Recruitment of two Opuntia species invading abandoned olive groves

In Europe, many agricultural areas are now abandoned and hence can be invaded by exotic species. The abundance and spatial distribution patterns of two Opuntia species were studied in old olive groves in the Parc Natural del Cap de Creus, Catalonia (Spain). Seedling recruitment (97.3% and 51.5% of juveniles for O. maxima and O. stricta, respectively) was higher than recruitment by cladodes. O. maxima had more seedlings recruited beneath olive trees and beneath Opuntia adults than expected. Most O. stricta seedlings were also located beneath Opuntia adult plants. However, although most seedlings were recruited beneath Opuntia, some (10–30%) were found away from putative parental plants. This may be due to seed dispersal by birds and wild boars. Seeds dispersed by wild boars were not significantly more viable than seeds from intact fruits. Seedlings grow very slowly but have a high survival rate. In conclusion, Opuntia seedling recruitment is very successful and ensures the persistence of these species within old olive groves. Consequently, it prevents restoration from an agricultural land-use back to the native community.     

Cheek-pouch dispersal of seeds by Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan

Seed dispersal by Japanese monkeys (Macaca fuscata yakui) via cheek-pouch was studied in a warm temperate evergreen forest on Yakushima Island. Plant list was compiled based on a study during 1986–1995, of which troops of monkeys have been habituated without artificial feeding. We followed the well-habituated monkeys in 1993 and 1994 to observe the feeding behavior and their treatments of fruits and seeds, and collected seeds dispersed by monkeys to record the distance carried from the mother trees. We checked the difference of germination ratio between seeds dispersed via cheek-pouch and seeds taken from mother trees by sowing experiments. Seeds and acorns of 22 species were observed to be dispersed via cheek-pouch of monkeys. Among them, three species with acorns were never dispersed via feces, and 15 species with drupes were seldom dispersed via feces. Plant species of which seeds are dispersed only via cheek-pouch had larger seeds than those of dispersed both via cheek-pouch and via feces, and typically had only one or two seeds in a fruit. As for one of cheek-pouch dispersal species,Persea thunbergii, the mean distance when seeds were carried from the mother trees via cheek-pouch was 19.7 m, and the maximum distance was as long as 105 m although more than 80% of seeds were dispersed within 30 m from mother trees. And 82% of seeds dispersed via cheek-pouch germinated. The easy separation of seeds from other parts of the fruit seems to facilitate cheek-pouch dispersal more than dispersal via feces. Cheek-pouch dispersal by monkeys has possibly enhanced the natural selection for larger seeds which bring forth larger seedlings with high shade-tolerance. In conclusion, cheek-pouch dispersal by monkeys is quite an important mode for trees in the mature stand in a warm temperate evergreen forest on Yakushima Island.     

Population structure and fruit production of Pyrus bourgaeana D. are affected by land-use

The Iberian wild pear (Pyrus bourgaeana D.) is a rare, fleshy-fruited tree restricted to dehesas and evergreen sclerophyllous Mediterranean forests in the southwestern Iberian Peninsula. It produces palatable fruits and leaves attractive to different species groups, playing an important trophic role in the ecological networks of Mediterranean ecosystems. However, the intensification in the traditional land-use linked to these areas could threaten the stability of the wild pear populations in the short/medium-term. In order to determine the population dynamics of this relevant species in relation to the land-use history, we selected two populations (southern Spain) subjected to different land-use management, dehesa (D) and abandoned olive grove (AOG). An analysis of 122 adult trees reported an overall density of 0.6 trees ha⁻¹. The tree age was estimated by tree-rings analysis in all adult trees. Dendrometric parameters, reproductive features, and germination rates were also measured. Regeneration was clearly biased, as evidenced by the truncated age structure. A low correlation (R² = 34%) between age and DBH (diameter at breast height) (244 cores analysed) showed that diameter seems not to be a reliable predictor of tree age. Trees from AOG populations had significantly-higher values of DBH, height and crown diameter, but were less productive in terms of fruits and seeds. Nested analysis of variance showed significant variation in fruit production, fruit size, dry mass, water content and seed viability. There were also significant differences in masting. No evidence was found to demonstrate that fruit production, seed viability, or germination rate influence the low natural recruitment of this species. These findings indicate that the traditional agrosilvopastoral practices carried out in the study area for decades, and its subsequent intensification, have strongly influenced the ecological structure of the Iberian wild pear populations at the local scale, which could compromise its stability in the near future.     

