The cost of movement: assessing energy expenditure in a long-distant ectothermic migrant under climate change.

Migration is an energetically taxing phenomenon as animals move across vast, heterogeneous landscapes where the cost of transport is impacted by permissible ambient conditions. In this study, we assessed the energetic demands of long-distance migration in a multigenerational ectothermic migrant, the monarch butterfly (Danaus plexippus). We tested the hypotheses that temperature-dependent physiological processes reduce energy reserves faster during migration than previously estimated, and that increasing climatic temperatures resulting from the climate crisis will intensify baseline daily energy expenditure. First, we reared monarchs under laboratory conditions to assess energy and mass conversion from fifth instar to adult stages, as a baseline for migratory adult mass and ontogenetic shifts in metabolic rate from larvae to adult. Then, using historical tag-recapture data, we estimated the movement propensity and migratory pace of autumn migrants using computer simulations and subsequently calculated energy expenditure. Finally, we estimated the energy use of monarchs based on these tag-recapture data and used this information to estimate daily energy expenditure over a 57 year period. We found support for our two hypotheses, noting that incorporating standard metabolic rate into estimates of migratory energy expenditure shows higher energy demand and that daily energy expenditure has been gradually increasing over time since 1961. Our study shows the deleterious energetic consequences under current climate change trajectories and highlights the importance of incorporating energetic estimates for understanding migration by small, ectothermic migrants.

Hidden Markov Models reveal a clear human footprint on the movements of highly mobile African wild dogs.

Large carnivores have experienced considerable range contraction, increasing the importance of movement across human-altered landscapes between small, isolated populations. African wild dogs (Lycaon pictus) are exceptionally wide-ranging, and recolonization is an important element of their persistence at broad scales. The competition-movement-connection hypothesis suggests that adaptations to move through areas that are unfavorable due to dominant competitors might promote the ability of subordinate competitors (like wild dogs) to move through areas that are unfavorable due to humans. Here, we used hidden Markov models to test how wild dog movements were affected by the Human Footprint Index in areas inside and outside of South Luangwa National Park. Movements were faster and more directed when outside the National Park, but slowed where the human footprint was stronger. Our results can be directly and quantitatively applied to connectivity planning, and we use them to identify ways to better understand differences between species in recent loss of connectivity.

So far, so good Similar fitness consequences and overall energetic costs for short and long-distance migrants in a seabird.

Although there is a consensus about the evolutionary drivers of animal migration, considerable work is necessary to identify the mechanisms that underlie the great variety of strategies observed in nature. The study of differential migration offers unique opportunities to identify such mechanisms and allows comparisons of the costs and benefits of migration. The purpose of this study was to compare the characteristics of short and long-distance migrations, and fitness consequences, in a long-lived seabird species. We combined demographic monitoring (survival, phenology, hatching success) of 58 Northern Gannets (Morus bassanus) breeding on Bonaventure Island (Canada) and biologging technology (Global Location Sensor or GLS loggers) to estimate activity and energy budgets during the non-breeding period for three different migration strategies: to the Gulf of Mexico (GM), southeast (SE) or northeast (NE) Atlantic coast of the U.S. Survival, timing of arrival at the colony and hatching success are similar for short (NE, SE) and long-distance (GM) migrants. Despite similar fitness consequences, we found, as expected, that the overall energetic cost of migration is higher for long-distance migrants, although the daily cost during migration was similar between strategies. In contrast, daily maintenance and thermoregulation costs were lower for GM migrants in winter, where sea-surface temperature of the GM is 4-7o C warmer than SE and NE. In addition, GM migrants tend to fly 30 min less per day in their wintering area than other migrants. Considering lower foraging effort and lower thermoregulation costs during winter for long-distance migrants, this suggests that the energetic benefits during the winter of foraging in the GM outweigh any negative consequences of the longer-distance migration. These results support the notion that the costs and benefits of short and long-distance migration is broadly equal on an annual basis, i.e. there are no apparent carry-over effects in this long-lived bird species, probably because of the favourable conditions in the furthest wintering area.

Do atmospheric events explain the arrival of an invasive ladybird (Harmonia axyridis) in the UK?

