Seed dispersal of alien and native plants by vertebrate herbivores

Seed dispersal is crucial for the success and spread of alien plants. Herbivores often establish a dual relationship with plants: antagonist, through herbivory, and mutualist, through seed dispersal. By consuming plants, herbivores may disperse large amounts of seeds, and can facilitate the spread of alien plants. However, seed dispersal of alien plants by herbivores has been largely uninvestigated. I studied factors associated with dispersal of alien and native seeds by the three most important vertebrate herbivores in SW Australia: emus (Dromaius novaehollandia), western grey kangaroos (Macropus fuliginosus) and European rabbits (Oryctolagus cuniculus). Overall frequencies of alien and native seeds dispersed by these herbivores were determined by differences among them in (1) the plant groups they predominantly disperse, that differed in frequencies of aliens versus natives, and (2) the predominant dispersal of aliens or natives within those plant groups. Emus and kangaroos (natives) tended to disperse predominantly alien seeds within plant groups (defined by life forms, dispersal syndromes, and diaspore size), whereas rabbits (alien) tended to disperse predominantly natives. This agrees with the hypothesis that herbivores will use predominantly plants that have evolved in different areas, because of less effective defences against new enemies. Overall frequencies were consistent with this pattern in kangaroos and rabbits, but not in emus. Kangaroos dispersed mostly plant groups that were mainly aliens (herbaceous species and small and medium sized dispersal units and seeds), which together with their predominant use of aliens over natives within groups resulted in the highest overall frequency of alien seeds (73%). Rabbits were similar to kangaroos in the type of plants dispersed, but their predominant use of natives over aliens within groups contributed to an overall predominance of native seeds in their pellets (88%). Emus dispersed mostly plant groups that were mainly natives (e.g. woody species with big diaspores), resulting in low overall frequency of alien seeds (11%), despite their predominant use of aliens over natives within plant groups. Thus, the within-groups trend pointed to a facilitative role of native herbivores of plant invasions through seed dispersal, but was obscured by the different use by herbivores of plant groups with different frequency of aliens.     

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.     

Myrmecochorous dispersal distances: a world survey

Abstract. Myrmecochorous dispersal distances are reviewed; the seed dispersal curve generated by ants shows a characteristic peak at short distances and a long tail, a shape suited to small densities of safe sites. Mean global distance is of 0.96 m (n= 2524) with a range of 0.01–77 m. Data have been broken down by geography (Northern hemisphere v. Southern hemisphere), taxonomy (ant subfamilies) and ecology (vegetation: sclerophyllous v. mesophyllous). Although a statistical difference exists between dispersal curves from the Northern hemisphere and the Southern hemisphere, this may be an artefact of lack of data from mesophyllous myrmecochores from this hemisphere. The four ant subfamilies do show also numerical differences but could not be subjected to statistical analysis. A difference between the shape of dispersal curve for sclerophyllous myrmecochores and mesophyllous myrmecochores has also been detected. We hypothesize that this difference is related to the myrmecological communities from both types of vegetation: dispersing ants from sclerophyllous vegetation would have smaller nest densities and/or bigger foraging areas than dispersing ants from mesic environments.     

Small Tent-Roosting Bats Promote Dispersal of Large-Seeded Plants in a Neotropical Forest

In Neotropical regions, fruit bats are among the most important components of the remaining fauna in disturbed landscapes. These relatively small-bodied bats are well-known dispersal agents for many small-seeded plant species, but are assumed to play a negligible role in the dispersal of large-seeded plants. We investigated the importance of the small tent-roosting bat Artibeus watsoni for dispersal of large seeds in the Sarapiquí Basin, Costa Rica. We registered at least 43 seed species > 8 mm beneath bat roosts, but a species accumulation curve suggests that this number would increase with further sampling. Samples collected beneath bat feeding roosts had, on average, 10 times more seeds and species than samples collected 5 m away from bat feeding roosts. This difference was generally smaller in small, disturbed forest patches. Species-specific abundance of seeds found beneath bat roosts was positively correlated with abundance of seedlings, suggesting that bat dispersal may influence seedling recruitment. Our study demonstrates a greater role of small frugivorous bats as dispersers of large seeds than previously thought, particularly in regions where populations of large-bodied seed dispersers have been reduced or extirpated by hunting.     

