Factors Limiting Tropical Rain Forest Regeneration in Abandoned Pasture: Seed Rain,Seed Germination,Microclimate, and Soil1

Vegetation, seed rain, seed germination, microclimate, and soil physical and chemical parameters were measured in a recently abandoned pasture and adjacent primary rain forest in southern Costa Rica. The goal of this study was to assess the importance of these factors in limiting forest regeneration in abandoned pastures. Seed rain of animal dispersed species decreased dramatically in the pasture >5 m from the forest/pasture edge; fewer wind dispersed seeds fell in the pasture than in the forest, but the difference was much less than for animal dispersed seeds. Percent seed germination of most species studied was similar in the forest and in pasture with grasses; seed germination was lower during the dry season in areas of pasture cleared of grasses. Air temperature, vapor pressure deficit (VPD), and photon flux density (PFD) were much higher in the pasture than in the forest at 1 m above the ground. VPD and PFD at ground level and soil temperature were similar in the pasture and the forest, indicating that pasture grasses strongly modify microclimatic conditions near the soil surface. The lowest gravimetric water content recorded in the pasture during the dry season was 0.5 and leaf relative water contents of the two species measured in the forest and pasture were identical, suggesting that plants in the pasture were not water stressed. Levels of most soil nutrients were lower in the pasture as compared to the forest; however, aboveground and root biomass for seedlings grown in pasture and forest soils did not differ significantly. Although a number of factors impede forest recovery in abandoned pastures, these results suggest that the most imporrant limitation is lack of seed dispersal.     

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

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

Seed Availability as a Limiting Factor in Forest Recovery Processes in Costa Rica

Abandoned pastures and secondary forests are increasingly prominent features of tropical landscapes. Forest regrowth on abandoned pastures is generally slow and virtually limited to regeneration from seeds from external sources, since agricultural activities alter site conditions. We hypothesize that seed availability is a major limiting factor in forest recovery on abandoned pastures. This hypothesis was tested by studying the seed bank, seed rain, and seed predation in a small pasture (1 ha) situated in a forest‐pasture mosaic in northwestern Costa Rica. The tree seed density in the pasture seed bank was much lower (21/m²) than the density in the seed bank of a neighboring secondary forest (402/m²). Within a period of five weeks, 23 tree seeds entered the pasture by seed rain. This number is low compared to densities found in closed forests but higher than densities reported in other studies where virtually no seeds were found beyond 20 m from the forest edge. Possibly the small size of the pasture with seed sources nearby and the small‐scale landscape mosaic enhance seed dispersal. Predation limits the seed density in pastures, with 42% of the woody species consumed by predators. The low seed density in the seed bank, and hampered recruitment combined with significant losses, pose severe restrictions to forest recovery on abandoned pastures. Moderate land use, and small sized clearings with seed sources nearby may increase the pace of recovery. Nevertheless, forest establishment may still take a considerable time. Thus, enlarging the available pool of species may be a worthwhile management strategy.     

Ignorant seed predators and factors affecting the seed survival of a tropical palm

In addition to acting as seed predators, some terrestrial mammals bury seeds via scatter hoarding. This study system used two permanent plots in examining the interaction between small rodents and the seeds of the palm Astrocaryum mexicanum. We tested how experimental burial, and fruiting status of the parent, distance to the parent, seed size, and microsite characteristics affect the survival of these seeds. Up to 34% of the buried seeds that were exposed only to ignorant rodent foragers (individuals not responsible for burial) survived. In comparison, less than 1% of seeds buried by scatter hoarding rodents survived in previous studies, a percentage that is comparable to the low survival of unburied seeds in this study (<2%). Although unburied seeds had very low survival, increasing distance and/or seed density positively affected survival of unburied seeds. Distance to parent had no effect on buried seed survival.
Buried seed survival was most strongly and significantly determined by the fruiting status of the trees under which they occurred. Seeds experienced significantly greater predation if buried under “parent” trees that fruited during the experiment. Buried seed survival was also negatively affected by germination, as germination may signal the presence of a seed to foraging rodents. There was some indication of a positive effect of tree density on seed survival between the two plots, whereas differences in rodent abundance appear to have no effect on seed survival. Seed size and microsite characteristics had no significant effect on buried seed survival, likely due to the greater proportional effects of other factors and the longevity of A. mexicanum seeds. The results of this study were used to generate a hypothetical causal network showing how comparatively low recovery of buried seeds by ignorant foragers – combined with processes determining the removal of scatter hoarding foragers from their scattered seed caches – may affect seedling recruitment in A. mexicanum.     

