Survival and cause-specific mortality of female Rocky Mountain elk exposed to human activity

Animal populations are becoming increasingly exposed to human activity as human populations expand and demand for energy resources (e.g., coal, oil and natural gas) increases. We initiated this study to document survival and cause-specific mortality patterns of female Rocky Mountain elk (Cervus elaphus) exposed to increasing levels of human activity. We fitted 184 females with VHF or GPS collars over 4 years and used the Kaplan–Meier survival estimator to calculate annual survival rates. We used multinomial logistic regression to assess differences in cause-specific mortality and generalized linear mixed models to determine how probability of survival was structured during hunting season; both analyses examined a suite of 5 covariates (i.e., age, year, extent of space use, cover, and human footprint) as potentially influencing cause-specific mortality and survival probability. Annual probability of survival averaged 0.8 (±0.02 SE) over 4 years but averaged 0.91 (±0.03 SE) when harvest mortality was excluded, which was the most significant source of mortality in most years ( [`(x)] = 0.13 ±0.02 \textSE \bar{x} = 0.13 \pm 0.02\,{\text{SE}} ). We found no difference between cause-specific mortality sources relative to elk that survived during the hunting season (χ ₁₀² = 5.79, P = 0.832). The probability of a female surviving during hunting season was negatively influenced by age, year, extent of space use, cover, and human footprint. We found evidence that human activity may have influenced annual rates of natural survival (i.e., exclusive of hunting mortality) and probability of survival during the hunting season. We note that this study occurred largely on privately owned and managed residential and ranch land and focused on female elk; we acknowledge that survival rate and cause-specific patterns of mortality may vary as a function of land ownership (private vs. public), demographic status, and management and harvest practices. While temporal and spatial scales of 1 week may be sufficient to describe patterns of direct mortality during hunting season, broad temporal or spatial scale analyses may be needed to address natural mortality during other seasons.     

Seasonal survival estimation for a long-distance migratory bird and the influence of winter precipitation

Conservation of migratory animals requires information about seasonal survival rates. Identifying factors that limit populations, and the portions of the annual cycle in which they occur, are critical for recognizing and reducing potential threats. However, such data are lacking for virtually all migratory taxa. We investigated patterns and environmental correlates of annual, oversummer, overwinter, and migratory survival for adult male Kirtland’s warblers (Setophaga kirtlandii), an endangered, long-distance migratory songbird. We used Cormack–Jolly–Seber models to analyze two mark–recapture datasets: 2006–2011 on Michigan breeding grounds, and 2003–2010 on Bahamian wintering grounds. The mean annual survival probability was 0.58 ± 0.12 SE. Monthly survival probabilities during the summer and winter stationary periods were relatively high (0.963 ± 0.005 SE and 0.977 ± 0.002 SE, respectively). Monthly survival probability during migratory periods was substantially lower (0.879 ± 0.05 SE), accounting for ~44% of all annual mortality. March rainfall in the Bahamas was the best-supported predictor of annual survival probability and was positively correlated with apparent annual survival in the subsequent year, suggesting that the effects of winter precipitation carried over to influence survival probability of individuals in later seasons. Projection modeling revealed that a decrease in Bahamas March rainfall >12.4% from its current mean could result in negative population growth in this species. Collectively, our results suggest that increased drought during the non-breeding season, which is predicted to occur under multiple climate change scenarios, could have important consequences on the annual survival and population growth rate of Kirtland’s warbler and other Neotropical–Nearctic migratory bird species.     

Seasonal changes in serum dehydroepiandrosterone,androstenedione, and testosterone levels in the squirrel monkey (Saimiri boliviensis boliviensis)

