Experimental assessment of plant seed retention times in fur of cattle and horse

Epizoochorous dispersal of plant seeds is an important long-distance dispersal mechanism. Yet little is known about retention times of seeds in animal furs and hence about potential dispersal distances of the seeds. Here, we used marked seeds of 12 plant species to determine seed depletion curves on Galloway cattle and Haflinger horse in three vegetation types (forest, tall herbage vegetation and meadow), in both dry and rainy weather conditions. In the long fur of Galloway cattle, seeds were retained significantly longer than in the short fur of Haflinger horse. In general, seed retention times were not considerably affected by the structure of the surrounding vegetation. The impact of the weather was negligible, only affecting the retention of some plant species. Negative exponential functions were fitted to the seed depletion curves. Using the parameters of curve estimations in the different conditions of animal species and vegetation structure, half-life seed retention times of up to 13 h for Galloway cattle and up to more than 4 h for Haflinger horse could be calculated, with corresponding potential half-life dispersal distances in the order of magnitude of tens of metres to a few kilometres. Different seed traits correlated with seed retention times in the long cattle fur and in the short horse fur, respectively.     

Epizoochorous seed dispersal in relation to seed availability – an experiment with a red fox dummy

Questions: Is the red fox a potential vector for epizoochorous seed dispersal? Can seed attachment and retention be predicted from plant and seed traits? Location: Grasslands in southern Norway. Methods: Epizoochorous seed attachment on the red fox was studied by walking a dummy fox through the vegetation and comparing seeds found on the dummy with the estimated seed availability in the vegetation. Seed retention, i.e. the ability of different seeds to stay on the fox, was estimated in a separate experiment. Seed attachment and retention were related to plant and seed traits using statistical models that account for heteroscedasticity and zero‐inflated data. Results: The majority of seeds attached to the fox originated from a few species, but also species without specific seed traits that are supposed to enhance epizoochory attached at least some seeds to the fox. The probability of seed attachment was positively related to plant height, bristle and hooked seed appendages, and negatively related to winged appendages, seed mass, and seed sphericity. Seed retention was positively related to the seed traits bristles, hooks and pappus. For several species, the results indicate a high potential for dispersal over long distances. Conclusions: In modern agricultural landscapes, large herbivores are often restricted in their mobility or are found at low densities, and other animal vectors may therefore be important for seed dispersal. In our study, a range of plant species were able to disperse by attaching seeds to, and having their seeds retained in, the fox fur some distance. We suggest that the red fox may be an important vector for epizoochorous seed dispersal in the agricultural landscape.     

Large herbivores as mobile links between isolated nature reserves through adhesive seed dispersal

Question: Do large herbivores contribute to the dispersal of plant seeds between isolated habitats by epizoochory? Location: Nature reserves in Flanders, Belgium. Methods: Epizoochory was studied by brushing plant seeds from the fur of 201 domesticated large herbivores (Galloway cattle, donkeys and horses), grazing in 27 Flemish nature reserves. Several herbivores were examined after transport between different nature reserves as part of the seasonal grazing system in Flanders, allowing detection of seed dispersal both within and between reserves. The seedling emergence method was used to identify the dispersed plant species. Results: In total, 6385 epizoochorous seeds from 75 species germinated, yet the real seed quantity was underestimated by the seedling emergence method. A wide variety of seed morphology, seed weights and plant heights was represented among the dispersed species, many of which had a transient seed bank. There was a gradual turnover in epizoochorous species composition in the course of the vegetation season, and seed dispersal occurred both within and between different nature reserves. Conclusions: Domesticated large herbivores, as models for wild mammals in the present and the past, are important dispersers of many plant species. Through seasonal grazing, the herbivores function as ‘mobile link organisms’, connecting isolated nature reserves through seed dispersal, possibly influencing vegetation development and long‐term survival of plant populations. As such, large herbivores are important instruments in ecological restoration, especially in fragmented ecosystems.     

