GPS tracking analyses reveal finely-tuned shorebird space use and movement patterns throughout the non-breeding season in high-latitude austral intertidal areas.

BACKGROUND: Long-distance migratory birds spend most of their annual cycle in non-breeding areas. During this period birds must meet their daily nutritional needs and acquire additional energy intake to deal with future events of the annual cycle. Therefore, patterns of space use and movement may emerge as an efficient strategy to maintain a trade-off between acquisition and conservation of energy during the non-breeding season. However, there is still a paucity of research addressing this issue, especially in trans-hemispheric migratory birds. METHODS: Using GPS-tracking data and a recently developed continuous-time stochastic process modeling framework, we analyzed fine-scale movements in a non-breeding population of Hudsonian godwits (Limosa haemastica), a gregarious long-distance migratory shorebird. Specifically, we evaluated if these extreme migrants exhibit restricted, shared, and periodic patterns of space use on one of their main non-breeding grounds in southern South America. Finally, via a generalized additive model, we tested if the observed patterns were consistent within a circadian cycle. RESULTS: Overall, godwits showed finely-tuned range-residence and periodic movements (each 24-72 h), being similar between day and night. Remarkably, range-resident individuals segregated spatially into three groups. In contrast, a smaller fraction of godwits displayed unpredictable and irregular movements, adding functional connectivity within the population. CONCLUSIONS: In coastal non-breeding areas where resource availability is highly predictable due to tidal cycles, range-resident strategies during both the day and night are the common pattern in a long-distance shorebird population. Alternative patterns exhibited by a fraction of non-resident godwits provide functional connectivity and suggest that the exploratory tendency may be essential for information acquisition and associated with individual traits. The methodological approach we have used contributes to elucidate how the composition of movement phases operates during the non-breeding season in migratory species and can be replicated in non-migratory species as well. Finally, our results highlight the importance of considering movement as a continuum within the annual cycle.

Mediterranean songbirds show pronounced seasonal variation in thermoregulatory traits.

Addressing the patterns of variation in thermal traits is crucial to better predict the potential effects of climate change on organisms. Here, we assessed seasonal (winter vs summer) adjustments in key thermoregulatory traits in eight Mediterranean-resident songbirds. Overall, songbirds increased whole-animal (by 8%) and mass-adjusted (by 9%) basal metabolic rate and decreased (by 56%) thermal conductance below the thermoneutral zone during winter. The magnitude of these changes was within the lower values found in songbirds from northern temperate areas. Moreover, songbirds increased (by 11%) evaporative water loss within the thermoneutral zone during summer, while its rate of increase above the inflection point of evaporative water loss (i.e., the slope of evaporative water loss versus temperature) decreased by 35% during summer - a value well above that reported for other temperate and tropical songbirds. Finally, body mass increased by 5% during winter, a pattern similar to that found in many northern temperate species. Our findings support the idea that physiological adjustments might enhance the resilience of Mediterranean songbirds to environmental changes, with short-term benefits by saving energy and water under thermally stressful conditions. Nevertheless, not all species showed the same patterns, suggesting different strategies in their thermoregulatory adaptations to seasonal environments.

Heat tolerance limits of Mediterranean songbirds and their current and future vulnerabilities to temperature extremes.