Pigeons as seed predators and dispersers of figs in a Malaysian lowland forest

Seven pigeon species fed on figs from at least 22 Ficus taxa at a lowland forest site in Peninsular Malaysia. Forest pigeons of the genus Treron are fig-eating specialists. Although these are important fig-seed predators, at least some of the population, and pigeons from other genera, potentially disperse Ficus seeds to distant sites.     

Fruit size preference in the New Zealand pigeon (Hemiphaga novaeseelandiae)

Abstract We investigated whether the New Zealand pigeon Hemiphaga novaeseelandiae (Columbidae) exhibits size‐based preferences for fruits. We tested the hypothesis that in small‐fruited species, pigeons would prefer larger fruits, but in larger‐fruited species, this preference would reverse as the pigeons become increasingly limited by their gape size. We collected undispersed fruits and bird‐dispersed seeds of 10 plant species, some over several sites or years (13 datasets in total). We estimated the fruit size of dispersed seeds by fitting regressions of fruit diameter to seed diameter in intact fruits. We were able to predict fruit diameter from seed diameter in 12 of the 13 populations, although the relationship was stronger in single‐seeded species than in multi‐seeded species. Seven of the 12 populations tested showed a significant difference in seed diameter among undispersed and dispersed seeds. However, our results showed no consistent pattern in fruit size preference by the New Zealand pigeon and did not support our hypothesis. The large‐bodied New Zealand pigeon is generally not gape limited and fruit size preferences appear to be independent of mean fruit size.     

Plants,position and pollination: Planting arrangement and pollination limitation in a revegetated eucalypt woodland

The spatial arrangement of plants is a key determinant of pollination services in natural ecosystems. Despite this, plant arrangements are rarely considered an important characteristic of revegetated communities and this may be limiting successful pollination dynamics in these re‐created systems. We assessed if aggregated South Australian Blue Gum (Eucalyptus leucoxylon, F. Muell) had greater pollination success than more dispersed trees by measuring fruit set, seed production and germination in a revegetated eucalypt woodland. The proportion of buds that developed into fruit (fruit set) was similar between aggregated (five flowering Blue Gum within 30 m) and dispersed (no flowering Blue Gum within 30 m) trees (<14%). Aggregated and dispersed trees produced a similar number of seeds per fruit in 2015, when fewer trees flowered and those that flowered produced fewer flowers (ca 8 seeds/fruit). In 2016, when flowering was more abundant, aggregated trees produced more seeds per fruit than dispersed trees (12 seeds/fruit compared to 4 seeds/fruit). Despite differences in seed number, there was no difference in the quality of seed produced, indicating outcrossed pollen was received by both aggregated and dispersed trees. Although outcrossing was likely, this did not prevent pollination limitation, with the addition of outcrossed pollen increasing fruit and seed set for trees in both arrangements. Consequently, we propose that planting individuals of the same species (conspecifics) in clumps, rather than intermixing species, can increase seed set in revegetated eucalypts, particularly in good flowering years. However, aggregated plantings will be most effective if the needs and foraging behaviour of pollinators are also considered, to attract and sustain pollinators in revegetated systems, thereby minimizing the risk of pollination limitation.     

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.     