Seed dispersal by ungulates as an ecological filter: a trait‐based meta‐analysis

Plant communities are often dispersal‐limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. We predicted that seed dispersal by ungulates acts as an ecological filter – which differentially affects individuals according to their characteristics and shapes species assemblages – and that the filter varies according to the dispersal mechanism (endozoochory, fur‐epizoochory and hoof‐epizoochory). We conducted two‐step individual participant data meta‐analyses of 52 studies on plant dispersal by ungulates in fragmented landscapes, comparing eight plant traits and two habitat indicators between dispersed and non‐dispersed plants. We found that ungulates dispersed at least 44% of the available plant species. Moreover, some plant traits and habitat indicators increased the likelihood for plant of being dispersed. Persistent or nitrophilous plant species from open habitats or bearing dry or elongated diaspores were more likely to be dispersed by ungulates, whatever the dispersal mechanism. In addition, endozoochory was more likely for diaspores bearing elongated appendages whereas epizoochory was more likely for diaspores released relatively high in vegetation. Hoof‐epizoochory was more likely for light diaspores without hooked appendages. Fur‐epizoochory was more likely for diaspores with appendages, particularly elongated or hooked ones. We thus observed a gradient of filtering effect among the three dispersal mechanisms. Endozoochory had an effect of rather weak intensity (impacting six plant characteristics with variations between ungulate‐dispersed and non‐dispersed plant species mostly below 25%), whereas hoof‐epizoochory had a stronger effect (eight characteristics included five ones with above 75% variation), and fur‐epizoochory an even stronger one (nine characteristics included six ones with above 75% variation). Our results demonstrate that seed dispersal by ungulates is an ecological filter whose intensity varies according to the dispersal mechanism considered. Ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales. Synthesis Plant communities are often dispersal‐limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. Our analysis is the first synthesis of ungulate seed dispersal that compares characteristics from both non‐dispersed and dispersed diaspores, distinguishing the three zoochory mechanisms ungulates are involved in: endozoochory, hoof‐epizoochory and fur‐epizoochory. We confirmed that seed dispersal by ungulates is an ecological filter whose intensity increases from endozoochory, then hoof‐epizoochory to finally fur‐epizoochory. By filtering seed traits through dispersal, ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales.     

Herbivore-induced expansion of <Emphasis Type="Italic">Helianthemum nummularium</Emphasis> in grassland–scrub mosaic vegetation: circumstantial evidence for zoochory and indirect grazing impact

Extensive grazing often has a strong influence on the structure and composition of herbaceous plant communities with increasing population sizes for some species and decreasing presence in others. Herbivores affect plant communities directly by selective grazing of plant species, and indirectly by either epizoochory or endozoochory. Helianthemum nummularium is considered an increasing species because its distribution increased after the introduction of large, free-ranging grazers in at least two coastal dune grassland areas in Belgium. However, its seeds lack any obvious adaptations for epizoochory, and direct observations of plant/seed consumption are scarce. Through field and lab experiments, we assessed the dispersal ability of H. nummularium via endozoochory and epizoochory. In a differentiated grazer exclusion experiment, evidence was found that plants are grazed by large domestic ungulates and small wild herbivores although these incidences were rare. Direct endozoochory evidence remained scarce. No seeds were found germinating in field-collected dung, and only few seedlings emerged following a seed feeding experiment. However, once deposited, we found higher growth rates when seeds were mixed with dung and decreased establishment success when seeds were sown in combination with competitively superior species. Epizoochory was plausible because both fur and hooves of cattle and horses were potentially capable of contributing to the transport of H. nummularium seeds. We conclude that herbivores play a role in seed dispersal, while their selective grazing behaviour most probably creates an appropriate environment for Helianthemum establishment and maintenance.     

Do plant-community-based grazing regimes lead to epizoochorous dispersal of high proportions of target species?

The contribution of large-herbivore epizoochory to the transfer of seeds within and between areas is thought to be significant. But often seeds of ubiquitous species are dispersed, which may enhance ruderalization processes. In order to study the dispersal of target species by sheep, we employed a community-based grazing approach followed by intra- and inter-area sheep transfers (maximum transfer distance 3 km). In case of inter-area transfers, well-developed target communities of an open inland sand ecosystem are used as “source”, linked to less-developed sand habitats (“sink”) via sheep. Also other factors determining which species become dispersed under field conditions were tested: seed surface structure, seed mass, plant seed-releasing height and animal behaviour. Finally the influence of animal movement on seed detachment and the actual arrival of seeds within a “sink” were studied.