The squirrel monkey (Saimiri boliviensis boliviensis) has a well-defined breeding season during which adult males undergo androgen-dependent morphological changes, with acquisition of active spermatogenesis. To assess the hormonal events of this annual cycle, blood samples were obtained weekly from ten adult males, and serum was assayed for testosterone (T), androstenedione (ΔA), and dehydroepiandrosterone (DHEA). A significant seasonal variation was noted in mean serum T (P < 0.02), ΔA (P < 0.02), and DHEA (P < 0.001) concentrations. Mean ΔA concentrations increased from a nonbreeding season nadir of 91.4 ± 12.9 ng/ml (mean ± standard error) to a prebreeding concentration of 139 ± 10.5 ng/ml and breeding season peak of 167.5 ± 15.4 ng/ml (P < 0.05). Mean DHEA concentrations increased from a nonbreeding season nadir of 8.3 ± 0.8 to a breeding season peak of 14.3 ± 1.2 (P < 0.001). Mean T levels in the nonbreeding (52.2 ± 11.6 ng/ ml) and prebreeding season (48.6 ± 7.4) were similar. However, T significantly increased during the breeding season to 103.5 ± 12.8 ng/ml (P < 0.05). Progressive changes in body weight and morphology paralleled the rise in serum ΔA levels. The pattern of peripheral serum androgen concentrations throughout the year would suggest annual activation of the hypothalamic-pituitary-adrenal and/or hypothalamic-pituitary-gonadal axes.     

Fruit fall in tropical and temperate forests: implications for frugivore diversity

There have been few attempts to compare fruit productivity throughout the world, although this is indispensable for understanding the global variations in frugivore diversity. The purposes of this study are (1) to reveal the patterns in fruit fall in tropical and temperate forests, (2) to examine the environmental factors (location, climate, and total litterfall) affecting these patterns, and (3) to assess the effect of fruit fall on frugivore diversity by using bird and primate data. Fruit fall was compared among 53 forests, from around the equator to the cool-temperate zone at 62°N, in Asia, Africa, North and South America, and Australia. Average ± SD of fruit fall (kg/ha/year) was 454 ± 258 in tropical, and 362 ± 352 in temperate forests. Fruit fall was exceptionally high in Australia (812 ± 461). When Australia was excluded, fruit fall significantly decreased with increasing absolute latitude and altitude, and fruit fall in tropical forest was 1.7 times larger than that in temperate forests (265 ± 227). Total litterfall affected fruit fall significantly, explaining 32, 28, and 64% of the variations of fruit fall in the entire data, tropical data, and temperate data, respectively. The fruit fall/litterfall ratio did not differ between temperate and tropical forests but was significantly higher in Australia than in other regions. Among climatic parameters (annual temperature, precipitation, actual evapotranspiration), a positive relationship was found between temperature and fruit fall in the entire dataset and within temperate forests. Fruit fall seemed to explain the temperate/tropical difference in frugivorous primate diversity to some extent, but not for frugivorous bird diversity. This study shows that the difference in fruit fall in tropical and temperate forests is smaller than that in frugivore diversity, and that it could explain at least part of the frugivore diversity.     

Rainfall-induced changes in food availability modify the spring departure programme of a migratory bird

Climatic warming has intensified selection for earlier reproduction in many organisms, but potential constraints imposed by climate change outside the breeding period have received little attention. Migratory birds provide an ideal model for exploring such constraints because they face warming temperatures on temperate breeding grounds and declining rainfall on many tropical non-breeding areas. Here, we use longitudinal data on spring departure dates of American redstarts (Setophaga ruticilla) to show that annual variation in tropical rainfall and food resources are associated with marked change in the timing of spring departure of the same individuals among years. This finding challenges the idea that photoperiod alone regulates the onset of migration, providing evidence that intensifying drought in the tropical winter could hinder adaptive responses to climatic warming in the temperate zone.     

Sex-bias and timing of natal dispersal in cooperatively breeding Puff-throated Bulbuls Alophoixus pallidus

While natal dispersal can have a significant impact on population dynamics, it is typically difficult to quantify. We investigated timing of natal dispersal of the cooperatively breeding Puff-throated Bulbul Alophoixus pallidus in a tropical evergreen forest by modelling the probability of staying in or dispersing from their natal territory whilst taking into account the effects of sex, group size, and the presence of helper(s). Birds did not disperse until the beginning of and during the breeding season following the hatching year. Dispersal was strongly female-biased both in frequency and distance: most females (95%) dispersed away from their natal territories, and of those relocated, traversed 2–7 territories. In contrast, 50% of males remained in the natal territory as helpers in their second year, while relocated dispersing males crossed 1–2 territories. Natal dispersal was not influenced by either group size or the presence of helpers. Males that fledged earlier in the breeding season exhibited higher rates of philopatry than the males that fledged later, but no correlation between fledging date and philopatry was observed in females. The probability of staying in the natal territory during the second year was 0.58 ± 0.14 SE and 0.05 ± 0.04 for males and females, respectively. These findings may add to our understanding of how natal dispersal can reflect social patterns and kin structure in cooperative breeding species from a little-studied tropical forest region.     

