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.     

Recent population declines in Afro‐Palaearctic migratory birds: the influence of breeding and non‐breeding seasons

Aim

Recent, rapid population declines in many Afro‐Palaearctic migratory bird species have focussed attention on changing conditions within Africa. However, processes influencing population change can operate throughout the annual cycle and throughout migratory ranges. Here, we explore the evidence for impacts of breeding and non‐breeding conditions on population trends of British breeding birds of varying migratory status and wintering ecology.

Location

Great Britain (England & Scotland).

Methods

Within‐ and between‐species variation in population trends is quantified for 46 bird species with differing migration strategies.

Results

Between 1994 and 2007, rates of population change in Scotland and England differed significantly for 19 resident and 15 long‐distance migrant species, but were similar for 12 short‐distance migrant species. Of the six long‐distance migrant species that winter in the arid zone of Africa, five are increasing in abundance throughout Britain. In contrast, the seven species wintering in the humid zone of Africa are all declining in England, but five of these are increasing in Scotland. Consequently, populations of both arid and humid zone species are increasing significantly faster in Scotland than England, and only the English breeding populations of species wintering in the humid zone are declining.

Main conclusions

Population declines in long‐distance migrants, especially those wintering in the humid zone, but not residents or short‐distance migrants suggest an influence of non‐breeding season conditions on population trends. However, the consistently less favourable population trends in England than Scotland of long‐distance migrant and resident species strongly suggest that variation in the quality of breeding grounds is influencing recent population changes. The declines in humid zone species in England, but not Scotland, may result from poorer breeding conditions in England exacerbating the impacts of non‐breeding conditions or the costs associated with a longer migration, while better conditions in Scotland may be buffering these impacts.

     

The rise and fall of local populations of ortolan buntings Emberiza hortulana: importance of movements of adult males

Changes in population size of local populations of birds have usually been interpreted in relation to adult return rate and recruitment of young individuals after natal dispersal. Little is known about the importance of redistribution of adult individuals through breeding dispersal. The small Norwegian population of ortolan buntings Emberiza hortulana has a patchy distribution with about 30 long‐term local populations. During a period of general population decline (29% decrease over 7 years), the population trends of local populations (measured as number of males recorded) were highly variable, with some even increasing four‐fold. Comparisons of demographic parameters showed that adult immigration rate (i.e. dispersal of adult males) explained both yearly changes in male population size and population trends over the whole study period better than adult return rate or adult emigration rate, or a measure of recruitment of young males. Adult immigration rates and recruitment rates were correlated, suggesting that both young and adult males find the same places attractive. In the study area, adult sex ratio was strongly male‐biased, and immigration rate was higher when local sex ratio was less skewed. In addition, less skewed sex ratio was related to higher adult return rate and lower emigration rate. We found no relationships between measures of breeding success and population change. We suggest that conspecific attraction may explain the observed patterns. Some local populations may act as hot‐spots attracting adult males from other populations. Thus, local population changes need not reflect overall population growth rate, but may be a consequence of redistribution of adult birds.     

GPS‐tracking reveals non‐breeding locations and apparent molt migration of a Black‐headed Grosbeak

