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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.

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. 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. 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 mm³), 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. 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.

1. Dispersal is essential for species to survive the threats of habitat destruction and climate change. Combining descriptions of dispersal ability with those of landscape structure, the concept of functional connectivity has been popular for understanding and predicting species' spatial responses to environmental change. 2. Following recent advances, the functional connectivity concept is now able to move beyond landscape structure to consider more explicitly how other external factors such as climate and resources affect species movement. We argue that these factors, in addition to a consideration of the complete dispersal process, are critical for an accurate understanding of functional connectivity for plant species in response to environmental change. 3. We use recent advances in dispersal, landscape and molecular ecology to describe how a range of external factors can influence effective dispersal in plant species, and how the resulting functional connectivity can be assessed. 4. Synthesis. We define plant functional connectivity as the effective dispersal of propagules or pollen among habitat patches in a landscape. Plant functional connectivity is determined by a combination of landscape structure, interactions between plant, environment and dispersal vectors, and the successful establishment of individuals. We hope that this consolidation of recent research will help focus future connectivity research and conservation.

Northern peatlands store large amounts of carbon. Observations indicate that forests and peatlands in northern biomes can be alternative stable states for a range of landscape settings. Climatic and hydrological changes may reduce the resilience of peatlands and forests, induce persistent shifts between these states, and release the carbon stored in peatlands. Here,we present a dynamic simulation model constrained and validated by a wide set of observations to quantify how feedbacks in water and carbon cycling control resilience of both peatlands and forests in northern landscapes. Our results show that 34% of Europe (area) has a climate that can currently sustain existing rainwater-fed peatlands (raised bogs). However, raised bog initiation and restoration by water conservation measures after the original peat soil has disappeared is only possible in 10% of Europe where the climate allows raised bogs to initiate and outcompete forests. Moreover, in another 10% of Europe, existing raised bogs (concerning ∼20% of the European raised bogs) are already affected by ongoing climate change. Here, forests may overgrow peatlands, which could potentially release in the order of 4% (∼24 Pg carbon) of the European soil organic carbon pool. Our study demonstrates quantitatively that preserving and restoring peatlands requires looking beyond peatland-specific processes and taking into account wider landscape-scale feedbacks with forest ecosystems.

Riparian ecosystems along streams naturally harbour a high plant diversity with many increasingly endangered species. In our current heavily modified and fragmented catchments, many of these species are sensitive to dispersal limitation. Better understanding of riparian plant dispersal pathways is required to predict species (re-)colonization potential and improve success rates of stream and riparian zone conservation and restoration. Dispersal by water (hydrochory) is an important mechanism for longitudinal and lateral dispersal of riparian species. Crucially for recruitment potential, it also influences the elevation along the riparian hydrological gradient where seeds become deposited. Due to the complex interplay between abiotic and biotic factors, however, it remains unclear how exactly patterns in seed deposition are formed. We compared hydrochorous and non-hydrochorous seed deposition, and quantified patterns of seed deposition along the bare substrate of newly created stream riparian gradients. Water levels were monitored and seed deposition was measured with seed traps along the full range of riparian hydrological conditions (from permanently flooded to never flooded). Average seed numbers and species richness were significantly higher in flooded than in non-flooded seed traps (5.7 and 1.5 times higher, respectively). Community-weighted trait means indicated that typically water-dispersed seeds were more dominant in flooded than in non-flooded seed traps and gradually decreased in concentration from the channel to the upland. Moreover, highly buoyant seeds accumulated at the average water line, and clear elevational sorting of non-buoyant seeds occurred within the floodplain. These results establish a critical role of flooding in shaping patterns of seed deposition along the riparian gradient, delivering many seeds of typical riparian species to riparian zones and depositing them at species-specific elevations as influenced by seed traits, suggesting species-specific dispersal pathways. This shows that hydrochory likely has important consequences for riparian vegetation development and that flooding forms a key process for successful restoration.

Animal migration is highly sensitised to environmental cues, but plant dispersal is considered largely passive. The common dandelion, Taraxacum officinale , bears an intricate haired pappus facilitating flight. The pappus enables the formation of a separated vortex ring during flight; however, the pappus structure is not static but reversibly changes shape by closing in response to moisture. We hypothesised that this leads to changed dispersal properties in response to environmental conditions. Using wind tunnel experiments for flow visualisation, particle image velocimetry, and flight tests, we characterised the fluid mechanics effects of the pappus morphing. We also modelled dispersal to understand the impact of pappus morphing on diaspore distribution. Pappus morphing dramatically alters the fluid mechanics of diaspore flight. We found that when the pappus closes in moist conditions, the drag coefficient decreases and thus the falling velocity is greatly increased. Detachment of diaspores from the parent plant also substantially decreases. The change in detachment when the pappus closes increases dispersal distances by reducing diaspore release when wind speeds are low. We propose that moisture-dependent pappus-morphing is a form of informed dispersal allowing rapid responses to changing conditions.

