales and southern England, whilst 'short-billed' birds predominated from Kincardine to Yorkshire. The total sizes of the 'long-' and 'short-billed7' populations were c. 15 000 and 4000, respectively. 'Short-billed' birds started arriving from Norway in early July. 'Long-billed' birds did not arrive until late October. Their origins are as yet unknown. No annual variations in the population structure were detected.     

Evidence for Environmental Sex Determination in the American eel, Anguilla rostrata

Males have predominated among migrating silver eels in the Annaquatucket River, Rhode Island, for at least two decades, with no significant variation in mean total length in either sex. Because the species is panmictic (random breeding), this consistency suggests environmental sex determination (ESD). Most yellow (feeding phase) eels <300mm total length in the Annaquatucket are sexually undifferentiated, and in contrast to all other published sex ratios, males greatly outnumber females (3:1) among differentiated yellow eels. Estimates of yellow eel population densities are 4–10 times greater than published values for other habitats. We propose that this crowding results in a long period of undifferentiation and the suppression of femaleness. Published field and experimental evidence indicates that high population density results in high proportions of males in Atlantic Anguilla, and that low population density results in the predominance of females. This ESD may be adaptive, resulting in vast numbers of small males in coastal habitats, relatively close to the spawing area, and much larger and more fecund females that occupy most of the available eel habitat.     

Cold tolerance,and not earlier arrival on breeding grounds,explains why males winter further north in an Arctic‐breeding songbird

Sex biases in distributions of migratory birds during the non‐breeding season are widespread; however, the proximate mechanisms contributing to broad‐scale sex‐ratio variation are not well understood. We analyzed a long‐term winter‐banding dataset in combination with spring migration data from individuals tracked by using geolocators to test three hypotheses for observed variation in sex‐ratios in wintering flocks of snow buntings Plectrophenax nivalis. We quantified relevant weather conditions in winter (temperature, snowfall and snow depth) at each banding site each year and measured body size and condition (fat scores) of individual birds (n > 5500). We also directly measured spring migration distance for 17 individuals by using light‐level geolocators. If the distribution pattern of birds in winter is related to interactions between individual body size and thermoregulation, then larger bodied birds (males) should be found in colder sites (body size hypothesis). Males may also winter closer to breeding grounds to reduce migration distance for early arrival at breeding sites (arrival timing hypothesis). Finally, males may be socially dominant over females, and thus exclude females from high‐quality wintering sites (social dominance hypothesis). We found support for the body size hypothesis, in that colder and snowier weather predicted both larger body size and higher proportions of males banded. Direct tracking revealed that males did not winter significantly closer to their breeding site, despite being slightly further north on average than females from the same breeding population. We found some evidence for social dominance, in that females tended to carry more fat than males, potentially indicating lower habitat quality for females. Global climatic warming may reduce temperature constraints on females and smaller‐bodied males, resulting in broad‐scale changes in distributional patterns. Whether this has repercussions for individual fitness, and therefore population demography, is an important area of future research.     

Age- and sex-dependent spring arrival dates of Eastern Kingbirds

ABSTRACT.   Among migratory passerines, the first birds to arrive on the breeding grounds are usually older males. Early arrival by older birds may be driven by experience, age-dependent changes in body condition, age-dependent access to resources during the nonbreeding period, or latitudinal segregation by age. Males may arrive earlier than females (protandry) because males maintain better condition due to greater access to resources during the winter, or because selection favors early arriving males that acquire the best territories or experience enhanced mating opportunities. During a 4-yr study (2004–2007) in Oregon, we found that older Eastern Kingbirds ( Tyrannus tyrannus ), regardless of sex, arrived nearly a week before younger birds and that males arrived about 5 d before females. Age- and sex-dependent arrival dates do not appear to be related to differences in body condition, social dominance in winter, or latitudinal segregation, and protandry is unrelated to the ability of early-arriving males to acquire high-quality territories. Instead, we propose that young birds have less to gain from early arrival because of their probable inability to displace experienced birds from prime territories and that protandry evolved due to enhanced mating opportunities for early arriving males that arise from the high rates of extra-pair matings in our population of Eastern Kingbirds.     

