A multi-scale test for dispersal filters in an island plant community

Constraints on plant distributions resulting from seed limitation (i.e. dispersal filters) were evaluated on two scales of ecological organization on islands off the coast of British Columbia, Canada. First, island plant communities were separated into groups based on fruit morphology, and patterns in species diversity were compared between fruit‐type groups. Second, abundance patterns in several common fleshy‐fruited, woody angiosperm species were compared to species‐specific patterns in seed dispersal by birds. Results from community‐level analyses showed evidence for dispersal filters. Dry‐fruited species were rare on islands, despite being common on the mainland. Island plant communities were instead dominated by fleshy‐fruited species. Patterns in seed dispersal were consistent with differences in diversity, as birds dispersed thousands of fleshy‐fruited seeds out to islands, while dry fruited species showed no evidence of mainland‐island dispersal. Results from population‐level analyses showed no evidence for dispersal filters. Population sizes of common fleshy‐fruited species were unrelated to island isolation, as were rates of seed dispersal. Therefore, island isolation distances were not large enough to impose constraints on species’ distributions resulting from seed limitation. Rates of seed dispersal were also unrelated to island area. However, several species increased in abundance with island area, indicating post‐dispersal processes also help to shape species distributions. Overall results suggest that seed dispersal processes play an important role in determining the diversity and distribution of plants on islands. At the community‐level, dry‐fruited species were seed limited and island communities were instead dominated by fleshy‐fruited species. At the population‐level, common fleshy‐fruited species were not seed limited and showed few differences in distribution among islands. Therefore, although evidence for dispersal filters was observed, their effects on plant distributions were scale‐dependent.     

Microsites of seed arrival: spatio–temporal variations in complex seed‐disperser networks

Microsites where seeds arrive during the dispersal process determine plant reproductive success, affecting the quality of dispersal. Despite their crucial role for plant recruitment, very few studies have addressed spatio–temporal variations in microsites of seed arrival in complex seed‐disperser networks. Using an endozoochorous dispersal system, we characterized the microsites of seed arrival of eight fleshy‐fruited plant species dispersed by five mammal species during two consecutive seasons across three sites in a Mediterranean environment (n = 383 feces with seeds; 261 453 seeds). We evaluated spatial and temporal variations in the probability of a seed to arrive at open microsites or at microsites with varying plant cover, considering selection by frugivores and assessing the extent to which seeds of particular species arrived under conspecifics or heterospecifics. We found strong spatio–temporal variations in the amounts of seeds of the eight target species arriving at different microsites. These variations were strongly driven by frugivores’ selection of different landscape elements (i.e. open areas and microsites dominated by different plant species), which differed from expectations based on their local availability. In general, more seeds than expected arrived at vacant (open) microsites. Using bipartite network graphs to connect seeds with their arrival microsites, we found that the proportion of seeds of fleshy‐fruited species arriving near conspecifics or heterospecifics, or at vacant microsites, varied depending on the target plant species, but also on the frugivore species dispersing it, on the study site and on the dispersal season. Our study revealed marked spatio–temporal variations in the microsites of seed arrival, which will potentially have implications for the quality of dispersal effectiveness, ultimately affecting plant population dynamics and community structure. Such a strong context‐dependence in the microsites of seed arrival is likely to confer resilience against unpredictable environmental conditions, like those typical of Mediterranean ecosystems.     

Effects of seed dispersal on Juniperus communis recruitment on a Mediterranean mountain

