Seed rain and advanced regeneration in a tropical rain forest

By comparing seed rain, seedling and sapling community structures we assessed the possible role played by vertebrate seed dispersal as a structuring factor in advanced regeneration of closed-canopied sites in the tropical rain forest of Los Tuxtlas, Mexico. Seed weight, initial morphology of seedlings and species abundance were also analyzed to determine if these traits influenced the probability of establishment in the shade.About half of the seed species falling in five closed forest sites (25×25 m) during one year came from fruiting trees growing within the sites (local seeds) and half from fruiting trees found outside the sites (immigrant seeds). Seeds of liana and upper-canopy species were over-represented among immigrant seeds compared with seeds of understory tree species. This probably reflects the activity of frugivorous arboreal mammals, bats, and birds. Species with immigrant seeds had both a lower abundance and a narrower spatial distribution than locally produced seeds. Therefore, immigrant seeds showed higher diversity values than locally produced seeds.Average seed size and the proportion of epigeous seedlings were similar in local and immigrant species. Under closed-canopied sites, factors affecting community organization seem to operate selectively, favoring the establishment of large-seeded, local abundant species in the advanced regeneration. However, the fact that some saplings of immigrant species were found in the plots suggest that a slow species infiltration may be occurring leading to a slow shift in the advanced regeneration species composition. We propose that the influence of seed dispersal on advanced regeneration structure depends on the disturbance history of the patches where seeds land.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

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.     

Effects of redispersal of seeds by ants on the vegetation patternin a deciduous forest: A case study

The floristic composition and distribution of mature plants, seedlings, and soil-seed material in both ant territories and territory borders between colonies of two ant species, Lasius fuliginosus and Formica polyctena, were compared in a deciduous forest in central Ukraine. Additional seed samples were taken from ant nests. Workers of both species collect seeds of myrmecochores, but only individuals of F. polyctena relocate seeds to territory borders after the removal of elaiosomes. Borders of F. polyctena territories are thus ‘garbage dumps’ of waste organic material and probably also nutrient-enriched microsites. The floristic composition of L. fuliginosus’ territory differed from that of the territory borders and F. polyctena territory owing to a lower abundance of myrmecochores in the former. There were no great differences in the abundance or in the number of species (altogether fourteen species) in mature myrmecochores or non-myrmecochores among the sampling sites. However, the abundance and to some extent the number of species (altogether 21 species) of seedlings of myrmecochores (but not of non-myrmecochores) were lower in L. fuliginosus territory than in the other sampling sites. In particular, seedlings of two large-seeded myrmecochores, Asarum europaeum and Viola odorata, were more abundant in territory borders than in the territories. With a small-seeded myrmecochore, Ballota nigra, no such differences were found. Seeds of large-seeded myrmecochores, small-seeded myrmecochores and non-myrmecochores were more abundant than expected in the territory borders, in the nest of L. fuliginosus, in the nest of F. polyctena, respectively. It is suggested that seed flow from F. polyctena nests to the territory borders results in an increase in the seed-dispersal distances from the parent plant and among seeds dispersed in both small- and large-seeded myrmecochores. In large-seeded myrmecochores, this also results in a higher likelihood of reaching ‘garbage dumps’. The influence of ant behaviour on plant seed flow with different seed-dispersal strategies are discussed.     

Interannual variation in rainfall,drought stress and seedling mortality may mediate monodominance in tropical flooded forests