Postdispersal Seed Fate of Tropical Montane Trees in an Agricultural Landscape, Southern Costa Rica

Variation in postdispersal seed fate is an important factor driving patterns of forest regeneration. Because most previous studies have not tracked final seed fate and have commonly equated seed removal with predation without considering the possibility of secondary dispersal, little is known about individual seed mortality factors in successional and degraded habitats. This study tracked the postdispersal fate of large-seeded tropical montane trees in abandoned pasture, young secondary forests, and small forest fragments during two consecutive years in an agricultural landscape in southern Costa Rica. The incidence of secondary dispersal by animals, scatterhoarding in particular, and the effects of seed burial on germination were measured. Overall, seeds survived through germination more often in secondary forests with high levels of mortality occurring in abandoned pastures and forest fragments. The majority of seed mortality was caused by rodent predation in forest fragments, insects and fungal pathogens in secondary forests, and a combination of desiccation, insects, and fungal pathogens in pastures. Seeds were frequently secondarily dispersed in larger forest fragments, whereas they were only rarely moved in pastures and secondary forests. Burial tended to improve germination in pastures and was important for an often scatterhoarded species, Otoba novogranatensis, in all habitats. The results of this study suggest that: (1) seed mortality factors differ in response to the type of habitat degradation; (2) large-seeded species have high potential for survival when dispersed to young secondary forests; and (3) seed removal is not a reliable proxy for seed predation, particularly in less degraded forest fragments.     

Evidence for secondary seed dispersal by rodents in Panama

Summary The data presented show thatVirola nobilis (Myristicaceae), a bird/mammal-dispersed tree species in Panama, may also be dispersed by a terrestrial rodent, the agouti (Dasyprocta punctata). Using a thread-marking method, we observed that agoutis scatterhoardedV. nobilis seeds that they found both singly or in clumps. Seed removal and seed burial rates were strongly affected by features of forest habitats, such asV. nobilis tree richness (rich vs poor) and/or forest age (old vs young), but not by seed dispersal treatment (scattered vs clumped). Predation (mostly post-dispersal) of unburied seeds by weevils was independent of habitat and dispersal treatment. Seeds artificially buried in aVirola-rich area were more likely to escape predation and become established than unburied seeds under natural conditions. The food reward for agoutis is in the germinating seedlings. The seed dispersal syndrome ofV. nobilis involves long- and short-distance dispersers which both appear important for tree recruitment.     

Barriers to Forest Regeneration in an Abandoned Pasture in Puerto Rico

Sources of forest regeneration (soil seed bank, seed rain) and barriers to seedling establishment were examined in a recently abandoned pasture in eastern Puerto Rico. Few woody species were found in the soil seed bank or in the seed rain. The number of seeds and species in the seed rain and soil seed bank declined with distance from the adjacent secondary forest. Nine species naturally dispersed and colonized plots during the study, with the wind‐dispersed tree Tabebuia heterophylla being the predominant colonizer (91% of all seedlings). Barriers to seedling establishment were determined using a blocked field experiment with eleven woody species representative of a variety of life histories. Each species was planted under the pasture vegetation (control) or in areas where all vegetation was removed (removal). Germination was enhanced for four species in the control treatment, five species were not affected, and two species did not germinate under either treatment. Survival to 6 months was higher in the removal treatment for two species. Seedling biomass was greater in the removal treatment at 12 months for one species. Seed mass was a good predictor of germination success and final shoot biomass, but not survival. This study demonstrates that seeding recently abandoned pastures with a mix of known pioneer species may accelerate the rate of secondary succession, but some species will have to be planted in later successional stages in order to overcome strong barriers to establishment.     