Sheep transfers resulted in the dispersal of 56 seed species, dominated by Red List (seven species) and other target species. Quantitatively, most transported seeds belonged to target species, whereas graminoid competitors were highly under-represented. Morphological traits enhance the attachment probability regardless of seed mass. But for seeds without these epizoochory-facilitating traits, mass seems to affect attachment negatively. Plant height affected the number of species present in sheep coats but not the seed quantities. Probably certain species in the vegetation produced large numbers of seeds, e.g. low-growing Medicago minima with seed surface structures and high-growing species Verbascum phlomoides without seed surface structures. Also, although transfer half-times were three times lower than grazing half-times, naturally attached Stipa capillata and Agrimonia procera seeds showed no significant detachment rates during transfer, whereas considerable losses were found during grazing. Other 3-km sheep transfers did not result in significant losses of either epizoochorously transported seed or species quantities. Our study shows that community-based grazing can lead to the dispersal of especially target species. Besides that, different habitat fragments can be connected to each other via sheep without significant seed losses along the way.     

Complementarity of epi- and endozoochory of plant seeds by free ranging donkeys

Epizoochory and endozoochory are well-recognized long-distance seed dispersal mechanisms, yet their relative importance has hardly been studied before. Here, epi- and endozoochory were compared on donkeys foraging in a species-rich 100 ha coastal dune ecosystem, through in vitro germination of zoochorous material obtained by fur brushing and dung collection. We identified 6675 seedlings of 66 plant species, covering 20% of the species recorded in the study area. Of the 66 species, only 16 occurred in both epi- and endozoochory samples, demonstrating the complementarity of both dispersal mechanisms. The species composition in the zoochory samples reflected a strong seasonality, and seedling numbers were partly correlated with species abundance in the study area. The non-zoochorously dispersed species in the study area differed from the zoochorous species in seed size and weight, plant height, life span, dispersal strategy and seedbank persistence. Dispersal-relevant plant traits were used to derive dispersal-functional plant types for all species in the study area. Epizoochory showed to be more specific than endozoochory and was associated with a more narrow range of dispersal-functional plant types.     

Long-distance seed dispersal in plant populations

Long-distance seed dispersal influences many key aspects of the biology of plants, including spread of invasive species, metapopulation dynamics, and diversity and dynamics in plant communities. However, because long-distance seed dispersal is inherently hard to measure, there are few data sets that characterize the tails of seed dispersal curves. This paper is structured around two lines of argument. First, we argue that long-distance seed dispersal is of critical importance and, hence, that we must collect better data from the tails of seed dispersal curves. To make the case for the importance of long-distance seed dispersal, we review existing data and models of long-distance seed dispersal, focusing on situations in which seeds that travel long distances have a critical impact (colonization of islands, Holocene migrations, response to global change, metapopulation biology). Second, we argue that genetic methods provide a broadly applicable way to monitor long-distance seed dispersal; to place this argument in context, we review genetic estimates of plant migration rates. At present, several promising genetic approaches for estimating long-distance seed dispersal are under active development, including assignment methods, likelihood methods, genealogical methods, and genealogical/demographic methods. We close the paper by discussing important but as yet largely unexplored areas for future research.     

Seed dispersal by white-tailed deer: implications for long-distance dispersal,invasion, and migration of plants in eastern North America

For many plant species in eastern North America, short observed seed dispersal distances (ranging up to a few tens of meters) fail to explain rapid rates of invasion and migration. This discrepancy points to a substantial gap in our knowledge of the mechanisms by which seeds are dispersed long distances. We investigated the potential for white-tailed deer (Odocoileus virginianus Zimm.), the dominant large herbivore in much of eastern North America, to disperse seeds via endozoochory. This is the first comprehensive study of seed dispersal by white-tailed deer, despite a vast body of research on other aspects of their ecology. More than 70 plant species germinated from deer feces collected over a 1-year period in central New York State, USA. Viable seeds included native and alien herbs, shrubs, and trees, including several invasive introduced species, from the full range of habitat types in the local flora. A mean of >30 seeds germinated per fecal pellet group, and seeds were dispersed during all months of the year. A wide variety of presumed dispersal modes were represented (endo- and exozoochory, wind, ballistic, ant, and unassisted). The majority were species with small-seeded fruits having no obvious adaptations for dispersal, underscoring the difficulty of inferring dispersal ability from diaspore morphology. Due to their broad diet, wide-ranging movements, and relatively long gut retention times, white-tailed deer have tremendous potential for effecting long-distance seed dispersal via ingestion and defecation. We conclude that white-tailed deer represent a significant and previously unappreciated vector of seed dispersal across the North American landscape, probably contributing an important long-distance component to the seed shadows of hundreds of plant species, and providing a mechanism to help explain rapid rates of plant migration.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Distribution of myrmecochorous species over the landscape and their potential long-distance dispersal by emus and kangaroos