Flood tolerance is commonly regarded as the main factor explaining low diversity and monodominance in tropical swamps. In this study we examined seedling mortality in relation to seasonality, i.e., flooding versus drought, of the dominant tree species (Prioria copaifera), and three associated species (Pterocarpus officinalis, Carapa guianensis and Pentaclethra macroloba), in seasonally flooded forests (SFF) in Darien, Panama. Seedling mortality differed among species, years and seasons. Prioria seedlings experienced the lowest overall mortality, and after 3 years many more Prioria seedlings remained alive than those of any of the associated species. In general, within species, larger seedlings had greater survival. Seed size, which can vary by close to 2 orders of magnitude in Prioria, had a confounding effect with that of topography. Large-seeded Prioria seedlings experienced 1.5 times greater mortality than small-seeded seedlings, as large-seeded Prioria seedlings were more likely to be located in depressions. This finding suggests that seed size, plant size and topography are important in understanding SFF regeneration. For all species, seedling mortality was consistently greater during the dry season than during flooding. For Prioria, dry season seedling mortality was correlated with drought stress, that is, high mortality during the long El Niño dry season of 1998 and the normal dry season of 2000, but very low dry season mortality during the mild dry season of 1999. Prioria’s ability to dominate in seasonally flooded forest of Central America is partly explained by its low drought-related mortality in comparison to associated species.     

Seed size and plant strategy across the whole life cycle

We compiled information from the international literature to quantify the relationships between seed mass and survival through each of the hazards plants face between seed production and maturity. We found that small-seeded species were more abundant in the seed rain than large-seeded species. However, this numerical advantage was lost by seedling emergence. The disadvantage of small-seeded species probably results from size-selective post-dispersal seed predation, or the longer time small-seeded species spend in the soil before germination. Seedlings from large-seeded species have higher survival through a given amount of time as seedlings. However, this advantage seems to be countered by the greater time taken for large-seeded species to reach reproductive maturity: our data suggested no relationship, or perhaps a weak negative relationship, between seed size and survival from seedling emergence through to adulthood. A previous compilation showed that the inverse relationship between seed mass and the number of seeds produced per unit canopy area per year is countered by positive relationships between seed mass, plant size and plant longevity. Taken together, these data show that our old understanding of a species' seed mass as the result of a trade–off between producing a few large offspring, each with high survival probability, versus producing many small offspring each with a lower chance of successfully establishing was incomplete. It seems more likely that seed size evolves as part of a spectrum of life history traits, including plant size, plant longevity, juvenile survival rate and time to reproduction.     

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.     

Non-native grass invasion suppresses forest succession

Multiple factors can affect the process of forest succession including seed dispersal patterns, seedling survival, and environmental heterogeneity. A relatively understudied factor affecting the process of succession is invasions by non-native plants. Invasions can increase competition, alter abiotic conditions, and provide refuge for consumers. Functional traits of trees such as seed size and life history stage may mediate the effects of invasions on succession. We tested the effects of the forest invader Microstegium vimineum on planted and naturally regenerating trees in a multi-year field experiment. We established plots containing nine species of small- and large-seeded tree species planted as seeds or saplings, and experimentally added Microstegium to half of all plots. Over 3 years, Microstegium invasion had an overall negative effect on small-seeded species driven primarily by the effect on sweetgum, the most abundant small-seeded species, but did not affect large-seeded species such as hickory and oak species, which have more stored seed resources. Natural regeneration was over 400% greater in control than invaded plots for box elder, red maple, and spicebush, and box elder seedlings were 58% smaller in invaded plots. In contrast to the effects on tree seedlings, invasion did not affect tree sapling survival or growth. Microstegium may be directly reducing tree regeneration through competition. Invaded plots had greater overall herbaceous biomass in 2006 and 2008 and reduced light availability late in the growing season. Indirect effects may also be important. Invaded plots had 120% more thatch biomass, a physical barrier to seedling establishment, and significantly greater vole damage to tree saplings during 2006 and 2007. Our results show that two tree functional traits, seed size and life history stage, determined the effects of Microstegium on tree regeneration. Suppression of tree regeneration by Microstegium invasions may slow the rate of forest succession and alter tree species composition.     