Abstract. The recruitment of the relict shrub Juniperus communis on a mountain in SE Spain was studied during the period 1994–1998. The main objective was to determine both the quantitative and qualitative effects of bird dispersal on seedling establishment. Seed removal by birds, seed rain, post‐dispersal seed predation, germination, and seedling emergence and survival were analysed in different microhabitats. Birds removed 53 ‐ 89% of the seeds produced by plants. Seed rain was spatially irregular as most seeds accumulated near stones used by birds as perches and below mother plants while a few seeds were dropped in wet meadows and open ground areas. Post‐dispersal seed predation by rodents affected < 10% of dispersed seeds but varied significantly among microhabitats. Only 3.6 ‐ 5.5% of dispersed seeds appeared viable, as many seeds had aborted or showed wasp damage. Seeds germinated in the second and third springs after sowing, reaching a germination percentage of 36%. Seedling emergence was concentrated in wet meadows. Seedling mortality was high (75–80%), but significantly lower in wet meadows, the only microhabitat where seedlings could escape from summer drought, the main mortality cause. Seed abortion, germination and seedling mortality proved to be the main regeneration constraints of J. communis on Mediterranean mountains. Birds exerted a strong demographic effect, although their qualitative effect was limited by abiotic factors which caused the pattern of seed rain to differ from the final pattern of recruitment between microhabitats.     

Colonization of Abandoned Land by Juniperus thurifera Is Mediated by the Interaction of a Diverse Dispersal Assemblage and Environmental Heterogeneity

Land abandonment is one of the most powerful global change drivers in developed countries where recent rural exodus has been the norm. Abandonment of traditional land use practices has permitted the colonization of these areas by shrub and tree species. For fleshy fruited species the colonization of new areas is determined by the dispersal assemblage composition and abundance. In this study we showed how the relative contribution to the dispersal process by each animal species is modulated by the environmental heterogeneity and ecosystem structure. This complex interaction caused differential patterns on the seed dispersal in both, landscape patches in which the process of colonization is acting nowadays and mature woodlands of Juniperus thurifera, a relict tree distributed in the western Mediterranean Basin. Thrushes (Turdus spp) and carnivores (red fox and stone marten) dispersed a high amount of seeds while rabbits and sheeps only a tiny fraction. Thrushes dispersed a significant amount of seeds in new colonization areas, however they were limited by the presence of high perches with big crop size. While carnivores dispersed seeds to all studied habitats, even in those patches where no trees of J. thurifera were present, turning out to be critical for primary colonization. The presence of Pinus and Quercus was related to a reduced consumption of J. thurifera seeds while the presence of fleshy fruited shrubs was related with higher content of J. thurifera seeds in dispersers’ faeces. Therefore environmental heterogeneity and ecosystem structure had a great influence on dispersers feeding behaviour, and should be considered in order to accurately describe the role of seed dispersal in ecological process, such as regeneration and colonization. J. thurifera expansion is not seed limited thanks to its diverse dispersal community, hence the conservation of all dispersers in an ecosystem enhance ecosystems services and resilience.     

Spatiotemporal patterns of seed dispersal in a wind-dispersed Mediterranean tree (Acer opalus subsp. granatense): implications for regeneration

Seed dispersal can severely limit the quantity of plant recruits and their spatial distribution. However, our understanding of the role of dispersal in regeneration dynamics is limited by the lack of knowledge of seed deposition patterns in space and time. In this paper, we analyse the spatiotemporal variability of seed dispersal patterns in the Mediterranean maple, Acer opalus subsp. granatense, by monitoring seed rain along two years at a broad spatial scale (2 mountain ranges, 2 populations per range, 4 microhabitats per population). We quantified seed limitation and its components (source and dispersal limitation), and explored dispersal limitation in space by analysing dispersal distances, seed aggregation, and microhabitat seed distribution. Acer opalus subsp. granatense was strongly seed‐limited throughout the gradients explored, being always dispersal limitation much higher than source limitation. The distribution of seeds with distance from adult individuals was leptokurtic and right‐skewed in all populations, being both kurtosis and skewness higher the year of the highest seed production. Dispersal distances were shorter than expected by random in the four populations, which suggests distance‐limited dispersal. Dispersal patterns were highly aggregated and showed a preferential direction around adults. At the microhabitat scale, most seeds accumulated under adult maples. However, there were no more seeds under trees and shrubs other than maple than in open interspaces, implying that established vegetation does not disrupt patterns of seed deposition by physically trapping seeds. When compared with patterns of seedling establishment, limited dispersal ability and inter‐annual spatial concordance in seed rain patterns suggest that several potentially safe sites for recruitment have a very low probability of receiving seeds in most maple populations. These findings are especially relevant for rare species such as Acer opalus subsp. granatense, and illustrate how dispersal studies are not only crucial for our understanding of plant population dynamics but also to provide conservation directions.     

