Summary1. Dabbling ducks (Anatinae) are omnivorous birds that are widespread, numerous, highly mobile and often migratory, and therefore have great potential for (long distance) dispersal of other organisms, including plants. However, their ability to act as plant dispersal vectors has received little attention compared to frugivores and is often assumed to be relevant only for wetland species. 2. To evaluate the potential for plant dispersal by dabbling ducks, we collated and analysed existing data. We identified all plant species whose seeds have been recorded in the diets of the seven dabbling duck (Anas) species in the Western Palaearctic, as reported from gut content analyses. We then analysed the habitats and traits of these plant species to identify general patterns, and related these to data on gut passage survival and duck movements. 3. A large number of plant species (> 445 species of 189 genera and 57 families) have been recorded in the diet of dabbling ducks. These plant species represent a very wide range of habitats, including almost the full range of site fertility, moisture and light conditions, excluding only very dry and deeply shaded habitats. The ducks prefer seeds of intermediate sizes (1-10 mm 3 ), which have good chances to survive gut passage, but also ingest smaller and larger seeds. Ingested seeds represent a wide range of dispersal syndromes, including fleshy fruits. Many species (62%) were not previously considered animal-dispersed in plant data bases, and 66% were not identified as bird-dispersed. Rarefaction analyses suggest that our analysis still greatly underestimates the total number of plant species ingested. 4. Synthesis. Dabbling ducks do not exclusively ingest seeds of wetland plants, which make up only 40% of the ingested species. Rather, they feed opportunistically on a wide cross-section of plant species available across the landscapes they inhabit. Given the millions of ducks, the hundreds to thousands of seeds ingested per individual on a daily basis, and known gut passage survival rates, this results in vast numbers of seeds dispersed by ducks per day. Internal seed dispersal by dabbling ducks appears to be a major dispersal pathway for a far broader spectrum of plant species than previously considered.

During autumn–winter 2016–2017, highly pathogenic avian influenza A(H5N8) viruses caused mass die-offs among wild birds in the Netherlands. Among the ≈13,600 birds reported dead, most were tufted ducks (Aythya fuligula) and Eurasian wigeons (Anas penelope). Recurrence of avian influenza outbreaks might alter wild bird population dynamics.

Summary1. Dispersal via animals (zoochory) is a primary mechanism for seed exchange between habitat patches. Recent studies have established that many plant species can survive waterbird gut passage. To quantify the patterns and consequences of waterbird-mediated dispersal, information on ingestion and gut passage must be combined with bird movement data. Such analysis has recently revealed seed dispersal kernels by migrating waterbirds. However, since many waterbird populations are largely resident, and migrating populations spend only a minor part of the main dispersal season (autumn-winter) on active migration, daily regional-scale movements probably cause more frequent dispersal. 2. We synthesized high-resolution empirical data on landscape-scale movements and seed gut passage times in a key disperser species, the mallard (Anas platyrhynchos), using a spatially explicit, mechanistic model to quantify dispersal distributions resulting from daily autumn-winter movements. We evaluated how landscape composition and seed traits affect these dispersal patterns. 3. The model indicates that mallards generate highly clumped seed deposition patterns, dispersing seeds primarily between core areas used for foraging and resting. Approximately 34% of all dispersed seeds are transported to communal roost areas, which may function as reservoirs for mallarddispersed species, and 7% are transported between foraging areas. Landscape-dependent movement patterns strongly affect the dispersal distributions, resulting in multi-modal dispersal kernels, with dispersal distances increasing with fragmentation of freshwater foraging habitat. Seed size-related gut retention times determine the proportion of seeds being dispersed away from the ingestion area, with larger seeds (20 mm 3 ) having a 8-10% higher potential for long-distance dispersal than smaller seeds (0Á2 mm 3 ), if surviving gut passage. However, twice as many small seeds will finally accomplish long-distance dispersal due to their higher gut passage survival. 4. Synthesis. Firstly, this study reveals how seed dispersal patterns resulting from daily waterfowl movements are shaped by landscape-dependent differences in movement patterns. Secondly, seed survival appears more important than retention time in determining the scale of long-distance dispersal by non-migrating mallards. We conclude that the frequent flights of staging waterbirds result in directed dispersal over distances inversely related to wetland availability, indicating that they maintain landscape connectivity across a range from wet to increasingly dry landscapes.