Bushmeat Hunting Alters Recruitment of Large-seeded Plant Species in Central Africa

We investigated the effect of local hunting on plant recruitment at the community level in the Ngotto Forest, Central African Republic. It is the first study of this kind in the afrotropics. To compare our results with previous studies conducted in the neotropics, we used the protocol proposed byNuñez-Iturri and Howe. We compared animal relative abundances and seed length, density and diversity of seedlings at two sites with contrasting anthropogenic pressures: one with low hunting pressure (LH) and one with high hunting pressure (HH). Furthermore, we investigated how density and diversity of recruitment vary with seed length in the two sites. Both sites exhibit similar soil, climate and tree species composition, and have never been logged. Large mammals (frugivores and herbivores) were extirpated from HH and relative abundance of medium-sized frugivores drastically decreased in HH compared with LH. In HH, diversity of seedlings was reduced compared with LH, especially for large-seeded plant species dispersed by large game mammals. The approach used in this study shows promising perspectives to investigate the effects of human disturbances on the regeneration of tropical forests. Furthermore, in the afrotropical context of a lack of data on plant–animal ecological relationships, seed length appears to be a good predictor of the strength of the deficit of plant recruitment due to large mammal extirpation because of hunting. Thus, more efforts are needed to study the ecological roles of large-seeded plants in order to better understand the effects of their disappearance on tropical forest resilience.     

Tree recovery and seed dispersal by birds: Comparing forest, agroforestry and abandoned agroforestry in coastal Ecuador

We used a highly replicated study to examine vegetation characteristics between patches of intervened forest, abandoned agroforestry systems with coffee and actively managed agroforestry systems with coffee in a tropical landscape. In all habitats, plant structural characteristics, individual abundance, species richness and composition were recorded for the three plant size classes: adult trees, saplings and seedlings. Furthermore, bird species richness and composition, and seeds dispersed by birds were recorded. Tree abundance was higher in forest habitats while saplings and seedlings were more abundant in abandoned coffee sites. Although species richness of adult trees was similar in the three habitats, species richness of saplings and seedlings was much higher in forest and abandoned coffee than in managed coffee sites. However, in spite of their relatively low species richness, managed coffee sites are an important refuge for tree species common to the almost disappeared mature forest in the area. Floristic similarity for adult trees was relatively low between land use types, but clearly higher for seedlings, indicating homogenizing processes at the landscape level. More than half of the saplings and seedling were not represented by adults in the canopy layer, suggesting the importance of seed dispersal by birds between habitats. Our results show that each of the studied ecosystems plays a unique and complementary role as seed source and as habitat for tree recovery and tree diversity.     

The Influence of Variable Rainfall Frequency on Germination and Early Growth of Shade-Tolerant Dipterocarp Seedlings in Borneo

Climate change induced alterations to rainfall patterns have the potential to affect the regeneration dynamics of plant species, especially in historically everwet tropical rainforest. Differential species response to infrequent rainfall may influence seed germination and seedling establishment in turn affecting species distributions. We tested the role of watering frequency intervals (from daily to six-day watering) on the germination and the early growth of Dipterocarpaceae seedlings in Borneo. We used seeds that ranged in size from 500 to 20,000 mg in order to test the role of seed mass in mediating the effects of infrequent watering. With frequent rainfall, germination and seedling development traits bore no relationship to seed mass, but all metrics of seedling growth increased with increasing seed mass. Cumulative germination declined by 39.4% on average for all species when plants were watered at six-day intervals, and days to germination increased by 76.5% on average for all species from daily to six-day intervals. Final height and biomass declined on average in the six-day interval by 16% and 30%, respectively, but the percentage decrease in final size was greater for large-seeded species. Rooting depth per leaf area also significantly declined with seed mass indicating large-seeded species allocate relatively more biomass for leaf production. This difference in allocation provided an establishment advantage to large-seeded species when water was non-limiting but inhibited their growth under infrequent rainfall. The observed reduction in the growth of large-seeded species under infrequent rainfall would likely restrict their establishment in drier microsites associated with coarse sandy soils and ridge tops. In total, these species differences in germination and initial seedling growth indicates a possible niche axis that may help explain both current species distributions and future responses to climate change.     