Epizoochory by large herbivores: merging data with models

The dispersal of plant seeds in the fur of large herbivores (epizoochory) is an important but complex long-distance dispersal mechanism. We developed a spatially explicit simulation model of epizoochorous seed dispersal, which was parameterized based on empirical studies of the movement and behaviour of donkeys, and the distribution, seed production, seed accessibility, seed adhesion, and seed retention on donkey fur of selected plant species in a coastal dune nature reserve in Flanders, Belgium. We compared predicted and observed seed numbers of the 14 plant species on donkey fur.

Modelled seed shadows indicate that for most species about half of all seeds dispersed by donkeys should travel a net distance of >100 m, and about 1% should travel >500 m within this more or less isodiametric 100 ha nature reserve. Seeds with longer retention times are expected to travel further than those with short retention times. Enlarging the reserve area had little impact on the forecasted dispersal distances.

Variation among plant species in the observed seed numbers found on donkey fur were well predicted by the model (R²=0.56, P=0.002), though the predictions relied on relatively crude estimates of seed production and accessibility to donkeys, indicating that more accurate estimates of these parameters are needed.

Our model confirms the important role of epizoochory in affecting long-distance seed dispersal, and provides a modelling framework for integrating the multiple components of the dispersal process.     

Importance of demography and dispersal for the resilience and restoration of a critically endangered tropical conifer Araucaria nemorosa

Aims In situ species survival involves persistence at current sites and/or colonization of new locations. Determining the likelihood of these events requires an understanding of population dynamics and dispersal. We address these issues in populations of a critically endangered tropical conifer Araucaria nemorosa and provide conservation recommendations. Location Port Boise, island of Grand Terre, New Caledonia, Pacific Ocean. Methods We characterize the demographic structure of six populations of A. nemorosa based upon size class frequencies and relative basal area compared to competing angiosperm trees. Using genotype data from 280 adult individuals at seven microsatellite loci, we indirectly estimate dispersal distances and project a maximal dispersal envelope around the extant populations. Results Our survey detected marked differences in demographic parameters including the proportion of basal area occupied by A. nemorosa versus angiosperm trees (ranging from 25% to 77%), the size class frequency distributions of A. nemorosa and seedling densities (ranging from 500 to 37300 seedlings ha^?1) among some populations. Wright’s genetic neighbourhood ranged from 22 to 876 trees, and historic gene dispersal ranged from 10.8 to 82.4 m, indicating that most seed dispersal is <100 m. Main conclusions This study indicates that the risk of recruitment failure in remnant populations of Araucaria nemorosa is low in most of the stands but is high in the inland population Forêt Nord. In situ natural regeneration appears to be constrained in this population, most likely as a consequence of competitive exclusion. Our results also suggest that the majority of seed dispersal is too short to allow A. nemorosa to disperse to new more hospitable sites within an ecologically relevant time‐scale. Our findings have implications not only for this emblematic tree species but also for a wide range of fragmented and degraded plant species populations with limited dispersal that are vulnerable to competitive exclusion.     