Species introduced outside their natural range threaten global biodiversity and despite greater awareness of invasive species risks at ports and airports, control measures in place only concern anthropogenic routes of dispersal. Here, we use the Harlequin ladybird, Harmonia axyridis, an invasive species which first established in the UK from continental Europe in 2004, to test whether records from 2004 and 2005 were associated with atmospheric events. We used the atmospheric- chemistry transport model SILAM to model the movement of this species from known distributions in continental Europe and tested whether the predicted atmospheric events were associated with the frequency of ladybird records in the UK. We show that the distribution of this species in the early years of its arrival does not provide substantial evidence for a purely anthropogenic introduction and show instead that atmospheric events can better explain this arrival event. Our results suggest that air flows which may assist dispersal over the English Channel are relatively frequent; ranging from once a week from Belgium and the Netherlands to 1-2 times a week from France over our study period. Given the frequency of these events, we demonstrate that atmospheric-assisted dispersal is a viable route for flying species to cross natural barriers.

Multi-method assessment of whale shark (Rhincodon typus) residency, distribution, and dispersal behavior at an aggregation site in the Red Sea.

Whale sharks (Rhincodon typus) are typically dispersed throughout their circumtropical range, but the species is also known to aggregate in specific coastal areas. Accurate site descriptions associated with these aggregations are essential for the conservation of R. typus, an Endangered species. Although aggregations have become valuable hubs for research, most site descriptions rely heavily on sightings data. In the present study, visual census, passive acoustic monitoring, and long range satellite telemetry were combined to track the movements of R. typus from Shib Habil, a reef-associated aggregation site in the Red Sea. An array of 63 receiver stations was used to record the presence of 84 acoustically tagged sharks (35 females, 37 males, 12 undetermined) from April 2010 to May 2016. Over the same period, identification photos were taken for 76 of these tagged individuals and 38 were fitted with satellite transmitters. In total of 37,461 acoustic detections, 210 visual encounters, and 33 satellite tracks were analyzed to describe the sharks' movement ecology. The results demonstrate that the aggregation is seasonal, mostly concentrated on the exposed side of Shib Habil, and seems to attract sharks of both sexes in roughly equal numbers. The combined methodologies also tracked 15 interannual homing-migrations, demonstrating that many sharks leave the area before returning in later years. When compared to acoustic studies from other aggregations, these results demonstrate that R. typus exhibits diverse, site-specific ecologies across its range. Sightings-independent data from acoustic telemetry and other sources are an effective means of validating more common visual surveys.

Cost of dispersal in a social mammal: body mass loss and increased stress.

Dispersal is a key process influencing the dynamics of socially and spatially structured populations. Dispersal success is determined by the state of individuals at emigration and the costs incurred after emigration. However, quantification of such costs is often difficult, due to logistical constraints of following wide-ranging individuals. We investigated the effects of dispersal on individual body mass and stress hormone levels in a cooperative breeder, the meerkat ( Suricata suricatta). We measured body mass and faecal glucocorticoid metabolite (fGCM) concentrations from 95 dispersing females in 65 coalitions through the entire dispersal process. Females that successfully settled lost body mass, while females that did not settle but returned to their natal group after a short period of time did not. Furthermore, dispersing females had higher fGCM levels than resident females, and this was especially pronounced during the later stages of dispersal. By adding information on the transient stage of dispersal and by comparing dispersers that successfully settled to dispersers that returned to their natal group, we expand on previous studies focusing on the earlier stages of dispersal. We propose that body mass and stress hormone levels are good indicators to investigate dispersal costs, as these traits often play an important role in mediating the effects of the environment on other life-history events and individual fitness.

Haematological traits co-vary with migratory status, altitude and energy expenditure: a phylogenetic, comparative analysis.

Aerobic capacity is assumed to be a main predictor of workload ability and haematocrit (Hct) and haemoglobin (Hb) have been suggested as key determinants of aerobic performance. Intraspecific studies have reported increases in Hct and Hb in response to increased workload. Furthermore, Hct and Hb vary markedly among individuals and throughout the annual cycle in free-living birds and it has been suggested that this variation reflects adaptive modulation of these traits to meet seasonal changes in energy demands. We used a comparative dataset of haematological traits, measures of metabolic rate (57 species), and life-history traits (160 species) to test several hypotheses for adaptive variation in haematology in relation to migration and altitude. We then extended these general ideas to test relationships between Hct and basal metabolic rate, daily energy expenditure and activity energy expenditure, using the 57 species that we have metabolic rate information for. We found that at the interspecific level, full migrants have higher Hct and Hb than partial migrants and non-migrants, and that altitude is positively correlated with Hb but not Hct. Hct is positively associated with activity energy expenditure (energy spent specifically on costly activities), suggesting that haematological traits could be adaptively modulated based on life-history traits and that Hct is a potential physiological mediator of energetic constraint.