Comparison of three seed trap types in a chalk grassland: toward a standardised protocol

In a chalk grassland in north-western France, vegetation and seed rain were studied along two transects. We compared the efficiency of three types of seed traps (funnel traps, sticky traps, pots exposed in the field) to estimate the annual seed rain and to reveal various processes involved in plant community regeneration. Data were analysed to compare seed density, species richness and composition across seed traps and vegetation. Geostatistical analyses (semivariograms. were used to detect possible autocorrelation and to examine patterns of spatial variation in seed rain and vegetation. The results show that (1) funnel traps are most efficient (56 species, 8079 seeds.m^–2). (2) Each type of seed traps brings different information about the processes involved in seed dispersal and seedling establishment. Sticky traps help to understand import and export of seeds in a community through anemochory. Funnel traps may provide a measure of the auto-regeneration capacity of the vegetation through its local seed production. Pot traps integrate various processes involved between seed dispersal and seedling emergence. (3.) The exposition to environmental conditions generates marked spatial patterns in species richness measured in pots and in vegetation, selecting species able to survive. Pot traps indicate that numerous species producing seeds in standing vegetation are unable to establish in the field through seedling recruitment. Our study emphasizes the need of using different types of seed traps and spatial sampling configurations to assess the regenerative potential of standing vegetation and the various mechanisms involved in seed dispersal. A standardised protocol for seed rain sampling was finally developed.     

Facilitation by herbivore-mediated nurse plants in a threatened tree, Taxus baccata: local effects and landscape level consistency

We investigated the regeneration of a threatened tree, the yew Taxus baccata, in relation to the presence of fleshy‐fruited woody plants acting as seed dispersal foci as well as protecting yew recruits against ungulate herbivores. We seek to determine if local facilitative effects are consistent across landscape in the Cantabrian range (NW Spain). Yew seed rain by birds mostly concentrated under yew trees and beneath hollies Ilex aquifolium. Seedling emergence distributed similarly to seed rain, but first‐year seedling survival was higher beneath hollies. In one site where woody vegetation was structured as nucleation centres (multispecific patches of fleshy‐fruited plants acting as foci for seed rain) yew recruits mostly occurred in yew‐dominated centres, suggesting dispersers‐mediated facilitation. However, holly was the main nurse plant for most of these recruits, considering the nurse as the species whose canopy covered directly the yew recruit. Living beneath nurse plants reduced herbivore damage on saplings and enhanced seedling survival. A planting experiment with yew rooted‐cuttings beneath different spiny shrubs corroborated this effect. Additional evidence on yew recruitment limitation by herbivory emerged from one population where ungulates were fence‐excluded. Our results suggest that nurse plants mitigate the negative effect of herbivores on yew regeneration, by providing defence against browsing and trampling. Shelter ability related to nurse structure, cone‐shaped shrubs with branches at their bases acting better as a barrier. Paradoxically, this structure resulted from heavy browsing on nurse plants. The study of yew regeneration and habitat structure in seven sites provided evidence for the consistency of facilitation by holly at the landscape level, since local values of yew recruitment positively related to nurse ground cover. Range‐scale yew management must consider the local functioning of the interaction among avian seed‐dispersers, nurse fleshy‐fruited plants and ungulate herbivores, in combination with regional measures, targeting the habitats where facilitation emerges.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

Artificial Bird Perches for the Regeneration of Degraded Tropical Peat Swamp Forest: A Restoration Tool with Limited Potential

