Positive relationship between fruit removal by animals and seedling recruitment in a tropical forest

Fleshy-fruited plants rely on animal frugivores to disperse their seeds, and seed removal by frugivores may leave an imprint on seedling recruitment. However, to what extent plant–frugivore interactions are related to seedling recruitment has rarely been quantified at the community level, especially in species-rich tropical forests. In this study, we tested the effect of different plant traits on fruit removal by frugivores and tested the relative importance of fruit removal, plant traits and abiotic factors for seedling recruitment. We quantified plant–frugivore interactions of 22 fleshy-fruited plant species consumed by 56 diurnal frugivore species, and counted the number of seedlings that emerged along an elevational gradient in the Colombian Andes. We measured a set of plant traits (i.e., crop size; fruit size; seed load and mass; fruit nutritional contents), estimated the density of adult plants and recorded relevant abiotic factors (light, temperature and humidity). We found that fruit removal by frugivores was positively associated with crop size, but negatively associated with fruit length and unrelated to seed load and fruit nutritional content. Seedling densities were positively related to the density of adult plants, seed mass and fruit removal by animals. We found no relationship between abiotic factors and seedling recruitment. Our results indicate that fruit abundance and morphology are important determinants of fruit removal and that fruit removal is positively associated with seedling recruitment accounting for effects of species abundance and plant traits. We conclude that plant traits shape fruit removal and seedling recruitment at the community level, while these two crucial processes of forest regeneration are directly linked by seed dispersal of animals.     

Seeds in droppings of tropical fruit-eating birds: importance of considering seed composition

Summary The probability of interspecific seedling competition from bird droppings is high because fruit-eating birds, on average, deposit more than two species in a single dropping. Moreover, birds vary both in the number of plant species they deposit in a given dropping and in the seed composition of those droppings. In a preliminary experiment, I examined effects of interspecific seedling competition from seeds found in bird droppings. Certain plant species were competitively superior in pairwise growth experiments using six common second-growth species. The survival of certain shrubs depended on which species it was disseminated with in bird droppings. Rapid germination time may promote competitive superiority in some cases. Birds affect plant fitness not only because of their behavior following dispersal, but also because they deposit seeds in different densities and combinations.     

Ecology of seed germination of eight non-pioneer tree species from a tropical seasonal rain forest in southwest China

We compared various aspects of the seed biology of eight non-pioneer tree species from a tropical seasonal rain forest in Xishuangbanna, SW China, that differ in time of dispersal, size and fresh seed moisture content (MC). Seeds were tested for germination under laboratory conditions after dehydration to different moisture levels and under 3.5, 10 and 30% solar irradiances in neutral-shade houses. For six species, germination was also compared in forest understory (3.5% light) and center of a forest gap (32.5% light). Under continuous dehydration over activated silica gel, 100% of seeds of four species had lost the ability to germinate after 48 h, and those of all species except Castanopsis hystrix (decreased from >90 to 30% germination) had lost the ability to germinate after 120 h. Four species did not differ in final germination percentages at the three irradiances (i.e. uniform germination). However, final germination percentages of Horsfieldia pandurifolia and Litsea pierrei var. szemaois were significantly lower in 30% than in 10 or 3.5% light, and seeds of Antiaris toxicaria and C. hystrix germinated to higher percentages in 30 and 10% than in 3.5% light. Mean time to germination (MTG) of the eight species (forest and shade house data combined) ranged from 5–5 days for Pometia tomentosa to 72–207days for L. pierrei; MTG for four species was ≤21 days. There was no obvious relationship between relative desiccation resistance and either time of dispersal, MTG or uniformity of germination at the three light levels, or between seed size and MC or MTG. However, the relationship between seed MC at maturity (25–60% fresh mass basis) and MC at 50% loss of seed viability (12.4–42.5%) was significant. Seven of the species fit Garwood’s (Ecol Monogr 53:159–181, 1983) rapid-rainy germination syndrome and one, L. pierrei, either her delayed-rainy or intermediate-dry germination syndrome. However, fresh, non-dehydrated seeds of all eight species germinated in ≤30 days at constant 30°C in light.     