Topographical heterogeneity can create a mosaic of substrate types leading to the formation of isolated plant populations. Seed dispersal then becomes crucial for the colonization of such suitable but remote substrate types. We surveyed the distribution of seven elaiosome-bearing species (myrmecochores) over 5 km² of natural heathland in southwestern Australia. Ants are the standard means of dispersal of these species, which provide limited dispersal (usually of a few metres). Six species were associated with particular substrate types (dune or swale) and all occurred as discrete populations, on average 270–500 m apart, with closest dune edges 280 m apart. We evaluated the possible roles of emus and kangaroos as alternative agents of long-distance seed dispersal between substrate types. Their droppings contained viable seeds of three of the target species, as well as other myrmecochores, and were evenly distributed over the substrate types. While migration of these plant species between preferred substrate types seems unlikely when considering only their standard dispersal agents (ants), it is highly likely in the presence of emus (in particular) and kangaroos that act as non-standard dispersers. This may have important consequences for plant species conservation by increasing habitat connectivity and favouring regional persistence.     

Epizoochorous dispersal by ungulates depends on fur,grooming and social interactions

The transport phase of the animal‐mediated plant dispersal process is critical to dispersal effectiveness as it determines the spatial distribution of the diaspores released and their chance for further recruitment. Assessing this specific phase of the dispersal process generally requires combining diaspore retention times with the associated distances covered. Here, we specifically tested the effect of grooming behavior, interindividual contacts and ungulate fur on diaspore retention times and associated dispersal distances for the hooked diaspores of Xanthium strumarium L. experimentally attached to tamed individuals of three ungulate species. We used a comparative approach based on differing fur quality on different body zones of these three ungulates. During 6‐hr sessions, we monitored for grooming and social interactions that may induce intended or inadvertent diaspore detachment. Additionally, we proposed innovative approaches to directly assessing diaspore dispersal distances by red deer in situ. Fat‐tailed functions fitted diaspore retention time, highlighting the potential for long‐distance dispersal events. The longer the hair, the higher the retention capacity of diaspores in the animal's fur. As predicted, donkey retained diaspores longer than red deer and dwarf goat; and we also confirmed that diaspores attached to the short hair of the head fell off more quickly than did those on the other body zones. Dwarf goat groomed more often than both red deer and donkey, but also when it carried diaspores. Up to 14% of the diaspores detached from animal fur after specific grooming behavior. We observed, in controlled conditions, for the first time and for each ungulate species, interindividual transfers of diaspores, representing 5% of the diaspores attached to animals’ fur. Our results militate for incorporating animal behavior into plant dispersal modeling approaches.     

Standard yet unusual mechanisms of long-distance dispersal: seed dispersal of Corymbia torelliana by bees

Mellitochory, seed dispersal by bees, has been implicated in long-distance dispersal of the tropical rain forest tree, Corymbia torelliana (Myrtaceae). We examined natural and introduced populations of C. torelliana for 4 years to determine the species of bees that disperse seeds, and the extent and distance of seed dispersal. The mechanism of seed dispersal by bees was also investigated, including fruit traits that promote dispersal, foraging behaviour of bees at fruits, and the fate of seeds. The fruit structure of C. torelliana , with seed presented in a resin reward, is a unique trait that promotes seed dispersal by bees and often results in long-distance dispersal. We discovered that a guild of four species of stingless bees, Trigona carbonaria, T. clypearis, T. sapiens , and T. hockingsi, dispersed seeds of C. torelliana in its natural range. More than half of the nests found within 250 m of fruiting trees had evidence of seed transport. Seeds were transported minimum distances of 20–220 m by bees. Approximately 88% of seeds were dispersed by gravity but almost all fruits retained one or two seeds embedded in resin for bee dispersal. Bee foraging for resin peaked immediately after fruit opening and corresponded to a peak of seed dispersal at the hive. There were strong correlations between numbers of seeds brought in and taken out of each hive by bees ( r =  0.753–0.992, P  < 0.05), and germination rates were 95 ± 5%. These results showed that bee-transported seeds were effectively dispersed outside of the hive soon after release from fruits. Seed dispersal by bees is a non-standard dispersal mechanism for C. torelliana, as most seeds are dispersed by gravity before bees can enter fruits. However, many C. torelliana seeds are dispersed by bees, since seeds are retained in almost all fruits, and all of these are dispersed by bees.     

