Logs and Fern Patches as Recruitment Sites in a Tropical Pasture

Forest recovery in degraded pastures is often highly variable, possibly due to variation in the availability of adequate recruitment sites. In an actively grazed pasture in northeastern Costa Rica, this study examines how recruitment of woody species in patches of the fern Nephrolepsis sp. and near logs compares with recruitment in grassy areas. Fern patches and logs had five and eight times higher densities of woody recruits, respectively, as grassy areas. They also had more than twice the species richness and growth as grassy areas. Grass apparently presents a competitive barrier against invading woody recruits, and also attracts cattle that may trample and/or consume recruits. Both logs and patches of fern appeared to provide superior conditions for establishment and growth of woody species, and they did not attract cattle. This study suggests that fern patches and logs can be managed to accelerate forest succession.     

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.     

Spatial distribution of understory fruit-eating birds and fruiting plants in a neotropical lowland wet forest

Spatial distribution of fruit-eating birds and fruiting shrubs of the Melastomataceae and Rubiaceae were examined on a 10 ha plot in tropical lowland wet forest of Costa Rica. Many plant species and most birds exhibited considerable spatial variation in their occurrence on the plot, as indicated by the distribution patterns of shrubs with ripe fruits and captures in mist nets, respectively. In many cases, captures of fruit-eating birds were correlated with abundance of fruiting plants, particularly for species that rely heavily on fruits. In general, fruit-eating birds concentrated their use of the plot to areas rich in fruiting shrubs. This differential use of certain areas likely results in differential visitation to fruiting plants located in these areas and in a heterogeneous dissemination of seeds into the habitat.     

Species Richness,Spatial Variation,and Abundance of the Soil Seed Bank of a Secondary Tropical Rain Forest1

Despite the importance of the soil seed bank in tropical forest regeneration, little is known about spatial variability in species composition and abundance of seeds stored in the soil. To develop sampling methods for comparative studies, we examined species richness, spatial variation, and abundance of germinants from the soil seed bank in a 16 year old secondary, tropical wet forest at La Selva Biological Station, Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at the intersection points of a gridded 1 ha plot (10 × 10-m grid, 121 samples) and in a nested 100 m² subplot (2 × 2-m grid, 36 samples). The 1 ha plot had a density of 4535 seeds/m² with 34 species observed. Based on a series of 100 randomized species accumulation curves, a Michaelis-Menten fit predicted a mean species richness of 36.3 species; the number of observed species was close to the predicted asymptote. A nonparametric, first-order jackknife species richness estimator predicted a species richness of 37.0 species. Eighty-five and 95 percent of the observed species richness is contained, on average, within 41 and 74 pooled samples, respectively. Within the 100 m² nested subplot, a density of 5476 seeds/m² was observed, comprising 26 species with an estimated species richness (Michaelis-Menten fit) of 29.1 species. The jackknife species richness estimator predicted a species richness of 36.7 species. For species richness and abundance of both plots, spatial autocorrelation statistics (Moran's I) were not significantly different from zero at lag distances from 2 to 100 m, indicating a random distribution at these spatial scales. For this site, accurate estimates of species composition depend upon the number of samples collected as well as the spatial distribution of sampling effort. Many small samples distributed over a large area provide greater accuracy and precision for estimating species richness of the soil seed bank.     

Growth of Mantled Howler Groups in a Regenerating Costa Rican Dry Forest

We examined population dynamics in mantled howlers (Alouatta palliata palliata) in a regenerating tropical dry forest in Santa Rosa National Park (SRNP), Costa Rica. The population has grown at a rate of about 7% per annum during the past decade. The growth in numbers from 342 in 1984 to 554 in 1992 reflects an increase in the number of groups (from 25 to 34) and a slight increase in their average size (from 13.6 to 16.3). Population density has increased from 4.9 to 7.9 individuals per km ². Santa Rosa's population density and group compositions are similar to those at several other mantled howler sites, but densities of mantled howlers are much higher at two other well-studied sites: La Pacifica and Barro Colorado Island (BCI). We relate the low density of howlers at Santa Rosa to local historical and ecological factors. Howler populations at high and low densities differ in average group size and sex ratio. At high population densities, groups are larger and include more adult females. The number of male howlers per group appears to be more strictly limited and less variable than the number of females is. However, there is greater variation in male group membership at Santa Rosa than at La Pacifica or BCI, and at Santa Rosa there are more generating forests available into which males and females can disperse and form new groups. We present case studies describing two ways in which new howler groups are formed, and we suggest that, compared to females and compared to males at high density sites, males are relatively advantaged in the uncrowded habitats at Santa Rosa and other low density sites.     

