Climate,trophic interactions,density dependence and carry‐over effects on the population productivity of a migratory Arctic herbivorous bird

Several driving forces can affect recruitment rates in bird populations. However, our understanding of climate‐induced effects or bottom–up vs top–down biological processes on breeding productivity typically comes from small‐scale studies, and their relative importance is rarely investigated at the population level. Using a 31‐year time series, we examined the effects of selected environmental parameters on the annual productivity of a key Arctic herbivore, the greater snow goose Anser caerulescens atlanticus. We determined the extent to which breeding productivity, defined as the percentage of juveniles in the fall population, was affected by 1) climatic conditions, 2) fluctuations in predation pressure caused by small rodent oscillations, and 3) population size. Moreover, we took advantage of an unplanned large‐scale manipulation (i.e. management action) to examine the potential non‐lethal carry‐over effects caused by disturbance on spring staging sites. The most parsimonious model explained 66% of the annual variation in goose productivity. The spring North Atlantic Oscillation and Arctic snow depth were the primary climatic parameters inversely affecting the production of juveniles, likely through bottom–up processes. Indirect trophic interactions generated by fluctuations in lemming abundance explained 18% of the variation in goose productivity (positive relationship). Mean temperature during brood‐rearing and disturbance on staging sites (carry‐over effects) were the other important factors affecting population recruitment. We observed a strong population increase, and found no evidence of density‐dependent effects. Spatially restricted studies can identify factors linking environmental parameters to local bird reproduction but if these factors do not act synchronously over the species range, they may fail to identify the relative importance of mechanisms driving large‐scale population dynamics.     

Watershed‐scale climate influences productivity of Chinook salmon populations across southcentral Alaska

The ecosystems supporting Pacific salmon (Oncorhynchus spp.) are changing rapidly as a result of climate change and habitat alteration. Understanding how—and how consistently—salmon populations respond to changes at regional and watershed scales has major implications for fisheries management and habitat conservation. Chinook salmon (O. tshawytscha) populations across Alaska have declined over the past decade, resulting in fisheries closures and prolonged impacts to local communities. These declines are associated with large‐scale climate drivers, but uncertainty remains about the role of local conditions (e.g., precipitation, streamflow, and stream temperature) that vary among the watersheds where salmon spawn and rear. We estimated the effects of these and other environmental indicators on the productivity of 15 Chinook salmon populations in the Cook Inlet basin, southcentral Alaska, using a hierarchical Bayesian stock‐recruitment model. Salmon spawning during 2003–2007 produced 57% fewer recruits than the previous long‐term average, leading to declines in adult returns beginning in 2008. These declines were explained in part by density dependence, with reduced population productivity following years of high spawning abundance. Across all populations, productivity declined with increased precipitation during the fall spawning and early incubation period and increased with above‐average precipitation during juvenile rearing. Above‐average stream temperatures during spawning and rearing had variable effects, with negative relationships in many warmer streams and positive relationships in some colder streams. Productivity was also associated with regional indices of streamflow and ocean conditions, with high variability among populations. The cumulative effects of adverse conditions in freshwater, including high spawning abundance, heavy fall rains, and hot, dry summers may have contributed to the recent population declines across the region. Identifying both coherent and differential responses to environmental change underscores the importance of targeted, watershed‐specific monitoring and conservation efforts for maintaining resilient salmon runs in a warming world.     

Asynchrony in population dynamics of sockeye salmon in southwest Alaska

Ecologists have examined the synchronization of population dynamics across space as a means to understand how populations respond to climate variation. However, response diversity may reflect important variation among local population dynamics driven by population‐specific responses to regional environmental change. We used long‐term data on sockeye salmon Oncorhynchus nerka from pristine watersheds of southwestern Alaska to show that populations spawning in close proximity (<40 km) to one another have a limited degree of synchrony in their dynamics, even after accounting for density‐dependent processes. In fact, the dynamics of local populations of stream‐spawning sockeye salmon were no more coherent than those of stocks at a much coarser resolution across this region of Alaska. We examined four hypotheses to explain the observed patterns of asynchrony among stream‐spawning populations, and found that populations spawning in dissimilar habitats, and using different nursery lakes were less synchronized in their productivity. Similarity in the age structure of spawning adults was less correlated with synchrony in productivity. These results emphasize the importance of maintaining diverse spawning and rearing habitat for the conservation of Pacific salmon, and should guide conservation planning for Pacific salmon populations in regions where natural dynamics have been altered by habitat loss, hatchery practices, and over‐fishing.     