Low energy expenditure and resting behaviour of humpback whale mother-calf pairs highlights conservation importance of sheltered breeding areas.

Understanding the behaviour of humpback whale mother-calf pairs and the acoustic environment on their breeding grounds is fundamental to assessing the biological and ecological requirements needed to ensure a successful migration and survival of calves. Therefore, on a breeding/resting ground, Exmouth Gulf, Western Australia, we used animal-borne DTAGs to quantify the fine-scale behaviour and energetic expenditure of humpback whale mothers and calves, while sound recorders measured the acoustic environment. We show that: (i) lactating humpback whales keep their energy expenditure low by devoting a significant amount of time to rest, and their use of energy, inferred from respiration rates, is ~half than that of adults on their foraging grounds; (ii) lactating females mainly rest while stationary at shallow depths within reach of the hull of commercial ships, thus increasing the potential for ship strike collisions; (iii) the soundscape is dominated by biological sources; and (iv) even moderate increases of noise from vessels will decrease the communication range of humpback whales. Planned commercial infrastructure in Exmouth Gulf will cause a substantial increase in shipping traffic with the risk of ship strikes and acoustic disturbance potentially compromising energy reserves for the southern migration of humpback whales.

Fin whale movements in the Gulf of California, Mexico, from satellite telemetry.

Fin whales (Balaenoptera physalus) have a global distribution, but the population inhabiting the Gulf of California (GoC) is thought to be geographically and genetically isolated. However, their distribution and movements are poorly known. The goal of this study was to describe fin whale movements for the first time from 11 Argos satellite tags deployed in the southwest GoC in March 2001. A Bayesian Switching State-Space Model was applied to obtain improved locations and to characterize movement behavior as either "area-restricted searching" (indicative of patch residence, ARS) or "transiting" (indicative of moving between patches). Model performance was assessed with convergence diagnostics and by examining the distribution of the deviance and the behavioral parameters from Markov Chain Monte Carlo models. ARS was the predominant mode behavior 83% of the time during both the cool (December-May) and warm seasons (June-November), with slower travel speeds (mean = 0.84 km/h) than during transiting mode (mean = 3.38 km/h). We suggest ARS mode indicates either foraging activities (year around) or reproductive activities during the winter (cool season). We tagged during the cool season, when the whales were located in the Loreto-La Paz Corridor in the southwestern GoC, close to the shoreline. As the season progressed, individuals moved northward to the Midriff Islands and the upper gulf for the warm season, much farther from shore. One tag lasted long enough to document a whale's return to Loreto the following cool season. One whale that was originally of undetermined sex, was tagged in the Bay of La Paz and was photographed 10 years later with a calf in the nearby San Jose Channel, suggesting seasonal site fidelity. The tagged whales moved along the western GoC to the upper gulf seasonally and did not transit to the eastern GoC south of the Midriff Islands. No tagged whales left the GoC, providing supporting evidence that these fin whales are a resident population.

A simple, cost-effective emitter for controlled release of fish pheromones: Development, testing, and application to management of the invasive sea lamprey.

Semiochemicals that elicit species-specific attraction or repulsion have proven useful in the management of terrestrial pests and hold considerable promise for control of nuisance aquatic species, particularly invasive fishes. Because aquatic ecosystems are typically large and open, use of a semiochemical to control a spatially dispersed invader will require the development of a cost-effective emitter that is easy to produce, environmentally benign, inexpensive, and controls the release of the semiochemical without altering its structure. We examined the release properties of five polymers, and chose polyethylene glycol (PEG) as the best alternative. In a series of laboratory and field experiments, we examined the response of the invasive sea lamprey to PEG, and to a partial sex pheromone emitted from PEG that has proven effective as a trap bait to capture migrating sea lamprey prior to spawning. Our findings confirm that the sea lamprey does not behaviorally respond to PEG, and that the attractant response to the pheromone component was conserved when emitted from PEG. Further, we deployed the pheromone-PEG emitters as trap bait during typical control operations in three Great Lakes tributaries, observing similar improvements in trap performance when compared to a previous study using mechanically pumped liquid pheromone. Finally, the polymer emitters tended to dissolve unevenly in high flow conditions. We demonstrate that housing the emitter stabilizes the dissolution rate at high water velocity. We conclude the performance characteristics of PEG emitters to achieve controlled-release of a semiochemical are sufficient to recommend its use in conservation and management activities related to native and invasive aquatic organisms.