Tree Seed Fate in a Logged and Fragmented Forest Landscape, Northeastern Costa Rica1

We compared the seed fate of two animal‐dispersed, large‐seeded timber species (Dipteryx panamensis [Fabaceae] and Carapa guianensis [Meliaceae]) in logged and fragmented forests with that for continuous forest in northeastern Costa Rica. For both species, we quantified rates of seed removal (an index of vertebrate predation) and the fate of dispersed seeds (those carried away from their original location that either germinated or were not subsequently removed within three months). We predicted that (1) fewer seeds would be dispersed by vertebrates in fragmented forest than in continuous forest due to low population abundances after hunting and/or loss of suitable habitat, and (2) seed predation rates would be higher in forest fragments than in continuous forest due to high abundance of small‐bodied seed consumers. We compared three forest fragments currently managed for timber (140–350 ha) and a large reserve of continuous forest (La Selva, 1500 ha and connected to a national park). An exclusion experiment was performed (seeds placed in the open vs. seeds within semipermeable wire cages; 5 cm mesh size) to evaluate the relative roles of large and small animals on seed removal. Seed germination capacity did not differ among all four sites for both species. Removal of Dipteryx seeds was higher in forest fragments (50% removal within 10 days and related to the activity of small rodents) compared to La Selva (50% removal after 50 days). Also, more Dipteryx seeds were dispersed at La Selva than in fragmented forests. Contrary to our predictions, removal of Carapa seeds was equally high among all four sites, and there was a trend for more seeds of Carapa to be dispersed in fragments than in La Selva. Our results suggest that fragmentation effects on tree seed fate may be specific to species in question and contingent on the animal biota involved, and that management strategies for timber production based on regeneration from seed may differ between forest patches and extensive forests.     

Seed characteristics and susceptibility to pathogen attack in tree seeds of the Peruvian Amazon

Many studies now suggest that pathogens can cause high levels of mortality in seeds and seedlings. Recruitment from seed to sapling is an important bottleneck for many tree species, and if specialist or generalist pathogens have differential negative effects among species of juvenile trees, then they may have a significant impact on forest community structure. To explore the effects of differential pathogen attack among tropical tree species, we quantified pathogen attack on the seeds of 16 tree species from the southeastern Peruvian Amazon and asked which seed characteristics, including size, hardness, germination time and mode, shade tolerance, and fruit type, were most closely correlated with susceptibility to pathogens. Shade tolerance and seed weight were positively and significantly correlated with susceptibility to pathogen attack by ecological trait regressions (ETRs), and correspondence analysis indicated that there might be increased attack rates in species with brightly colored, pulpy fruits (often dispersed by primates). Only shade tolerance was significantly correlated with pathogen attack when the analyses accounted for phylogenetic relatedness between species. Thus, contrary to standard predictions of size-defense ratios, our results suggest that larger, shade-tolerant seeds tend to be more susceptible to pathogen attack than smaller, light-dependent seeds. Moreover, differential pathogen attack may shape seed community composition, which may affect the successful recruitment of adults.     

Diversity,composition, and structure of tropical dry forests in Central America

Tropical dry forests have been reduced to less than 0.1% of their original expanse on the Pacific side of Central America and are considered by some to be the most endangered ecosystem in the lowland tropics. Plots 1000 m² were established in seven tropical dry forests in Costa Rica and Nicaragua in order to compare levels of species richness to other Neotropical dry forest sites and to identify environmental variables associated with species richness and abundance. A total of 204 species and 1484 individuals 2.5 cm were encountered. Santa Rosa National Park was the richest site with the highest family (33), genera (69), and species (75) diversity of all sites. Species richness and forest structure were significantly different between sites. Fabaceae was the dominant tree and shrub family at most sites, but no species was repeatably dominant based on number of stems in all fragments of tropical dry forest. Central American dry forests had similar species richness when compared to other Neotropical forests. There was no correlation between forest cover within reserves, or precipitation and plant species richness. There was a significant correlation between anthropogenic disturbance (intensity and frequency of fire, wood collection, grazing) and total species richness, tree and shrub species richness, and liana abundance. These results suggest controlling levels on anthropogenic disturbance within reserves should be a high priority for resource managers in Central America. Further research in forest fragments which examine individual and a combination of disturbance agents would help clarify the importance of anthropogenic disturbance on species richness and abundance.     