Community seed rain patterns and a comparison to adult community structure in a West African tropical forest

We examined the seed rain throughout a twelve month period in a lowlandtropical forest in Cameroon, West Africa, 1996–97. Traps (0.5m², n = 216) were erected throughout a 25km² area in 24 randomly placed clusters of nine trapseach. Fruits and seeds that landed in traps were collected every 7–10daysand classified by species and dispersal type. More than 32,000 seeds fromapproximately 200 species fell into the traps, an average of 297 seedsm^–2 yr^–1 Thirty species represent 82%of the total seed rain while an additional 175 species comprise the remaining18%. When we compared the adult community to the seed rain community within thesame plots, we found no apparent correlation between seed rain patterns andadult community structure for this year of study. Furthermore, only 49% of theadult tree community produced and dispersed seed into traps in this year. Morethan 100 species (52%) found in the seed rain represented long-distanceimmigrant seed rain. Seed rain was highly variable at several scales, bothspatially and seasonally, although seeds arrived in traps during eachcollectionperiod. Cluster analyses showed that traps within plots were seldom moresimilarto one another than traps between plots. While 82% of the tree species in thecommunity are thought to be animal dispersed, only 28% of all seeds that fellinto traps had been obviously handled (bitten, chewed, or passed)by animals. Tests for fruit and seed removal by predators or dispersers found5%or less removal rate from traps.     

Small-scale patterning in the recruitment of forest species during succession in tropical dry forest,Mozambique

Within a few years of field abandonment on the coastal dunes, a considerable number of species of trees and woody lianes of the climax evergreen forest enter the succession. At a relatively large-scale of analysis (a 100 m² plot), there is no evidence of soil development during the early succession, with no accumulation of soil organic matter nor nutrients being detected. At this scale there is, therefore, little support for the facilitation model of succession. However, when smaller scales of analysis are employed (e.g. a plot equivalent to the canopy area of a sapling) a very different picture emerges; most recruitment of forest woody species occurs beneath saplings found in early succession. Nutrient concentrations were higher in soils under sapling canopies than in soils beyond sapling canopies. With increasing sapling size, an increasing number of seedlings of forest woody species were found and there is progressive soil amelioration. Thus, during the early succession, there is considerable small-scale patterning in soil development and recruitment. This pattern is largely established through bird dispersal, with the intensity of recruitment being dependent on the distance to seed sources.     

都江堰林区取食樱桃果实(种子)的鸟类及其种子扩散作用

樱桃(Prunus pseudocerasus)是广泛分布于我国亚热带常绿阔叶林内的一种重要核果植物。为了解食果鸟类在樱桃种群更新中的作用,于2007年和2008年在四川都江堰亚热带常绿阔叶林内研究了取食樱桃果实(种子)的鸟类及其种子扩散作用。研究表明,樱桃成熟果实的下落高峰发生在4月下旬至5月上旬;2007年的种子扩散率为4.0%±1.0%,明显低于2008年(27.7%±5.7%)。在研究地内,发现至少有16种鸟取食樱桃果实或种子,根据其对果实和种子的处理方式分为3个功能群:白头鹎(Pycnontus sinensis)、领雀嘴鹎(Spizixos semitorques)、黑鹎(Hypisipetes leucocephalus)、白颊噪鹛(Garrulax snnio)、红嘴蓝鹊(Urocissa erythorhyncha)等10种鸟吞食樱桃果实,而种子通过消化道末端排出并将种子携至远离母树的地方,是重要的种子扩散者;暗绿绣眼鸟(Alcippe morrisonia)和灰眶雀鹛(Zosterops iaponicus)等4种鸟主要啄食果肉而将种子丢弃在母树下,为啄食果肉者;而普通朱雀(Carpopacus erythrinus)和灰头鸦雀(Paradoxornis gularis)则主要取食种子,为纯粹的种子捕食者。在吞食樱桃果实的食果鸟中,3种鹎科鸟类访问频次所占的比例达55.3%(2007年)和35.3%(2008年),说明鹎科鸟类是都江堰林区樱桃种子的主要扩散者,对樱桃种群的空间格局和自然更新可能有重要影响。     

Seed predation and seedling recruitment in plants: the effect of the distance between parents

We present a graphic model that explores the effect of distance between parent plants on seed predation and seedling recruitment. Based on the assumption that distance between parents may affect the shape of the seed shadow, the model predicts that seed predators may affect seedling recruitment curves under isolated plants but they are unable to affect these curves under close parent plants. The predictions of the model are tested experimentally in Cryptocarya alba (Lauraceae), a common tree of the Mediterranean forest, Central Chile. Results show that predictions are not met under isolated parent plants. Although seed density decreases significantly away from parent plants, this effect is not relevant for seed predation and seedling recruitment. The biotic/abiotic contrast existing under the canopy vs outside the canopy, plus the shade-tolerance of this tree, better explains the seedling recruitment observed under isolated parent plants. Nevertheless, the predictions of the model are corroborated under close parent plants. Seed shadows overlap to the extent that they generate a homogeneous seed distribution, homogeneous seed predation and homogeneous seedling recruitment as well. We discuss the implications of the model in terms of the spatial pattern of seedlings and the benefits of dispersal from isolated and close parent plants.     

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.     