Seed dispersal spectra: a comparison of temperate plant communities

Abstract. We compare the dispersal spectra of diaspores from varied plant communities in Australia, New Zealand, and North America, assigning dispersal mode to each diaspore type on the basis of apparent morphological adaptations. Species with ballistic and external dispersal modes were uncommon in most communities we surveyed. Ant dispersal was also rather uncommon, except in some Australian sclerophyll vegetation types. The frequency of vertebrate dispersal ranged up to 60% of the flora, the highest frequencies occurring in New Zealand forests. Wind dispersal ranged as high as 70% of the flora, with the highest values in Alaska, but usually comprised 10–30% of the flora. Many species in most communities had diaspores with no special morphological device for dispersal. Physiognomically similar vegetation types indifferentbiogeographic regions usually had somewhat dissimilar dispersal spectra. The frequency of dispersal by vertebrates often increased and the frequency of species with no special dispersal device decreased along gradients of increasing vertical diversity of vegetation structure. Elevation and moisture gradients also exhibited shifts in dispersal spectra. Within Australia, vertebrate- and wind-dispersal increased in frequency along a soil-fertility gradient, and dispersal by ants and by no special device decreased. Habitat breadths (across plant communities) and microhabitat breadths (within communities) for species of each major dispersal type did not show consistent differences, in general. Ant-dispersed species often had lower cover-values than other species in several Australian vegetation types. We discuss the ecological bases of these differences in dispersal spectra in terms of the availability of dispersal agents, seed size, and other ecological constraints. Seed size is suggested to be one ecological factor that is probably of general relevance to the evolution of dispersal syndromes.     

Alongshore dispersal and site fidelity of juvenile plaice from tagging and transplants

Juvenile plaice Pleuronectes platessa ( n  = 1281) were tagged and released at two locations 300 m apart on a 1 km long sandy beach. Most (>90%) of the fish were recaptured within 100 m of the release site (shown by the colour of the tag), with very few caught >200 m distance after 6 weeks. The changing spatial distribution of marked fish was adequately reproduced by a simple dispersal model with a single parameter: a 78% probability of remaining in a 100 m wide zone from one day to the next, with a 22% chance that fish move into an adjacent zone. In a subsequent similar study at the same beach, fish were either released at the point of capture ( n  = 881) or transplanted to the alternate site ( n  = 910) 100 m distant. After 6 weeks, transplanted fish moved along the shore towards their sites of original capture. Fish replaced at the point of capture showed no such movement along the shore. Further modification of the dispersal model to allow for a distinction between dispersal from home sites and from sites away from the original point of collection, was sufficient to reproduce the behaviour of the populations of both transplanted and control treatment groups. The likelihood of dispersal from home sites was much less than that seen at sites away from home. Juvenile plaice thus have a degree of long‐shore site fidelity not expected of a fish with strong depth‐related migration behaviour in a relatively homogenous habitat.     

The effect of isolation and history on colonization patterns of plant species in secondary woodland

Colonization success of woodland originating after 1850 was determined for seventeen forest plant species having different dispersal strategies. Colonization rate of the studied endo-and exozoochorous species apparently was considerable higher than that of species having short distance dispersal like myrmecochores and species lacking dispersal mechanisms. The occurrence of eight species in this young forest habitat was related to the distance to the nearest source patch (DNS), as well as to the age of the young patches and to their former land use. DNS calculated to old, existent and occupied source patches affected most analysed species. Only Ilex aquifolium L. had a significant higher occurrence in woodland originating before 1916 than in those originating after 1916. Former land use showed significant effects for three species. Although colonization rate and effects of studied parameters on occurrence were different for the studied species, no clear differences were found between different dispersal groups. The consequences of these results for the understanding of colonization processes of the species studied is discussed.     

Contemporary pollen and seed dispersal in a Prunus mahaleb population: patterns in distance and direction

Pollination and seed dispersal determine the spatial pattern of gene flow in plant populations and, for those species relying on pollinators and frugivores as dispersal vectors, animal activity plays a key role in determining this spatial pattern. For these plant species, reported dispersal patterns are dominated by short-distance movements with a significant amount of immigration. However, the contribution of seed and pollen to the overall contemporary gene immigration is still poorly documented for most plant populations. In this study we investigated pollination and seed dispersal at two spatial scales in a local population of Prunus mahaleb (L.), a species pollinated by insects and dispersed by frugivorous vertebrates. First, we dissected the relative contribution of pollen and seed dispersal to gene immigration from other parts of the metapopulation. We found high levels of gene immigration (18.50%), due to frequent long distance seed dispersal events. Second, we assessed the distance and directionality for pollen and seed dispersal events within the local population. Pollen and seed movement patterns were non-random, with skewed distance distributions: pollen tended moved up to 548 m along an axis approaching the N-S direction, and seeds were dispersed up to 990 m, frequently along the SW and SE axes. Animal-mediated dispersal contributed significantly towards gene immigration into the local population and had a markedly nonrandom pattern within the local population. Our data suggest that animals can impose distinct spatial signatures in contemporary gene flow, with the potential to induce significant genetic structure at a local level.     