Forest rehabilitation activities have been initiated on degraded peatland at several sites in Southeast Asia. In order to achieve rehabilitation efficiently and on the largest possible scale, cost‐effective, transferable methods need to be established. One potential method, which has previously proven successful in both temperate and neotropical forest ecosystems, is the construction of artificial bird perches outside the forest edge. These provide resting perches for frugivorous birds, encouraging them to fly out of the forest, thus increasing seed dispersal and subsequent seedling recruitment into the degraded area. This method was trialled for the first time in degraded tropical peat swamp forest in Indonesia. The results show that the perches were used by frugivorous birds, leading to a significant increase in seed dispersal; however, seedling recruitment was not increased. The frugivorous birds using the perches were degraded zone species and dispersed mainly tree species from the degraded area. Furthermore, while some seeds of forest‐area tree species were dispersed, largely only the degraded area tree species survived to seedling stage. Neither seasonality nor distance from forest edge proved to be significant factors influencing seed dispersal or seedling recruitment, the latter highlighting that seeds were principally being dispersed from within the degraded area rather than from the forest. Although artificial perches did increase seed dispersal, their use as an actual restoration tool in the process of forest regeneration on degraded tropical peatland is limited. Furthermore, when the cost‐ and effort‐to‐area factors are considered, this method is shown to be inefficient.     

Seed rain of fleshy and dry propagules in different habitats in the temperate rainforests of Chiloé Island,Chile

In temperate rainforests on Chiloé Island in southern Chile (42°S), most canopy trees bear fleshy, avian‐dispersed propagules, whereas emergent tree species have dry, wind‐borne propagules. In the present study, the following hypothesis was tested: regardless of species, fleshy propagules are deposited in greater numbers in canopy gaps and in forest margins and hence have a more heterogeneous seed shadow than wind‐dispersed propagules. To test this hypothesis, the seed rains of these two types of propagules were compared in the following forest habitats: (i) tree‐fall gaps (edges and centre); (ii) forest margins with adjacent pastures; and (iii) under closed canopy (forest interior). Seed collectors (30‐cm diameter) were placed in two (15 and 100 ha) remnant forest patches (n = 60–100 seed collectors per patch) distributed in the four habitats. Seeds were retrieved monthly from each collector during two reproductive seasons (1996, 1997). In both years, the seed rain was numerically dominated by two species with dry propagules (Laureliopsis philippiana and Nothofagus nitida) and three species with fleshy fruits (Drimys winteri, Amomyrtus luma, and Amomyrtus meli). The seed shadows of the two species with dry, wind‐dispersed seeds differed markedly. Seeds of L. philippiana were deposited predominantly in canopy openings, whereas N. nitida seeds fell almost entirely in the forest interior. The fleshy‐fruited species, Drimys and Amomyrtus spp., had similar seed deposition patterns in the various habitats studied, but the between‐year differences in seed rain were greater in Drimys winteri than in Amomyrtus spp. Although no more than 10% of fleshy‐fruited propagules reached the margins of the patch, approximately 7% of these were carried there by birds. Every year, canopy gaps (pooling data from edges and centres) concentrated approximately 60% of the total seed rain of both propagule types in both forest patches. Forest margins received less than 20% of the total seed rain, which was largely dominated by fleshy‐fruited species. Seed shadows were a species‐specific attribute rather than a trait associated with propagule type and dispersal mode.     

Rainforest seed rain into abandoned tropical Australian pasture is dependent on adjacent rainforest structure and extent

It is well known that the recovery of abandoned tropical pastures to secondary rainforest benefits from the arrival of seeds from adjacent rainforest patches. Less is known, however, about how the structural attributes of adjacent rainforest (e.g. tree density, canopy cover and tree height) impact seed rain patterns into abandoned pastures. Between 2011 and 2013, we used seed traps and ground seed surveys to track the richness and abundance of rainforest seeds entering abandoned pastures in Australia's wet tropics. We also tested how seed rain diversity is related to the distance from forest, the proportion of forest cover in the landscape and several structural attributes of adjacent forest patches, specifically average tree height, canopy cover, tree species richness and density. Almost no seeds were captured in elevated pasture seed traps, even near forest remnants. Abundant forest seeds were found in ground surveys but only within 10 m of forest edges. In ground surveys, seeds from wind‐dispersed species were more abundant, but less species rich, than animal‐dispersed species. A survey of pasture seedling recruits suggested that some forest seeds must be dispersing more than 10 m into pasture at very low frequencies, but only a few species are establishing there. Recruits were predominantly animal‐dispersed not wind‐dispersed species. In addition to distance from forest and the proportion of forest within a 100‐ to 200‐m radius of sampling sites, the richness and density of adjacent forest trees were the most important factors for explaining the probability of seed occurrence in abandoned pastures. Results suggest that without some restoration assistance, the recovery of abandoned pastures into secondary rainforest in Australia's tropical rainforests will likely be limited, at least in part, by a very low rate of seed dispersal away from forest edges and by the diversity and density of trees in adjacent remnant forests.     