Soil seed banks near rubbing trees indicate dispersal of plant species into forests by wild boar

Current knowledge about processes that generate long-distance dispersal of plants is still limited despite its importance for persistence of populations and colonization of new potential habitats. Today wild large mammals are presumed to be important vectors for long-distance transport of diaspores within and between European temperate forest patches, and in particular wild boars recently came into focus. Here we use a specific habit of wild boar, i.e. wallowing in mud and subsequent rubbing against trees, to evaluate epizoochorous dispersal of vascular plant diaspores. We present soil seed bank data from 27 rubbing trees versus 27 control trees from seven forest areas in Germany. The mean number of viable seeds and the plant species number were higher in soil samples near rubbing trees compared with control trees. Ten of the 20 most frequent species were more frequent, and many species exclusively appeared in the soil samples near rubbing trees. The large number of plant species and seeds – more than 1000 per tree – in the soils near rubbing trees is difficult to explain unless the majority were dispersed by wild boar. Hooked and bristly diaspores, i.e. those adapted to epizoochory, were more frequent; however, many species with unspecialized diaspores occurred exclusively near rubbing trees. As opposed to plant species closely tied to forests species which occur both in forest and open vegetation and non-forest species were more frequent near rubbing trees compared with controls. These findings are consistent with previous studies on diaspore loads in the coats and hooves of shot wild boars. However, our method allows to identify the transport of diaspores from the open landscape into forest stands, where they might especially emerge after disturbance, and a clustered distribution of epizoochorically dispersed seeds. Moreover, accumulation of seeds of wetness indicators near rubbing trees demonstrates directed dispersal of plant species inhabiting wet places among remote wallows.     

Seed dispersal by woolly monkeys (Lagothrix lagothricha) at Tinigua National Park, Colombia: dispersal distance, germination rates, and dispersal quantity

The purpose of this study was to describe seed dispersal patterns of woolly monkeys (Lagothrix lagothricha) in terms of dispersal quantity and two factors related to dispersal quality: germination rates of dispersed seeds and the distance of dispersal to parental trees. The possible influence of retention time, travel distance, seed size, activity patterns, and fruit abundance on dispersal distance was also analyzed. Observations on activity, diet, daily movements, and seed dispersal were made on focal individuals of a group of woolly monkeys at a tropical rain forest in Tinigua National Park (Colombia). Sixty hours of focal samples per month were completed during 1 year. A total of 753 depositions were collected during the study. Each dropping contained seeds from an average of 2.68 different species (range 0 to 9). Collected depositions contained an underestimated total of 50,168 seeds (>1 mm). Given a population density of 30 individuals/km2, the woolly monkeys in the study area disperse more than 25,000 seeds/km2/day. These seeds belong to 112 different plant species. Germination rates of dispersed seeds are usually similar or higher than those of non-swallowed seeds. It was possible to determine dispersal distance in 264 cases when the focal animal was continuously followed from ingestion at the parental tree to deposition. Only 1% of these depositions landed in close proximity (<15 m) of the parental tree. It was very common that the droppings were deposited between 100 and 500 m from the parent tree, and up to 1.5 km. Higher retention times and longer travel distances were not correlated with increased dispersal distance. Two main reasons for this result were the prolonged and variable passage rates (avg=11.2 hr+/-6.5 SD.) and the circuitous routes of monkeys in this forest.     