Shifting dynamic forces in fish stock fluctuations triggered by age truncation?

Accumulating evidence shows that environmental fluctuations and exploitation jointly affect marine fish populations, and understanding their interaction is a key issue for fisheries ecology. In particular, it has been proposed that age truncation induced by fisheries exploitation may increase the population's sensitivity to climate. In this study, we use unique long‐term abundance data for the Northeast Arctic stock of cod (Gadus morhua) and the Norwegian Spring‐Spawning stock of herring (Clupea harengus), which we analyze using techniques based on age‐structured population matrices. After identifying time periods with different age distributions in the spawning stock, we use linear models to quantify the relative effect of exploitation and temperature on the population growth rates. For the two populations, age truncation was found to be associated with an increasing importance of temperature and a relatively decreasing importance of exploitation, while the population growth rate became increasingly sensitive to recruitment variations. The results suggested that the removal of older age classes reduced the buffering capacity of the population, thereby making the population growth rate more dependent on recruitment than adult survival and increasing the effect of environmental fluctuations. Age structure appeared as a key characteristic that can affect the response of fish stocks to climate variations and its consequences may be of key importance for conservation and management.     

Reproductive strategy and recruitment dynamics of amphidromous goby Sicyopterus japonicus as revealed by otolith microstructure

The daily ages of 312 of 879 newly recruited postlarvae of Sicyopterus japonicus, collected from the Shuang‐Chi Estuary in north‐eastern Taiwan during February 1996 to April 1997, were determined from daily growth increments in their otoliths. Pelagic larval duration, growth rate of the marine larval stage and hatching dates were estimated, and recruitment timing was linked to environmental factors. The mean ±s .d . total length (L_T) and daily ages of S. japonicus at recruitment to the estuary were estimated to be 33·95 ± 1·31 mm (range 30·7 to 38·1, n = 317) and 163·72 ± 12·79 days (range 130 to 198, n = 312), respectively. The recruitment of S. japonicus larvae is size dependent not age dependent because L_T of the larvae is independent of age at recruitment. Periodic analysis indicated that L_T and growth rate of the larvae were inversely correlated with the age at recruitment, which means that the fast‐growing individuals recruited earlier. The growth rate of S. japonicus in the marine larval stage was synchronous with marine productivity in this subtropical area, i.e. the spring cohort recruited in the autumn had a higher growth rate than the autumn cohort recruited the following spring. The main spawning season of S. japonicus as backcalculated from otolith daily increments was in autumn, a relatively low productivity period compared with spring. During this season, there were fewer competitors and predators than in the more productive spring. The recruitment of 95% of postlarva coincided with low salinity (14) and low water temperature (23° C) in the river mouth that provided a buffer area for the adaptation of the larvae for upstream migration. This unique reproduction strategy and prolonged larval duration facilitated the widespread distribution of the fish along the coasts of East Asia.     

Inter-annual plasticity of squid life history and population structure: ecological and management implications

Population size and structure, as well as individual growth rates, condition, and reproductive output, respond to environmental factors, particularly in short-lived and fast-growing squid species. We need to understand the mechanisms through which populations respond to environmental conditions, to predict when or if established relationships, used as management tools to forecast recruitment strength, might break down completely. Identifying characteristics of successful recruits who have grown under different environmental scenarios may improve our understanding of the mechanistic connections between environmental conditions and the temporal variation in life history characteristics that ultimately affect recruitment. This 5-year study sought to determine the association between key life history characteristics of southern calamary Sepioteuthis australis (growth rate, body size, and patterns of repro-somatic energy allocation) and the environmental conditions experienced by individuals on the east coast of Tasmania, Australia. Among years, all population and individual parameters examined were highly variable, despite the environmental regime during the study not encompassing the extremes that may occur in this dynamic region. Temperature was not clearly associated with any of the individual or population differences observed. Populations of apparently similar abundance were composed of individuals with strikingly different biological characteristics, therefore seeking relationships between abundance and environmental parameters at gross levels did not shed light on the mechanisms responsible for population size. Importantly, inter-annual differences in squid size, condition, reproductive investment, and possibly growth rate, were sex-specific, indicating that males and females responded differently to similar factors. Among years differences in body size were extreme, both among the male component of the population and between genders. The relative importance of many size-based processes that contribute to population size and structure (e.g. predation, starvation, competition, and reproductive success) will therefore vary inter-annually.An erratum to this article can be found at     