Tug of war between continental gene flow and rearing site philopatry in a migratory bird: the sex-biased dispersal paradigm reconsidered

Nonrandom dispersal has been recently advanced as a mechanism promoting fine-scale genetic differentiation in resident populations, yet how this applies to species with high rates of dispersal is still unclear. Using a migratory species considered a classical example of male-biased dispersal (the greater snow goose, Chen caerulescens atlantica ), we documented a temporally stable fine-scale genetic clustering between spatially distinct rearing sites (5–30 km apart), where family aggregates shortly after hatching. Such genetic differentiation can only arise if, in both sexes, dispersal is restricted and nonrandom, a surprising result considering that pairing occurs among mixed flocks of birds more than 3000 km away from the breeding grounds. Fine-scale genetic structure may thus occur even in migratory species with high gene flow. We further show that looking for genetic structure based on nesting sites only may be misleading. Genetically distinct individuals that segregated into different rearing sites were in fact spatially mixed during nesting. These findings provide new, scale-dependent links between genetic structure, pairing, and dispersal and show the importance of sampling different stages of the breeding cycle in order to detect a spatial genetic structure.     

Long‐term consistency in spatial patterns of primate seed dispersal

Seed dispersal is a key ecological process in tropical forests, with effects on various levels ranging from plant reproductive success to the carbon storage potential of tropical rainforests. On a local and landscape scale, spatial patterns of seed dispersal create the template for the recruitment process and thus influence the population dynamics of plant species. The strength of this influence will depend on the long‐term consistency of spatial patterns of seed dispersal. We examined the long‐term consistency of spatial patterns of seed dispersal with spatially explicit data on seed dispersal by two neotropical primate species, Leontocebus nigrifrons and Saguinus mystax (Callitrichidae), collected during four independent studies between 1994 and 2013. Using distributions of dispersal probability over distances independent of plant species, cumulative dispersal distances, and kernel density estimates, we show that spatial patterns of seed dispersal are highly consistent over time. For a specific plant species, the legume Parkia panurensis, the convergence of cumulative distributions at a distance of 300 m, and the high probability of dispersal within 100 m from source trees coincide with the dimension of the spatial–genetic structure on the embryo/juvenile (300 m) and adult stage (100 m), respectively, of this plant species. Our results are the first demonstration of long‐term consistency of spatial patterns of seed dispersal created by tropical frugivores. Such consistency may translate into idiosyncratic patterns of regeneration.     

How important is long‐distance seed dispersal for the regional survival of plant species?

Long‐distance seed dispersal is generally assumed to be important for the regional survival of plant species. In this study, we quantified the importance of long‐distance seed dispersal for regional survival of plant species using wind dispersal as an example. We did this using a new approach, by first relating plant species’ dispersal traits to seed dispersal kernels and then relating the kernels to regional survival of the species. We used a recently developed and tested mechanistic seed dispersal model to calculate dispersal kernels from dispersal traits. We used data on 190 plant species and calculated their regional survival in two ways, using species distribution data from 36,800 1 km²‐grid cells and 10,754 small plots covering the Netherlands during the largest part of the 20th century. We carried out correlation and stepwise multiple regression analyses to quantify the importance of long‐distance dispersal, expressed as the 99‐percentile dispersal distance of the dispersal kernels, relative to the importance of median‐distance dispersal and other plant traits that are likely to contribute to the explanation of regional survival: plant longevity (annual, biennial, perennial), seed longevity, and plant nutrient requirement. Results show that long‐distance dispersal plays a role in determining regional survival, and is more important than median‐distance dispersal and plant longevity. However, long‐distance dispersal by wind explains only 1–3% of the variation in regional survival between species and is equally important as seed longevity and much less important than nutrient requirement. In changing landscapes such as in the Netherlands, where large‐scale eutrophication and habitat destruction took place in the 20th century, plant traits indicating ability to grow under the changed, increasingly nutrient‐rich conditions turn out to be much more important for regional survival than seed dispersal.     