Soil seed banks in different environments: initial forest,mature forest,Pinus and Eucalyptus abandoned stands

Abstract

Different environments (initial forest, mature forest, Pinus and Eucalyptus stands) found in Seasonal Semideciduous Forest fragments affect the density of viable seeds, as well as the floristic similarity, diversity, and richness of tree species in soil seed banks. This hypothesis was tested in the current study. Soil seed bank samples were collected in the aforementioned environments during rainy and dry seasons, and taken to a shade house, where they remained under favorable seed germination conditions. Tree seedling emergence was measured, and sample species were identified every 15?days, for six months, in each sampling period. In total, 97 individuals m^?2 and 23 species emerged in all environments and periods. The highest density of viable seeds of tree species in the soil seed bank was found in the initial forest stretch, mature forest stretch and abandoned Eucalyptus stand. Only the Pinus stand seed bank in the dry season had different floristic and lower viable seed density than the mature forest seed bank. Thus, all environments, except the abandoned Pinus stand, can preserved Seasonal Semideciduous Forest fragments.     

Consequences of habitat disturbance on seed fate of a Brazilian tropical dry forest tree Cavanillesia arborea (Malvaceae)

We compared seed fate (survival, mortality by rodent predators, desiccation and other causes) of the tree Cavanillesia arborea (Malvaceae) in preserved old‐growth tropical dry forests and in nearby abandoned pasturelands in Brazil. For this purpose, we performed an experiment where 15 seeds were placed in the surroundings of 15 parental individuals in each habitat. These 450 seeds were monitored over four months. At the end of the experiment, seed predation by rodents was higher in forest (56.5%) than in pasture (8.9%) areas, but seed desiccation showed the opposite pattern (8.9% vs. 80.4%). Mortality by desiccation was also faster in pasture than in forest areas, probably reducing their attractiveness to predators in these areas. None of the seeds placed in the pasture survived, whereas 26.2% of seeds became seedlings in forest areas. The absence of seedling recruitment of Cavanillesia arborea in pastures is likely a consequence of their incapacity to tolerate the harsh abiotic conditions in this habitat. Thus, forest conversion imposes a strong limitation to the long‐term population viability of this species. However, restoration and natural regeneration of abandoned pastures can recreate the forest structure and microclimatic conditions favourable to seed germination and seedling establishment. Remnant individuals of Cavanillesia arborea in agricultural landscapes may have a negligible contribution to current recruitment, but they can attract potential dispersers of pioneer species, with positive feedbacks to future recruitment during secondary succession.     

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.     

Post‐Dispersal Seed Removal and Germination Selected Tree Species Dispersed by Cebus capucinus on Barro Colorado Island,Panama1

Dispersal quality, an important component of seed disperser effectiveness, may strongly affect the rate of plant recruitment. Here we evaluated the quality of Cebus monkey dispersal by comparing the secondary removal fate and germination of fresh and Cebus‐ingested seeds of nine tree species on Barro Colorado Island, Panama. Overall, rates of secondary seed removal by vertebrates were low, with most Cebus defecations remaining undisturbed for extended periods on the forest floor. Only four of 30 feces were completely buried by dung beetles, and we found significantly higher vertebrate removal of defecated seeds than control seeds for only one species, Cordia bicolor. Seed germination varied greatly between plant taxa. Seeds of 3 out of 9 species showed significantly higher percent germination after monkey gut passage than control fresh seeds. Germination times tended to be shorter for defecated than for control seeds but were significantly different only for one of nine species, Cecropia insignis. Low rates of seed removal from Cebus feces, coupled with high germination probabilities, suggest high dispersal effectiveness for Cebus and contrasts strongly with patterns of post‐dispersal seed fate recorded for other primate species.     