Seasonal reproductive costs contribute to reduced survival of female greater sage‐grouse

Tradeoffs among demographic traits are a central component of life history theory. We investigated tradeoffs between reproductive effort and survival in female greater sage‐grouse breeding in the American Great Basin, while also considering reproductive heterogeneity by examining covariance among current and future reproductive success. We analyzed survival and reproductive histories from 328 individual female greater sage‐grouse captured between 2003 and 2011, and examined the effect of reproductive effort on survival and future reproduction. Monthly survival of females was variable within years, and this within‐year variation was associated with distinct biological seasons. Monthly survival was greatest during the winter (November–March; Φ_W= 0.99 ± 0.001 SE), and summer (June–July; Φ_S= 0.98 ± 0.01 SE), and lower during nesting (April–May; Φ_N= 0.93 ± 0.02 SE) and fall (August–October; Φ_F= 0.92 ± 0.02 SE). Successful reproduction was associated with reduced monthly survival during summer and fall, and this effect was greatest during fall. Females that successfully fledged chicks had lower annual survival (0.47 ± 0.05 SE) than females who were not successful (0.64 ± 0.04 SE). Annual survival did not vary across years, consistent with a slow‐paced life history strategy in greater sage‐grouse. In contrast, reproductive success varied widely, and was positively correlated with annual rainfall. We found evidence for heterogeneity among females with respect to reproductive success; compared with unsuccessful females, females that raised a brood successfully in year t were more than twice as likely to be successful in year t+ 1. Female greater sage‐grouse incur costs to survival associated with reproduction, however, variation in quality among females may override costs to subsequent reproductive output.     

Haldane's rule revisited: do hybrid females have a shorter lifespan? Survival of hybrids in a recent contact zone between two large gull species

Haldane's rule predicts that particularly high fitness reduction should affect the heterogametic sex of interspecific hybrids. Despite the fact that hybridization is widespread in birds, survival of hybrid individuals is rarely addressed in studies of avian hybrid zones, possibly because of methodological constraints. Here, having applied capture–mark–recapture models to an extensive, 19‐year‐long data set on individually marked birds, we estimate annual survival rates of hybrid individuals in the hybrid zone between herring (Larus argentatus) and Caspian (Larus cachinnans) gulls. In both parental species, males have a slightly higher survival rate than females (model‐weighted mean ± SE: herring gull males 0.88 ± 0.01, females 0.87 ± 0.01, Caspian gull males 0.88 ± 0.01, females 0.87 ± 0.01). Hybrid males do not survive for a shorter time than nonhybrid ones (0.88 ± 0.01), whereas hybrid females have the lowest survival rate among all groups of individuals (0.83 ± 0.03). This translates to a shorter adult (reproductive) lifespan (on average by 1.7–1.8 years, i.e. ca 25%) compared with nonhybrid females. We conclude that, in line with Haldane's rule, the lower survival rate of female hybrids may contribute to selection against hybrids in this hybrid zone.     

Home-ranges of tropical Red-capped Larks are influenced by breeding rather than vegetation,rainfall or invertebrate availability

Home-range studies have received considerable attention from ecologists but are greatly skewed towards the north temperate areas. Tropical areas offer an ideal setting to tease apart hypotheses about weather, food availability and social interactions as important factors influencing home-range. In this study, we investigated home-range and movement patterns of the tropical Red-capped Lark Callandrella cineria, a year-round breeding bird with a dynamic social structure. We tracked 56 individuals using radiotransmitters and colour-ring readings over a 23-month period. Our objective was to understand year-round variation in home-range size in the context of the highly aseasonal and unpredictable variation in weather and resources typical of many equatorial habitats, in addition to the birds’ changing social structure and year-round breeding. The mean composite monthly home-range of Red-capped Larks was 58.0 ha, and the mean individual home-range size was 19.9 ha, but this varied considerably between individuals. The total number of nests found per month (breeding intensity) best predicted home-range size of non-breeding birds, and of breeding and non-breeding birds combined. We show for the first time that breeding intensity decreases the home-range size of non-breeding individuals. Our study also underlines the relevance of conducting more studies in aseasonal tropical areas in order to disentangle effects of weather, food availability and breeding that vary in parallel, peaking simultaneously in most seasonal areas.     