Arborescent palm seed morphology and seedling distribution.

This study examines how the seed morphology of two large arborescent palms, Attalea maripa (Aubl.) Mart. and Astrocaryum aculeatum G. Mey, may affect their seed shadow in a seasonally dry Amazonian forest. In addition to being smaller and produced in larger numbers than those of A. aculeatum, A. maripa seeds also presented a substantially lower amount of nutritional reserves available for the embryo. However, A. maripa seedlings were found in much higher numbers than those of A. aculeatum. The results suggest that, within the spatial scale considered, the seed rain of A. maripa is more restricted to the area surrounding around reproductive conspecifics than that of A. aculeatum. Furthermore, in comparison with those of A. aculeatum, the smaller seeds of A. maripa might be less attractive to scatterhoarding rodents (e.g. Dasyprocta aguti). The pattern observed emphasizes the importance of scatterhoarding rodents as dispersers of large-seeded plant species in Neotropical forests.     

Effects of seed size on seedling size in Virola surinamensis; a within and between tree analysis

Summary We conducted a greenhouse study of the effects of initial seed mass on seedling characteristics in a Panamanian population of Virola surinamensis, a canopy tree in which mean seed mass of different individuals ranges from 1.34 to 4.04g. The system is of particular interest because birds preferentially eat fruits of small-seeded plants, leaving seedlings of large-seeded individuals under conditions of potentially severe sibling competition (Howe and Vande Kerckhove 1980).Effects of differences of mean seed mass between trees are explored using an analysis of variance, while effects of seed-mass variation within crops are demonstrated with a regression analysis. A two-way analysis of variance decisively shows effects of parental source and light condition on seedling height, leaf length, and dry shoot mass (all P<0.0001). A posteriori tests show that differences in seedling characteristics reflect differences in initial seed mass, with especially strong differences apparent in shoot mass. Regression of seedling characteristics on initial seed mass shows that variation of seed size within a crop is sufficient to influence shoot mass at 15 weeks (P<0.0001).Effects of size differences of seeds that land adjacent to each other, either under the parent or in monkey droppings, are documented with growth of pairs of seedlings in pots. Differences in shoot height and mass at 15 weeks are evident when seeds of average size differ by only 0.2 g, and dramatic differences are evident when paired seeds differ by an average of 1.5 g. Seedlings grow more when isolated than when planted with conspecifics.These experimental results offer indirect support for the hypothesis that small-seeded Virola parents secure an advantage in reproduction through differential dispersal, while large-seeded plants produce more competitive seedlings under their own crowns — an advantage most likely to be of importance when frugivores are scarce.     

Seed dispersal, seedling establishment and gap partitioning among tropical pioneer trees

1 We examined the abundance and distribution patterns of pioneer seeds in the soil seed bank, and of pioneer seedlings in 53 recently formed gaps, in a 50‐ha forest dynamics plot on Barro Colorado Island (BCI), Panama. The aim was to assess the importance of dispersal limitation (failure of seeds to arrive at all sites suitable for their germination) and establishment limitation (failure of seeds having reached a site to germinate successfully and establish as seedlings) in determining patterns of gap occupancy.
2 The abundance of seeds in the soil seed bank was strongly negatively correlated with seed size, but was not correlated with the abundance of reproductive‐sized adult trees in the plot. In contrast, the abundance of pioneer seedlings > 10 cm height in natural gaps was strongly correlated with adult abundance, but was not correlated with seed size.
3 Seedlings were non‐randomly distributed among gaps, but seedling abundance was not directly related to gap size, and there was no evidence of partitioning of the light environment of gaps by small seedlings. Large differences in growth and mortality rates among species were observed after 1 year, and this may result in the gap size partitioning previously found in saplings of the same species.
4 Seedlings of most species, particularly those with large seeds, were relatively more abundant than expected in gaps close to their conspecific adults. Proximity to reproductives, and by inference dispersal limitation, therefore exerts some effect on seedling distribution. None the less, large differences between seed and seedling abundances for some species, and low seedling occupancy rates in some gaps close to adult conspecifics, suggest that seedling emergence probabilities and species‐specific establishment requirements may also be important determinants of local abundance.     