Movement Patterns of a Native and Non‐native Frugivore in Hawaii and Implications for Seed Dispersal

Avian frugivores historically played important roles as seed dispersers across the Hawaiian Islands, but presently, the ‘ōma‘o (Myadestes obscurus) is the only extant native frugivore in the wild on the Island of Hawaii. During recent decades, the introduced generalist Japanese White‐eye (Zosterops japonicus) has become the most common bird in Hawaii. The movements of avian frugivores largely dictate how far seeds get dispersed and into what kinds of microhabitats. This study compares the movement patterns and diet of the ‘ōma‘o to the Japanese White‐eye to understand how a native differs from a non‐native frugivore in the type and distances of seeds dispersed. Radiotelemetry was conducted on nine ‘ōma‘o and nine Japanese White‐eyes in a system of natural forest fragments (kīpuka) created by lava flows. Japanese White‐eyes disperse seeds approximately twice as far as ‘ōma‘o; during the time of gut passage, ‘ōma‘o move a mean distance of 98.1 m, and Japanese White‐eyes move 170.1–194.8 m. However, the ‘ōma‘o disperses the seeds of at least seven different native fruit species compared with two dispersed by Japanese White‐eyes. Japanese White‐eyes were found to disperse seeds smaller than 1.5 mm, whereas the ‘ōma‘o dispersed seeds up to 6 mm in diameter. Despite their ecological differences, both birds distribute certain seeds within and among kīpuka and likely facilitate primary succession of fruiting plants in the young lava matrix. However, this study suggests that if the ‘ōma‘o were extirpated, a smaller‐bodied generalist cannot entirely substitute for the ecological role played by the native frugivore.     

The effect of bat (Rousettus aegyptiacus) dispersal on seed germination in eastern Mediterranean habitats

The fruit-bat Rousettus aegyptiacus (Pteropodidae) in Israel consumes a variety of cultivated and wild fruits. The aim of this study was to explore some of its qualities as a dispersal agent for six fruit-bearing plant species. The feeding roosts of the fruit-bat are located an average of 30 m from its feeding trees and thus the bats disperse the seeds away from the shade of the parent canopy. The bat spits out large seeds but may pass some (2%) of the small seeds (<4 mg) through its digestive tract. However, neither the deposited seeds nor the ejected seeds (except in one case) had a significantly higher percentage germinating than intact seeds. Although the fruit-bat did not increase the percentage germinating, seeds of three plant species subject to different feeding behaviors (deposited in feces or spat out as ejecta) had a different temporal pattern of germination from the intact seeds. The combined seed germination distribution generated by these different treatments is more even over time than for each treatment alone. It is sugested that this increases asynchronous germination and therefore enhances plant fitness by spreading the risks encountered during germination, especially in eastern Mediterranean habitats where the pattern of rainfall is unpredictable.     

Microsatellite analysis of the maternal origin of <Emphasis Type="Italic">Myrica rubra</Emphasis> seeds in the feces of Japanese macaques

Identifying the maternal origin of dispersed seeds is a challenging task because it is impossible to directly track seed movement once an animal has ingested them. However, recent development of molecular techniques allows us to identify the maternal origin of dispersed seeds in natural plant populations. Here we analyzed the maternal origin of Myrica rubra seeds found in the feces of Yakushima macaques. We detected a high level of diversity among the dispersed seeds, with an average of 3.0 maternal origins (range 1–9) in each feces. The average dispersal distance was 270.0 m (range 20.4–634.0 m) and the average of distance between maternal trees of dispersed seeds in the same feces was 161.5 m (range 0–573.9 m). These results suggest that seed dispersal by macaques plays an important role in maintaining the genetic diversity of Myrica rubra populations.     

Artificial selection, naturalization, and fitness: Darwin's pigeons revisited

Central in biology is the view that the commonly observed association between the phenotype and the ecology of organisms is the result of the process of natural selection. There are numerous examples where the current and/or past ecological pressures in which a trait presumably arose have been demonstrated. Less well documented, however, are cases where a species that has artificially been moved from its adaptive peak is seen to return to its ancestral state when exposed to the ancient selective regime. One of the few suggested cases is that of the feral pigeon ( Columba livia ), the free-living descends from domesticated, artificially selected rock pigeons. Contemporary feral pigeons are significantly closer in morphology to wild rock pigeons than to their more direct domestic ancestors, suggesting that natural selection has been reconstituting their wild phenotype. Because they still preserve some characters engendered through artificial selection during their ancestral period of domesticity, notably a tarsus that is disproportionally long for a terrestrial pigeon of its size, feral pigeons provide a rare opportunity to test whether natural selection is the process behind the reconstitution of the wild phenotype. In the present study, it is shown that feral pigeons with tarsi that depart the most from the wild phenotype appear to have limitations in the use of some food resources and, as a result, experience lower survival fitness. These findings support the view that natural selection is reconstituting the wild phenotype in feral pigeons. The possibility nonetheless remains that this process will never be fully completed.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 657–665.     