Songbirds are one of the groups most vulnerable to extreme heat events. Although several recent studies have assessed their physiological responses to heat, most of them have focused solely on arid-zone species. We investigated thermoregulatory responses to heat in eight small-sized songbirds occurring in the Mediterranean Basin, where heatwaves are becoming more frequent and intense. Specifically, we determined their heat tolerance limits (HTLs) and evaporative cooling efficiency, and evaluated their current and future vulnerabilities to heat in southwestern Iberia, a Mediterranean climate warming hotspot. To do this, we exposed birds to an increasing profile of air temperatures (Ta) and measured resting metabolic rate (RMR), evaporative water loss (EWL), evaporative cooling efficiency (the ratio between evaporative heat loss and metabolic heat production) and body temperature (Tb). HTL ranged between 40 and 46°C across species, and all species showed rapid increases in RMR, EWL and Tb in response to increasing Ta. However, only the crested lark (Galerida cristata) achieved an evaporative cooling efficiency greater than 1. The studied songbirds currently experience summer Ta maxima that surpass the upper critical temperatures of their thermoneutral zone and even their HTL. Our estimates indicate that five of the eight species will experience moderate risk of lethal dehydration by the end of the century. We argue that the limited heat tolerance and evaporative cooling efficiency of small-sized Mediterranean songbirds make them particularly vulnerable to heatwaves, which will be exacerbated under future climate change scenarios.

The thermoregulatory role of relative bill and leg surface areas in a Mediterranean population of Great tit (Parus major).

There is growing evidence on the role of legs and bill as 'thermal windows' in birds coping with heat stress. However, there is a lack of empirical work examining the relationship between the relative bill and/or leg surface areas and key thermoregulatory traits such as the limits of the thermoneutral zone (TNZ) or the cooling efficiency at high temperatures. Here, we explored this relationship in a Mediterranean population of Great tit (Parus major) facing increasing thermal stress in its environment. The lower and upper critical limits of the TNZ were found to be 17.7 ± 1.6ºC and 34.5 ± 0.7°C, respectively, and the basal metabolic rate was 0.96 ± 0.12 ml O2 min-1 on average. The evaporative water loss (EWL) inflection point was established at 31.85 ± 0.27°C and was not significantly different from the value of the upper critical limit. No significant relationship was observed between the relative bill or tarsi size and TNZ critical limits, breadth, mass-independent VO2, or mass-independent EWL at any environmental temperature (from 10 to 40°C). However, Great tit males (but not females) with larger tarsi areas (a proxy of leg surface area) showed higher cooling efficiencies at 40°C. We found no support for the hypothesis that the bill surface area plays a significant role as a thermal window in Great tits, but the leg surface areas may play a role in males' physiological responses to high temperatures. On the one hand, we argue that the studied population occupies habitats with available microclimates and fresh water for drinking during summer, so active heat dissipation by EWL might be favored instead of dry heat loss through the bill surface. Conversely, male dominance behaviors could imply a greater dependence on cutaneous EWL through the upper leg surfaces as a consequence of higher exposure to harsh environmental conditions than faced by females.

Urohidrosis as an overlooked cooling mechanism in long-legged birds.

Behavioural thermoregulation could buffer the impacts of climate warming on vertebrates. Specifically, the wetting of body surfaces and the resulting evaporation of body fluids serves as a cooling mechanism in a number of vertebrates coping with heat. Storks (Ciconiidae) frequently excrete onto their legs to prevent overheating, a phenomenon known as urohidrosis. Despite the increasingly recognised role of bare and highly vascularised body parts in heat exchange, the ecological and evolutionary determinants of urohidrosis have been largely ignored. We combine urohidrosis data from a scientifically curated media repository with microclimate and ecological data to investigate the determinants of urohidrosis in all extant stork species. Our phylogenetic generalised linear mixed models show that high temperature, humidity and solar radiation, and low wind speed, promote the use of urohidrosis across species. Moreover, species that typically forage in open landscapes exhibit a more pronounced use of urohidrosis than those mainly foraging in waterbodies. Substantial interspecific variation in temperature thresholds for urohidrosis prevalence points to different species vulnerabilities to high temperatures. This integrated approach that uses online data sources and methods to model microclimates should provide insight into animal thermoregulation and improve our capacity to make accurate predictions of climate change's impact on biodiversity.

Mate selection based on labile traits affects short-term fitness in a long-lived seabird.