Establishment of Fabaceae Tree Species in a Tropical Pasture: Influence of Seed Size and Weeding Methods

The successful establishment of trees in pastures may be related to species, seed size, and weed control treatments. This study experimentally evaluated the emergence, establishment after 2 years, and growth of individuals of 7 tree species with different seed sizes sowed under three weeding methods (no weeding, grasses clipped, and grasses removed). The experiment was carried out in a tropical abandoned pasture in Midwestern Brazil, dominated by the exotic invasive grass Urochloa brizantha. The effect of seed size on seedling emergence was significant and was more intense in the no weeding and weeding treatment plots than in grasses clipped. Furthermore, an increase in seed size resulted in an increase in the probability of establishment of tree species, although this pattern also differed among weeding treatments. The increase in seed size reflected negatively on tree growth in the absence of grasses, whereas in the presence of grasses there was an inverse relationship. The experimental results suggested that the use of seeds of different sizes for direct seeding in pastures is a possible strategy of Fabaceae tree species reintroduction in pastures within the Cerrado. Fabaceae species with seeds larger than 100 mg can establish in areas with no weeding, whereas species with seeds smaller than 50 mg can establish preferentially in areas with weeding, growing faster than larger ones. Thus, weeding strategies can determine the success of direct seeding of different species in restoration projects of tropical pastures.     

The effect of frugivory on postdispersal seed removal and germination in the pantropical forest tree Antiaris toxicaria Leschenault

The quality of seed treatment by frugivores has an effect on seed removal after dispersal, seed germination and tree recruitment. We provide information on postdispersal seed removal, germination and subsequent recruitment in tropical forest tree species Antiaris toxicaria in Ghana. We tested whether postdispersal seed removal and germination rates were differentially affected by the following seed treatments: seeds that were spat out by monkeys with all fruit pulp removed and spitting seeds with fruit pulp partially removed as observed in some birds and bats. We used seeds of intact ripened fruits as control. Frugivore seed treatment and distance from bole affected seed removal patterns, whereas intact seeds were significantly removed from all seed stations. The germination success was greater for seeds that were spat out by monkeys and poor for seeds with fruit pulp partially removed and intact fruits. More recruits were recorded at the edge of the adult A. toxicaria canopy radius. There was weak relationship (r² = 0.042) between the number of recruits and distance away from the adult tree. Results suggest that the subsequent recruitment in tropical forest tree species may be enhanced by some frugivore fruit‐handling behaviour where fruit pulp is removed from the seeds without destroying the seeds.     

Seed Dynamics of Early and Late Successional Tree Species in Tropical Abandoned Pastures: Seed Burial as a Way of Evading Predation

We explored different treatments to enhance the probability of sowed seeds of two early successional (ES, Cecropia obtusifolia and Ochroma pyramidale) and two late successional (LS, Brosimum costaricanum and Dialium guianense) species to escape predation and germinate in abandoned cattle‐raising pasture fields in Southeastern Mexico. ES species were sown in groups of 50 seeds under three treatments: invertebrate exclusion, burial, and exposition to seedeaters. LS species were sown in groups of 10 seeds under three treatments: vertebrate exclusion, burial, and exposition to seedeaters. We registered seed predation and germination 2, 4, 8, 16, 32, and 64 days after the initial sowing. Overall, ES showed higher predation rates (mean ± SE = 0.45 ± 0.07 seed seed^?1 day^?1; n = 3) than LS species (0.09 ± 0.02 seed seed^?1 day^?1). Cecropia obtusifolia was completely predated in all treatments after 8 days. Burial and exclusion treatments reduced final predation in circa 6% for O. pyramidale, relative to that of exposed seeds (85% after 8 days); most germination occurred in buried seeds (3.7%). In B. costaricanum, burial enabled germination by 10%; exposed and excluded seeds were removed 100%. Dialium guianense showed 12% germination in buried seeds and circa 20% of the seeds were not removed after 64 days. Direct sowing would be a recommended rainforest restoration practice for species with relatively large seeds if deposited in groups and buried. Studies which address variation across numerous sites are necessary in order to generate more consistent seed predation patterns and rainforest restoration principles in tropical pastures.     