An Advantage of Large Seed Size: Tolerating Rather than Succumbing to Seed Predators1

Although large seeds might be more attractive and apparent to seed predators, large seed size could enable tolerance of seed predators. If seeds are large enough to sustain damage that would kill smaller seeds yet still produce viable seedlings, investment above the minimum by the maternal plant could be advantageous. I tested this hypothesis by removing 0–80 percent of the cotyledons of four large-seeded (4–180 g) tree species from Papua New Guinea and monitoring germination and seedling growth for eight months. All species showed little negative effect on seedling size with up to 50 percent removal of cotyledons and the larger species showed a less serious effect on growth than smaller-seeded species above 50 percent removal. Large-seeded species clearly have more than the minimum-required cotyledonary reserves. Observations of viable seedlings with heavily damaged cotyledons suggest that these species withstand attack by rodents and beetles by virtue of their large size.     

Seed size,establishment sites and species coexistence in a Chilean rain forest

Abstract. Seedling densities on the forest floor and on elevated microsites (logs and stumps) were compared for eight woody species in a temperate rain forest in southern Chile. Degree of association with elevated microsites varied significantly between species, showed no systematic relationship with reported shade tolerance, but was significantly negatively correlated with seed mass. Large-seeded Podocarpus nubigena established preferentially on undisturbed forest floor sites, whereas seedlings of small-seeded species such as Nothofagus nitida and Laurelia philippiana were found mainly on fallen logs and stumps. The abundance of large seedlings and saplings of N. nitida on logs/stumps, and the growth forms of canopy trees, confirm that recruitment of this species occurs mainly on decaying wood. The relationship between seed size and microsite preferences may be caused by effects of seed size on (1) ability to establish in forest floor litter and (2) retention of seeds on logs. Seedling occupancy of logs and stumps varied with state of decay. Few seedlings of any species were present on logs in the early stages of decay. N. nitida established earlier than the other species, attaining maximum abundance on wood in the middle decay classes. Species richness and overall seedling abundance were highest on wood in advanced stages of decay. Seed size differences are suggested as a determinant of differential utilization of forest floor heterogeneity, and hence of plant species coexistence.     

Influence of litter and weather on seedling recruitment in a mixed oak-pine woodland

The effects of regular litter removal and annual variation in temperature and precipitation on seedling recruitment of species differing in their seed size and mode of dispersal were studied in a 16-year (1984-1999) experiment in a mixed oak-pine wood in southern Poland. Litter was the most important factor in determining spatial variability in seedling recruitment, whereas differences in climatic conditions among years, especially temperature fluctuations in late winter and early spring, determined the temporal variability in seedling recruitment. Compared with control plots, significantly more new individuals of bryophytes and seedlings as well as a number of new species of vascular plants were noted in the litter-removal plots over the 16-year study. Litter strongly impeded seedling emergence of small-seeded species. The negative effect of litter on seedling recruitment of large-seeded species and the recruitment of new shoots in species growing clonally was much weaker. There was a significant positive correlation between the numbers of seedlings in the litter-removal and control plots and temperatures in January to March. In the litter-removal plots this mainly affected small-seeded species. Seedling recruitment was less consistently related to variation in precipitation. Positive relationships were found only between the number of seedlings of large-seeded species in the litter-removal plots and precipitation in July of the current year and in September of the previous year, and between the number of seedlings in the control plots and precipitation in September and November of the previous year.     