Black‐headed Grosbeaks (Pheucticus melanocephalus) have been observed to undergo prebasic molt during fall in the North American Monsoon region of the southwestern United States and northwestern Mexico, but it is unknown whether molt migration is pervasive across populations of the species. During the 2014 breeding season, we GPS‐tagged (where GPS is global positioning system) nine adult Black‐headed Grosbeaks in Yosemite National Park with archival GPS tags to determine specific locations where grosbeaks breeding in Yosemite spent portions of the non‐breeding season, and to assess whether those locations were consistent with molt migration. On 2 June 2015, one of these birds, a male GPS‐tagged on 19 June 2014, was recaptured with its GPS unit still attached. Data downloaded from the unit revealed that, by 20 August 2014, the bird had moved 1300 km from Yosemite National Park to Sonora, Mexico, where it remained until at least 15 October 2014. By 24 November 2014, the grosbeak had moved >1300 m from Sonora to the Michoacán‐Jalisco border region, where it remained until the last GPS‐determined location was obtained on 24 March 2015. The seasonal timing of these movements and the length of stay in Sonora are consistent with the expected behavior of a molt‐migrating bird. Remote‐sensed enhanced vegetation index (EVI) data indicated that the grosbeak arrived in the monsoon region near the area's annual peak in EVI, and then, as the index was declining sharply, departed for the Michoacán‐Jalisco region, where the index also declined during the same period, but substantially less so than in Sonora. Climate change in the coming decades is expected to delay the annual onset of the monsoon while also accelerating the initiation of arid, summer‐like conditions throughout much of western North America, possibly yielding a temporal mismatch between fall migration and the monsoon‐driven conditions that may be critical for molt‐migrating birds.     

Foraging behavior of Cerulean Warblers during the breeding and non‐breeding seasons: evidence for the breeding currency hypothesis

Birds require additional resources for raising young, and the breeding currency hypothesis predicts that insectivorous species exploit large soft‐bodied prey during the breeding season, but shift to small, likely hard‐bodied, prey during the non‐breeding season. To test this hypothesis, we examined prey use by Cerulean Warblers (Setophaga cerulea), foliage‐gleaning Nearctic‐Neotropical migrants, during the breeding and non‐breeding seasons. We collected data on foraging behavior during the breeding season (including observations of prey items fed to young) in upland mixed‐oak forest in southeastern Ohio in 2009 and 2010 and, during the non‐breeding season, in shade coffee in the Cordillera de Merida, Venezuela, in 2008–2009. Cerulean Warblers captured 7% more large prey (visible prey extending beyond the bill) during the breeding than the non‐breeding season, but foraged at similar rates during both seasons. Large, soft‐bodied prey appeared to be especially important for feeding young. We found that adults delivered large prey on >50% of provisioning visits to nests and 69% of identifiable large prey fed to nestlings were greenish larvae (likely Lepidoptera or caterpillars) that camouflage against leaves where they would tend to be captured by foliage‐gleaning birds. Availability of specific taxa appeared to influence tree species foraging preferences. As reported by other researchers, we found that Cerulean Warblers selected trees in the genus Carya for foraging and our examination of caterpillar counts from the central Appalachian Mountains (Butler and Strazanac 2000 ) showed that caterpillars with greenish coloration, especially Baileya larvae, may be almost twice as abundant on Carya than Quercus. Our results provide evidence for the breeding currency hypothesis, and highlight the importance of caterpillars to a foliage‐gleaning migrant warbler of conservation concern.     

Annual variation in long‐distance dispersal driven by breeding and non‐breeding season climatic conditions in a migratory bird

Long‐distance dispersal is a fundamental process in ecology and evolution but the factors that influence these movements remain poorly understood in most species. We used stable hydrogen isotopes to quantify the rate and direction of long‐distance immigration in a breeding population of American redstarts and to test whether the settlement decisions that result in long‐distance dispersal are driven by habitat saturation or by the phenology of breeding‐season resources. Our results provide evidence that both natal dispersal and breeding dispersal were influenced by the timing of breeding‐season phenology, with both age classes more likely to disperse north in years when the onset of breeding‐season phenology occurs earlier than normal. Yearlings were also more likely to disperse north following winters with poor habitat quality on their non‐breeding grounds, demonstrating that carry‐over effects from the non‐breeding season influence natal dispersal in this species. Collectively, these results are consistent with the hypothesis that American redstarts use the phenology of breeding season resources as a cue to select breeding sites. Our results suggest that long‐distance dispersal may allow individuals to rapidly respond to advancing phenology caused by global climate change, though their ability to do so may be constrained by long‐term decline in habitat quality predicted for their tropical non‐breeding grounds.     

