Non-breeding season habitat quality mediates the strength of density-dependence for a migratory bird

Our understanding of when natural populations are regulated during their annual cycle is limited, particularly for migratory species. This information is needed for parametrizing models that can inform management and conservation. Here, we use 14 years of data on colour-marked birds to investigate how conspecific density and habitat quality during the tropical non-breeding period interact to affect body condition and apparent annual survival of a long-distance migratory songbird, the American redstart (Setophaga ruticilla). Body condition and survival of birds in high-quality mangrove habitat declined as density increased. By contrast, body condition improved and survival did not vary as density increased in adjacent, lower quality scrub habitat, although mean condition and survival were almost always lower than in mangrove. High rainfall enhanced body condition in scrub but not in mangrove, suggesting factors such as food availability outweighed consequences of crowding in lower quality habitat. Thus, survival of overwintering redstarts in mangrove habitat, disproportionately males, appears to be regulated by a crowding mechanism based on density-dependent resource competition. Survival of individuals in scrub, mostly females, appears to be limited by density-independent environmental factors but not regulated by crowding. The contrasting effects of density and food limitation on individuals overwintering in adjacent habitats illustrate the complexity of processes operating during the non-breeding period for migratory animals, and emphasize the need for long-term studies of animals in multiple habitats and throughout their annual cycles.     

Seasonal mortality and sequential density dependence in a migratory bird

Migratory bird populations may be limited during one or more seasons, and thus at one or more places, but there is a dearth of empirical examples of this possibility. We analyse seasonal survival in a migratory shellfish‐eating shorebird (red knot Calidris canutus islandica) during a series of years of intense food limitation on the nonbreeding grounds (due to overfishing of shellfish stocks), followed by a relaxation period when destructive harvesting had stopped and food stocks for red knots recovered. For the estimation of seasonal survival from the 15 yr‐long near‐continuous capture–resight dataset, we introduce a ‘rolling window’ approach for data exploration, followed by selection of the best season definition. The average annual apparent survival over all the years was 0.81 yr^?1. During the limitation period, survival probability of adult red knots was low in winter (0.78 yr^?1), but this was compensated by high survival in summer (0.91 yr^?1). During the relaxation period survival rate levelled out with a winter value of 0.81 yr^?1 and a summer survival of 0.82 yr^?1. The fact that during the cockle‐dredging period the dip in survival in winter was completely compensated by higher survival later in the annual cycle suggests sequential density dependence. We conclude that seasonal compensation in local survival (in concert with movements to areas apparently below carrying capacity) allowed the islandica population as a whole to cope, in 1998–2003, with the loss of half of the suitable feeding habitat in part of the nonbreeding range, the western Dutch Wadden Sea. As a more general point, we see no reason why inter‐seasonal density dependence should not be ubiquitous in wildlife populations, though its limits and magnitude will depend on the specific ecological contexts. We elaborate the possibility that with time, and in stable environments, seasonal mortality evolves so that differences in mortality rates between seasons would become erased.     

Corticosterone levels as indicators of habitat quality: effects of habitat segregation in a migratory bird during the non-breeding season

During the non-breeding season, many species of territorial migratory birds exhibit a non-random pattern of habitat distribution, with males and females occupying different habitats. In this study, we examined possible physiological consequences arising from such habitat segregation in one migrant passerine species, the American redstart (Setophaga ruticilla), on its non-breeding grounds in Jamaica, West Indies. For 2 years, we measured concentrations of corticosterone, at the time of capture (baseline) and 30 min after capture (profile of acute corticosterone secretion), in redstarts in two distinct habitats, one occupied predominately by males and one mostly by females. All redstarts in both habitat types exhibited similar concentrations of baseline corticosterone levels in fall (October), whereas in spring (March–April), redstarts in female-biased habitat exhibited significantly higher baseline levels regardless of age or sex. In fall, all individuals in both habitats exhibited significant increases in corticosterone concentration with capture and handling, but in spring only redstarts (both sexes) in male-biased habitat continued to exhibit acute corticosterone secretion. Redstarts in female-biased habitat had elevated baseline corticosterone levels and reduced acute corticosterone secretion. In spring, baseline corticosterone concentration was negatively correlated with body mass, suggesting muscle catabolism associated with high corticosterone concentrations or possibly that birds are leaner as a result of increased foraging effort. These results indicate that redstarts (primarily females) in female-biased habitats suffered a decline in physiological condition, which could in turn influence their departure schedules, migration patterns and even their condition and arrival schedules on the breeding grounds. Thus, segregation of populations into habitats of different quality during the non-breeding period may have ramifications throughout the annual cycle of such migratory species. Furthermore, these results show the usefulness of plasma corticosterone levels as indicators of physiological condition and thus habitat quality for birds during the non-breeding period. Received: 14 November 1997 / Accepted: 9 March 1998     