Unravelling seed dispersal through fragmented landscapes: Frugivore species operate unevenly as mobile links

Seed dispersal constitutes a pivotal process in an increasingly fragmented world, promoting population connectivity, colonization and range shifts in plants. Unveiling how multiple frugivore species disperse seeds through fragmented landscapes, operating as mobile links, has remained elusive owing to methodological constraints for monitoring seed dispersal events. We combine for the first time DNA barcoding and DNA microsatellites to identify, respectively, the frugivore species and the source trees of animal‐dispersed seeds in forest and matrix of a fragmented landscape. We found a high functional complementarity among frugivores in terms of seed deposition at different habitats (forest vs. matrix), perches (isolated trees vs. electricity pylons) and matrix sectors (close vs. far from the forest edge), cross‐habitat seed fluxes, dispersal distances and canopy‐cover dependency. Seed rain at the landscape‐scale, from forest to distant matrix sectors, was characterized by turnovers in the contribution of frugivores and source‐tree habitats: open‐habitat frugivores replaced forest‐dependent frugivores, whereas matrix trees replaced forest trees. As a result of such turnovers, the magnitude of seed rain was evenly distributed between habitats and landscape sectors. We thus uncover key mechanisms behind “biodiversity–ecosystem function” relationships, in this case, the relationship between frugivore diversity and landscape‐scale seed dispersal. Our results reveal the importance of open‐habitat frugivores, isolated fruiting trees and anthropogenic perching sites (infrastructures) in generating seed dispersal events far from the remnant forest, highlighting their potential to drive regeneration dynamics through the matrix. This study helps to broaden the “mobile‐link” concept in seed dispersal studies by providing a comprehensive and integrative view of the way in which multiple frugivore species disseminate seeds through real‐world landscapes.     

Spatial dynamics of <Emphasis Type="Italic">Ilex aquifolium</Emphasis> populations seed dispersal and seed bank: understanding the first steps of regeneration

The objective of this study was to analyse quantitatively the spatial distribution of holly (Ilex aquifolium L.) seed rain and seed bank, and to detect the relationships between these consecutive processes. We measured seed dispersal by birds and fallen fruits, and also density and viability of seed bank in two Ilex populations in central Spain. Analysis was made distinguishing the following microhabitats: holly woodland, edge of holly woodland, open grassland 10 m and 100 m from the woodland, fleshy fruit shrubs, dry fruit shrubs, and adjacent non-holly woodland. Spatial distribution of dispersed and in-soil seeds was measured by the clumping index. Seed rain and seed bank under holly woodlands were significantly higher than in the other microhabitats. Forest edges and fleshy fruit shrubs were the next microhabitats with the highest seed rain and seed bank density. Interannual and interlocality variations were not significant. The relative importance of the different dispersal methods varied between microhabitats, with a similar support of bird dispersed seeds and fallen fruits within the woodland and a greater influence of cattle dispersal in open areas. Seed spatial aggregation was significant in both dispersed seeds and soil seeds from holly woodlands and the edge of the forest. Aggregation under shrubs, grasslands and the adjacent forests evidenced a general random distribution of holly seeds (only in some cases clumping index was significant). Quantitative differences between seed rain and seed bank are important. Post-dispersal seed predation did not modify seed rain distribution, which was mirrored in the seed bank pattern. These two phases of holly regeneration had a heavy spatial influence, determined by the landscape structure and activity of the dispersal agents, that reflects a differential recruitment potential. Comparisons between both populations suggest that in the southern locality (Robregordo) holly has a weaker capacity to colonize open areas, and a stronger recruitment limitation due to propagule availability.     