1. In plants, long-distance dispersal (LDD) is a widespread phenomenon of great importance, especially in spatial dynamics such as in metapopulations, invasions and migration. Much has become known about the mechanisms underlying LDD by wind, but selective release mechanisms have received little attention. Recent papers call for particular effort in clarification of the seed release stage of the dispersal process, which is our aim here. 2. We studied non-random seed release at a range of spatial scales, from the individual plant to species' spatial dynamics. We did this by combining wind tunnel experiments, field trap data and simulation and analytical models, using two common wind-dispersed heathland plants (Calluna vulgaris and Erica cinerea) as study species. 3. In both species, seed release from the plant increases with increasing wind velocity. There is a minimum release threshold, above which the probability of seed release increases with increasing wind velocity. The drag caused by the wind is the motive force for seed release. 4. Simulations of seed dispersal with non-random seed release match measured dispersal kernels better, especially in the tails, than 'conventional' simulations with random seed release. Seed release during gusts gives the most realistic kernel. This result matches previous findings that seeds are released primarily at the start of gusts and during turbulent wind flow. 5. Dispersal kernels assuming non-random seed release had large impacts on analytically modelled population migration rates. Wavespeeds assuming seed release during gusts were, on average, more than double those assuming random seed release. Increases in wavespeeds under different seed release assumptions were due mostly to increases in the tail of the dispersal kernel. 6. Synthesis. This study shows how the small-scale process of a seed being released from the plant could affect the large-scale spatial dynamics of plant species in landscapes. The mechanism of non-random seed release, i.e., seed release during gusts, is an important mechanism for attaining LDD by wind. As such, non-random seed release influences landscape-scale species dynamics such as invasions and migration.

Background Plant dispersal is a critical factor driving ecological responses to global changes. Knowledge on the mechanisms of dispersal is rapidly advancing, but selective pressures responsible for the evolution of dispersal strategies remain elusive. Recent advances in animal movement ecology identified general strategies that may optimize efficiency in animal searches for food or habitat. Here we explore the potential for evolution of similar general movement strategies for plants. Methods We propose that seed dispersal in plants can be viewed as a strategic search for suitable habitat, where the probability of finding such locations has been optimized through evolution of appropriate dispersal kernels. Using model simulations, we demonstrate how dispersal strategies can optimize key dispersal trade-offs between finding habitat, avoiding kin competition, and colonizing new patches. These trade-offs depend strongly on the landscape, resulting in a tight link between optimal dispersal strategy and spatiotemporal habitat distribution. Results Our findings reveal that multi-scale seed dispersal strategies that combine a broad range of dispersal scales, including Lévy-like dispersal, are optimal across a wide range of dynamic and patchy landscapes. At the extremes, static and patchy landscapes select for dispersal strategies dominated by short distances, while uniform and highly unpredictable landscapes both select for dispersal strategies dominated by long distances. Conclusions By viewing plant seed dispersal as a strategic search for suitable habitat, we provide a reference framework for the analysis of plant dispersal data. Consideration of the entire dispersal kernel, including distances across the full range of scales, is key. This reference framework helps identify plant species’ dispersal strategies, the evolutionary forces determining these strategies and their ecological consequences, such as a potential mismatch between plant dispersal strategy and altered spatiotemporal habitat dynamics due to land use change. Our perspective opens up directions for future studies, including exploration of composite search behaviour and ‘informed searches’ in plant species with directed dispersal.

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 affect 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, affects 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 flume tank at different 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 significantly with mallard activity (up to 80% in the fastest swimming treatment compared to the control), identifying reduced digestive efficiency 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 effect 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 findings 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 effects of physical activity on digestive efficiency of mammals suggests that endozoochorous dispersal of plant seeds by vertebrates is more effective and plays a quantitatively more important ecological role in both terrestrial and aquatic ecosystems than previously thought.

Biodiversity in wetlands is threatened by habitat loss and fragmentation, of which agricultural activities often are a cause. Dispersal of plant seeds via wind and ditches (water) may contribute to connecting remnant wetland plant populations in modern agricultural landscapes, and help to maintain and restore biodiversity. We developed a spatially explicit model to assess the relative importance of dispersal by wind and dispersal by water through drainage ditches for two wetland plant species in agricultural landscapes: a typical wind disperser and a typical water-disperser. Simulation results show that the typical wind disperser had a much higher capability to disperse by wind (90th percentile <30 m) than the typical water-disperser (90th percentile <2 m). Surprisingly, the capability to disperse via water was similar for the two species: 90th percentile dispersal distances following a combination of wind and water dispersal were between approximately 100 and 1000 m. Dispersal by water transported more seeds over long distances for both species. The main determinants for dispersal distance by water were roughness of the ditch (determined by, for example, bank vegetation) and the presence of obstructions (for example, culverts). Density or direction of the ditch network did not seem to affect water dispersal distances substantially. From a biodiversity conservation perspective, it would be most useful if areas with suitable riparian wetland habitat were intersected with a network of shallow ditches with a high roughness promoting seed deposition. These areas should then be connected to other suitable areas by a few regularly cleaned ditches with no obstructions and low seed trapping probability.