Dispersal of aquatic and terrestrial organisms by waterbirds: A review of current knowledge and future priorities

1. We review progress in our understanding of the importance of waterbirds as dispersal vectors of other organisms, and identify priorities for further research. 2. Waterbirds are excellent for long-distance dispersal (LDD), whereas other vectors such as fish and mammals disperse similar propagules, but over shorter distances. Empirical studies of internal and external transport by waterbirds have shown that the former mechanism generally is more important. Internal transport is widely recognised for aquatic plants and aquatic invertebrates with resting eggs, but also is important for other organisms (e.g., terrestrial flowering plants not dispersed by frugivores, bryophytes, tardigrades, fish eggs). 3. Waterbird vectors also are important in terrestrial habitats, and provide connectivity across terrestrial-aquatic boundaries. There are important differences in the roles of different waterbird species, especially those using different habitats along the aquatic-terrestrial gradient. Early attempts to predict zoochory based on propagule morphology have been found wanting, and more research is needed into how the traits of vectors and vectored organisms (including life history, dormancy and growth traits) explain dispersal interactions. Experimental studies have focused on the potential of propagules to survive internal or external transport, and research into factors determining the establishment success of propagules after dispersal is lacking. 4. Recent spatially explicit models of seed dispersal by waterbirds should be expanded to include invertebrate dispersal, and to compare multiple bird species in the same landscape. Network approaches have been applied to plant-waterbird dispersal interactions, and these are needed for invertebrates. Genetic studies support effective LDD of plants and invertebrates along waterbird flyways, but there remains a lack of examples at a local scale. Next Generation Sequencing and genomics should be applied to waterbird-mediated dispersal across the landscape. More studies of biogeography, community ecology, or population genetics should integrate waterbird movements at the design stage. 5. Zoochory research has paid little attention to the dispersal of non-pathogenic microbes (both eukaryotic and prokaryotic). Nevertheless, there is evidence that dispersal via avian guts can be central to the connectivity of aquatic microbial metacommunities. More work on microbial dispersal by waterbirds should explore its implications for biogeochemistry, and the interchange with gut flora of other aquatic organisms. In the Anthropocene, the role of migratory waterbirds in LDD of plants and other organisms is particularly important, for example in compensating for loss of large migratory mammals and fish, allowing native species to adjust their distributions under global warming, and spreading alien species along flyways after their initial introductions by human vectors. Recent technological advances have opened exciting opportunities that should be fully exploited to further our understanding of dispersal by waterbirds.