Plant populations in fragmented ecosystems rely largely on internal dispersal by animals. To unravel the mechanisms underlying this mode of dispersal, an increasing number of experimental feeding studies is carried out. However, while physical activity is known to aff ect vertebrate digestive processes, almost all current knowledge on mechanisms of internal seed dispersal has been obtained from experiments with resting animals. We investigated how physical activity of the mallard Anas platyrhynchos , probably the quantitatively most important biotic dispersal agent in aquatic habitats in the entire Northern Hemisphere, aff ects gut passage survival and retention time of ingested plant seeds. We fed seeds of nine common wetland plants to mallards trained to subsequently swim for six hours in a fl ume tank at diff erent swimming speeds (activity levels). We compared gut passage survival and retention times of seeds against a control treatment with mallards resting in a conventional dry cage. Intact gut passage of seeds increased signifi cantly with mallard activity (up to 80% in the fastest swimming treatment compared to the control), identifying reduced digestive effi ciency due to increased metabolic rates as a mechanism enhancing the dispersal potential of ingested seeds. Gut passage speed was modestly accelerated (13% on average) by increased mallard activity, an eff ect partly obscured by the interaction between seed retention time and probability of digestion. Gut passage acceleration will be more pronounced in digestion-resilient seed species, thereby modulating their dispersal distances. Our fi ndings imply that seed dispersal potential by mallards calculated from previous experiments with resting birds is highly underestimated, while dispersal distances may be overestimated for some plant species. Similar eff ects of physical activity on digestive effi ciency of mammals suggests that endozoochorous dispersal of plant seeds by vertebrates is more eff ective and plays a quantitatively more important ecological role in both terrestrial and aquatic ecosystems than previously thought.

IntroductionHighly pathogenic avian influenza (HPAI) viruses of subtype H5N8 were re-introduced into the Netherlands by late 2016, after detections in south-east Asia and Russia. This second H5N8 wave resulted in a large number of outbreaks in poultry farms and the deaths of large numbers of wild birds in multiple European countries. Methods: Here we report on the detection of HPAI H5N8 virus in 57 wild birds of 12 species sampled during active (32/5,167) and passive (25/36) surveillance activities, i.e. in healthy and dead animals respectively, in the Netherlands between 8 November 2016 and 31 March 2017. Moreover, we further investigate the experimental approach of wild bird serology as a contributing tool in HPAI outbreak investigations. Results: In contrast to the first H5N8 wave, local virus amplification with associated wild bird mortality has occurred in the Netherlands in 2016/17, with evidence for occasional gene exchange with low pathogenic avian influenza (LPAI) viruses. Discussion: These apparent differences between outbreaks and the continuing detections of HPAI viruses in Europe are a cause of concern. With the current circulation of zoonotic HPAI and LPAI virus strains in Asia, increased understanding of the drivers responsible for the global spread of Asian poultry viruses via wild birds is needed.

Field studies have shown that waterbirds, especially members of the Anatidae family, are major vectors of dispersal by endozoochory for a broad range of plants lacking a fleshy fruit, yet whose propagules can survive gut passage. Widely adopted dispersal syndromes ignore this dispersal mechanism, and we currently have little understanding of what traits determine the potential of angiosperms for endozoochory by waterbirds. Results from previous experimental studies have been inconsistent as to how seed traits affect seed survival and retention time in the gut and have failed to control for the influence of plant phylogeny. Using 13 angiosperm species from aquatic and terrestrial habitats representing nine families, we examined the effects of seed size, shape, and hardness on the proportion of seeds surviving gut passage through mallards (Anas platyrhynchos) and their retention time within the gut. We compiled a molecular phylogeny for these species and controlled for the nonindependence of taxa due to common descent in our analyses. Intact seeds from all 13 species were egested, but seed survival was strongly determined by phylogeny and by partial effects of seed mass and hardness (wet load): species with seeds harder than expected from their size, and smaller than expected from their loading, had greater survival. Once phylogeny was controlled for, a positive partial effect of seed roundness on seed survival was also revealed. Species with seeds harder than expected from their size had a longer mean retention time, a result retained after controlling for phylogeny. Our study is the first to demonstrate that seed shape and phylogeny are important predictors of seed survival in the avian gut. Our results demonstrate that the importance of controlling simultaneously for multiple traits and relating single traits (e.g., seed size) alone to seed survival or retention time is not a reliable way to detect important patterns, especially when phylogenetic effects are ignored.