Recruitment and growth of two small‐bodied resident fish species (Gobiidae and Atherinidae) in oligohaline,seasonally open lagoons

Spatio‐temporal recruitment patterns, growth and survival of the Swan River goby Pseudogobius olorum and western hardyhead Leptatherina wallacei are described from two small, coastal lagoons on the south coast of Western Australia. In these lagoons, estuarine salinity dynamics were relatively stable over much of the autumn–spring period when freshwater inputs from rivers were reduced and there was no oceanic connection. Preflexion and flexion stages of both fish species contributed strongly to population size structure in downstream reaches, whereas upstream reaches were dominated by postflexion larvae and juvenile stages. Spawning of both species was protracted and largely asynchronous, although the episodic presence of stronger preflexion and flexion cohorts suggested some synchronized spawning had occurred. Comparison with estuarine conditions over this period provided evidence that synchronized spawning may be related to temperature and salinity variations from a combination of freshwater inputs and periods of marine exchange. Uninterrupted growth and the progression of cohorts through to juvenile stages were consistent with the generally stable estuarine conditions. Larval and juvenile stages of both species were also tolerant of abrupt changes in salinity and temperature, which occurred due to a non‐seasonal oceanic connection. These findings were consistent with the euryhaline nature of adults of both species.     

Negative or positive density‐dependence in movements depends on climatic seasons: The case of a Neotropical marsupial

One of the major challenges in animal ecology is to understand the factors and processes driving movement behaviour. Although density may influence movement patterns, the occurrence and nature of density‐dependence in animal movements are still unclear, particularly whether it may vary among populations of a species, or across time within a population. Here, we evaluate the occurrence and nature of density‐dependence in the movements of a Neotropical marsupial, the Grey four‐eyed opossum Philander frenatus (Didelphidae, Didelphimorphia). We quantified fine‐scale path tortuosity of individuals inhabiting continuous forest areas and forest fragments, in different climatic seasons (humid vs. super‐humid). We also determined the relative importance of population size compared to sex and body mass on movements, using a model‐selection approach. In forest fragments, path tortuosity increased with population size in the super‐humid season, but decreased in the humid season. In the continuous forest, path tortuosity was affected only by sex and body mass, being slightly higher in males and negatively related to body mass. The occurrence of density‐dependence on movements only in forest fragments is likely to reflect the higher overall density of P. frenatus in small forest fragments. The variation in the nature of density‐dependence between climatic seasons is likely to reflect a trade off between foraging over large areas (humid season, low resource availability) versus avoiding agonistic encounters (super‐humid season, high resource availability). Our results show that (i) density‐dependence in movements may be context‐dependent occurring only in areas of relatively high overall population density; and (ii) density may affect movements in different ways at different climatic seasons.     

What if it gets crowded? Density‐dependent tortuosity in individual movements of a Neotropical mammal

Effects of density dependence on animal movements have received much attention in ecology, but it is still debated to what extent dispersal and movements in general are density dependent, and their potential contribution to population regulation processes. Here, we determine the occurrence and nature of density dependence in the movements of a Neotropical marsupial, the black‐eared opossum Didelphis aurita Wied‐Neuwied 1826. Using spool‐and‐line tracking devices, we estimated the tortuosity of fine‐scale movements of 149 individuals by their fractal dimension D. We evaluated the relative importance of population size, reproductive or climatic seasons and reproductive maturity of individuals as determinants of movement tortuosity, using a model selection approach. Population size was the most important determinant of movement tortuosity, with season (climatic seasons for females, reproductive seasons for males) and reproductive maturity as secondary but also important variables. We detected a positive density‐dependent effect on movement tortuosity, resulting in more intensive use of areas by individuals during periods of high population size. This positive association between movement tortuosity and population size is more likely to result from intraspecific competition, which forces individuals to explore their environment more intensively during high‐density periods. Therefore, despite being density dependent, movements in D. aurita apparently do not contribute to population regulation mechanisms.     