Climate anomalies affect annual survival rates of swifts wintering in sub‐Saharan Africa

Several species of migratory swifts breed in the Western Palearctic, but they differ in reproductive traits and nonbreeding areas explored in Africa. We examined survival and recapture probabilities of two species of swifts by capture–mark–recapture data collected in northern Italy (Pallid Swift Apus pallidus in Carmagnola, Turin, and Common Swift Apus apus in Guiglia, Modena) in the breeding season (May–July). Apparent survival rates were relatively high (>71%), comparable to other studies of European swifts, but showed marked annual variations. We used geolocators to establish the exact wintering areas of birds breeding in our study colonies. Common Swifts explored the Sahel zone during migration and spent the winter in SE Africa, while the Pallid Swifts remained in the Sahel zone for a longer time, shifting locations southeast down to Cameroun and Nigeria later in winter. These movements followed the seasonal rains from north to south (October to December). In both species, we found large yearly differences in survival probabilities related to different climatic indices. In the Pallid Swift, wintering in Western Africa, the Sahel rainfall index best explained survival, with driest seasons associated with reduced survival. In the Common Swift, wintering in SE Africa, the El Niño–Southern Oscillation (ENSO) cycle performed significantly better than Sahel rainfall or North Atlantic Oscillation (NAO). Extreme events and precipitation anomalies in Eastern Africa during La Niña events resulted in reduced survival probabilities in Common Swifts. Our study shows that the two species of swifts have similar average annual survival, but their survival varies between years and is strongly affected by different climatic drivers associated with their respective wintering areas. This finding could suggest important ecological diversification that should be taken into account when comparing survival and area use of similar species that migrate between temperate breeding areas and tropical wintering areas.     

Divergent patterns of telomere shortening in tropical compared to temperate stonechats

Telomeres have emerged as important biomarkers of health and senescence as they predict chances of survival in various species. Tropical birds live in more benign environments with lower extrinsic mortality and higher juvenile and adult survival than temperate birds. Therefore, telomere biology may play a more important role in tropical compared to temperate birds. We measured mean telomere length of male stonechats (Saxicola spp.) at four age classes from tropical African and temperate European breeding regions. Tropical and temperate stonechats had similarly long telomeres as nestlings. However, while in tropical stonechats pre‐breeding first‐years had longer telomeres than nestlings, in temperate stonechats pre‐breeding first‐years had shorter telomeres than nestlings. During their first breeding season, telomere length was again similar between tropical and temperate stonechats. These patterns may indicate differential survival of high‐quality juveniles in tropical environments. Alternatively, more favorable environmental conditions, that is, extended parental care, may enable tropical juveniles to minimize telomere shortening. As suggested by previous studies, our results imply that variation in life history and life span may be reflected in different patterns of telomere shortening rather than telomere length. Our data provide first evidence that distinct selective pressures in tropical and temperate environments may be reflected in diverging patterns of telomere loss in birds.     

Survival and population growth of a long-lived threatened snake species, Drymarchon couperi (Eastern Indigo Snake)

Demographic data provide a basis for understanding the life history and ecology of species, factors which are vital for informing conservation efforts; however, little is known regarding the population ecology of most snake species, including the threatened Eastern Indigo Snake (Drymarchon couperi). We used 11 years (1999–2009) of capture-mark-recapture (CMR) and 2.5 years (2003–2005) of radiotelemetry data from southeastern Georgia, USA, in a CMR modeling framework to estimate apparent survival, capture and transition probabilities, and evaluate factors influencing these parameters. The model-averaged estimate of overall apparent annual survival probability was 0.700 (±0.030 SE) and is comparable to that obtained from known fate analysis (radiotelemetry) at the same site. Body size positively influenced survival, regardless of sex. Capture probability differed seasonally by sex, suggesting lower capture probability for females in fall and males in winter. There was no evidence for effect of precipitation or site-specific differences in survival. Model averaged estimate of annual adult survival estimated using multistate CMR models was 0.738 ± 0.030 and 0.515 ± 0.189 for subadults. We estimated population growth rate (λ) and elasticity (proportional sensitivity) of λ to vital rates using a stage-structured matrix population model. Population growth rate ranged from 0.96 to 1.03 depending on the value of the probability of transitioning from subadult to adult stage. The λ was proportionally most sensitive to changes in adult survival rate, followed by subadult survival. Our results suggest that protecting adult snakes and their habitats would result in the highest likelihood of long-term population stability and growth.     