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.     

Movement patterns of Rhyssomatus lineaticollis Say (Coleoptera: Curculionidae) within and among Asclepias syriaca (Asclepiadaceae) patches in a fragmented landscape

Abstract.  1. Dispersal capabilities of organisms are critical in determining the landscape population structure of species as well as their likelihood of survival in fragmented landscapes. Using mark–recapture techniques on the monophagous weevil Rhyssomatus lineaticollis Say (Curculionidae), within- and between-patch dispersal capabilities, landscape level population structure, and the role of beetle density and host patch characteristics in setting distances, amounts, and timing of dispersal were studied.
2. The data indicate that R. lineaticollis is sedentary, with 50% of recaptured beetles moving < 1 m and the maximum distance moved < 1 km. Within- and between-patch movement of beetles was unrelated to host plant patch characteristics and beetle densities.
3. Despite limited dispersal, R. lineaticollis probably functions as a patchy population in east-central Iowa, U.S.A. because dispersals between patches are common and because all host patches surveyed contained this herbivore, indicating a lack of suitable vacant patches, a prerequisite for metapopulation structure.
4. Between-patch distances are well within the dispersal capabilities of R. lineaticollis , although this may be the result of an increase in the density of patches of its host, Asclepias syriaca , in the landscape over the last 150 years as a result of human disturbance and this species' weedy habit.
5. Metapopulation structure in monophagous prairie herbivores may be most likely in species whose non-weedy host plants form highly predictable resources in space and time, but which are now widely scattered in habitat fragments.     

Species,Diaspore Volume and Body Mass Matter in Gastropod Seed Feeding Behavior

Background

Seed dispersal of ant-dispersed plants (myrmecochores) is a well studied ecosystem function. Recently, slugs have been found to act as seed dispersers of myrmecochores. The aim of our study was to (1) further generalize the finding that gastropods feed on seeds of myrmecochores and hence may act as seed dispersers, (2) to test whether gastropod body mass and the volume of diaspores have an influence on the seed dispersal potential.

Methodology and Principal Findings

We assessed the seed dispersal potential of four slug and snail species with a set of seven myrmecochorous plant species from seven different plant families common to Central European beech forests. Diaspores differed in shape and size. Gastropods differed in their readiness to feed on diaspores and in the proportion of seeds that were swallowed as a whole, and this readiness generally decreased with increasing diaspore size. Smaller Arionid slugs (58 mm body length; mean) mostly fed on the elaiosome but also swallowed small diaspores and therefore not only act as elaiosome consumers, a nutrient rich appendage on myrmecochorous diaspores, but may also disperse seeds. Large Arionid slugs (>100 mm body length) swallowed diaspores of all sizes. Diaspores swallowed by gastropods were defecated without damage. Within-species variability in body size also affect seed dispersal potential, as larger individuals of the red slug (Arion rufus) swallowed more diaspores of wood anemone (Anemone nemorosa) than smaller ones.

Conclusions and Significance

Our results help to generalize the finding that gastropods consume and potentially disperse seeds of myrmecochores. The dispersal potential of gastropods is strongly influenced by diaspore size in relation to gastropod size.     

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.     

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.     

Plant colonization of a bare peat surface: population changes and spatial patterns

Abstract. Changes in size and spatial arrangement of plant populations established on an initially bare peat surface were described over a period of 5 yr by following plant individuals on a 1-cm grid in an area of 10 m x 25 m. The spatial pattern of populations and association between species was analyzed statistically. The study site was very slowly colonized by 14 perennial plant species. The early successional stage was dominated by Carex rostrata, with a clumped spatial distribution, and the homogeneously distributed Eriophorum vaginatum and Pinus sylvestris. Both the growth in size of populations and changes in their spatial distribution were interpreted as a result of species dispersal ability, tolerance to severity of the substrate and pattern of reproduction.