Divergence, convergence, and the ancestry of feral populations in the domestic rock pigeon

Domestic pigeons are spectacularly diverse and exhibit variation in more traits than any other bird species [1]. In The Origin of Species, Charles Darwin repeatedly calls attention to the striking variation among domestic pigeon breeds-generated by thousands of years of artificial selection on a single species by human breeders-as a model for the process of natural divergence among wild populations and species [2]. Darwin proposed a morphology-based classification of domestic pigeon breeds [3], but the relationships among major groups of breeds and their geographic origins remain poorly understood [4, 5]. We used a large, geographically diverse sample of 361 individuals from 70 domestic pigeon breeds and two free-living populations to determine genetic relationships within this species. We found unexpected relationships among phenotypically divergent breeds as well as convergent evolution of derived traits among several breed groups. Our findings also illuminate the geographic origins of breed groups in India and the Middle East and suggest that racing breeds have made substantial contributions to feral pigeon populations.     

Fruit tracking,frugivore satiation,and their consequences for seed dispersal

Vertebrate frugivore communities are easily satiated by abundant fruit supplies and, contrary to abiotic dispersal agents, typically disperse only part of the available seed pool. This frugivore satiation is likely to be a widespread phenomenon and should be an influential predictor of plants’ ability to disperse their offspring to suitable establishment sites; yet it has never been systematically quantified. Here I investigate patterns of fruit abundance, frugivore activity and frugivore satiation, and their consequences for seed dispersal in the fleshy-fruited tree Frangula alnus. Based on constant-effort seed trapping conducted over 3 years, I assess densities of total and frugivore-consumed seedfall across two spatial (within/between populations) and two temporal (within/between ripening seasons) scales. Furthermore, I examine relationships between fruit abundance and the amount of seeds that are actually dispersed away from fruiting trees. Frugivore activity tightly matched fruit abundance, although some differences existed between scales. This marked fruit tracking did not prevent a significant frugivore satiation, however, and only 53% of the available fruit crops were actually consumed. The extent of satiation varied most at the within-population level, likely due to the territorial behaviour of important frugivore species. In contrast, levels of satiation remained remarkably invariable through time, suggesting that frugivores behave as opportunists and closely adjust the composition of their diet to the available food supply. Overall, greater fruit abundance resulted in a higher proportion of seeds falling beneath fruiting trees, but it also helped increase the (absolute) number of seeds dispersed. This study shows that frugivore satiation can be an important phenomenon even when frugivores tightly track fruit abundance. Its negative effects on recruitment may be attenuated, however, if greater fruit crops help increase population-wide frugivore activity and the amount of seeds being dispersed to suitable establishment sites.     

Seed retention times in the New Zealand pigeon,

We offered ripe fruits of tawa (Beilschmiedia tawa), taraire (B. tarairi), and pūriri (Vitex lucens) to captive New Zealand pigeons (Hemiphaga novaeseelandiae) and recorded seed retention times. We also recorded seed retention times while radio-tracking wild pigeons in Taranaki and Canterbury. We report wild pigeon retention times for tawa, pūriri, miro (Prumnopitys ferruginea), fivefinger (Pseudopanax arboreus), and kahikatea (Dacrycarpus dacrydioides) seeds. Where data were available for the same plant species from wild and captive pigeons, retention times were similar. Seed retention time differed significantly among fruit species, and was positively related to seed mass. Mean retention times ranged from 37-45 min for the two smallest-seeded species (fivefinger and kahikatea) up to 109-181 min for the three largest species (pūriri, taraire, and tawa). We also report the second published instance of regurgitation by the New Zealand pigeon.     