Changes in seed rain across Atlantic Forest fragments in Northeast Brazil

The objectives of this study were to characterize the distribution of seeds in remnant fragments of the Atlantic Coastal Forest and to determine whether the species diversity, seed weight, and species composition of plant communities are altered by forest fragmentation. A transect of 100 m was established in the core of each of nine fragments of Atlantic Coastal Forest in a private sugarcane plantation in the state of Alagoas, NE Brazil, and ten seed-traps were distributed at intervals of 10 m each along the transects. For 12 consecutive months seeds were collected, dried, counted, weighed, and identified to species. Seeds were assigned to categories according to their size, dispersal mode, and shade tolerance. Multiple regression models and Mantel correlation tests were used to detect the effects of fragment size, percent forest cover nearby, distance from the source area, and distance from the nearest fragment on species diversity, mean seed weight, and species similarity. Analyses were carried out for all species and for subsets corresponding to each seed category. A total of 21,985 diaspores of 190 species were collected. Most seeds were small, shade-intolerant, and zoochoric, which corroborates other studies of fragmented forest landscapes and reflects the high disturbance levels in isolated forest remnants. Our data indicate that fragmentation processes such as habitat loss can alter species diversity and species composition by reducing habitat availability and increasing fragment isolation. We also found that large-seeded species are more affected by fragment isolation, possibly because their seed dispersers rarely cross non-forested areas between fragments, while zoochoric species are more strongly affected by fragment size and apparently more strongly associated with local edaphic conditions than with distance from seed sources.     

Seed germination in Chrysophyllum sp. nov., a large-seeded rainforest species in north Queensland: effects of seed size,litter depth and seed position

Leaf litter affects seed germination in many ways and past studies have shown greater impacts on relatively small seeds, both within and among species. In this shade-house experiment I examined the impact of forest litter on seed germination in Chrysophyllum sp. nov. (Sapotaceae), a large-seeded (2.4 g) rainforest tree from north Queensland. Seed mass varies more than 30-fold in this species, making it useful for studying the role of litter as a possible selective pressure in the evolution of seed size in large-seeded species. Seeds of varying size (small, medium, large) were sown in planting boxes containing one of three litter levels (low, medium, high) and placed either below or on top of the litter. Seed size and litter biomass had no significant impact on the number of germinating seeds or the time to germination but seeds placed below the litter germinated around twice as frequently, and 20% sooner, than seeds placed on top of the litter. There were no significant interactions between any of the three factors. This shade-house experiment suggests that leaf litter is not an important selective pressure in the evolution of seed size in this species. However, if litter disturbance under field conditions differentially affects the probability of seed germination in relation to seed size and/or litter biomass, then litter could still act as a selective pressure in the evolution of seed size in Chrysophyllum and other large-seeded species.     

Habitat disturbance reduces seed dispersal of a forest interior tree in a fragmented African cloud forest

Habitat fragmentation and disturbance are known to impact animals and plants in different ways, depending on species' characteristics and the type and scale of habitat modification involved. In contrast, direct or indirect ramifications on mutualistic relationships between plants and animals are less clear, possibly because general patterns are confounded by the diffuse nature of many of these interactions. Here, we examine how fragment size and/or severe disturbance of a Kenyan mountain cloud forest affects the frugivore community and seed removal of a large-seeded, bird-dispersed tree of the forest interior, for three consecutive years. Forest deterioration reduced avian visitation and seed removal rates independent of fragment size, consistently so despite strong temporal variation in fruit production over the three-year study. In disturbed forest fragments, seed removal rates were on average 3.5 times lower than in more intact ones. Strong differences in both visitation and seed removal rates were largely attributable to shifts in frugivore assemblages, characterized by loss or reduced abundance of the most effective seed dispersers, most of which were forest specialists. Although some disturbed fragments benefited from visits of non-forest dependent seed dispersers, such 'resilience' was not predictable or reliable in time or space. We conclude that disruption of seed disperser-seed interactions in highly fragmented and disturbed tropical forests may be persistent in time when resiliency is inadequate, possibly posing long-term effects on tree communities.