In long-lived monogamous social species, partner compatibility can play a crucial role in reproductive success. We evaluated assortative mating based on body condition (plasma triglyceride concentration), diet (δ15N), and foraging habitat (δ13C) in the blue-footed booby Sula nebouxii, a long-lived monogamous seabird. We investigated the effects of assortative mating (sum of triglycerides in a pair) and asymmetry within pairs (residuals from regression of female-male triglycerides) on reproductive performance and offspring growth (alkaline phosphatase, ALP). We found that strong assortative mating determined by body condition and diet seemed to be related to a signalling mechanism (nutritional state). This mating pattern had a substantial effect on the breeding parameters and influenced offspring ALP. Within-pair asymmetry did not influence any reproductive parameters, but the ALP of offspring was related to the within-pair relative female condition. Overall, our results indicate that individuals seek the best possible match to maximize their breeding investment and/or individuals are limited in their mate options by their current body condition, which has consequences for offspring fitness in the short term. Our findings show that assortative mating based on body condition produces notable variation in the joint condition of the pair, which determines their breeding success.

Going to sleep with a full belly: Thermal substitution by specific dynamic action in shorebirds.

Many bird species occupy habitats where environmental temperatures fall well below their thermoneutral zone (TNZ), so they must deal with high energy costs of thermoregulation to keep in heat balance. In such circumstances, specific dynamic action (SDA) - also referred to as heat increment of feeding - could be used to substitute for these high thermoregulatory costs. If birds ingest food before going to roost in cold environments, the SDA will be beneficial as an energy-conserving mechanism by thermal substitution. We investigated the magnitude and duration of SDA in a small-sized shorebird, the dunlin Calidris alpina, while feeding on living prey. We simulated in the aviary the food availability of a semidiurnal tidal cycle, and calculated the thermal substitution by SDA below their TNZ at the beginning of the "high tide" (resting period), after feeding ad libitum during the "low tide" (feeding period). Within TNZ (25 °C), dunlins consumed 12% (2.15 kJ) of the gross energy intake in excess by the SDA, with a duration of ~95 min. At 10 °C, i.e. below the lower critical limit of TNZ, SDA magnitude and duration were reduced by 29% and 31%, respectively. The amount of food ingested significantly affected the duration and magnitude of SDA, as well as the dunlin's body temperature. Thermal substitution by SDA saved 11% of the dunlin's theoretical daily energy requirement during winter. This thermal substitution could be commonly used by birds going to roost in cold climates. Interacting with other different behavioral and/or physiological strategies would help to maintain lower energetic costs and enhance survival in cold environments.

Understanding how birds rebuild fat stores during migration: insights from an experimental study.

Mechanisms underlying fat accumulation for long-distance migration are not fully understood. This is especially relevant in the context of global change, as many migrants are dealing with changes in natural habitats and associated food sources and energy stores. The continental Black-tailed godwit Limosa limosa limosa is a long-distance migratory bird that has undergone a considerable dietary shift over the past few decades. Historically, godwits fed on an animal-based diet, but currently, during the non-breeding period godwits feed almost exclusively on rice seeds. The latter diet may allow building up of their fuel stores for migration by significantly increasing de novo lipogenesis (DNL) activity. Here, we performed an experiment to investigate lipid flux and the abundance of key enzymes involved in DNL in godwits, during fasting and refueling periods at the staging site, while feeding on rice seeds or fly larvae. Despite no significant differences found in enzymatic abundance (FASN, ME1, ACC and LPL) in stored fat, experimental godwits feeding on rice seeds presented high rates of DNL when compared to fly-larvae fed birds (~35 times more) and fasted godwits (no DNL activity). The increase of fractional DNL in godwits feeding on a carbohydrate-rich diet can potentially be enhanced by the fasting period that stimulates lipogenesis. Although requiring further testing, these recent findings provide new insights into the mechanisms of avian fat accumulation during a fasting and refueling cycle and associated responses to habitat and dietary changes in a migratory species.

Day and night use of habitats by northern pintails during winter in a primary rice-growing region of Iberia.