The evolution of a rare mammalian trait - benefits and costs of male philopatry in proboscis bats.

While inbreeding avoidance is widely accepted as the major driver of female natal dispersal, the evolution of male philopatry is still poorly understood and discussed to be driven by male mating strategy, mate competition among male kin and kin cooperation. During a twelve-year study, we gathered detailed genetic and observational data of individually marked proboscis bats to assess the degree of male philopatry as well as its costs and benefits to improve the understanding of its evolution. Our results reveal several patrilines with simultaneous presence of closely related males and a small proportion of unrelated immigrant males in their colonies. Philopatric males benefit from avoiding the costs of immigration into foreign colonies through significantly longer tenure, better integration (i.e. frequent nocturnal presence in the colonies) and consequently significantly higher reproductive success compared to immigrant males. Finally, we illustrate that despite a high proportion of philopatric males in the groups, the number of closely related competing males is low. Thus, the hypothesised costs of mate competition among male kin seem to be low in promiscuous mammalian societies with unrelated females and a small degree of male immigration and are readily outweighed by the benefits of staying in the natal group.

Smartphone-Based Distributed Data Collection Enables Rapid Assessment of Shorebird Habitat Suitability.

Understanding and managing dynamic coastal landscapes for beach-dependent species requires biological and geological data across the range of relevant environments and habitats. It is difficult to acquire such information; data often have limited focus due to resource constraints, are collected by non-specialists, or lack observational uniformity. We developed an open-source smartphone application called iPlover that addresses these difficulties in collecting biogeomorphic information at piping plover (Charadrius melodus) nest sites on coastal beaches. This paper describes iPlover development and evaluates data quality and utility following two years of collection (n = 1799 data points over 1500 km of coast between Maine and North Carolina, USA). We found strong agreement between field user and expert assessments and high model skill when data were used for habitat suitability prediction. Methods used here to develop and deploy a distributed data collection system have broad applicability to interdisciplinary environmental monitoring and modeling.

Spatial Distribution of a Large Herbivore Community at Waterholes: An Assessment of Its Stability over Years in Hwange National Park, Zimbabwe.

The spatial structuring of populations or communities is an important driver of their functioning and their influence on ecosystems. Identifying the (in)stability of the spatial structure of populations is a first step towards understanding the underlying causes of these structures. Here we studied the relative importance of spatial vs. interannual variability in explaining the patterns of abundance of a large herbivore community (8 species) at waterholes in Hwange National Park (Zimbabwe). We analyzed census data collected over 13 years using multivariate methods. Our results showed that variability in the census data was mostly explained by the spatial structure of the community, as some waterholes had consistently greater herbivore abundance than others. Some temporal variability probably linked to Park-scale migration dependent on annual rainfall was noticeable, however. Once this was accounted for, little temporal variability remained to be explained, suggesting that other factors affecting herbivore abundance over time had a negligible effect at the scale of the study. The extent of spatial and temporal variability in census data was also measured for each species. This study could help in projecting the consequences of surface water management, and more generally presents a methodological framework to simultaneously address the relative importance of spatial vs. temporal effects in driving the distribution of organisms across landscapes.

Costs of migratory decisions: A comparison across eight white stork populations.

Annual migratory movements can range from a few tens to thousands of kilometers, creating unique energetic requirements for each specific species and journey. Even within the same species, migration costs can vary largely because of flexible, opportunistic life history strategies. We uncover the large extent of variation in the lifetime migratory decisions of young white storks originating from eight populations. Not only did juvenile storks differ in their geographically distinct wintering locations, their diverse migration patterns also affected the amount of energy individuals invested for locomotion during the first months of their life. Overwintering in areas with higher human population reduced the stork's overall energy expenditure because of shorter daily foraging trips, closer wintering grounds, or a complete suppression of migration. Because migrants can change ecological processes in several distinct communities simultaneously, understanding their life history decisions helps not only to protect migratory species but also to conserve stable ecosystems.

Guidelines for Using Movement Science to Inform Biodiversity Policy.