BackgroundMovement behaviour is fundamental to the ecology of animals and their interactions with other organisms, and as such contributes to ecosystem dynamics. Waterfowl are key players in ecological processes in wetlands and surrounding habitats through predator-prey interactions and their transportation of nutrients and other organisms. Understanding the drivers of their movement behaviour is crucial to predict how environmental changes affect their role in ecosystem functioning. Mallards (Anas platyrhynchos) are the most abundant duck species worldwide and important dispersers of aquatic invertebrates, plants and pathogens like avian influenza viruses. By GPS tracking of 97 mallards in four landscape types along a gradient of wetland availability, we identified patterns in their daily movement behaviour and quantified potential effects of weather conditions and water availability on the spatial scale of their movements.ResultsWe demonstrate that mallard movement patterns were highly predictable, with regular commuting flights at dusk and dawn between a fixed day roost and one or several fixed nocturnal foraging sites, linked strongly to surface water. Wind and precipitation hardly affected movement, but flight distances and home range sizes increased when temperatures dropped towards zero. Flight distances and home range sizes increased exponentially with decreasing availability of freshwater habitat. Total shoreline length and the number of water bodies in the landscape surrounding the roost were the best predictors of the spatial scale of daily mallard movements.ConclusionsOur results show how mallards may flexibly adjust the spatial scale of their movements to wetland availability in the landscape. This implies that mallards moving between discrete habitat patches continue to preserve biotic connectivity in increasingly fragmented landscapes. The high predictability of mallard movement behaviour in relation to landscape features makes them reliable dispersal vectors for organisms to adapt to, and allows prediction of their ecological role in other landscapes.Electronic supplementary materialThe online version of this article (doi:10.1186/s40462-016-0092-7) contains supplementary material, which is available to authorized users.

Waterbirds disperse a wide range of plant seeds via their guts, promoting biotic connectivity between isolated habitat patches. However, the intensity of digestive forces encountered by seeds, and therefore their potential to survive digestive tract passage, varies within and between waterbird species. Here, we investigate under controlled conditions how the interaction between seed traits and digestive strategies affect the germinability of seeds following waterbird-mediated dispersal. We exposed seeds of 30 wetland plant species to the main digestive processes in the dabbling duck digestive system: mechanical, chemical and intestinal digestion. These were simulated by 1) a pressure test and scarification treatment, 2) incubation in simulated gastric juice, and 3) incubation in intestinal contents of culled mallards (Anas platyrhynchos). We evaluated their separate and combined effects on seed germination, and identified the role of seed size and seed coat traits in resisting the digestive forces. Seeds were generally resistant to separate digestive processes, but highly sensitive to a combination. Resistance to mechanical break-down was reduced by up to 80% by chemical pre-treatment, especially for seeds with permeable coats. Scarified seeds were 12–17% more vulnerable to chemical and intestinal digestive processes than undamaged seeds. Large seeds and seeds with thin, permeable coats were particularly sensitive to chemical and intestinal digestion. These results indicate that efficient digestion of seeds requires multiple digestive processes. The gizzard, responsible for mechanical digestion, plays a key role in seed survival. Omnivorous birds, which have relatively light gizzards compared to pure herbivores or granivores, are thus most likely to disperse seeds successfully. Regardless of digestive strategy, small seeds with tough seed coats are most resistant to digestion and may be adapted to endozoochorous dispersal by waterbirds.