Loss of natural wetlands is a global phenomenon that has severe consequences for waterbird populations and their associated ecosystem services. Although agroecosystems can reduce the impact of natural habitat loss, drivers of use of such artificial habitats by waterbirds remain poorly understood. Using the cosmopolitan northern pintail Anas acuta as a model species, we monitored home-range and fine-scale resource selection across the agricultural landscape. Individuals were tracked using GPS-GSM transmitters, and a suite of environmental and landscape features were measured throughout the winter seasons. Spatial patterns of habitat use were analysed using generalized linear mixed effect models by integrating field-observations with GPS telemetry. All birds used rice fields as foraging grounds at night and commuted to an adjacent reservoir to roost during daylight. Home-ranges and maximum foraging distances of nocturnally foraging birds increased with decreasing availability of flooded fields, and were positively correlated with moonlight levels. Birds selected flooded rice paddies (water depth range: 9-21 cm) with standing stubble and substrate with pebbles smaller than 0.5 cm in diameter. Density of rice seeds, rice paddy size, and other environmental and landscape features did not emerge as significant predictors. Our findings indicate that nocturnal foraging of northern pintails within rice fields is driven primarily by straw manipulation, water level and substrate pebble size. Thus, the presence of standing stubble in flooded paddies with soft bottoms should be prioritized to improve foraging areas for dabbling ducks. These management procedures in themselves would not increase economic costs or affect rice production and could be applied for dabbling-duck conservation throughout the world.

Generational shift in spring staging site use by a long-distance migratory bird.

In response to environmental change, species have been observed altering their migratory behaviours. Few studies, however, have been able to determine whether these alterations resulted from inherited, plastic or flexible changes. Here, we present a unique observation of a rapid population-level shift in migratory routes-over 300 km from Spain to Portugal-by continental black-tailed godwits Limosa limosa limosa This shift did not result from adult godwits changing staging sites, as adult site use was highly consistent. Rather, the shift resulted from young godwits predominantly using Portugal over Spain. We found no differences in reproductive success or survival among individuals using either staging site, indicating that the shift resulted from developmental plasticity rather than natural selection. Our results therefore suggest that new migratory routes can develop within a generation and that young individuals may be the agents of such rapid changes.

Wetland salinity induces sex-dependent carry-over effects on the individual performance of a long-distance migrant.

Salinization is having a major impact on wetlands and its biota worldwide. Specifically, many migratory animals that rely on wetlands are increasingly exposed to elevated salinity on their nonbreeding grounds. Experimental evidence suggests that physiological challenges associated with increasing salinity may disrupt self-maintenance processes in these species. Nonetheless, the potential role of salinity as a driver of ecological carry-over effects remains unstudied. Here, we investigated the extent to which the use of saline wetlands during winter - inferred from feather stable isotope values - induces residual effects that carry over and influence physiological traits relevant to fitness in black-tailed godwits Limosa limosa limosa on their northward migration. Overwintering males and females were segregated by wetland salinity in West Africa, with females mostly occupying freshwater wetlands. The use of these wetlands along a gradient of salinities was associated with differences in immune responsiveness to phytohaemagglutinin and sized-corrected body mass in godwits staging in southern Europe during northward migration - 3,000 km from the nonbreeding grounds - but in males only. These findings provide a window onto the processes by which wetland salinity can induce carry-over effects and can help predict how migratory species should respond to future climate-induced increases in salinity.

Metabolic plasticity for subcutaneous fat accumulation in a long-distance migratory bird traced by 2H2O.