Disentangling the effects of environmental conditions on wintering and breeding grounds on age‐specific survival rates in a trans‐Saharan migratory raptor

Migratory species are subject to environmental variability occurring on breeding and wintering grounds. Estimating the relative contribution of environmental factors experienced sequentially during breeding and wintering, and their potential interaction, to the variation of survival is crucial to predict population viability of migratory species. Here we investigated this issue for the Montagu's harrier Circus pygargus, a trans‐Saharan migrant. We analysed capture–recapture data from a 29‐year long monitoring of wing‐tagged offspring and adults at two study sites in France (Rochefort‐RO and Maine‐et‐Loire‐ML). The study period covers a climatic shift occurring in the Sahel with increasing rainfall following a period of droughts (Sahel greening). We found that harriers’ adult survival in RO (between 1988 and 2005) varied over time and was sensitive to the interaction between the amount of rainfall in the Sahel and the annual mean breeding success, two proxies of prey availability. The occurrence of adverse conditions on breeding and wintering grounds in the same year decreased survival from 0.70–0.77 to 0.48 ± 0.05. Juvenile survival in RO was slightly more sensitive to conditions in Europe than in the Sahel. Unexpectedly, lower survival rates were found in years with higher mean breeding success, suggesting compensatory density feedbacks may operate. By contrast, adult survival in ML, monitored between 1999 and 2017, was higher compared to RO (0.76 ± 0.03 versus 0.66 ± 0.02), remained constant and unaffected by any proxy of prey availability. This difference seems consistent with the fact that harriers in ML experienced better and especially less variable environmental conditions during breeding and wintering seasons compared to RO. Overall, we showed that survival of a migratory bird is sensitive to the level of variability in environmental conditions and that adverse conditions on wintering grounds can amplify the negative effects of conditions during the previous breeding season on birds’ survival.     

Potential interactions between humans and non‐breeding shorebirds on a subtropical intertidal flat

Abstract The primary aim of this study was to investigate whether bait harvesting, with all its inherent effects, occurring in the intertidal zone of a subtropical estuary, had an impact on a migratory shorebird, the eastern curlew Numenius madagascariensis. In a large‐scale manipulative study (units of experiment were 1 ha plots), callianassid shrimp Trypaea australiensis populations were harvested simulating the technique (manual pumping) and the levels of harvesting intensity per unit area (347 shrimp per hectare per harvesting event) exhibited by bait‐collectors in SE Australia and South Africa. It was found that at present levels of harvesting intensity per unit area (approximately 1% of standing stock removed per harvesting event) there is no threat to the stocks of Trypaea exploited by the curlews in Moreton Bay, Australia. However, the results show that the curlews themselves apply a considerable predation pressure on Trypaea. Based on the birds' foraging rates and densities, it was estimated that they would consume up to 100% of the initial Trypaea stock over the course of a non‐breeding season (October to March). However, the stable seasonal trend in the density of the size‐cohort of Trypaea preyed upon by the curlews indicates that the existing rates of predation are easily counterbalanced, e.g. through continuous density‐dependent recruitment of these crustaceans. We suggest that this mechanism will provide for a stable foraging environment for both the shorebirds and bait collectors.     

Body condition and feather molt of a migratory shorebird during the non‐breeding season

Migratory shorebirds have some of the highest fat loads among birds, especially species which migrate long distances. The upland sandpiper Bartramia longicauda makes long‐distance migrations twice a year, but variation in body condition or timing of feather molt during the non‐breeding season has not been studied. Molt is an important part of the annual cycle of migratory birds because feather condition determines flight performance during migration, and long‐distance movements are energetically costly. However, variation in body condition during molt has been poorly studied. The objective of our field study was to examine the timing and patterns of feather molt of a long distance migratory shorebird during the non‐breeding season and test for relationships with body size, fat depots, mass, and sex. Field work was conducted at four ranches in the Northern Campos of Uruguay (Paysandú and Salto Departments). We captured and marked 62 sandpipers in a 2‐month period (Nov–Jan) during four non‐breeding seasons (2008–2012). Sex was determined by genetic analyses of blood samples taken at capture. Molt was measured in captured birds using rank scores based on published standards. Body mass and tarsus length measurements showed female‐biased sexual size dimorphism with males smaller than females. Size‐corrected body mass (body condition) showed a U‐shaped relationship with the day of the season, indicating that birds arrived at non‐breeding grounds in relatively good condition. Arriving in good body condition at non‐breeding grounds is probably important because of the energetic demands due to physiological adjustments after migration and the costs of feather molt.     