Edge effects in a tropical montane forest mosaic: experimental tests of post-dispersal acorn removal

Forest edges created by scattered-patch clear-cutting have become common in tropical montane cloud forests in the highlands of Chiapas, Mexico. It was hypothesised that forest edges may influence regeneration of oak species, which are canopy dominants in these forests, by affecting the activities of small mammal species. Acorns of different oak species varying in germination timing were offered to predators and/or dispersers at different positions along replicated forest edges during 2 consecutive years. We investigated the effects of (1) edge type (hard and soft), (2) distance from the edge (0, 15, 30, 45 and 60 m inside forest fragments) and (3) oak species, on the rate of acorn removal mainly by small mammals. During a non-masting year, acorn removal was affected by the interaction of edge type and distance from the edge (P<0.05), with acorn removal being highest near hard edges compared to adjacent forest interiors. As predicted, acorn removal was greater along soft (100%) than along hard edges (82%), but this pattern was recorded only during the non-masting year. This study partly supports previous studies of rodents preferentially consuming acorns with early germination rather than acorns exhibiting dormancy, however these patterns may change with variation in acorn abundance. These results suggest that patch clear-cutting affects regeneration processes within forest fragments by influencing the activities of small mammals, but the nature of this effect also depends on acorn abundance and the characteristics of the forest edge created.     

Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

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

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

Seedling establishment in a masting desert shrub parallels the pattern for forest trees

The masting phenomenon along with its accompanying suite of seedling adaptive traits has been well studied in forest trees but has rarely been examined in desert shrubs. Blackbrush (Coleogyne ramosissima) is a regionally dominant North American desert shrub whose seeds are produced in mast events and scatter-hoarded by rodents. We followed the fate of seedlings in intact stands vs. small-scale disturbances at four contrasting sites for nine growing seasons following emergence after a mast year. The primary cause of first-year mortality was post-emergence cache excavation and seedling predation, with contrasting impacts at sites with different heteromyid rodent seed predators. Long-term establishment patterns were strongly affected by rodent activity in the weeks following emergence. Survivorship curves generally showed decreased mortality risk with age but differed among sites even after the first year. There were no detectable effects of inter-annual precipitation variability or site climatic differences on survival. Intraspecific competition from conspecific adults had strong impacts on survival and growth, both of which were higher on small-scale disturbances, but similar in openings and under shrub crowns in intact stands. This suggests that adult plants preempted soil resources in the interspaces. Aside from effects on seedling predation, there was little evidence for facilitation or interference beneath adult plant crowns. Plants in intact stands were still small and clearly juvenile after nine years, showing that blackbrush forms cohorts of suppressed plants similar to the seedling banks of closed forests. Seedling banks function in the absence of a persistent seed bank in replacement after adult plant death (gap formation), which is temporally uncoupled from masting and associated recruitment events. This study demonstrates that the seedling establishment syndrome associated with masting has evolved in desert shrublands as well as in forests.     

The effect of a seed source on primary succession in a forest ecosystem

An analysis of the Relative Importance Value Index (RIVI) of later successional tree species and the Shannon diversity indexH of all tree and shrub species was undertaken on previously mined sites within a sparsely forested area in central Florida, USA. A comparative analysis of the distance to a seed source and the age of a site suggested that the distance to a seed source was the best predictor (R ²=0.85) of the regeneration of the later successional species and a good predictor of species diversity. Both theRIVI of the later successional species and the diversity index decreased with the distance from the seed source. The lack of a seed source containing climax species resulted in arrested succession at some sites. It is suggested that biological information is the causative means of succession and the dispersal of this information entails spatial dependency while its introduction and development are time dependent processes.Research was part of a project of the Center for Wetlands, University of Florida, Gainesville under contract with the Florida Institute of Phosphate Research. Interactions of Wetlands with Phosphate Mining, H. T. Odum and G. R. Best, principal investigators. B. T. Rushton, J. Butner, G. R. Best, R. S. Schnoes and S. R. Humphrey are thanked for the use of their data.     

Decreased frugivory and seed germination rate do not reduce seedling recruitment rates of <Emphasis Type="Italic">Aristotelia chilensis</Emphasis> in a fragmented forest

Habitat fragmentation reduces frugivorous bird abundance. Such a reduction may lead to a reduction in seed dispersal, thereby compromising seedling recruitment rate with far reaching consequences for plant population persistence. We assessed frugivory, seed germination, and seedling recruitment rates in a fragmented forest of central Chile by comparing a continuous forest with four forest fragments surrounded by pine plantations. Frugivory was 2.4 times higher in continuous forest than in forest fragments. Seeds eaten by birds germinated 1.7 and 3.7 times higher than non-eaten seeds from continuous forest and fragments respectively. Non-eaten seeds from continuous forest germinated 2.2 times higher than those from forest fragments, suggesting inbreeding depression. However, seedling recruitment rates at forest fragments were far higher than in continuous forest where no seedling recruited in the five years analysed. Therefore, despite forest fragmentation negatively affected frugivory, it did not translate into a decreased fitness of plants, thus highlighting the importance of considering the overall processes leading the reproductive success of plants following anthropogenic disturbances.     