Seed Dispersal Distances and Plant Migration Potential in Tropical East Asia

Most predictions of vegetation responses to anthropogenic climate change over the next 100 yr are based on plant physiological tolerances and do not account for the ability of plant species to migrate over the distances required in the time available, or the impact of habitat fragmentation on this ability. This review assesses the maximum routine dispersal distances achievable in tropical East Asia and their vulnerability to human impacts. Estimates for various plant–vector combinations range from < 10 m, for species dispersed by ants or mechanical means, to > 10 km for some species dispersed by wind (tiny seeds), water, fruit pigeons, large fruit bats (tiny seeds), elephants, rhinoceroses, and people. Most plant species probably have maximum dispersal distances in the 100–1000 m range, but the widespread, canopy-dominant Dipterocarpaceae and Fagaceae are normally dispersed < 100 m. Large fruit bats and fruit pigeons are particularly important for long-distance dispersal in fragmented landscapes and should be protected from hunting. The maximum seed dispersal distances estimated in this study are potentially sufficient for many plant species to track temperature changes in steep topography, but are far too small for a significant role in mitigating climate change impacts in the lowlands, where temperature and rainfall gradients are much more shallow.     

Differences in endozoochorous dispersal between aquatic plant species, with reference to plant population persistence in rivers

Summary 1. In river ecosystems, populations are continuously subjected to unidirectional downstream currents resulting in a downstream movement of populations. To ensure long-term population persistence in rivers, organisms must have a mechanism for upstream dispersal, which allows them to re-colonise upstream areas.
2. In this study we assessed differences in the potential for endozoochorous seed dispersal of Sparganium emersum and Sagittaria sagittifolia , two aquatic plant species with different seed morphologies, by mallard ( Anas platyrhynchos ) and teal ( Anas crecca ), two duck species with different body weights.
3. We found no significant differences in seed retrieval (the proportion of ingested seeds retrieved after gut passage) and seed retention time (time between seed ingestion and retrieval), between mallard and teal, despite the difference in body weights. We did find a significantly higher germination (%) over retention time of S. emersum seeds retrieved from teal compared with mallard, most likely related to a more efficient removal of the seed coat during passage through the gut of teal.
4. There were large differences between S. emersum versus S. sagittifolia in: (i) seed retrieval (22.65 ± 20.8% versus 1.60 ± 2.4%, respectively); (ii) seed retention time in duck gut, with a maximum of 60 h versus 12 h; (iii) the effect of gut passage on seed germination, with an increase of approximately 35% versus a decrease of 25%; and (iv) the effect of gut passage on seed germination rate, with an acceleration of 10 days versus a delay of 3 days on average. The results show that S. emersum has a higher potential for endozoochorous dispersal by ducks and postdispersal establishment than S. sagittifolia .
5. We propose that, in rivers, bird-mediated seed dispersal may promote re-colonisation of upstream areas, enabling long-term plant population persistence.     

First evidence of epizoochorous seed dispersal by golden snub-nosed monkeys (<Emphasis Type="Italic">Rhinopithecus roxellana</Emphasis>) in temperate forest

Although many primates are identified as effective endozoochorous seed dispersers in forest ecosystems, epizoochorous seed dispersal by primates has generally been overlooked. In this study, we report epizoochorous seed dispersal in a group of free-ranging golden snub-nosed monkeys (Rhinopithecus roxellana) residing in the temperate forests of central China by identifying seeds carried by 12 well-habituated individuals. The results showed a total of 1920 seeds (five families, seven genera) belonging to eight plant species attached to the monkeys’ fur; the three most abundant species were Geum aleppicum (54% of the total), Torilis japonica (17%), and Agrimonia pilosa (14%). The majority of seeds (95.3%) that attached to the monkeys’ fur had special morphologies such as hairs, hooks, or awns. We also found that the quantity of seeds attached to the fur was not significantly different by age-sex class, even though adult males have very long fur. Our study provides the first empirical evidence of epizoochorous seed dispersal by primates and suggests the role of primates in seed dispersal may be greater than previously assumed.     