Integrated modeling predicts shifts in waterbird population dynamics under climate change

Climate change has been identified as one of the most important drivers of wildlife population dynamics. The in‐depth knowledge of the complex relationships between climate and population sizes through density dependent demographic processes is important for understanding and predicting population shifts under climate change, which requires integrated population models (IPMs) that unify the analyses of demography and abundance data. In this study we developed an IPM based on Gaussian approximation to dynamic N‐mixture models for large scale population data. We then analyzed four decades (1972–2013) of mallard Anas platyrhynchos breeding population survey, band‐recovery and climate data covering a large spatial extent from North American prairies through boreal habitat to Alaska. We aimed to test the hypothesis that climate change will cause shifts in population dynamics if climatic effects on demographic parameters that have substantial contribution to population growth vary spatially. More specifically, we examined the spatial variation of climatic effects on density dependent population demography, identified the key demographic parameters that are influential to population growth, and forecasted population responses to climate change. Our results revealed that recruitment, which explained more variance of population growth than survival, was sensitive to the temporal variation of precipitation in the southern portion of the study area but not in the north. Survival, by contrast, was insensitive to climatic variation. We then forecasted a decrease in mallard breeding population density in the south and an increase in the northwestern portion of the study area, indicating potential shifts in population dynamics under future climate change. Our results implied that different strategies need to be considered across regions to conserve waterfowl populations in the face of climate change. Our modelling approach can be adapted for other species and thus has wide application to understanding and predicting population dynamics in the presence of global change.     

Dry season watering alters the significance of climate factors influencing phenology and growth of saplings of savanna woody species in central Zambia,southern Africa

Although tree growth in southern African savannas is correlated with rainfall in the wet season, some studies have shown that tree growth is controlled more by rainfall in the dry season. If more rainfall occurred in the dry season in future climates, it would affect the growth of savanna trees, especially saplings that have shallower roots which limit access to subsoil water during the dry season when leaf flush and shoot extension occur. Recent paleobotanical evidence has revealed that there was relatively more precipitation in the dry season in eastern Africa in the Eocene than under the current climate. Saplings therefore can be expected to respond more to water addition during the dry season than mature trees that have more stored water and deeper roots that access subsoil water. Accordingly, I hypothesized that irrigation in the dry season should (i) advance the onset of the growing season, (ii) increase growth rates and (iii) alter the growth responses of saplings to climate factors. To test these hypotheses saplings of five savanna woody species were irrigated during the hot‐dry season at a site in central Zambia and their monthly and annual growth rates compared to those of conspecifics growing under control conditions. Although the responses among the species were variable, all irrigated saplings had significantly higher monthly and annual growth rates than control plants. In addition, dry season watering significantly altered the climatic determinants of sapling growth by either strengthening the role of the same climatic factors that were important under control conditions or displacing them altogether. In conclusion, more precipitation during the hot‐dry season is likely to have significant positive effects on sapling growth and consequently reduce the sapling‐tree transition periods and promote future tree population recruitment in some southern African savanna tree species.     

Plastic reproductive allocation as a buffer against environmental stochasticity – linking life history and population dynamics to climate

Empirical work suggest that long‐lived organisms have adopted risk sensitive reproductive strategies where individuals trade the amount of resources spent on reproduction versus survival according to expected future environmental conditions. Earlier studies also suggest that climate affects population dynamics both directly by affecting population vital rates and indirectly through long‐term changes in individual life histories. Using a seasonal and state‐dependent individual‐based model we investigated how environmental variability affects the selection of reproductive strategies and their effect on population dynamics. We found that: (1) dynamic, i.e. plastic, reproductive strategies were optimal in a variable climate. (2) Females in poor and unpredictable climatic regimes allocated fewer available resources in reproduction and more in own somatic growth. This resulted in populations with low population densities, and a high average female age and body mass. (3) Strong negative density dependence on offspring body mass and survival, along with co‐variation between climatic severity and population density, resulted in no clear negative climatic effects on reproductive success and offspring body mass. (4) Time series analyses of population growth rates revealed that populations inhabiting benign environments showed the clearest response to climatic perturbations as high population density prohibited an effective buffering of adverse climatic effects as individuals were not able to gain sufficient body reserves during summer. Regularly occurring harsh winters ‘harvested’ populations, resulting in persistent low densities, and released them from negative density dependent effects, resulting in high rewards for a given resource allocation.     