High survival rate of a critically endangered species, the Azores Bullfinch Pyrrhula murina, as a contribution to population recovery

This paper reports analyses of a capture–mark–recapture (CMR) dataset of 149 Azores Bullfinches ringed on S?o Miguel island (Azores) between 2005 and 2007, and recaptured–resighted on a monthly basis over a 4-year period (2005–2008) throughout their breeding range. We examined the effect of time, age (adults vs. juveniles), gender (adult males and females), and environmental covariates (temperature, rainfall, NAO index) on survival probabilities. The modelling found a high and constant monthly survival probability (mean ± SE) estimated at 0.96 ± 0.01, similar between both adults and juveniles and independent of environmental conditions and gender. These findings agree with expectations from island-based life-history theory where relatively mild conditions and lack of predators should favour high survival rates to compensate for the low reproductive output. The annual survival rate was estimated at 0.62, which was also consistent with this pattern when compared with survival estimates of mainland bullfinch and passerine species on other subtropical islands obtained in similar CMR studies. Based on a canonical estimator, the size of the studied population (mean ± SE) was estimated at 1608 ± 326 individuals. Given that the population size was only around 120–400 individuals in the early 1990s, we suggest that the high survival probabilities currently applying to this critically endangered species may have substantially contributed to the recent recovery of this population. Future research studies on the species’ demography should continue to monitor survival in order to measure the effect of management interventions currently taking place within the range of the Azores Bullfinch, including the restoration of the biodiversity rich laurel forest, but also focusing on nest success, which is important for understanding population dynamics.     

Corticosterone,testosterone and life-history strategies of birds

Steroid hormones have similar functions across vertebrates, but circulating concentrations can vary dramatically among species. We examined the hypothesis that variation in titres of corticosterone (Cort) and testosterone (T) is related to life-history traits of avian species. We predicted that Cort would reach higher levels under stress in species with higher annual adult survival rates since Cort is thought to promote physiological and behavioural responses that reduce risk to the individual. Conversely, we predicted that peak T during the breeding season would be higher in short-lived species with high mating effort as this hormone is known to promote male fecundity traits. We quantified circulating hormone concentrations and key life-history traits (annual adult survival rate, breeding season length, body mass) in males of free-living bird species during the breeding season at a temperate site (northern USA) and a tropical site (central Panama). We analysed our original data by themselves, and also combined with published data on passerine birds to enhance sample size. In both approaches, variation in baseline Cort (Cort0) among species was inversely related to breeding season length and body mass. Stress-induced corticosterone (MaxCort) also varied inversely with body mass and, as predicted, also varied positively with annual adult survival rates. Furthermore, species from drier and colder environments exhibited lower MaxCort than mesic and tropical species; T was lowest in species from tropical environments. These findings suggest that Cort0, MaxCort and T modulate key vertebrate life-history responses to the environment, with Cort0 supporting energetically demanding processes, MaxCort promoting survival and T being related to mating success.     

Latitudinal gradients in some,but not all,avian life history traits extend into the Arctic

Latitudinal variation in avian life history strategies is well documented. Clutch size and nest success tend to increase with latitude, whereas longevity and developmental periods have been argued to decrease with latitude. However, these patterns are largely based on interspecific comparisons of species breeding at tropical and temperate latitudes. We compared the life history of Yellow Warblers Setophaga petechia breeding in arctic habitat at the northern extent of their range, in Inuvik, NWT (68°N), Canada, with those breeding in temperate habitat in Revelstoke, BC (50°N), and use data from 21 populations spanning 0–68°N to evaluate latitudinal trends in life history traits from tropical to arctic habitats. Females breeding in Inuvik laid first clutches that were slightly (although not significantly) larger and had higher nest success, which resulted in higher annual productivity compared with their low- latitude counterparts. Apparent adult survival rates were only marginally lower in Inuvik than in Revelstoke, whereas incubation and nestling periods in the arctic were similar to our temperate site. When comparing life history traits across the Yellow Warbler breeding range, we observed increases in clutch sizes and nest success with increasing latitude that appeared to be associated with declines in adult survival, though this relationship was weakened by the addition of our arctic site. We detected more moderate declines in incubation and nestling periods with increasing latitude. As we observed latitudinal variation in some life history traits, but not a consistent transition of traits associated with a shift from a slow to fast life history from tropical to arctic latitudes, our study suggests that the expectation for a general shift in life history traits may be over-simplified.     