Die Taube – eine Erfolgsgeschichte. Vom Liebling der Götter zur Eroberung der Städte

The pigeon – a success story The pigeon is one of the first animals that man domesticated and it holds a special place in man's affections. The pigeon is far more than an ordinary bird since it is a symbol of different gods and human virtues. In the beginning, the pigeon used to be an ordinary biological creature until it found it's way into man's spiritual world. These associations contribute to man's love and admiration of pigeons. The feral pigeon is a wild descendant of dovecote pigeons and domestic breeds. All experiences up to now lead to the conclusion that reduction of pigeon populations can only be achieved by reducing their food supply. Because of the enormous reproduction rate of the pigeons, killing attempts without any simultaneous food reduction do not have any influence on the number of birds in the population.     

Agriculture as matchmaker of an unexpected mutualism: Great bustard disperses and enhances emergence of domestic olive seeds

By changing the habitats and altering plant traits, agriculture has severely disrupted many plant–animal mutualisms. Interestingly, however, the intensification of agricultural practices could also facilitate mutualistic relationships between species with naturally mismatching phenotypes. We illustrate the potential of the great bustard (Otis tarda), a large steppe bird, as disperser of domestic olive (Olea europaea) seeds, originally a forest species. In an area of southwestern Spain, 30% of bustard faeces included olive stones (from 1 to 13). Only 1.7% of the bustard-ingested olive seeds were broken. Moreover, using a sowing experiment, we show bustard ingestion enhanced seedling emergence, which reached 8.8%, 3.4% and 0.0% for bustard-ingested, hand-depulped, and control seeds, respectively. As expected for Mediterranean habitats, seedling mortality was very high in the first summer for all seed treatments. In 6 out of 19 non-plowed patches within our study area, we found olive saplings of different ages likely to emerge from bustard-dispersed seeds. Given the large size of domestic olive fruits, bustards are among the few local animals able to disperse their seeds and thus to assist in the forestation of field boundaries and abandoned lands. Paradoxically, because bustards are rather restricted to open habitats, their success in shaping the habitat (i.e., ‘planting’ olive trees) could lead to their own removal from the resulting forested landscape.     

Mammalian seed dispersal in Cantabrian woodland pastures: Network structure and response to forest loss

We evaluated the role of wild large mammals as dispersers of fleshy-fruited woody plants in woodland pastures of the Cantabrian range (N Spain). By searching for seeds in mammal scats across four localities, we addressed how extensive seed dispersal was in relation to the fleshy-fruited plant community, and applied a network approach to identify the relative role of mammal species in the seed dispersal process. We also tested the response of mammalian dispersers to forest availability at increasing spatial scales. Five carnivores and three ungulates dispersed seeds of eight fleshy-fruited trees and shrubs. Mammalian seed dispersal did not mirror community-wide fruit availability, as abundant fruiting trees were scarce whereas thorny shrubs were over-represented among dispersed species. The dispersal network was dominated by bramble (Rubus ulmifolius/fruticosus), the remaining plants being rarer and showing more restricted disperser coteries. Fox (Vulpes vulpes), badger (Meles meles), and wild boar (Sus scrofa) dispersed mostly bramble, whereas martens (Martes sp.) dispersed mostly wild rose (Rosa sp.). Ungulates occasionally dispersed holly (Ilex aquifolium) and hawthorn (Crataegus monogyna). The empirical network reflected a skewed distribution of interactions and some functional complementarity (as judged from the low levels of connectance and nestedness), but also some degree of specialization. Mammals overused uncovered microsites for seed deposition, and increased their disperser activity in those landscape sectors devoid of forest. Combined with previous findings on avian seed dispersal, this study suggest a strong functional complementarity coming from the low overlap in the main plant types that mammals and birds disperse – thorny shrubs and trees, respectively – and the differential patterns of seed deposition, with mammals mostly dispersing into deforested areas, and birds into forest-rich landscapes.     