Fine‐scale temporal adaptation within a salmonid population: mechanism and consequences

We demonstrate a clear example of local adaptation of seasonal timing of spawning and embryo development. The consequence is a population of pink salmon that is segmented into spawning groups that use the same limited habitat. We synthesize published observations with results of new analyses to demonstrate that genetic variation of these traits results in survival differentials related to that variation, and that density‐dependent embryo mortality and seasonally variable juvenile mortality are a mechanism of selection. Most examples of local adaptation in natural systems depend on observed correlations between environments and fitness traits, but do not fully demonstrate local adaptation: that the trait is genetically determined, exhibits different fitness in common environments or across different environments, and its variation is mechanistically connected to fitness differences. The geographic or temporal scales of local adaptation often remain obscure. Here, we show that heritable, fine‐scale differences of timing of reproductive migration in a pink salmon (Oncorhynchus gorbuscha) resulted in temporal structure that persisted several generations; the differences enable a density‐dependent population to pack more spawners into limited spawning habitat, that is, enhance its fitness. A balanced trade‐off of survivals results because embryos from early‐migrating fish have a lower freshwater survival (harsh early physical conditions and disturbance by late spawners), but emigrant fry from late‐migrating fish have lower marine survivals (timing of their vernal emergence into the estuarine environment). Such fine‐scale local adaptations increase the genetic portfolio of the populations and may provide a buffer against the impacts of climate change.     

Changing environmental gradients over forty years alter ecomorphological variation in Guadalupe Bass Micropterus treculii throughout a river basin

Understanding the degree of intraspecific variation within and among populations is a key aspect of predicting the capacity of a species to respond to anthropogenic disturbances. However, intraspecific variation is usually assessed at either limited temporal, but broad spatial scales or vice versa, which can make assessing changes in response to long‐term disturbances challenging. We evaluated the relationship between the longitudinal gradient of changing flow regimes and land use/land cover patterns since 1980 and morphological variation of Guadalupe Bass Micropterus treculii throughout the Colorado River Basin of central Texas. The Colorado River Basin in Texas has experienced major alterations to the hydrologic regime due to changing land‐ and water‐use patterns. Historical collections of Guadalupe Bass prior to rapid human‐induced change present the unique opportunity to study the response of populations to varying environmental conditions through space and time. Morphological differentiation of Guadalupe Bass associated with temporal changes in flow regimes and land use/land cover patterns suggests that they are exhibiting intraspecific trait variability, with contemporary individuals showing increased body depth, in response to environmental alteration through time (specifically related to an increase in herbaceous land cover, maximum flows, and the number of low pulses and high pulses). Additionally, individuals from tributaries with increased hydrologic alteration associated with urbanization or agricultural withdrawals tended to have a greater distance between the anal and caudal fin. These results reveal trait variation that may help to buffer populations under conditions of increased urbanization and sprawl, human population growth, and climate risk, all of which impose novel selective pressures, especially on endemic species like Guadalupe Bass. Our results contribute an understanding of the adaptability and capacity of an endemic population to respond to expected future changes based on demographic or climatic projection.     

Extreme individual flexibility of heterothermy in free-ranging Malagasy mouse lemurs (<Emphasis Type="Italic">Microcebus griseorufus</Emphasis>)

Flexibility in physiological processes is essential to adequately respond to changes in environmental conditions. Madagascar is a particularly challenging environment because climatic conditions seem less predictable than in comparative ecosystems in other parts of the world. We used the reddish-gray mouse lemur (Microcebus griseorufus) from the most unpredictable environment in Madagascar as a model to investigate the flexibility of energy saving strategies to cope with the unpredictability of their habitat. For this we measured T _sk of free-ranging mouse lemurs throughout the year using temperature data loggers. M. griseorufus showed a very strong seasonal as well as an individual flexibility in thermoregulation. During the rainy season all M. griseorufus remained normothermic. At the beginning of the dry season individuals started to exhibit different energy saving strategies: irregular short torpor bouts, regular daily torpor, prolonged torpor of a few days, and hibernation over several weeks. The accumulation of sufficient seasonal body fat was the crucial factor determining the thermal behavior of individuals. The observed intraspecific and sex independent variation in thermoregulatory patterns within one population inhabiting the same small geographical area is exceptional and gives M. griseorufus the ability to respond to current environmental as well as individual conditions. This thermal plasticity might be seen as a key to success and survival for M. griseorufus in an extremely unpredictable environment.     