Delayed plumage maturation increases overwinter survival in North Island robins

Many bird species show delayed plumage maturation (DPM), retaining sub-adult plumage until after their first breeding season. Most explanations assume that DPM increases fitness over the breeding season. However, unless birds undergo a full moult before breeding, DPM could also be an adaptation to increase survival over the previous winter. The winter adaptation hypothesis has never been tested owing to the difficulty of measuring overwinter survival. We experimentally tested this hypothesis in North Island robins (Petroica longipes) using a closed island population where we could accurately estimate survival. The experiment involved dyeing 41 juveniles to mimic adult males, and comparing their survival with 41 control juveniles treated with the same peroxide base minus the pigment. The population was monitored with a series of resighting surveys, and mark-recapture analysis used to estimate overwinter survival. Survival probability was estimated to be 10% for dyed birds versus 61% for control birds in 2001, and 29% for dyed birds versus 40% for control birds in the winter of 2002, supporting the winter adaptation hypothesis for DPM. Access to suitable habitat is the key factor limiting juvenile survival in this population, and the locations where dyed juveniles were sighted suggest that they were often excluded from suitable areas.     

First pairing in Snow Geese Anser caerulescens: at what age and at what time of year does it occur?

In migratory birds, the place and time of pair formation are important parameters for population structure and dynamics. Geese are not only migratory but also exhibit long-term monogamy, and therefore the first pairing event in a bird's lifetime is of particular importance. Through behavioural observations of young, known-age, marked birds conducted on the wintering grounds during three winter seasons we investigated two aspects of the timing of first pair formation in the Wrangel Island population of Snow Geese Anser caerulescens : (1) the age at which birds first form pair bonds, and (2) the seasonal pattern of first pair formation. Wrangel Island Snow Geese paired considerably later in life than Snow Geese from a low-Arctic population: almost none of the birds formed pairs in their second winter, and many were still in sibling groups for at least part of that season. The proportion of birds in pairs continued to increase until at least 5 years of age. Most pairing took place during the observation periods, and in general the proportion of birds in pair bonds increased gradually throughout the winter season. The amount of pairing during spring migration or summer varied annually and among cohorts, indicating that even very young birds may be able to form pair bonds quickly if conditions on the breeding grounds are unusually favourable. Pairing later and remaining in family groups longer may be a response to breeding conditions in this high-Arctic colony. Here, productivity is typically low due to harsh weather and predation, whereas Snow Geese breeding in the low Arctic are less restricted and form pairs and start to breed when younger. The fact that most, but not all, pairing takes place on the wintering grounds helps explain why a previous study found a certain amount of gene flow between the two Wrangel Island subpopulations with separate wintering grounds.     

Latitudinal variation in population structure of wintering Pacific Black Brant

ABSTRACT.   Latitudinal variation in population structure during the winter has been reported in many migratory birds, but has been documented in few species of waterfowl. Variation in environmental and social conditions at wintering sites can potentially influence the population dynamics of differential migrants. We examined latitudinal variation in sex and age classes of wintering Pacific Black Brant ( Branta bernicla nigricans ). Brant are distributed along a wide latitudinal gradient from Alaska to Mexico during the winter. Accordingly, migration distances for brant using different wintering locations are highly variable and winter settlement patterns are likely associated with a spatially variable food resource. We used resightings of brant banded in southwestern Alaska to examine sex and age ratios of birds wintering at Boundary Bay in British Columbia, and at San Quintin Bay, Ojo de Liebre Lagoon, and San Ignacio Lagoon in Baja California from 1998 to 2000. Sex ratios were similar among wintering locations for adults and were consistent with the mating strategy of geese. The distribution of juveniles varied among wintering areas, with greater proportions of juveniles observed at northern (San Quintin Bay and Ojo de Liebre Lagoon) than at southern (San Ignacio Lagoon) locations in Baja California. We suggest that age-related variation in the winter distribution of Pacific Black Brant is mediated by variation in productivity among individuals at different wintering locations and by social interactions among wintering family groups.     