Dispersal traits determine passive restoration trajectory of a Nigerian montane forest

Passive restoration methods offer great promise for tropical regions where resources are limited but the success of such efforts can be variable. Using trait-based theory, we investigated the likely trajectories of passive restoration efforts in a degraded Nigerian montane forest system recently protected from burning and cattle grazing. We quantified the density, species richness, and functional trait dispersion of dispersed seeds and seedling communities at increasing distances from the forest edge. We then determined which plant traits are responsible for colonisation by quantifying changes in functional-trait dispersion and relative frequencies of dispersal-linked traits with increasing distance from the forest. We found a rapid decrease in density and species richness, and significant species turnover in both seeds and seedlings just beyond the forest edge. This was mirrored by a significant decline in functional-trait dispersion and a shift in the relative frequencies of dispersal-linked traits. These findings suggest that the reassembly of plant communities adjacent to remnant forest is dependent on functional traits present in these remnant source populations, providing support for the incorporation of trait-based theory in restoration management.     

Seed demography of three co-occurring Acer species in a Japanese temperate deciduous forest

Abstract. Seed demography of three co-occurring Acer species in an old-growth mixed deciduous forest in Japan was studied. Almost all of the seeds of A. mono germinated in the first spring, while those of A. palmatum var. amoenum showed a delay in germination of almost one year. A. rufinerve showed a rather opportunistic germination habit. Both A. palmatum var. amoenum and A. rufinerve form short-term persistent seed banks, but without input of newly dispersed seeds they may become extinct in about one year. The seed bank for these two species is not as significant as for a typical pioneer species, and the seedling bank is important for all three species. Only a small proportion of the dispersed propagules contained viable embryos, mainly due to pollination failure or abortion (A. mono and A. palmatum var. amoenum), and invertebrate predation (A. rufinerve). For all three species, larger seed crops had a higher percentage of viable seeds. Even for these relatively small, wind-dispersed seeds, the predation pressure was very high. A large part of the dispersed seeds was eaten by wood mice during the first winter (30–80 %). Estimation from the 5-yr average of seed dispersal and seedling emergence showed that only 7–16 % of the dispersed viable seeds succeeded in germinating.     

Avian frugivory and seed dispersal of a large fruited tree in an Indian moist deciduous forest

Seed dispersal systems in degraded areas can be compromised following the decline of large-bodied frugivore populations responsible for their dispersal. In this context we examined the seed dispersal ecology of a large fruited deciduous tree (Dillenia pentagyna) along a forest degradation gradient in India. We examined the effect of structural components of vegetation and frugivore foraging behavior on D. pentagyna seed dispersal. Depauperate mammalian community and declined large avian frugivores e.g. hornbills in our study site make this system a specialized one and currently dependent on only two large bodied avian frugivores. Seed dispersal followed an overall leptokurtic pattern and the seed dispersal kernels were best explained by an inverse power function. Seed dispersal kernels in dense forest indicated longer dispersal distances than moderately dense forest and degraded forest. In degraded areas, no dispersal away from the crown was recorded for D. pentagyna and it occurred at low density. Canopy foliage abundance of the surrounding vegetation of the focal trees was best explained by quantity of seed dispersal by large avian frugivores. The number of avian frugivore species those are effective disperser of D. pentagyna decreased along the degradation gradient. Avian frugivore behavior in terms of visitation and seed swallowed is a determining factor that controls quantity of seed dispersal. Our study underscores deleterious impact of forest degradation on avian disperser community which in turn would affect regeneration capacity of degraded forest.     

Seedling recruitment patterns in a tropical dry forest in Ghana

Abstract. Studies of seedling demography and dynamics were carried out in a dry forest at Pinkwae, Ghana (rainfall 1100 mm). Seedlings of tree and liana species in permanent transects were tagged, identified, measured and recensused at bimonthly intervals. A total of 1931 seedlings in 53 species were followed over 14 censuses during a 2-yr period. Germination and mortality were seasonal in their distribution; mortality was highest in dry periods. Seedling density varied seasonally, with the highest densities in comparatively wet periods. Diversity of seedlings was low, corresponding to that of the adult assemblage. Survivorship of tagged seedlings was recorded during the study period. The cohort tagged at the first census, comprising 1033 individuals of mixed ages and species, lost 61.9% during a 24-month period. Many seedlings were found to be rather large at the initial post-germination stage, with strong root development. Vegetative reproduction by root suckers or epicormic shoots was quite common at the site, although the majority of regeneration was by seeds. The current seedling assemblage appears to be representative, in terms of composition and diversity, of the adult species assemblage at Pinkwae.