Maladaptive habitat selection of a migratory passerine bird in a human-modified landscape

In human-altered environments, organisms may preferentially settle in poor-quality habitats where fitness returns are lower relative to available higher-quality habitats. Such ecological trapping is due to a mismatch between the cues used during habitat selection and the habitat quality. Maladaptive settlement decisions may occur when organisms are time-constrained and have to rapidly evaluate habitat quality based on incomplete knowledge of the resources and conditions that will be available later in the season. During a three-year study, we examined settlement decision-making in the long-distance migratory, open-habitat bird, the Red-backed shrike (Lanius collurio), as a response to recent land-use changes. In Northwest Europe, the shrikes typically breed in open areas under a management regime of extensive farming. In recent decades, Spruce forests have been increasingly managed with large-size cutblocks in even-aged plantations, thereby producing early-successional vegetation areas that are also colonised by the species. Farmland and open areas in forests create mosaics of two different types of habitats that are now occupied by the shrikes. We examined redundant measures of habitat preference (order of settlement after migration and distribution of dominant individuals) and several reproductive performance parameters in both habitat types to investigate whether habitat preference is in line with habitat quality. Territorial males exhibited a clear preference for the recently created open areas in forests with higher-quality males settling in this habitat type earlier. Reproductive performance was, however, higher in farmland, with higher nest success, offspring quantity, and quality compared to open areas in forests. The results showed strong among-year consistency and we can therefore exclude a transient situation. This study demonstrates a case of maladaptive habitat selection in a farmland bird expanding its breeding range to human-created open habitats in plantations. We discuss the reasons that could explain this decision-making and the possible consequences for the population dynamics and persistence.     

Plumage quality mediates a life-history trade-off in a migratory bird

Background

Moult is one of the most costly activities in the annual cycle of birds and most avian species separate moult from other energy-demanding activities, such as migration. To this end, young birds tend to undergo the first post-juvenile moult before the onset of migration, but in some species the time window for the pre-migratory feather replacement is too narrow. We hypothesized that in such species an increased investment in the structural quality of juvenile feathers may allow to retain juvenile plumage throughout the entire migratory period and delay moult until arriving at wintering grounds, thus avoiding a moult-migration overlap.

Methods

The effect of juvenile plumage quality on the occurrence of moult-migration overlap was studied in a migratory shorebird, the common snipe Gallinago gallinago. Ca. 400 of first-year common snipe were captured during their final stage of autumn migration through Central Europe. The quality of juvenile feathers was assessed as the mass-length residuals of retained juvenile rectrices. Condition of migrating birds was assessed with the mass of accumulated fat reserves and whole-blood hemoglobin concentration. Path analysis was used to disentangle complex interrelationships between plumage quality, moult and body condition.

Results

Snipe which grew higher-quality feathers in the pre-fledging period were less likely to initiate moult during migration. Individuals moulting during migration had lower fat loads and hemoglobin concentrations compared to non-moulting birds, suggesting a trade-off in resource allocation, where energetic costs of moult reduced both energy reserves available for migration and resources available for maintenance of high oxygen capacity of blood.

Conclusions

The results of this study indicate that a major life-history trade-off in a migratory bird may be mediated by the quality of juvenile plumage. This is consistent with a silver spoon effect, where early-life investment in feather quality affects future performance of birds during migration period. Our results strongly suggest that the juvenile plumage, although retained for a relatively short period of time, may have profound consequences for individuals’ fitness.