Recruitment in Japanese sea bass,Lateolabrax japonicas (Cuvier, 1828): effects of timing on spawning and larval quality and quantity

This study aimed to elucidate the causes of variability in larval survival and juvenile abundance (recruitment) within and among cohorts of Japanese sea bass (JSB; Lateolabrax japonicus), a winter‐spawning temperate coastal marine fish. Larvae and settled individuals (settlers) belonging to four cohorts were collected from Tango Bay (the Sea of Japan coast) during eight sampling cruises in 2007 and 2008. Larvae were sampled in January and February each year using an ichthyoplankton net, and settlers were collected in February and March each year using a beam trawl. Age of individual larva and settlers was determined and growth history was back‐calculated from otolith microstructure, and the hatch date distribution was computed. Temperature, daily growth rate, size‐at‐age, hatch date, and density data of larvae and settlers allowed elucidating the effects of the timing of spawning and larval quantity and quality (growth rate and body size) on larval survival and recruitment within and among cohorts of JSB. Results showed that cohorts that hatched earlier in the season had higher quantity of larvae, experienced higher mean temperatures and survived better than cohorts hatched later. Recruitment variability among cohorts is determined largely by the initial quantity of larvae, as this explained >97% of the variability in recruitment among cohorts. Within cohorts, larger hatched larvae grew faster than their smaller conspecifics, and the bigger and faster growing larvae survived and settled. Results from this study suggest the following scenarios for recruitment of JSB: (i) earlier spawning in the season promotes larval survival since earlier cohorts are likely to encounter a better temperature and perhaps food conditions, and therefore recruit better than later cohorts; (ii) the initial quantity of larvae appears to be an important determinant of recruitment variability among cohorts; and (iii) the size‐ and growth‐related mechanisms operating during the larval phase appear to start at the time of the hatch.     

Multi-species response to rapid environmental change in a large estuary system: A biochronological approach

The sensitivity of species to environmental change is dependent on their ecological requirements (i.e. specialist v. generalist), and hence likely to be species-specific. Identifying species level variation in environmental sensitivity informs assessments of community vulnerability and assists in developing adaptive management strategies. We investigated species-specific sensitivity in fish to understand the vulnerability of differing life histories and ecological requirements to rapid environmental alteration (i.e. drought). Biochronologies of fish growth, based on increment widths in otoliths, were analysed using a mixed modelling approach. We assessed multi-decadal responses in fish growth to environmental variation in the terminal system of Australia’s largest river, for three long-lived fish species with differing life histories and ecological requirements: a freshwater specialist and two estuarine generalists. Biochronologies were between 20 and 38 years long, spanned a decade of severe drought and showed considerable inter-annual variation in growth. Precipitation influenced the growth of the obligate freshwater specialist, Macquaria ambigua ambigua. Temperature and salinity influenced the growth of the two estuarine generalists: Argyrosomus japonicus (estuarine opportunist) and Acanthopagrus butcheri (estuarine dependent), respectively. These results suggest that generalisations about how species respond to environmental change may mask species-specific responses to dependent on the constraints of their ecological requirements (i.e. specialist v. generalist). These findings also highlight the importance of considering the diversity of life history strategies that inhabit an ecosystem when developing conservation and management strategies.     

Density‐dependent survival and recruitment in a long‐distance Palaearctic migrant,the Sand Martin Riparia riparia

Long‐term studies can provide powerful insights into the relative importance of different demographic and environmental factors determining avian population dynamics. Here we use 23 years of capture–mark–recapture data (1981–2003) to estimate recruitment and survival rates for a Sand Martin Riparia riparia population in Cheshire, NW England. Inter‐annual variation in recruitment and adult survival was positively related to rainfall in the sub‐Saharan wintering grounds, but unrelated to weather conditions on the breeding grounds. After allowing for the effects of African rainfall, both demographic rates were negatively density‐dependent: adult survival was related to the size of the western European Sand Martin population (probably reflecting competition for resources in the shared wintering grounds) while recruitment was related to the size of the local study population in Cheshire (potentially reflecting competition for nesting sites or food). Local population size was more sensitive to variation in adult survival than to variation in recruitment, and an increase in population size after 1995 was driven mainly by the impact of more favourable conditions in the African wintering grounds on survival rates of adults. Overwinter survival in this long‐distance Palaearctic migrant is determined partly by the amount of suitable wetland foraging habitat in the sub‐Saharan wintering grounds (which is limited by the extent of summer rainfall) and partly by the number of birds exploiting that habitat.