Reproductive phenology of Melastomataceae species with contrasting reproductive systems: contemporary and historical drivers

• Flowering and fruiting are key events in the life history of plants, and both are critical to their reproductive success. Besides the role of evolutionary history, plant reproductive phenology is regulated by abiotic factors and shaped by biotic interactions with pollinators and seed dispersers. In Melastomataceae, a dominant Neotropical family, the reproductive systems vary from allogamous with biotic pollination to apomictic, and seed dispersal varies from dry (self‐dispersed) to fleshy (animal‐dispersed) fruits. Such variety in reproductive strategies is likely to affect flowering and fruiting phenologies.
• In this study, we described the reproductive phenology of 81 Melastomataceae species occurring in two biodiversity hotspots: the Atlantic rain forest and the campo rupestre. We aim to disentangle the role of abiotic and biotic factors defining flowering and fruiting times of Melastomataceae species, considering the contrasting breeding and seed dispersal systems, and their evolutionary history.
• In both vegetation types, pollinator‐dependent species had higher flowering seasonality than pollinator‐independent ones. Flowering patterns presented phylogenetic signal regardless of vegetation type. Fruiting of fleshy‐fruited species was seasonal in campo rupestre but not in Atlantic rain forest; the fruiting of dry‐fruited species was also not seasonal in both vegetation types. Fruiting showed a low phylogenetic signal, probably because the influence of environment and dispersal agents on fruiting time is stronger than the phylogenetic affinity.
• Considering these ecophylogenetic patterns, our results indicate that flowering may be shaped by the different reproductive strategies of Melastomataceae lineages, while fruiting patterns may be governed mainly by the seed dispersal strategy and flowering time, with less phylogenetic influence.

     

Can salvage logging affect seed dispersal by birds into burned forests?

The recovery of vegetation in Mediterranean ecosystems after wildfire is mostly a result of direct regeneration, since the same species existing before the fire regenerate on-site by seeding or resprouting. However, the possibility of plant colonization by dispersal of seeds from unburned areas remains poorly studied. We addressed the role of the frugivorous, bird-dependent seed dispersal (seed rain) of fleshy-fruited plants in a burned and managed forest in the second winter after a fire, before on-site fruit production had begun. We also assessed the effect on seed rain of different microhabitats resulting from salvage logging (erosion barriers, standing snags, open areas), as well as the microhabitats of unlogged patches and an unburned control forest, taking account of the importance of perches as seed rain sites. We found considerable seed rain by birds in the burned area. Seeds, mostly from Olive trees Olea europaea and Evergreen pistaches Pistacia lentiscus, belonged to plants fruiting only in surrounding unburned areas. Seed rain was heterogeneous, and depended on microhabitat, with the highest seed density in the unburned control forest but closely followed by the wood piles of erosion barriers. In contrast, very low densities were found under perches of standing snags. Furthermore, frugivorous bird richness seemed to be higher in the erosion barriers than elsewhere. Our results highlight the importance of this specific post-fire management in bird-dependent seed rain and also may suggest a consequent heterogeneous distribution of fleshy-fruited plants in burned and managed areas. However, there needs to be more study of the establishment success of dispersed seeds before an accurate assessment can be made of the role of bird-mediated seed dispersal in post-fire regeneration.     

Incorporating seed fate into plant–frugivore networks increases interaction diversity across plant regeneration stages