     

Climate change and fitness components of a migratory bird breeding in the Mediterranean region

The increase in spring temperatures in temperate regions over the last two decades has led to an advancing spring phenology, and most resident birds have responded to it by advancing their onset of breeding. The pied flycatcher (Ficedula hypoleuca) is a long‐distance migrant bird with a relatively late onset of breeding with respect to both resident birds and spring phenology in Europe. In the present correlational study, we show that some fitness components of pied flycatchers are suffering from climate change in two of the southernmost European breeding populations. In both montane study areas, temperature during May increased between 1980 and 2000 and an advancement of oak leafing was detected by using the normalized difference vegetation index (NDVI) to assess tree phenology. This might result in an advancement of the peak in availability of caterpillars, the main prey during the nestling stage. Over the past 18 yr, the time of egg laying and clutch size of pied flycatchers were not affected by the increase in spring temperatures in these Mediterranean populations. However, this increase seems to have an adverse effect on the reproductive output of pied flycatchers over the same period. Our data suggest that the mismatch between the timing of peak food supply and nestling demand caused by recent climate change might result in a reduction of parental energy expenditure that is reflected in a reduction of nestling growth and survival of fledged young in our study populations. The data seem to indicate that the breeding season has not shifted and it is the environment that has shifted away from the timing of the pied flycatcher breeding season. Mediterranean pied flycatchers were not able to advance their onset of breeding, probably, because they are constrained by their late arrival date and their restricted high altitude breeding habitat selection near the southern border of their range.     

Migration phenology and winter habitat quality are related to circulating androgen in a long‐distance migratory bird

In migratory birds, the timing of departure from wintering grounds is often dependant on the quality of habitat on an individual's territory and may influence individual fitness, resulting in an interaction of life history stages across large geographical distances. American redstart Setophaga ruticilla males who overwinter in high quality habitats arrive early to breed and subsequently produce more offspring than late arrivers. Since many migratory species overlap vernal migration with the physiological transition to breeding, we examined if breeding preparation plays a role in this seasonal interaction. We tested the hypothesis that early arriving male redstarts from high quality winter habitats are in superior breeding condition by simultaneously measuring winter habitat quality (stable‐carbon isotopes) and breeding preparation (circulating androgen, cloacal protuberance (CP) diameter) upon arrival at breeding grounds. Compared with late arrivers, early arriving males were from higher quality winter habitats and had higher androgen, but smaller CPs. Males arriving with higher androgen were in more advanced physiological migratory condition, as measured by haematocrit. Early arrivers were more likely to successfully breed, but there was no significant relationship between androgen upon arrival and breeding success. One possible explanation for these relationships is that androgen measured during arrival is most relevant in a migratory context, such that birds with high androgen may benefit from effects on migratory condition, positively influencing fitness through earlier arrival.     

Carry-over effects and habitat quality in migratory populations

Determining the factors that influence migratory population abundance has been constrained by the inability to connect events in different periods of the annual cycle. Carry-over effects are events that occur in one season but influence individual success the following season and recent empirical evidence suggests that they may play an important role in migratory population dynamics. Using a long distance migratory shorebird as an example, I incorporate carry-over effects and changes in the relative amount of habitat quality into a density-dependent equilibrium population model. The model uses the example where the quality of habitat on the wintering grounds (nonbreeding season) influences breeding output the following summer (breeding season). Carry-over effects, however, may be manifested in a number of other ways that could influence population dynamics. In the simulations, population declines occur when habitat is lost on the wintering grounds. However, results show that carry-over effects can magnify these declines when a disproportionate amount of high quality habitat is lost the previous winter. Simulations also show that carry-over effects can have a relative, positive impact on population size when the majority of habitat that is lost in the previous season is low quality. In this case, the carry-over interacts with density-dependence the following season producing an additive and positive effect, buffering the population from severe declines. To predict changes in population size of migratory animals, it will be important to determine (i) which demographic factors in which season produce strong carry-over effects and, (ii) not just the amount, but the relative quality of habitat that is lost. If carry-over effects are significant, they could potentially mitigate 'seasonal compensation effects' from density-dependence, leading to exacerbated population declines.     

Demographic responses to climate‐driven variation in habitat quality across the annual cycle of a migratory bird species

The demography and dynamics of migratory bird populations depend on patterns of movement and habitat quality across the annual cycle. We leveraged archival GPS‐tagging data, climate data, remote‐sensed vegetation data, and bird‐banding data to better understand the dynamics of black‐headed grosbeak (Pheucticus melanocephalus) populations in two breeding regions, the coast and Central Valley of California (Coastal California) and the Sierra Nevada mountain range (Sierra Nevada), over 28 years (1992–2019). Drought conditions across the annual cycle and rainfall timing on the molting grounds influenced seasonal habitat characteristics, including vegetation greenness and phenology (maturity dates). We developed a novel integrated population model with population state informed by adult capture data, recruitment rates informed by age‐specific capture data and climate covariates, and survival rates informed by adult capture–mark–recapture data and climate covariates. Population size was relatively variable among years for Coastal California, where numbers of recruits and survivors were positively correlated, and years of population increase were largely driven by recruitment. In the Sierra Nevada, population size was more consistent and showed stronger evidence of population regulation (numbers of recruits and survivors negatively correlated). Neither region showed evidence of long‐term population trend. We found only weak support for most climate–demographic rate relationships. However, recruitment rates for the Coastal California region were higher when rainfall was relatively early on the molting grounds and when wintering grounds were relatively cool and wet. We suggest that our approach of integrating movement, climate, and demographic data within a novel modeling framework can provide a useful method for better understanding the dynamics of broadly distributed migratory species.     