西双版纳热带季节雨林四种珍稀濒危树种种子萌发对脱水和光的响应

绒毛番龙眼(Pometia tomentosa)、琴叶风吹楠(Horsfieldia pandurifolia)、箭毒木(Antiaris toxicaria)和思茅木姜子(Litseapierrei var. szemois)是西双版纳热带季节雨林乔木层的优势树种,也属于珍稀濒危物种。实验研究了4个树种的种子在经过不同脱水水平、人工遮荫条件下的不同光照水平以及热带季节雨林之林下和林窗的萌发特性。结果显示:脱水对于种子萌发特征具有显著影响,绒毛番龙眼和琴叶风吹楠经48h的连续脱水后丧失了萌发能力;箭毒木和思茅木姜子具较强的脱水耐性,经96h的连续脱水后仍具有较高的种子含水量,且种子萌发率分别为15%和33%。干旱季散布的种子比雨季散布的种子具有更强的脱水耐性。遮荫及森林种子萌发试验中,除绒毛番龙眼外,其它3个树种的种子萌发率以及平均萌发周期在不同光水平或生境下差异显著。琴叶风吹楠和思茅木姜子在30%的光照以及林窗中心的萌发率要显著低于低光处理(10%和3.5%光照)和林下,而箭毒木在30%和10%光水平以及林窗中心获得较高萌发率。雨季传播的具有干燥敏感性的种子,可以减少种子由于干燥而引起的死亡。试验结果可以为4种珍稀濒危植物的保护和人工繁育提供一定理论依据。     

Seed Dormancy,Seedling Establishment and Dynamics of the Soil Seed Bank of Stipa bungeana (Poaceae) on the Loess Plateau of Northwestern China

Studying seed dormancy and its consequent effect can provide important information for vegetation restoration and management. The present study investigated seed dormancy, seedling emergence and seed survival in the soil seed bank of Stipa bungeana, a grass species used in restoration of degraded land on the Loess Plateau in northwest China. Dormancy of fresh seeds was determined by incubation of seeds over a range of temperatures in both light and dark. Seed germination was evaluated after mechanical removal of palea and lemma (hulls), chemical scarification and dry storage. Fresh and one-year-stored seeds were sown in the field, and seedling emergence was monitored weekly for 8 weeks. Furthermore, seeds were buried at different soil depths, and then retrieved every 1 or 2 months to determine seed dormancy and seed viability in the laboratory. Fresh seeds (caryopses enclosed by palea and lemma) had non-deep physiological dormancy. Removal of palea and lemma, chemical scarification, dry storage (afterripening), gibberellin (GA₃) and potassium nitrate (KNO₃) significantly improved germination. Dormancy was completely released by removal of the hulls, but seeds on which hulls were put back to their original position germinated to only 46%. Pretreatment of seeds with a 30% NaOH solution for 60 min increased germination from 25% to 82%. Speed of seedling emergence from fresh seeds was significantly lower than that of seeds stored for 1 year. However, final percentage of seedling emergence did not differ significantly for seeds sown at depths of 0 and 1 cm. Most fresh seeds of S. bungeana buried in the field in early July either had germinated or lost viability by September. All seeds buried at a depth of 5 cm had lost viability after 5 months, whereas 12% and 4% seeds of those sown on the soil surface were viable after 5 and 12 months, respectively.     

南亚热带森林不同演替阶段土壤种子库的初步研究

本文对鼎湖山不同演替阶段的森林（马尾松林、针阔叶混交林、季风常绿阔叶林）的土壤种子库进行了初步研究．分别从各个阶段的样地内抽取４或８个１×１ｍ的小样地,分三层（共１０ｃｍ厚）采集土样,带回实验室,通过萌发法观察记录其土壤种子库状况．通过统计分析,其结果如下：１．种子数量和物种多样性一般随演替发展而减少,种子数目以３—５ｃｍ厚土层为最多．２．各演替阶段土壤种子库的种类组成均以草本植物为主．３．在光照和湿度基本一致的情况下,萌发种子数与温度成正相关．４．雨季取的种子数目较旱季多,且种类组成有所不同．这是因为南亚热带森林内植物种子的休眠期短,在旱季采集土样时许多种的种子尚未下落．５．种子库组成与地上植物相关性不明显,但演替早期阶段的相关性比演替后期更密切。     

Tropical Montane Forest Restoration in Costa Rica: Overcoming Barriers to Dispersal and Establishment