The migrant black-tailed godwit (Limosa limosa) traditionally used natural wetlands in the Iberian Peninsula to prepare for migratory flights by feeding mainly in estuaries. In recent decades, this species has become increasingly dependent on rice fields, thereby relying on a plant-based diet for fuelling. Dietary fatty acids (FA) seem to be determinant to the composition of accumulated subcutaneous fat in migratory birds. It is still unclear whether metabolic plasticity allows for modification and/or synthesis of FA, contributing to a lipid profile that enables a successful migratory performance. Deuterated water was administered to captive black-tailed godwits submitted to two diets (fly larvae versus rice) and the incorporation of deuterium (2H) into subcutaneous triglycerides was analyzed by NMR. A recently developed localized biopsy method for sampling subcutaneous fat was employed with later successful release of all birds into the wild. The average chemical structure reflected mostly a mixture of saturated and monounsaturated 16- and 18-carbon FA, a profile frequently found in migrant birds. Significantly higher levels of polyunsaturated FA, as well as detectable levels of n-3 FA, were observed in fly-larvae-fed birds. Excess 2H-enrichments in FA revealed significantly higher rates of fractional de novo lipogenesis and FA desaturation capacity in rice-fed birds. This novel and non-lethal tracer method revealed the capacity of this species to alter its lipid metabolism to compensate for a poorer dietary lipid contribution. Because of its versatility, adapting this method to other scenarios and/or other migratory species is considered feasible and cost-effective.

Physiological, Morphological and Behavioural Responses of Self-Feeding Precocial Chicks Copying with Contrasting Levels of Water Salinity during Development.

Combined physiological and behavioural responses to salt loads during development have rarely been studied in air-breathing vertebrates able to inhabit hypersaline habitats, but they may be of particular importance in understanding, for example, the differences among species in patterns of habitat use or ontogenetic diet switches. Here, we compared the physiological and behavioural responses of self-feeding precocial chicks developed in contrasting levels of water salinity. The model species was the Black-winged Stilt (Himantopus himantopus) a precocial shorebird that breeds in a range of habitats from freshwater to hypersaline wetlands. Specifically, we compared resting metabolic rate (RMR), heat shock proteins (Hsp70), plasma ions, hematocrit, body mass, body size, growth rate and head-shaking behaviour of captive-reared Black-winged Stilt fledglings developed under fresh (0 ‰), saline (20 ‰), and hypersaline (60 ‰) water. Contrary to expectations, none of the physiological and morphological variables measured differed significantly among treatments. Likewise, the RMR of wild and captive-reared fledglings was similar. Surprisingly, the saltgland mass of wild fledglings from freshwater and those from hypersaline habitats was also similar. However, head-shaking, a behavioural response associated to minimize salt intake and to expel the secretions of salt glands, differed according to salinity source: head-shaking rate increased with increasing salinity. The results of this study support the key role of behavioural osmoregulation in avoiding salt stress during development.

First record of Babesia sp. in Antarctic penguins.

This is the first reported case of Babesia sp. in Antarctic penguins, specifically a population of Chinstrap penguins (Pygoscelis antarctica) in the Vapour Col penguin rookery in Deception Island, South Shetlands, Antarctica. We collected peripheral blood from 50 adult and 30 chick Chinstrap penguins. Examination of the samples by microscopy showed intraerythrocytic forms morphologically similar to other avian Babesia species in 12 Chinstrap penguin adults and seven chicks. The estimated parasitaemias ranged from 0.25×10(-2)% to 0.75×10(-2)%. Despite the low number of parasites found in blood smears, semi-nested PCR assays yielded a 274 bp fragment in 12 of the 19 positive blood samples found by microscopy. Sequencing revealed that the fragment was 97% similar to Babesia sp. 18S rRNA from Australian Little Penguins (Eudyptula minor) confirming presence of the parasite. Parasite prevalence estimated by microscopy in adults and chicks was higher (24% vs. 23.3%, respectively) than found by semi-nested PCR (16% vs. 13.3% respectively). Although sampled penguins were apparently healthy, the effect of Babesia infection in these penguins is unknown. The identification of Babesia sp. in Antarctic penguins is an important finding. Ixodes uriae, as the only tick species present in the Antarctic Peninsula, is the key to understanding the natural history of this parasite. Future work should address the transmission dynamics and pathogenicity of Babesia sp. in Chinstrap penguin as well as in other penguin species, such as Gentoo penguin (Pygoscelis papua) and Adélie penguin (Pygoscelis adeliae), present within the tick distribution range in the Antarctic Peninsula.