Seasonal Variation in Seed Dispersal by Tamarins Alters Seed Rain in a Secondary Rain Forest

Reduced dispersal of large seeds into degraded areas is one of the major factors limiting rain forest regeneration, as many seed dispersers capable of transporting large seeds avoid these sites with a limited forest cover. However, the small size of tamarins allows them to use small trees, and hence to disperse seeds into young secondary forests. Seasonal variations in diet and home range use might modify their contribution to forest regeneration through an impact on the seed rain. For a 2-yr period, we followed a mixed-species group of tamarins in Peru to determine how their role as seed dispersers in a 9-yr-old secondary-growth forest varied across seasons. These tamarins dispersed small to large seeds of 166 tree species, 63 of which were into a degraded area. Tamarins’ efficiency in dispersing seeds from primary to secondary forest varied across seasons. During the late wet season, high dietary diversity and long forays in secondary forest allowed them to disperse large seeds involved in later stages of regeneration. This occurred precisely when tamarins spent a more equal amount of time eating a high diversity of fruit species in primary forest and pioneer species in secondary forest. We hypothesized that well-balanced fruit availability induced the movement of seed dispersers between these 2 habitats. The noteworthy number of large-seeded plant species dispersed by such small primates suggests that tamarins play an important, but previously neglected, role in the regeneration and maintenance of forest structure.     

Chloroplast-DNA variation in cultivated and wild olive (Olea europaea L.)

Polymorphism in the lengths of restriction fragments of the whole cpDNA molecule was studied in cultivated olive and in oleaster (wild olive) over the whole Mediterranean Basin. Seventy two olive cultivars, 89 very old trees cultivated locally, and 101 oleasters were scored for ten endonucleases. Moreover, maternal inheritance of cpDNA in olive was shown by analysing the progeny of a controlled cross between two parents which differed in their cpDNA haplotypes. In the whole species, three site- and three length-mutations were observed, corresponding to five distinct chlorotypes. The same chlorotype (I) was predominant in both oleasters and cultivated olive trees, confirming that these are closely related maternally. Three other chlorotypes (II, III and IV) were observed exclusively in oleaster material and were restricted either to isolated forest populations or to a few individuals growing in mixture with olive trees possessing the majority chlorotype. An additional chlorotype (V) was characterised by three mutations located in distinct parts the cpDNA molecule but which were never observed to occur separately. This chlorotype, more widely distributed than the other three, in both cultivated and wild olive, and occurring even in distant populations, was observed exclusively in male-sterile trees showing the same specific pollen anomaly. However, in the present study, no evidence was provided for a direct relationship between the occurrence of the cpDNA mutations and male sterility. It is suggested that the large geographic distribution of chlorotype V may be related to the high fruit production usually observed on male-sterile trees. These may be very attractive for birds which are fond of olive fruit and spread the stones efficiently. Probably for the same reason, people preserved male-sterile oleasters for long periods and, in several places, used male-sterile cultivars over large areas. Received: 25 November 1998 / Accepted: 19 December 1998     

Disentangling the effects of facilitation on restoration of the Atlantic Forest

Facilitation is an important ecological mechanism with potential applications to forest restoration. We hypothesized that different facilitation treatments, distance from the forest edge and time since initiation of the experiment would affect forest restoration on abandoned pastures. Seed and seedling abundance, species richness and composition were recorded monthly during two years under isolated trees, bird perches and in open pasture. Seed arrival and seedling establishment were measured at 10 m and 300 m from the forest edge. We sampled a total of 131,826 seeds from 115 species and 487 seedlings from 46 species. Isolated trees and bird perches increased re-establishment of forest species; however, species richness was higher under isolated trees. Overall, abundance and richness of seeds and seedlings differed between sampling years, but was unaffected by distance from the forest edge. On the other hand, species composition of seeds and seedlings differed among facilitation treatments, distance from the forest edge and between years. Seedling establishment success rate was larger in large-seeded species than medium- and small-seeded species. Our results suggest that isolated trees enhance forest re-establishment, while bird perches provide a complementary effort to restore tree abundance in abandoned pastures. However, the importance of seed arrival facilitation shifts toward establishment facilitation over time. Arriving species may vary depending on the distance from the forest edge and disperser attractors. Efforts to restore tropical forests on abandoned pastures should take into account a combination of both restoration strategies, effects of time and proximity to forest edge to maximize regeneration.