Migration and diving activity in three non‐breeding flesh‐footed shearwaters Puffinus carneipes

The flesh‐footed shearwater Puffinus carneipes is a medium‐sized shearwater and transequatorial migrant within the Pacific Ocean. We used archival data loggers to study the non‐breeding migration and diving behaviour of three flesh‐footed shearwaters following breeding in New Zealand. In early April, the birds migrated to the western North Pacific Ocean in 23±2 days, occupying core distributions within the Kuroshio/Oyashio transition system for 91±17 days. Subsequent movements were made into the Sea of Okhotsk prior to return migrations to New Zealand in mid September (19±1 days). Diving depths during migration (2.5±2.4 m), and in the western North Pacific (2.4±2.6 m) were shallower than during the onset of breeding (4.8±8.7 m). Non‐breeding flesh‐footed shearwaters occupy a region of high fisheries activity and the impact of these fisheries on adult survival in this declining species warrant further study.     

Male‐mimicking females increase male‐male interactions,and decrease male survival and condition in a female‐polymorphic damselfly

Biologists are still discovering diverse and powerful ways sexual conflicts shape biodiversity. The present study examines how the proportion of females in a population that exhibit male mimicry, a mating resistance trait, influences conspecific males’ behavior, condition, and survival. Like most female‐polymorphic damselflies, Ischnura ramburii harbors both “andromorph” females, which closely resemble males, and sexually dimorphic “gynomorph” counterparts. There is evidence that male mimicry helps andromorphs evade detection and harassment, but males can also learn to target locally prevalent morph(s) via prior mate encounters. I hypothesized that the presence of male mimics could therefore predispose males to mate recognition errors, and thereby increase rates of costly male‐male interactions. Consistent with this hypothesis, male‐male interaction rates were highest in mesocosms containing more andromorph (vs. gynomorph) females. Males in andromorph‐biased mesocosms also had lower final body mass and higher mortality than males assigned to gynomorph‐majority treatments. Male survival and body mass were each negatively affected by mesocosm density, and mortality data revealed a marginally significant interaction between andromorph frequency and population density. These findings suggest that, under sufficiently crowded conditions, female mating resistance traits such as male mimicry could have pronounced indirect effects on male behavior, condition, and survival.     

Cost of reproduction and the evolution of deferred breeding in the western gull

Survival patterns in a population of western gulls (Larus occidentalis)of known age of first breeding, a, indicate a cost of reproductionrelated to the age of initial breeding. Among both sexes, birdsthat commenced breeding at the earliest ages (3 years in malesand 4 years in females) had higher annual mortality than thosethat deferred breeding one or more years. In addition, females(but not males) evidenced a cumulative cost of reproduction:holding age constant, females with more annual breeding attemptsdemonstrated poorer survival. These patterns of -specific survivalwere statistically significant after controlling for interannualvariation in food availability and are not explained simplyby variation in the intrinsic quality of individuals. To assessthe effects of these sex-specific costs on fitness, we combinedthe observed survival patterns with data on prebreeding survivorshipand -specific reproductive success to estimate rates of populationgrowth and lifetime reproductive success for different agesat first reproduction. Males showed a clearly defined fitnessoptimum at = 4 years, which coincided with the modal for malesin the population. Females showed no clear optimum, except thatbreeding at age 4 was suboptimal, hence females benefited fromdeferring breeding to ages 5-7 years. Observed age of firstbreeding also showed no clear mode for females, with slightpeaks at ages 5 and 7. As a result, in both sexes, the fitnesssurface for corresponded well with observed frequencies of. We suggest that stabilizing selection has acted to shape thephenotypic distribution of in males but, due to trade-offsbetween survival and early reproduction, stabilizing selectionis weak or absent in females     