Substantial advances have been made in our understanding of the movement of species, including processes such as dispersal and migration. This knowledge has the potential to improve decisions about biodiversity policy and management, but it can be difficult for decision makers to readily access and integrate the growing body of movement science. This is, in part, due to a lack of synthesis of information that is sufficiently contextualized for a policy audience. Here, we identify key species movement concepts, including mechanisms, types, and moderators of movement, and review their relevance to (1) national biodiversity policies and strategies, (2) reserve planning and management, (3) threatened species protection and recovery, (4) impact and risk assessments, and (5) the prioritization of restoration actions. Based on the review, and considering recent developments in movement ecology, we provide a new framework that draws links between aspects of movement knowledge that are likely the most relevant to each biodiversity policy category. Our framework also shows that there is substantial opportunity for collaboration between researchers and government decision makers in the use of movement science to promote positive biodiversity outcomes.

The cost of migration: spoonbills suffer higher mortality during trans-Saharan spring migrations only.

Explanations for the wide variety of seasonal migration patterns of animals all carry the assumption that migration is costly and that this cost increases with migration distance. Although in some studies, the relationships between migration distance and breeding success or annual survival are established, none has investigated whether mortality during the actual migration increases with migration distance. Here, we compared seasonal survival between Eurasian spoonbills (Platalea leucorodia leucorodia) that breed in The Netherlands and migrate different distances (ca 1000, 2000 and 4500 km) to winter in France, Iberia and Mauritania, respectively. On the basis of resightings of individually marked birds throughout the year between 2005 and 2012, we show that summer, autumn and winter survival were very high and independent of migration distance, whereas mortality during spring migration was much higher (18%) for the birds that wintered in Mauritania, compared with those flying only as far as France (5%) or Iberia (6%). As such, this study is the first to show empirical evidence for increased mortality during some long migrations, likely driven by the presence of a physical barrier (the Sahara desert) in combination with suboptimal fuelling and unfavourable weather conditions en route.

Ecological and scaling analysis of the energy expenditure of rest, activity, flight, and evaporative water loss in Passeriformes and non-Passeriformes in relation to seasonal migrations and to the occupation of boreal stations in high and moderate latitudes.

A unified system of bioenergetic parameters that describe thermal regulation and energy metabolism in many passerine and non-passerine species has been developed. These parameters have been analyzed as functions of ambient temperature, and bioenergetic models for various species have been developed. The level of maximum food energy or maximal existence metabolism (MPE) is 1.3 times higher in passerines than in non-passerines, which is consistent with the ratio of their basal metabolic rates (BMR). The optimal ambient temperature for maximizing productive processes (e.g., reproduction, molting) is lower for passerines than for non passerines, which allows passerines to have higher production rates at moderate ambient temperatures. This difference in the optimal ambient temperature may explain the variation in bioenergetic parameters along latitudinal gradients, such as the well-known ecological rule of clutch size (or mass) increase in the more northerly passerine birds. The increased potential for productive energy output in the north may also allow birds to molt faster there. This phenomenon allows passerine birds to occupy a habitat that fluctuates widely in ambient temperature compared with non-passerine birds of similar size. Passerines have a more effective system for maintaining heat balance at both high and low temperatures. The high metabolism and small body sizes of passerines are consistent with omnivore development and with ecological plasticity. Among large passerines, the unfavorable ratio of MPE to BMR should decrease the energy that is available for productive processes. This consequence limits both the reproductive output and the development of long migration (particularly in Corvus corax). The hypothesis regarding BMR increase in passerines was suggested based on an aerodynamic analysis of the flight speed and the wing characteristics. This allometric analysis shows that the flight velocity is approximately 20% lower in Passeriformes than in non-Passeriformes, which is consistent with the inverted ratio of their BMR level. The regressions for the aerodynamic characteristics of wings show that passerines do not change the morphological characteristics of their wings to decrease velocity. Passerine birds prefer forest habitats. The size range of 5-150 g for birds in forest habitats is almost exclusively occupied by passerines because of their large energetic capability.

Selection and assessment of reference genes for quantitative PCR normalization in migratory locust Locusta migratoria (Orthoptera: Acrididae).