Latitudinal-Related Variation in Wintering Population Trends of Greylag Geese (Anser Anser) along the Atlantic Flyway: A Response to Climate Change?

The unusually high quality of census data for large waterbirds in Europe facilitates the study of how population change varies across a broad geographical range and relates to global change. The wintering population of the greylag goose Anser anser in the Atlantic flyway spanning between Sweden and Spain has increased from 120 000 to 610 000 individuals over the past three decades, and expanded its wintering range northwards. Although population sizes recorded in January have increased in all seven countries in the wintering range, we found a pronounced northwards latitudinal effect in which the rate of increase is higher at greater latitudes, causing a constant shift in the centre of gravity for the spatial distribution of wintering geese. Local winter temperatures have a strong influence on goose numbers but in a manner that is also dependent on latitude, with the partial effect of temperature (while controlling for the increasing population trend between years) being negative at the south end and positive at the north end of the flyway. Contrary to assumptions in the literature, the expansion of crops exploited by greylag geese has made little contribution to the increases in population size. Only in one case (expansion of winter cereals in Denmark) did we find evidence of an effect of changing land use. The expanding and shifting greylag population is likely to have increasing impacts on habitats in northern Europe during the course of this century.

When Siberia came to the Netherlands: the response of continental black-tailed godwits to a rare spring weather event.

1. Extreme weather events have the potential to alter both short- and long-term population dynamics as well as community- and ecosystem-level function. Such events are rare and stochastic, making it difficult to fully document how organisms respond to them and predict the repercussions of similar events in the future. 2. To improve our understanding of the mechanisms by which short-term events can incur long-term consequences, we documented the behavioural responses and fitness consequences for a long-distance migratory bird, the continental black-tailed godwit Limosa limosa limosa, resulting from a spring snowstorm and three-week period of record low temperatures. 3. The event caused measurable responses at three spatial scales - continental, regional and local - including migratory delays (+19 days), reverse migrations (>90 km), elevated metabolic costs (+8·8% maintenance metabolic rate) and increased foraging rates (+37%). 4. There were few long-term fitness consequences, however, and subsequent breeding seasons instead witnessed high levels of reproductive success and little evidence of carry-over effects. 5. This suggests that populations with continued access to food, behavioural flexibility and time to dissipate the costs of the event can likely withstand the consequences of an extreme weather event. For populations constrained in one of these respects, though, extreme events may entail extreme ecological consequences.

How salinity and temperature combine to affect physiological state and performance in red knots with contrasting non-breeding environments.

Migratory shorebirds inhabit environments that may yield contrasting salinity-temperature regimes-with widely varying osmoregulatory demands, even within a given species-and the question is: by which physiological means and at which organisational level do they show adjustments with respect to these demands? Red knots Calidris canutus winter in coastal areas over a range of latitudes. The nominal subspecies winters in salty areas in the tropics, whereas the subspecies Calidris canutus islandica winters in north-temperate regions of comparatively lower salinities and temperatures. In this study, both subspecies of red knot were acclimated to different salinity (28/40‰)-temperature (5/35 °C) combinations for 2-week periods. We then measured food/salt intakes, basal metabolic rate (BMR), body mass and temperature, fat and salt gland scores, gizzard mass, heat-shock proteins, heterophils/lymphocytes (H/L) ratio and plasma Na(+) to assess the responses of each taxon to osmoregulatory challenges. High salinity (HS)-warm-acclimated birds reduced food/salt intake, BMR, body mass, fat score and gizzard mass, showing that salt/heat loads constrained energy acquisition rates. Higher salt gland scores in saltier treatments indicated that its size was adjusted to higher osmoregulatory demands. Elevated plasma Na(+) and H/L ratio in high-salinity-warm-acclimated birds indicated that salt/heat loads might have a direct effect on the water-salt balance and stress responses of red knots. Subspecies had little or no effect on most measured parameters, suggesting that most adjustments reflect phenotypic flexibility rather than subspecific adaptations. Our results demonstrate how salinity and temperature affect various phenotypic traits in a migrant shorebird, highlighting the importance of considering these factors jointly when evaluating the environmental tolerances of air-breathing marine taxa.