Range sizes and habitat use of non‐breeding Crested Caracaras in Florida

In Florida, habitats that include the breeding territories of Crested Caracaras (Caracara cheriway) are protected, but non‐breeding individuals may be vulnerable because they may occupy different areas and habitats. We captured and radio‐tagged 58 non‐breeding caracaras in Florida from July 2006–March 2009, determined their locations during weekly flights, and used GIS and compositional analysis to evaluate range sizes and habitat use. Non‐breeding caracaras (N = 58) ranged five times more widely during breeding seasons (N = 573 locations) than during non‐breeding seasons (N = 592 locations), and ranged >250 times more widely than breeding caracaras that defend territories year‐round. The large ranges of non‐breeders suggest they may be searching for and evaluating prospective territories or breeding opportunities (territory prospecting). Pasture occupied by cattle was the most used habitat relative to availability and was used more than pasture without cattle, likely because insects associated with cattle are an important food source for caracaras. Cattle numbers in Florida are declining and, because both breeding and non‐breeding caracaras primarily occupy pasture, this may present difficulties for long‐term management. Citrus groves were also used more than expected given availability by non‐breeding caracaras, but are rarely included in nesting territories. Because pasture and citrus were often adjacent, we suggest that citrus groves may function as refugia from socially dominant breeding caracaras. Conservation and recovery efforts for Florida's caracara population are needed throughout the range of non‐breeders, and should include management that ensures availability of habitat matrices of cattle pasture and citrus groves.     

Estimability of migration survival rates from integrated breeding and winter capture–recapture data

Long‐distance migration is a common phenomenon across the animal kingdom but the scale of annual migratory movements has made it difficult for researchers to estimate survival rates during these periods of the annual cycle. Estimating migration survival is particularly challenging for small‐bodied species that cannot carry satellite tags, a group that includes the vast majority of migratory species. When capture–recapture data are available for linked breeding and non‐breeding populations, estimation of overall migration survival is possible but current methods do not allow separate estimation of spring and autumn survival rates. Recent development of a Bayesian integrated survival model has provided a method to separately estimate the latent spring and autumn survival rates using capture–recapture data, though the accuracy and precision of these estimates has not been formally tested. Here, I used simulated data to explore the estimability of migration survival rates using this model. Under a variety of biologically realistic scenarios, I demonstrate that spring and autumn migration survival can be estimated from the integrated survival model, though estimates are biased toward the overall migration survival probability. The direction and magnitude of this bias are influenced by the relative difference in spring and autumn survival rates as well as the degree of annual variation in these rates. The inclusion of covariates can improve the model's performance, especially when annual variation in migration survival rates is low. Migration survival rates can be estimated from relatively short time series (4–5 years), but bias and precision of estimates are improved when longer time series (10–12 years) are available. The ability to estimate seasonal survival rates of small, migratory organisms opens the door to advancing our understanding of the ecology and conservation of these species. Application of this method will enable researchers to better understand when mortality occurs across the annual cycle and how the migratory periods contribute to population dynamics. Integrating summer and winter capture data requires knowledge of the migratory connectivity of sampled populations and therefore efforts to simultaneously collect both survival and tracking data should be a high priority, especially for species of conservation concern.     