Emus as non-standard seed dispersers and their potential for long-distance dispersal

Long-distance seed dispersal may have important consequences for species range, migration rates, metapopulation dynamics, and gene flow. Plants have evolved various adaptations for seed dispersal by standard agents, with typical dispersal distances associated with them. Seeds may also be dispersed by non-standard agents for which they do not show any apparent adaptation and may reach long distances. By sampling the droppings of emus Dromaius novaehollandiae at three localities in Western Australia, we investigated their potential to act as long-distance dispersers of seeds with adaptations for dispersal modes other than endozoochory, such as unassisted, ant, wind and exozoochory, for which they act as non-standard agents. Seventy-seven plant species with five types of dispersal syndromes were found in the 112 droppings analysed, with at least 68 having viable seeds. Although endozoochory was the most frequent syndrome, the presence of other syndromes was important in terms of number of species (61%) and seeds (50%). Estimates of species richness indicated that an increase in sampling effort would increase the number of species observed, especially among non-endozoochores. As a consequence of their long gut retention times and high mobility, emus can provide long-distance dispersal opportunities that may be especially relevant for species with dispersal modes of typically short distances (unassisted, ant).
Our results suggest that the role of emus as non-standard agents for long-distance dispersal should be taken into account for understanding current geographic ranges, gene flow and metapopulation dynamics of some plant species, as well as for predicting their future responses to climate change and fragmentation.     

Endozoochory largely outweighs epizoochory in migrating passerines

Fruits and seeds are critical food sources for many European passerines during the autumn migration, which in turn contribute to disperse seeds either internally, i.e. after ingestion (endozoochory), or externally, when seeds adhere to the body surface (epizoochory). Despite the recognized importance of birds as seed dispersers, the vast majority of studies focused on endozoochory while the external transport of seeds is frequently invoked as being potentially important, but remains largely unexplored. This is particularly important during the post‐breeding migration of passerines, the most ubiquitous and diverse movement of potential seed carriers across Europe and into Africa, which coincides with the fruiting peak of many plant species (August–October). Our aim was to evaluate the role of migrating birds as potential long‐distance seed dispersers, and comparing the prevalence of epizoochory and endozoochory during post‐breeding migration. We sampled 926 wild birds in nine locations in Portugal, and retrieved 1833 seeds of 19 plant species dispersed internally and only three seeds externally attached to three birds (Serinus serinus, Locustella naevia and Turdus merula), showing an endozoochory prevalence 85 times higher than that of epizoochory. Migrating and non‐migrating passerines dispersed seeds equally. While two of the seeds transported externally had specific adaptations to epizoochory, namely spines (Torilis arvensis) and hooks (Galium aparine), the third is a large seed from a fleshy‐fruited plant, Frangula alnus (i.e. typical endozoochorous syndrome). These seeds were found on bird species with different diets, but similar behaviour (ground foragers) and in similar habitats (open agro‐ecosystems). Our results highlight the strong role of migrating passerines as potentially long‐distance seed dispersers and show that, at least in the autumn, the prevalence of epizoochory is several orders of magnitude lower than that of endozoochory.     

Dispersal mechanisms of the narrow endemic <Emphasis Type="Italic">Polygala vayredae</Emphasis>: dispersal syndromes and spatio-temporal variations in ant dispersal assemblages

This study assesses the dispersal mechanisms of the narrow endemic Polygala vayredae, analysing the functioning of its dispersal syndromes (anemochory and myrmecochory), the spatio-temporal variability of the disperser assemblage, foraging behaviour and dispersal ability, and the role of the elaiosome in ant attraction and seed germination. The dispersion of diaspores begins when either (1) capsules or seeds fall beneath the mother plant (barochory) or (2) the seeds are directly collected in the suspended capsules by ants (myrmecochory). As capsules frequently open and expose/disseminate seeds before leaving the mother plant, the adaptation for anemochory appears to be reduced and rarely functional, possibly with only occasional events of long-distance dispersal (e.g. under extreme weather conditions). P. vayredae is essentially myrmecochorous and a diverse array of ant species are involved in seed manipulation, with the elaiosome playing a major role in ant attraction. From the plant’s perspective for dispersal, the majority of ant species had a positive interaction with the seeds, but negative and potential neutral interactions were also observed. Overall, dispersal distances were limited and were mainly determined by ant body size. The frequency of interactions and the ant assemblage varied significantly both spatially and temporally, and these factors may have an effect on directing or disrupting the selection of plant traits. Low seed predation and similar germination rates of intact seeds compared with seeds without elaiosome indicate that seed predator avoidance and seed germination improvement after ant manipulation are not among the selective advantages of myrmecochory operating at present. Dispersal mechanisms that enhance seed dispersal within the population and only occasionally lead to long-distance dispersal events, along with the rarity and patchiness of suitable habitats, may be the main factors explaining the actual density and narrow distribution of this species.