Seasonal distribution of a migratory bird: effects of local and regional resource tracking

Aim We studied how local and regional abundance of a migratory passerine (the blackcap Sylvia atricapilla) track resource availability in breeding and wintering grounds, in an attempt to understand the processes underlying the distribution and regulation of migratory bird populations in summer and in winter. Location Our study was conducted in Spain. In summer, we sampled five localities representing the diversity of environmental conditions met by breeding Spanish blackcaps. In winter, we sampled eight localities in the wintering range of the species including different habitat types (forests and shrublands). Methods Our approach was based on the matching rule, a model that predicts that any local variation in resource abundance between two adjacent habitat patches should be tracked by animals through a similar variation in population abundance. Eventually, this local process should conform to abundance distributions at regional scales. We sampled two habitat patches in each locality, each one including three to five line transects, 500‐m long and 50‐m wide, where we counted blackcaps and measured vegetation structure and fruit abundance. Results During the breeding season, the abundance of blackcaps was strongly correlated with the ground cover of brambles (Rubus spp.), a bush which grows in moist sectors in Mediterranean forests and is the commonest nesting substrate of Spanish blackcaps. Both local and regional changes in bramble cover were tracked by variations in blackcap abundance. However, the rate of increase in blackcap abundance with increasing bramble cover along the Spanish gradient was lower than the one predicted under resource matching. In winter, abundance of fruiting shrubs was the best predictor of blackcap abundance, although local abundance of blackcaps not always fitted local abundance of fruits. Notwithstanding this effect, the regional pattern of abundance tracked changes in fruit availability according to the matching rule. Main conclusions Our results support the strong effect of habitat quality on the abundance distribution of blackcaps and the tracking of different key resources along the year. Together with the different degrees of resource tracking by blackcaps at local and regional scales, these results also support the view that both breeding and wintering processes have to be studied, and studies have to be conducted at the appropriate spatial scales, if we are to understand the processes underlying the abundance distribution of migratory birds.     

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.     

Colonization and extinction dynamics of a declining migratory bird are influenced by climate and habitat degradation

Uncovering the mechanisms involved in the decline of long‐distance migrants remains one of the most pressing issues in European conservation. Since the 1980s, the British breeding population of Garden Warbler Sylvia borin has declined by more than 25%. Here we use data from repeated bird surveys of woodland sites in the 1980s and in 2003–2004 to show that, although the overall population declined between the two periods, the probability of occupancy for this species increased at high latitudes and decreased at low latitudes. Range shifts such as this arise from a change in the ratio of colonizations to extinctions at the range margins, and we therefore related colonization and local extinction at the patch level to concurrent changes in temperature and habitat. The probability of patch colonization by this species was significantly lower where the percentage cover of vegetation in the understorey had declined, reducing habitat quality for this species. The probability of local extinction was significantly correlated with increasing mean May temperature, which may reflect a change in phenology, making breeding conditions less suitable. Changed regimes of grazing and woodland management could be used to increase habitat suitability and thereby increase colonization probability at the local scale, which may in turn increase the probability of patch occupancy despite future climatic unsuitability.     

Tropical winter habitat limits reproductive success on the temperate breeding grounds in a migratory bird

Identifying the factors that control population dynamics in migratory animals has been constrained by our inability to track individuals throughout the annual cycle. Using stable carbon isotopes, we show that the reproductive success of a long-distance migratory bird is influenced by the quality of habitat located thousands of kilometres away on tropical wintering grounds. For male American redstarts (Setophaga ruticilla), winter habitat quality influenced arrival date on the breeding grounds, which in turn affected key variables associated with reproduction, including the number of young fledged. Based on a winter-habitat model, females occupying high-quality winter habitat were predicted to produce more than two additional young and to fledge offspring up to a month earlier compared with females wintering in poor-quality habitat. Differences of this magnitude are highly important considering redstarts are single brooded, lay clutches of only three to five eggs and spend only two-and-a-half months on the breeding grounds. Results from this study indicate the importance of understanding how periods of the annual cycle interact for migratory animals. Continued loss of tropical wintering habitat could have negative effects on migratory populations in the following breeding season, minimizing density-dependent effects on the breeding grounds and leading to further population declines. If conservation efforts are to be successful, strategies must incorporate measures to protect all the habitats used during the entire annual cycle of migratory animals.     