Tropical forests are being cleared at an alarming rate although our understanding of their ecology is limited. It is therefore essential to design restoration experiments that both further our basic knowledge of tropical ecology and inform management strategies to facilitate recovery of these ecosystems. Here we synthesize the results of research on tropical montane forest recovery in abandoned pasture in Costa Rica to address the following questions: (1) What factors limit tropical forest recovery in abandoned pasture? and (2) How can we use this information to design strategies to facilitate ecosystem recovery? Our results indicate that a number of factors impede tropical forest recovery in abandoned pasture land. The most important barriers are lack of dispersal of forest seeds and seedling competition with pasture grasses. High seed predation, low seed germination, lack of nutrients, high light intensity, and rabbit herbivory also affect recovery. Successful strategies to facilitate recovery in abandoned pastures must simultaneously overcome numerous obstacles. Our research shows that establishment of woody species, either native tree seedlings or early‐successional shrubs, can be successful in facilitating recovery, by enhancing seed dispersal and shading out pasture grasses. On the contrary, bird perching structures alone are not an effective strategy, because they only serve to enhance seed dispersal but do not reduce grass cover. Remnant pasture trees can serve as foci of natural recovery and may enhance growth of planted seedlings. Our results highlight the importance of: (1) understanding the basic biology of an ecosystem to design effective restoration strategies; (2) comparing results across a range of sites to determine which restoration strategies are most generally useful; and (3) considering where best to allocate efforts in large‐scale restoration projects.     

Experiments on the mechanism of tree and shrub establishment in temperate grassy woodlands: Seedling emergence

Field experiments were designed to examine tree and shrub seedling emergence in temperate grassy woodlands on the New England Tablelands. The effects of study sites, intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds and ant seed theft on seedling emergence were tested in two field experiments. Six tree and seven shrub species were used in the experiments and their cumulative emergence was compared with laboratory germination studies. All species used in field experiments had lower cumulative emergence than those in laboratory germination studies despite prolonged periods of above average rainfall before and after seeds were sown. Eucalypt species emerged faster in the field than the shrub species and generally attained higher cumulative emergence than the shrubs. Spatial effects of sites and patches within sites, and of previous grazing history did not strongly influence patterns of seedling emergence in most species. Ground and litter cover generally did not enhance or suppress the emergence of seedlings, although the removal of cover in recently grazed areas enhanced the emergence of some species. Burning enhanced the emergence of some tree and shrub species where plots had more fuel and intense fires, but this effect was not strong. Compared with other treatments, seedbed manipulations produced the strongest effects. In the absence of both invertebrate and vertebrate predators, seedling emergence was lower for surface‐sown seed, compared with seed sown on scarified soil surfaces. Higher seedling emergence of buried seeds in the presence of invertebrate predators probably resulted from the combined effects of predator escape and enhanced moisture status of the germination environment. Some promotion of emergence was achieved for all species in most sown treatments probably as a result of a prolonged above average rainfall. In contrast, the natural recruitment of trees and shrubs was negligible in experimental plots, highlighting the importance of seed supply and dispersal as ultimate determinants of recruitment.     

Seed rain from forest fragments into tropical pastures in Los Tuxtlas,Mexico

Deforestation has resulted in the fragmentation of forests. Remnant fragments are widely assumed to be sources of seeds for forest regeneration in abandoned pastures. The seed rain in 12 pastures at Los Tuxtlas, Mexico, their relationship with riparian vegetation adjacent to them, and their closeness to the reserve is described. For all the species found in the seed rain, we classified them by typical habitat (pasture or rain forest), life form, dispersal syndrome, and plant strategy (pioneer or non-pioneer species). In addition, germinability was evaluated for all seeds. We also assessed the correlation in composition of the seed rain and the riparian vegetation. Only 11% of the total species that occur in the Los Tuxtlas region reach pastures via dispersal from forest fragments. However, nearly 80% of the species in the seed rain did not come from the fruiting individuals in the adjacent riparian vegetation. The proportion of dispersal syndromes, life forms and plant strategies of the species in the seed rain was similar to those observed in the rain forest. On average, the germinability of forest species was less than 30%.The forest species richness was similar in the seed rain in pastures and inside the forest, but the pasture seed rain contained fewer seeds per species. Pastures have a high potential for natural regeneration because of seed dispersal from adjacent forest. However, the forest that regenerates in pastures close to the reserve is expected to contain different species than the forest regenerating far away from it.