Individual heterogeneity in reproductive rates and cost of reproduction in a long‐lived vertebrate

Individual variation in reproductive success is a key feature of evolution, but also has important implications for predicting population responses to variable environments. Although such individual variation in reproductive outcomes has been reported in numerous studies, most analyses to date have not considered whether these realized differences were due to latent individual heterogeneity in reproduction or merely random chance causing different outcomes among like individuals. Furthermore, latent heterogeneity in fitness components might be expressed differently in contrasted environmental conditions, an issue that has only rarely been investigated. Here, we assessed (i) the potential existence of latent individual heterogeneity and (ii) the nature of its expression (fixed vs. variable) in a population of female Weddell seals (Leptonychotes weddellii), using a hierarchical modeling approach on a 30‐year mark–recapture data set consisting of 954 individual encounter histories. We found strong support for the existence of latent individual heterogeneity in the population, with “robust” individuals expected to produce twice as many pups as “frail” individuals. Moreover, the expression of individual heterogeneity appeared consistent, with only mild evidence that it might be amplified when environmental conditions are severe. Finally, the explicit modeling of individual heterogeneity allowed us to detect a substantial cost of reproduction that was not evidenced when the heterogeneity was ignored.     

Large‐scale oceanographic fluctuations drive Antarctic petrel survival and reproduction

Polar Regions are experiencing environmental changes at unprecedented rates. These changes can spread throughout entire food webs from lower trophic levels to apex predators. As many top predators forage over large areas, these indirect effects may be associated with large‐scale patterns of climate variability. Using global climate indices that are known to impact the Southern Ocean ecosystem (the El Niño Southern Oscillation and Antarctic Oscillation Indices) we assessed their efficacy to predict variation in the demographic parameters of Antarctic seabirds. First, we used a long‐term dataset on adult survival (estimated from capture–mark–recapture data) and reproduction of Antarctic petrel Thalassoica antarctica, from the largest known breeding colony (Svarthamaren, Dronning Maud Land) and examined whether large‐scale oceanographic fluctuations impact survival and reproduction. Second, we conducted an exhaustive literature review to determine whether the effects of large‐scale environmental variability on Antarctic seabirds have a coherent fingerprint across the Antarctic continent and nearby islands. We found that most of the variation in both reproductive success, timing of hatching, and survival of Antarctic petrels can be accurately modeled using the two modes of large‐scale climate variability in Antarctica. The literature review, combined with the results from our field study, suggests that while the anticipated trends in the global patterns of climatic variability will generally have detrimental effects on populations of top predators in the Southwest Atlantic, these conclusions cannot be extrapolated to all seabird populations in Antarctica without additional data.     

Fine‐scale habitat use during the non‐breeding season suggests that winter habitat does not limit breeding populations of a declining long‐distance Palearctic migrant

For migrant birds, which habitats are suitable during the non‐breeding season influences habitat availability, population resilience to habitat loss, and ultimately survival. Consequently, habitat preferences during winter and whether habitat segregation according to age and sex occurs directly influences migration ecology, survival and breeding success. We tested the fine‐scale habitat preferences of a declining Palearctic migrant, the whinchat Saxicola rubetra, on its wintering grounds in west Africa. We explored the influence of habitat at the territory‐scale and whether dominance‐based habitat occupancy occurs by describing the variation in habitat characteristics across wintering territories, the degree of habitat change within territories held throughout winter, and whether habitat characteristics influenced territory size and space‐use within territories or differed with age and sex. Habitat characteristics varied substantially across territories and birds maintained the same territories even though habitat changed significantly throughout winter. We found no evidence of dominance‐based habitat occupancy; instead, territories were smaller if they contained more perching shrubs or maize crops, and areas with more perching shrubs were used more often within territories, likely because perches are important for foraging and territory defence. Our findings suggest that whinchats have non‐specialised habitat requirements within their wintering habitat of open savannah and farmland, and respond to habitat variation by adjusting territory size and space‐use within their territories instead of competing with conspecifics. Whinchats show a tolerance for human‐modified habitats and results support previous findings that some crop types may provide high‐quality wintering habitat by increasing perch density and foraging opportunities. By having non‐specialised requirements within broad winter habitat types, migrants are likely to be flexible to changing wintering conditions in Africa, both within and across winters, so possibly engendering some resilience to the rapid anthropogenic habitat degradation occurring throughout their wintering range.