Immunoreactive cortisone in droppings reflect stress levels, diet and growth rate of gull-billed tern chicks.

Blood levels of corticosterone have been traditionally analyzed to assess stress levels in birds; however, measuring steroid hormone metabolites in feces and droppings has gained much interest as a noninvasive technique successfully used for such purposed in vertebrates. Diet may affect these fecal metabolite levels (e.g., due to nutritional stress), however, this variable has not been taken into account in studies with chicks despite the great dietary flexibility of many avian species. In this study, we addressed for the first time this key issue and validated the technique in wild gull-billed tern chicks (Gelochelidon nilotica). Several enzyme immunoassays were used to determine the most appropriate test to measure the stress response. Subsequently, we performed an experiment in captivity to assess adrenocortical activity in gull-billed tern chicks fed with two diets: piscivorous vs. insectivorous. Finally, the relation between the chicks' growth rate and excreted immunoreactive glucocorticoid metabolites (EGMs) was also evaluated. We found the immunoreactive cortisone metabolites to be a good index of stress (as being an index of adrenocortical reactivity) in chicks of this species. Fish-fed chicks had higher levels of cortisone metabolites when comparing both concentration and total daily excreted metabolites. Within each treatment diet, cortisone metabolite levels and growth rates were negatively correlated. These findings suggest that the diet should be considered when using this technique for comparative purposes and highlight the trade-off between stress levels and chicks growth rates.

Unravelling trophic subsidies of agroecosystems for biodiversity conservation: food consumption and nutrient recycling by waterbirds in Mediterranean rice fields.

Waterbirds can reallocate a considerable amount of nutrients within agricultural fields and between agriculture sites and wetlands. However their effects on biogeochemical cycles have rarely been quantified. We estimated bird numbers, diet (from stable isotope analysis), food supply, and the food consumption on rice fields by overwintering waterbirds in one of the most important areas for rice production in southwestern Europe and a key area for various migrating and resident waterbird species. Herein, we modelled the nutrient (N and P) recycling in rice fields, and their transport to reservoirs. The energy consumption by waterbirds (96,605±18,311 individuals) on rice fields during winter averaged at 89.9±39.0 kJ·m(-2), with its majority (89.9%) belonging to foraging on rice seeds. Thus, the birds removed about 26% of rice seeds leftover after harvest (estimated in 932.5±504.7 seeds·m(-2) in early winter) wherein common cranes and dabbling ducks (four species) were the most important consumers. Waterbirds foraging and roosting in the rice fields recycled more than 24.1 (1.0 kg·ha(-1)) of N and an additional 5.0 tons (0.2 kg·ha(-1)) of P in the Extremadura's rice fields during winter. Additionally, we estimated that 2.3 tons of N and 550 kg of P were removed from rice fields and transported to reservoirs. The seasonal foraging of wildlife should result in a direct benefit for rice farmers by improving nutrient recycling through defecation by waterbirds with respect to artificial fertilisation. Additionally, rice fields located in the cranes' core wintering areas can provide sufficient food supply to induce habitat shift from their traditional wintering habitat in 'dehesas' to rice fields, which causes indirect socioeconomic benefit through reduced acorn consumption by cranes. Our modelling approach may thus be especially helpful for management decisions regarding rice agroecosystems in areas which are also important for the conservation of migratory waterbirds.