Plant–animal mutualistic interactions, such as pollination and seed dispersal, affect ecosystem functioning by driving plant population dynamics. However, little is known of how the diversity of interactions in these mutualistic networks determines plant regeneration dynamics. To fill this gap, interaction networks should not only account for the number of seeds dispersed by animals, but also for seed fate after dispersal. Here, we compare plant–animal networks at both the seed dispersal and seedling recruitment stage to evaluate how interaction diversity, represented by different network metrics, changes throughout the process of plant regeneration. We focused on a system with six species of frugivorous birds and three species of fleshy‐fruited trees in the temperate secondary forest of the Cantabrian Range (northern Iberian Peninsula). We considered two plant cohorts corresponding to two fruiting years showing strong differences in fruit and frugivore abundance. Seed dispersal interactions were estimated from a spatially‐explicit, field‐validated model predicting tree and bird species‐specific seed deposition in different microhabitats. These interactions were further transformed into interactions at the seedling recruitment stage by accounting for plant‐ and microhabitat‐specific seed fates estimated from field sampling. We found that network interaction diversity varied across plant regeneration stages and cohorts, both in terms of the evenness and the number of paired interactions. Tree–bird interactions were more evenly distributed across species pairs at the recruitment stage than at the seed deposition stage, although some interactions disappeared in the seed‐to‐seedling transition for one plant cohort. The variations in interaction diversity were explained by between‐plant differences in post‐dispersal seed fate and in inter‐annual fruit production, rather than by differences between frugivores in seed deposition patterns. These results highlight the need for integrating plant traits and disperser quality to predict the functional outcome of plant–animal mutualistic networks.     

Intercontinental long‐distance seed dispersal across the Mediterranean Basin explains population genetic structure of a bird‐dispersed shrub

Long‐distance dispersal (LDD) is a pivotal process for plants determining their range of distribution and promoting gene flow among distant populations. Most fleshy‐fruited species rely on frugivorous vertebrates to disperse their seeds across the landscape. While LDD events are difficult to record, a few ecological studies have shown that birds move a sizeable number of ingested seeds across geographic barriers, such as sea straits. The foraging movements of migrant frugivores across distant populations, including those separated by geographic barriers, creates a constant flow of propagules that in turn shapes the spatial distributions of the genetic variation in populations. Here, we have analysed the genetic diversity and structure of 74 populations of Pistacia lentiscus, a fleshy‐fruited shrub widely distributed in the Mediterranean Basin, to elucidate whether the Mediterranean Sea acts as a geographic barrier or alternatively whether migratory frugivorous birds promote gene flow among populations located on both sides of the sea. Our results show reduced genetic distances among populations, including intercontinental populations, and they show a significant genetic structure across an eastern‐western axis. These findings are consistent with known bird migratory routes that connect the European and African continents following a north‐southwards direction during the fruiting season of many fleshy‐fruited plants. Further, approximate Bayesian analysis failed to explain the observed patterns as a result of historical population migrations at the end of Last Glacial Maximum. Therefore, anthropic and/or climatic changes that would disrupt the migratory routes of frugivorous birds might have genetic consequences for the plant species they feed upon.     

Linking structure and functionality in mutualistic networks: do core frugivores disperse more seeds than peripheral species?

Seed dispersal networks are often organized in nested structures in a way that a few core species can disproportionally affect the remaining species in a network, in both the ecological and evolutionary sense. Yet, the relative importance of core and peripheral species has not been properly tested in seed dispersal network studies. We determined core species from 10 local seed dispersal networks composed by fleshy‐fruited plants and frugivorous birds. Each of those local quantitative networks was characterized with a core quality value, a core score for each species and a threshold value between core and peripheral species. From a total of 52 bird and 69 plant species that interacted in the study area, only 8 and 15, respectively, were identified as core. Each local network had a core that comprised a small number of birds and plants, always lower than 30% of the interacting species. There was no difference in the quantitative component of seed dispersal effectiveness (QC) provided by the frugivorous bird assemblage to plant functional groups clustered according to their growth form and fruit characteristics. Core birds dispersed seeds from each of these plant functional groups with a higher QC than peripheral species. Thus, we empirically demonstrate for the first time that seed dispersal networks at a regional scale have a small core set of fruit‐eating birds, upon which heavily rely most fleshy‐fruited plants for their seed removal. Hence, the activity of just a few core frugivores could deeply impact the demography of an entire assemblage of fleshy‐fruited plants.     