Relative vulnerability of female turtles to road mortality

Recent studies suggest that freshwater turtle populations are becoming increasingly male-biased. A hypothesized cause is a greater vulnerability of female turtles to road mortality. We evaluated this hypothesis by comparing sex ratios from published and unpublished population surveys of turtles conducted on- versus off-roads. Among 38 166 turtles from 157 studies reporting sex ratios, we found a consistently larger female fraction in samples from on-roads (61%) than off-roads (41%). We conclude that female turtles are indeed more likely to cross roadways than are males, which may explain recently reported skewed sex ratios near roadways and signify eventual population declines as females are differentially eliminated.     

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.     

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.     

Philopatry and natal dispersal in a sedentary population of western marsh harrier

Dispersal is a key life-history trait because it influences population dynamics and population genetic structure. From a behavioural perspective, the study of natal dispersal requires some understanding of the mechanisms that affect individual movements, because movements of an animal form a path that is continuous throughout its life. Our aim was to investigate juvenile dispersal strategies in the western marsh harrier Circus aeruginosus , between fledging and first breeding attempt (from 1 to 4 years later, depending on the sex and individual). Using radio tracking, we monitored dispersing juvenile harriers and their home-range size variations within a sedentary population in central western France from 2001 to 2007. Juvenile dispersal strategy was mainly characterized by a very high natal philopatry (i.e. birds that remained within the study area) and short-distance dispersal. All but one bird (out of 39) remained within the study area between their first winter and their first spring, and 96.0% during their first spring. The distance moved at 2 years of age was significantly larger for males than for females (3800± sd 3593 m and 935± sd 481 m for seven males and six females, respectively), in contrast to most bird species studied so far. Home-range size was not sex biased and significantly decreased with age. In addition, non-breeding birds had larger home ranges (1603± sd 2128 ha) than breeders (349± sd 185 ha). Using data obtained from other populations, juvenile marsh harrier dispersal strategies appeared to be determined by migratory status (migratory birds dispersing farther) and demographic parameters (juvenile survival or fecundity).     

Estimation of genetic parameters for monthly egg production in laying hens based on random regression models

Heritability and genetic correlations of monthly egg production under random regression models were estimated. Three layer lines (A22, A88, K66) in six consecutive generations were analysed. A22 (13, 770 recorded hens) and A88 (13, 950 recorded hens) are maternal lines of Rhode Island White birds selected on egg production and shell colour; K66 (9, 351 recorded birds) is a paternal line of Rhode Island Red birds selected on egg weight. Eight models with different orders of Legendre polynomials were applied. Adequacy of the models was checked by the Akaike Information Criterion. According to the most adequate model including second order Legendre polynomials for fixed effects and third order for additive genetic and permanent environmental effects, relatively high heritabilities were estimated in the first (h²=0.3) and final (h² above 0.3) periods of production with a substantial decrease in heritability during the egg production peak. Methodology based on random regression animal models can be recommended for genetic evaluation of laying hens.     

Winter survival of North American grassland birds is driven by weather and grassland condition in the Chihuahuan Desert

Populations of grassland birds that overwinter in the Chihuahuan Desert are declining more rapidly than other grassland birds, and survival during the non‐breeding season may have a strong influence on population trends of these species. Habitat loss and deterioration due to desertification may be contributing to these declines, and the winter ecology of grassland birds under these changing environmental conditions remains relatively unexplored. To fill this information gap, we estimated the survival of two grassland‐obligate sparrows, Baird's Sparrows (Ammodramus bairdii) and Grasshopper Sparrows (A. savannarum), on their wintering grounds in the Chihuahuan Desert, and investigated the role of habitat structure and weather on survival rates. We deployed radio‐transmitters on Baird's (N = 49) and Grasshopper (N = 126) sparrows near Janos, Chihuahua, and tracked birds from November to March during the winters of 2012–2013 and 2013–2014. Causes of mortality included avian predators, mammals, and possibly weather. We estimated an overall weekly winter survival probability of ? = 92.73% (95% CI[s] = 88.63–95.44%) for Baird's Sparrows in 2012–2013. We estimated a weekly winter survival probability of ? = 93.48% (95% CI[s] = 90.29–96.67%) and ? = 98.78% (95% CI[s] = 97.88–99.68%) for Grasshopper Sparrow in 2012–2013 and 2013–2014, respectively. Weekly winter survival was lower with colder daily minimum temperatures for both species and in areas with taller shrubs for Grasshopper Sparrows, with the shrubs potentially increasing predation risk by providing perches for Loggerhead Shrikes (Lanius ludovicianus). Our results highlight the need to maintain healthy grass structure in wintering areas to provide birds with food, protection from predators, and adequate cover from inclement weather. Our results also demonstrate that the presence of shrubs can lower winter survival, and suggest that shrub encroachment into the winter habitat of these sparrows may be an important driver of their population declines. Shrub removal could increase survival of wintering sparrows in the Chihuahuan Desert by reducing availability of perches for avian predators, thus reducing predation risk.     