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.     

Heritability of arrival date in a migratory bird

The behaviour of long-distance migratory birds is assumed to partly be under the influence of genes, as demonstrated by selection experiments. Furthermore, competition for early arrival among males may lead to condition-dependent migration associated with fitness benefits of early arrival achieved by individuals in prime condition. Here I present field data on the repeatability and the heritability of arrival date in a trans-equatorial migratory bird, the barn swallow Hirundo rustica, and I test for a genetic correlation between arrival date and the expression of a condition-dependent secondary sexual character. The repeatability was statistically significant and the heritability of arrival date was estimated to be 0.54 (s.e. = 0.15). There was no significant evidence of this estimate being inflated by environmental or maternal condition during rearing. Arrival date and migration are condition dependent in the barn swallow, with males with the most exaggerated secondary sexual characters also arriving the earliest. There was a significant genetic correlation between arrival date and tail length in male barn swallows, providing indirect evidence for a genetic basis of this condition dependence. Given the high level of heritability, arrival date could readily respond to selection caused by environmental change.     

Seasonal home range dynamics and sex differences in habitat use in a threatened,coastal marsh bird

A comprehensive understanding of spatiotemporal ecology is needed to develop conservation strategies for declining species. The king rail (Rallus elegans) is a secretive marsh bird whose range historically extended across the eastern United States. Inland migratory populations have been greatly reduced with most remaining populations inhabiting the coastal margins. Our objectives were to determine the migratory status of breeding king rails on the mid‐Atlantic coast and to characterize home range size, seasonal patterns of movement, and habitat use. Using radiotelemetry, we tracked individual king rails among seasons, and established that at least a segment of this breeding population is resident. Mean (±SE) home range size was 19.8 ± 5.0 ha (95% kernel density) or 2.5 ± 0.9 (50% kernel density). We detected seasonal variation and sex differences in home range size and habitat use. In the nonbreeding season, resident male home ranges coincided essentially with their breeding territories. Overwintering males were more likely than females to be found in natural emergent marsh with a greater area of open water. Females tended to have larger home ranges than males during the nonbreeding season. We report for the first time the use of wooded natural marsh by overwintering females. Brood‐rearing king rails led their young considerable distances away from their nests (average maximum distance: ~600 ± 200 m) and used both wooded natural and impounded marsh. King rails moved between natural marsh and managed impoundments during all life stages, but the proximity of these habitat types particularly benefitted brood‐rearing parents seeking foraging areas with shallower water in proximity to cover. Our results demonstrate the importance of interspersion of habitat types to support resident breeders. Summer draining of impounded wetlands that are seasonally flooded for wintering waterfowl allows regrowth of vegetation and provides additional habitat at a critical time for wading birds.     

Experimental evidence for a novel mechanism driving variation in habitat quality in a food-caching bird

Variation in habitat quality can have important consequences for fitness and population dynamics. For food-caching species, a critical determinant of habitat quality is normally the density of storable food, but it is also possible that quality is driven by the ability of habitats to preserve food items. The food-caching gray jay (Perisoreus canadensis) occupies year-round territories in the coniferous boreal and subalpine forests of North America, but does not use conifer seed crops as a source of food. Over the last 33 years, we found that the occupancy rate of territories in Algonquin Park (ON, Canada) has declined at a higher rate in territories with a lower proportion of conifers compared to those with a higher proportion. Individuals occupying territories with a low proportion of conifers were also less likely to successfully fledge young. Using chambers to simulate food caches, we conducted an experiment to examine the hypothesis that coniferous trees are better able to preserve the perishable food items stored in summer and fall than deciduous trees due to their antibacterial and antifungal properties. Over a 1–4 month exposure period, we found that mealworms, blueberries, and raisins all lost less weight when stored on spruce and pine trees compared to deciduous and other coniferous trees. Our results indicate a novel mechanism to explain how habitat quality may influence the fitness and population dynamics of food-caching animals, and has important implications for understanding range limits for boreal breeding animals.