Locusta migratoria is a classic hemimetamorphosis insect and has caused widespread economic damage to crops as a migratory pest. Researches on the expression pattern of functional genes in L. migratoria have drawn focus in recent years, especially with the release of genome information. Real-time quantitative PCR is the most reproducible and sensitive approach for detecting transcript expression levels of target genes, but optimal internal standards are key factors for its accuracy and reliability. Therefore, it's necessary to provide a systematic stability assessment of internal control for well-performed tests of target gene expression profile. In this study, twelve candidate genes (Ach, Act, Cht2, EF1α, RPL32, Hsp70, Tub, RP49, SDH, GAPDH, 18S, and His) were analyzed with four statistical methods: the delta Ct approach, geNorm, Bestkeeper and NormFinder. The results from these analyses aimed to choose the best suitable reference gene across different experimental situations for gene profile study in L. migratoria. The result demonstrated that for different developmental stages, EF1α, Hsp70 and RPL32 exhibited the most stable expression status for all samples; EF1α and RPL32 were selected as the best reference genes for studies involving embryo and larvae stages, while SDH and RP49 were identified for adult stage. The best-ranked reference genes across different tissues are RPL32, Hsp70 and RP49. For abiotic treatments, the most appropriate genes we identified were as follows: Act and SDH for larvae subjected to different insecticides; RPL32 and Ach for larvae exposed to different temperature treatments; and Act and Ach for larvae suffering from starvation. The present report should facilitate future researches on gene expression in L. migratoria with accessibly optimal reference genes under different experimental contexts.

Bats on a budget: torpor-assisted migration saves time and energy.

Bats and birds must balance time and energy budgets during migration. Migrating bats face similar physiological challenges to birds, but nocturnality creates special challenges for bats, such as a conflict between travelling and refueling, which many birds avoid by feeding in daylight and flying at night. As endothermic animals, bats and birds alike must expend substantial amounts of energy to maintain high body temperatures. For migratory birds refueling at stopovers, remaining euthermic during inactive periods reduces the net refuelling rate, thereby prolonging stopover duration and delaying subsequent movement. We hypothesized that bats could mitigate similar ambient-temperature dependent costs by using a torpor-assisted migration strategy. We studied silver-haired bats Lasionycteris noctivagans during autumn migration using a combination of respirometry and temperature-sensitive radiotelemetry to estimate energy costs incurred under ambient temperature conditions, and the energy that bats saved by using torpor during daytime roosting periods. All bats, regardless of sex, age, or body condition used torpor at stopover and saved up to 91% of the energy they would have expended to remain euthermic. Furthermore, bats modulated use of torpor depending on ambient temperature. By adjusting the time spent torpid, bats achieved a rate of energy expenditure independent of the ambient temperature encountered at stopover. By lowering body temperature during inactive periods, fuel stores are spared, reducing the need for refuelling. Optimal migration models consider trade-offs between time and energy. Heterothermy provides a physiological strategy that allows bats to conserve energy without paying a time penalty as they migrate. Although uncommon, some avian lineages are known to use heterothermy, and current theoretical models of migration may not be appropriate for these groups. We propose that thermoregulatory strategies should be an important consideration of future migration studies of both bats and birds.

Key features of intertidal food webs that support migratory shorebirds.

The migratory shorebirds of the East Atlantic flyway land in huge numbers during a migratory stopover or wintering on the French Atlantic coast. The Brouage bare mudflat (Marennes-Oléron Bay, NE Atlantic) is one of the major stopover sites in France. The particular structure and function of a food web affects the efficiency of carbon transfer. The structure and functioning of the Brouage food web is crucial for the conservation of species landing within this area because it provides sufficient food, which allows shorebirds to reach the north of Europe where they nest. The aim of this study was to describe and understand which food web characteristics support nutritional needs of birds. Two food-web models were constructed, based on in situ measurements that were made in February 2008 (the presence of birds) and July 2008 (absence of birds). To complete the models, allometric relationships and additional data from the literature were used. The missing flow values of the food web models were estimated by Monte Carlo Markov Chain--Linear Inverse Modelling. The flow solutions obtained were used to calculate the ecological network analysis indices, which estimate the emergent properties of the functioning of a food-web. The total activities of the Brouage ecosystem in February and July are significantly different. The specialisation of the trophic links within the ecosystem does not appear to differ between the two models. In spite of a large export of carbon from the primary producer and detritus in winter, the higher recycling leads to a similar retention of carbon for the two seasons. It can be concluded that in February, the higher activity of the ecosystem coupled with a higher cycling and a mean internal organization, ensure the sufficient feeding of the migratory shorebirds.