Age‐ratio bias among hunter‐based surveys of Eurasian Wigeon Anas penelope based on wing vs. field samples

We compared age and sex ratios among Eurasian Wigeon Anas penelope derived from Danish field observations and hunter‐based shot samples throughout an entire winter. Sex ratios did not differ significantly between the two samples. Overall, first‐year males were more than three times more likely to be represented than adult males in the hunter sample compared with field samples and were 7–20 times overrepresented in the hunting sample at the beginning of the season. These results confirm the need to account for such bias and its temporal variation when using the results of hunting surveys to model population parameters. Hunter‐shot age ratios may provide a long‐term measure of reproductive success of dabbling duck flyway populations given an understanding of such bias.     

The reproductive ecology of resident manta rays (Manta alfredi) off Maui, Hawaii, with an emphasis on body size

In resident manta rays (Manta alfredi) off Maui, sexual maturity appears delayed until growth exceeds 90% of maximum size, an indicator that large body size provides a reproductive advantage at the expense of a shorter reproductive time period. In this study, 286 surveys were conducted between 2005 and 2010 using photo-identification and photogrammetry to study the reproductive ecology of a resident population of manta rays off Maui, Hawaii, and investigate the reproductive benefits of large body size in each sex. Although reproductive activities occurred year-round, mating trains and late-term pregnant females were significantly more likely to be observed during the winter months. Some females were pursued by males during both winter and summer of the same year, suggesting multiple ovulations may be possible in a single year. Males likely detect a female’s reproductive state by positioning directly behind her, or passing through her bodily excretions. The mean pregnancy rate was estimated at 0.56 pregnancies/adult female/year with larger females pregnant more often, and more likely in consecutive years. The operational sex ratio was heavily skewed with 2.68 adult males per reproductively available female. Although males appear to compete with one another for females within a mating train, no direct physical competition was ever observed between males. Evidence of highly dynamic mating trains lasting more than one day suggests endurance rivalry may be the primary mating strategy among males, during which larger males may benefit from greater energy reserves. The study area appears to be an important staging area for mating individuals in this population.     

Elevated mortality among birds in Chernobyl as judged from skewed age and sex ratios

Background

Radiation has negative effects on survival of animals including humans, although the generality of this claim is poorly documented under low-dose field conditions. Because females may suffer disproportionately from the effects of radiation on survival due to differences in sex roles during reproduction, radiation-induced mortality may result in male-skewed adult sex ratios.

Methodology/Principal Finding

We estimated the effects of low-dose radiation on adult survival rates in birds by determining age ratios of adults captured in mist nets during the breeding season in relation to background radiation levels around Chernobyl and in nearby uncontaminated control areas. Age ratios were skewed towards yearlings, especially in the most contaminated areas, implying that adult survival rates were reduced in contaminated areas, and that populations in such areas could only be maintained through immigration from nearby uncontaminated areas. Differential mortality in females resulted in a strongly male-skewed sex ratio in the most contaminated areas. In addition, males sang disproportionately commonly in the most contaminated areas where the sex ratio was male skewed presumably because males had difficulty finding and acquiring mates when females were rare. The results were not caused by permanent emigration by females from the most contaminated areas because none of the recaptured birds had changed breeding site, and the proportion of individuals with morphological abnormalities did not differ significantly between the sexes for areas with normal and higher levels of contamination.

Conclusions/Significance

These findings are consistent with the hypothesis that the adult survival rate of female birds is particularly susceptible to the effects of low-dose radiation, resulting in male skewed sex ratios at high levels of radiation. Such skewed age ratios towards yearlings in contaminated areas are consistent with the hypothesis that an area exceeding 30,000 km² in Chernobyl’s surroundings constitutes an ecological trap that causes dramatic excess mortality.