Carryover effects and climatic conditions influence the postfledging survival of greater sage‐grouse

Prebreeding survival is an important life history component that affects both parental fitness and population persistence. In birds, prebreeding can be separated into pre‐ and postfledging periods; carryover effects from the prefledging period may influence postfledging survival. We investigated effects of body condition at fledging, and climatic variation, on postfledging survival of radio‐marked greater sage‐grouse (Centrocercus urophasianus) in the Great Basin Desert of the western United States. We hypothesized that body condition would influence postfledging survival as a carryover effect from the prefledging period, and we predicted that climatic variation may mediate this carryover effect or, alternatively, would act directly on survival during the postfledging period. Individual body condition had a strong positive effect on postfledging survival of juvenile females, suggesting carryover effects from the prefledging period. Females in the upper 25th percentile of body condition scores had a postfledging survival probability more than twice that (Φ = 0.51 ± 0.06 SE) of females in the bottom 25th percentile (Φ = 0.21 ± 0.05 SE). A similar effect could not be detected for males. We also found evidence for temperature and precipitation effects on monthly survival rates of both sexes. After controlling for site‐level variation, postfledging survival was nearly twice as great following the coolest and wettest growing season (Φ = 0.77 ± 0.05 SE) compared with the hottest and driest growing season (Φ = 0.39 ± 0.05 SE). We found no relationships between individual body condition and temperature or precipitation, suggesting that carryover effects operated independently of background climatic variation. The temperature and precipitation effects we observed likely produced a direct effect on mortality risk during the postfledging period. Conservation actions that focus on improving prefledging habitat for sage‐grouse may have indirect benefits to survival during postfledging, due to carryover effects between the two life phases.     

Seasonality in basal metabolic rate and thermal conductance in a long-distance migrant shorebird,the knot (Calidris canutus)

Knots Calidris canutus live highly seasonal lives, breeding solitarily on high arctic tundra and spending the non-breeding season in large social flocks in temperate to tropical estuaries. Their reproductive activities and physiological preparations for long flights are reflected in pronounced plumage and body mass changes, even in long-term captives of the islandica subspecies (breeding in north Greenland and northeast Canada and wintering in western Europe) studied in outdoor aviaries. The three to four fattening episodes in April-July in connection with the flights to and from the high arctic breeding grounds by free-living birds, are represented by a single period of high body mass, peaking between late May and early July in a sample of ten captive islandica knots studied over four years. There are consistent and synchronized annual variations in basal metabolic rate and thermal conductance in three islandica knots. Basal metabolic rate was highest during the summer body mass peak. Within the examined individuals, basal metabolic rate scales on body mass with an exponent of about 1.4, probably reflecting a general hypertrophy of metabolically expensive muscles and organs. Any potential effect of moult on basal metabolic rate was obscured by the large seasonal mass-associated variations. In breeding plumage, insulation (the inverse of thermal conductance) was a factor of 1.35 lower than in winter plumage. This was paralleled by the dry mass of contour feathers being a factor of 1.17 lower. In this subspecies the breeding season is indeed the period during which the costs of thermoregulation are lowest. In captive knots seasonal changes in basal metabolic rate and thermal conductance likely reflect an anticipatory programme adaptive to the variable demands made by the environment at different times of the year.     

Seasonal associations with novel climates for North American migratory bird populations

Determining the implications of global climate change for highly mobile taxa such as migratory birds requires a perspective that is spatiotemporally comprehensive and ecologically relevant. Here, we document how passerine bird species that migrate within the Western Hemisphere (n = 77) are associated with projected novel climates across the full annual cycle. Following expectations, highly novel climates occurred on tropical non‐breeding grounds and the least novel climates occurred on temperate breeding grounds. Contrary to expectations, highly novel climates also occurred within temperate regions during the transition from breeding to autumn migration. This outcome was caused by lower inter‐annual climatic variability occurring in combination with stronger warming projections. Thus, migrants are projected to encounter novel climates across the majority of their annual cycle, with a pronounced peak occurring when juveniles are leaving the nest and preparing to embark on their first migratory journey, which may adversely affect their chances of survival.     

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.