Unlike exercising mammals, migratory birds fuel very high intensity exercise (e.g., flight) with fatty acids delivered from the adipose tissue to the working muscles by the circulatory system. Given the primary importance of fatty acids for fueling intense exercise, we discuss the likely limiting steps in lipid transport and oxidation for exercising birds and the ecological factors that affect the quality and quantity of fat stored in wild birds. Most stored lipids in migratory birds are comprised of three fatty acids (16:0, 18:1 and 18:2) even though migratory birds have diverse food habits. Diet selection and selective metabolism of lipids play important roles in determining the fatty acid composition of birds which, in turn, affects energetic performance during intense exercise. As such, migratory birds offer an intriguing model for studying the implications of lipid metabolism and obesity on exercise performance. We conclude with a discussion of the energetic costs of migratory flight and stopover in birds, and its implications for bird migration strategies. Migration poses distinct physiological challenges for birds. For example, long-distance migrants rely virtually entirely on stored energy and nutrients to fuel each flight, and then must rapidly restore the necessary energy and nutrients at stopover sites along their migration route. Solutions to the physiological challenges associated with alternating intense exercise without feeding and then intense feeding and refueling at stopover sites are often physiologically incompatible. Determining how birds overcome these physiological challenges requires understanding how the physiological capabilities of exercising and fasting birds relate to the ecological conditions encountered during migration. This review focuses on the nutritional and physiological ecology of birds during migration. We discuss the current model of how birds fuel the costs of migration and the likely limiting steps in lipid transport and oxidation for exercising birds, and how lipid metabolism in birds is different from that in other vertebrates. We then place these aspects of the biochemistry and metabolism of birds during migration within an ecological context. We review selected aspects of the nutritional ecology of birds during migration with an emphasis on how birds accumulate fat stores, how fatty acid composition of diet influences composition of fat stores, and the effect of fatty acid composition of fat # JOURNAL OF AVIAN BIOLOGY REVIEW Reviews provide an opportunity to summarize existing knowledge within ornithological research, especially in areas where rapid and significant advances are occurring. Reviews should be concise and should cite all key references. An abstract is required.

Summary Long-distance dispersal (LDD) is important in plants of dynamic and ephemeral habitats.For plants of dynamic wetland habitats, waterfowl are generally considered to be important LDD vectors. However, in comparison to the internal (endozoochorous) dispersal of terrestrial plants by birds, endozoochorous dispersal of wetland plants by waterfowl has received little attention. We quantified the capacity for endozoochorous dispersal of wetland plants by waterfowl and identified the mechanisms underlying successful dispersal, by comparing the dispersal capacities of a large number of wetland plant species. 2. We selected 23 common plant species from dynamic wetland habitats and measured their seed characteristics. We fed seeds of all species to mallards ( Anas platyrhynchos ), a common and highly omnivorous duck species, and quantified seed gut survival, gut passage speed and subsequent germination. We then used a simple model to calculate seed dispersal distances. 3. In total 21 of the 23 species can be dispersed by mallards, with intact seed retrieval and subsequent successful germination of up to 32% of the ingested seeds. The species that pass fastest through the digestive tract of the mallards are retrieved in the greatest numbers (up to 54%) and germinate best (up to 87%). These are the species with the smallest seeds. Seed coat thickness plays only a minor role in determining intact passage through the mallard gut, but determines if ingestion enhances or reduces germination in comparison to control seeds. 4. Model calculations estimate that whereas the largest seeds can hardly be dispersed by mallards, most seeds can be dispersed up to 780 km, and the smallest seeds up to 3000 km, by mallards during migration. 5. Synthesis . This study demonstrates the mechanism underlying successful endozoochorous dispersal of wetland plant seeds by mallards: small seed size promotes rapid, and hence intact and viable, passage through the mallard gut. Mallards can disperse wetland plant seeds of all but the largest-seeded species successfully in relatively large numbers (up to 32% of ingested seeds) over long distances (up to thousands of kilometres) and are therefore important dispersal vectors.

It is increasingly acknowledged that migratory birds, notably waterfowl, play a critical role in the maintenance and spread of influenza A viruses. In order to elucidate the epidemiology of influenza A viruses in their natural hosts, a better understanding of the pathological effects in these hosts is required. Here we report on the feeding and migratory performance of wild migratory Bewick's swans (Cygnus columbianus bewickii Yarrell) naturally infected with low-pathogenic avian influenza (LPAI) A viruses of subtypes H6N2 and H6N8. Using information on geolocation data collected from Global Positioning Systems fitted to neck-collars, we show that infected swans experienced delayed migration, leaving their wintering site more than a month after uninfected animals. This was correlated with infected birds travelling shorter distances and fuelling and feeding at reduced rates. The data suggest that LPAI virus infections in wild migratory birds may have higher clinical and ecological impacts than previously recognised.