Biogeography of seed-dispersal syndromes, life-forms and seed sizes among woody rain-forest plants in Australia's subtropics

Aim To enhance our understanding of the evolutionary interactions between seed‐dispersal syndromes, life‐forms, seed size, and habitat characteristics by studying their association with the regional‐scale distributions of subtropical rain‐forest plants in the context of climatic gradients. Location South‐east Queensland, subtropical eastern Australia (152° E, 26° S). Methods We classified 250 rain‐forest sites into six floristic site‐groups based on their woody plant composition. The resulting classification was strongly associated with variation in rainfall. The distribution of species across the floristic site‐groups was used to assign 568 species to seven habitat classes (one class for ‘widespread’ species, with all other species classified according to the site‐group within which they were most frequent). Species were also classified for three other categorical life‐history factors: three dispersal syndromes based on diaspore morphology (fleshy, wind‐assisted, and unadorned); four life‐forms (trees, shrubs and small trees, tall climbers, and short and shrubby climbers); and four seed‐diameter classes (< 3 mm, ≥ 3 and < 4.5 mm, ≥ 4.5 and < 7 mm, and ≥ 7 mm). We used a basic comparative approach augmented by simple phylogenetically constrained comparisons to assess association between dispersal syndrome, seed size, life‐form, and habitat class. Results Across the rain forests of south‐east Queensland, the proportion of species with fleshy diaspores or of large stature increases with rainfall. High‐rainfall sites also have larger average seed sizes, but the difference in average seed size between high‐ and low‐rainfall sites is small compared with variation within sites. Among species, those with fleshy fruit tend to have larger seeds and to favour high‐rainfall sites. Very few small trees produce diaspores adapted for wind‐assisted dispersal. On average, species with unadorned diaspores have smaller seeds than those with fleshy diaspores. However, within sites, species with unadorned and fleshy diaspores have similar average seed sizes, and some species with unadorned diaspores from high‐rainfall habitats have extremely large seeds. Main conclusions Commonly observed associations between fleshy fruit, larger plants, larger seeds, and productive habitats are apparent within the rain‐forest flora of south‐east Queensland. However, these associations are generally weak and involve complex interactions. For example, the strong tendency for species with fleshy fruit to have larger seeds than those with unadorned diaspores concealed a significant group of species from wetter forests that produce extremely large seeds and unadorned diaspores. The most widespread species in this study tend to be large plants (particularly robust lianes) and to produce fleshy fruit, but they tend not to have relatively large seeds. The association between large seeds, large plants, fleshy fruit and productive habitats is discussed as part of an evolutionary strategy favouring fitness in populations close to carrying capacity. We review some problems with focusing on establishment chances per seed as the driver towards association between large seeds, large plants and productive rain‐forest habitats (the difficult‐establishment hypothesis). Instead we suggest that production of large, short‐lived seeds by long‐lived plants in temporally stable, closed habitats may reflect the limited evolutionary potential for strategies enhancing colonization (e.g. producing large numbers of dormant seeds), thus allowing the establishment benefits of large seeds greater selective influence (the slow‐replacement hypothesis). The association of fleshy fruit with large seeds probably reflects the difficulty of dispersing large seeds by other means (the difficult‐dispersal hypothesis).     

Spatial and Temporal Variation of Seed Rain in the Canopy and on the Ground of a Tropical Cloud Forest

Spatial and temporal patterns of seed rain impact plant fitness, genetic and demographic structure of plant populations, and species' interactions. Because plants are sessile, they rely on biotic and abiotic dispersal agents to move their seeds. The relative importance of these dispersal agents may shift throughout the year. In tropical forests, seed dispersal of epiphytes constitutes a major but hitherto unknown portion of seed rain ecology. For the first time, we report on patterns of seed rain for both epiphytic and terrestrial plants across an entire year in a Neotropical montane forest. To examine seed rain, we placed traps in the canopy and on the ground. We analyzed seed dispersal syndrome (bird, mammal, wind) and plant habit (epiphyte, liana, shrub, small tree, large tree) across all seasons of the year (dry, misty, wet). We found that the community of species collected in canopy traps was significantly different from the community in ground traps. Epiphytes were the most common plant habit found in canopy traps, while large trees were most common in ground traps. Species with bird‐dispersed seeds dominated all traps. Species richness was significantly higher during the dry season in ground traps, but did not vary across seasons in canopy traps. Our results highlight the distinct seed rain found in the canopy and on the ground and underscore the importance of frugivores for dispersing both arboreal and terrestrial plants in tropical ecosystems.