Aim Patterns of high biodiversity among less mobile organisms throughout isolated locations suggest that passive dispersal importantly contributes to biodiversity. We examined the contribution of waterbirds to the dispersal of plant seeds and macroinvertebrates between aquatic wetlands. Birds are renowned vectors for seeds of terrestrial plants, but less is known about their role in more dispersal-dependent aquatic systems. We therefore performed a meta-analysis on bird-mediated endozoochorous dispersal of aquatic species.Location Our review included studies that collected data world-wide.Methods We analysed data from 81 peer-reviewed publications on endozoochorous dispersal of aquatic plant seeds and macroinvertebrates by waterbirds. ResultsIn total, 36% of 1581 waterbird droppings collected in the field contained one or more intact propagules, with macroinvertebrates found almost as frequently as plant seeds. Positive droppings contained on average 3.3 intact propagules, of which one-third were viable. In 728 trials from 17 published feeding experiments 24% of the ingested propagules were retrieved intact, with c. 6.5% both viable and intact. As many as 17 species of Anatidae and Rallidae were involved in the dispersal of at least 39 species of macroinvertebrates and seeds from 97 species of plants across a wide taxonomic range. Smaller propagules seemed less affected by digestion than larger ones. We provide a first quantitative model that can be used to estimate waterbird-mediated dispersal of propagules between wetlands. This model indicates that an average waterbird has the potential to disperse five viable propagules after flying more than 100 km, and one additional propagule after flying 300 km. Main conclusionsWe demonstrate that waterbirds have the potential to transport a wide variety of aquatic plants and animals over several hundreds of kilometres. High survival of propagules might be explained by propagule adaptations or by the digestive adaptations of birds, whereby energy absorption is thought to be maximized rather than assimilation efficiency. Our meta-analysis suggests that waterbirds might contribute significantly to wetland biodiversity around the world, despite several limitations to our current knowledge. We outline avenues for future research to address these knowledge gaps. KeywordsAnatidae, aquatic propagules, digestive physiology, long-distance dispersal, macroinvertebrates, plant seeds, Rallidae.2006). Species distributions often range across geographical barriers, such as deserts and oceans (Schabetsberger et al., 2009;Jocque et al., 2010), suggesting high species mobility in the landscape (Lester et al., 2007). However, for many species it is still unknown how, and how often, they disperse ª 2012 Blackwell Publishing Ltd

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We tested whether the spatial variation in resource depletion by Tundra Swans (Cygnus columbianus) foraging on belowground tubers of sago pondweed (Potamogeton pectinatus) was caused by differences in net energy intake rates. The variation in givingup densities within the confines of one lake was nearly eightfold, the giving-up density being positively related to water depth and, to a lesser extent, the silt content of the sediment. The swans' preference (measured as cumulative foraging pressure) was negatively related to these variables. We adjusted a model developed for diving birds to predict changes in the time allocation of foraging swans with changes in power requirements and harvest rate. First, we compared the behavior of free-living swans foraging in shallow and deep water, where they feed by head-dipping and up-ending, respectively. Up-ending swans had 1.3-2.1 times longer feeding times than head-dipping swans. This was contrary to our expectation, since the model predicted a decrease in feeding time with an increase in feeding power. However, up-ending swans also had 1.9 times longer trampling times than headdipping swans. The model predicted a strong positive correlation between trampling time and feeding time, and the longer trampling times may thus have masked any effect of an increase in feeding power. Heart rate measurements showed that trampling was the most energetically costly part of foraging. However, because the feeding time and trampling time changed concurrently, the rate of energy expenditure was only slightly higher in deep water (1.03-1.06 times). This is a conservative estimate since it does not take into account that the feeding costs of up-ending are possibly higher than that of head-dipping. Second, we compared captive swans foraging on sandy and clayey sediments. We found that the harvest rate on clayey sediment was only 0.6 times that on sandy sediment and that the power requirements for foraging were 1.2-1.4 times greater. Our results are in qualitative agreement with the hypothesis that the large spatial variation in giving-up densities was caused by differences in net rates of energy intake. This potentially has important implications for the prey dynamics, because plant regrowth has been shown to be related to the same habitat factors (water depth and sediment type).