Soluble reactive phosphorus (SRP) transport and retention in tropical, rain forest streams draining a volcanic landscape in Costa Rica: in situ SRP amendment to streams and laboratory studies

Soluble reactive phosphorus (SRP) transport/retention was determined in two rain forest streams (Salto, Pantano) draining La Selva Biological Station, Costa Rica. There, SRP levels can be naturally high due to groundwater enriched by geothermal activity within the surfically dormant volcanic landscape, and subsequently discharged at ambient temperature. Combined field and laboratory approaches simulated high but natural geothermal SRP input with the objective of estimating the magnitude of amended SRP retention within high and low SRP settings and determining the underlying mechanisms of SRP retention. First, we examined short-term SRP retention/transport using combined SRP-conservative tracer additions at high natural in situ concentrations. Second, we attempted to observe a DIN response during SRP amendment as an indicator of biological uptake. Third, we determined SRP release/retention using laboratory sediment assays under control and biologically inhibited conditions. Short-term in situ tracer-SRP additions indicated retention in both naturally high and low SRP reaches. Retention of added SRP mass in Upper Salto (low SRP) was 17% (7.5 mg-P m⁻² h⁻¹), and 20% (10.9 mg-P m⁻² h⁻¹) in Lower Salto (high SRP). No DIN response in either nitrate or ammonium was observed. Laboratory assays using fresh Lower Salto sediments indicated SRP release (15.4 ± 5.9 μg-P g dry wt.⁻¹ h⁻¹), when incubated in filter sterilized Salto water at ambient P concentration, but retention when incubated in filter sterilized river water amended to 2.0 mg SRP l⁻¹ (233.2 ± 5.8 μg-P g dry wt.⁻¹ h⁻¹). SRP uptake/release was similar in both control- and biocide-treated sediments indicating predominantly abiotic retention. High SRP retention even under biologically saturated conditions, absence of a DIN response to amendment, patterns of desorption following amendment, and similar patterns of retention and release under control and biologically inhibited conditions all indicated predominantly abiotic P flux.     

Effects of Single and Recurrent Wildfires on Fruit Production and Large Vertebrate Abundance in a Central Amazonian Forest

Wildfires are an increasing threat to tropical rainforests, yet little is known about their effects on fruit production and forest wildlife. We examined the effects of both single and recurrent wildfires on fruit production and large vertebrate abundance in a central Amazonian terra firme forest for 3 years following a large fire event. The estimated mortality of 42 and 74% of stems ≥10 cm in once- and twice-burnt forest led to a substantial loss of fruiting tree basal area (29 and 62% were lost in once- and twice-burnt forest, respectively) and crown coverage of fruiting woody lianas (89 and 97% were lost in once- and twice-burnt forest, respectively). Some important tree families producing fleshy fruits were less abundant than expected in once- and twice-burnt forest, suggesting that tree mortality was non-random in terms of species composition. Asynchronous fruit production was affected, and burnt forest transects sustained a much lower fruiting basal area, and fewer fruiting species during the dry season period of fruit scarcity. The number of fruiting trees in once- and twice-burnt forest was higher than the number predicted from actual levels of tree mortality recorded in each fire disturbance treatment, suggesting some surviving trees which may have benefited from higher irradiance levels and lower competition for resources. Many large frugivores and other vertebrate species declined in response to single fires, and most primary forest specialists were extirpated from twice-burnt forest, which sustained a higher number of species associated with second growth and other disturbed habitats.     

Long-Term Effects of Forest Regrowth and Selective Logging on the Seed Bank of Tropical Forests in NE Costa Rica1.

To investigate long-term effects of land use on the soil seed bank, we compared the abundance/density, species richness, life form distribution, and species composition of seeds stored in the soil of four 15–20 yr-old second-growth stands, two old-growth stands, and two previously selectively-logged stands in the Caribbean lowlands of Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at 10 m intervals along three 120–160 m long transects in each stand (44–48 soil cores, 22–24 combined seed bank samples per site). Seed density was highest but variable in second-growth stands (8331–14535 seeds/m²), low and homogeneous in old-growth stands (2258–2659 seeds/m²), and intermediate and highly variable in selectively-logged stands (1165–6854 seeds/m²), which also had contrasting logging intensities. Species richness was strongly dependent on seed density, but showed less variation. Life form distribution did not differ statistically among or within land-use categories. In each stand, herbs-forbs, shrubs, and vines dominated the seed bank (> 75% of the species richness and abundance), whereas trees were a minor component (< 20% of the species richness and < 5% of the abundance) and were predominandy early successional. Shrubs and vines were most abundant in second-growth stands where regrowth vegetation was repeatedly cut before abandonment, whereas grasses and sedges were most abundant in the only forest stand that was completely surrounded by pastures. In terms of species composition, old-growth stands were more similar to selectively-logged stands than to second-growth stands, but across stands, selectively-logged forests were most distinct from the other two forest types. An inventory of the standing woody vegetation in each site showed little representation of the woody taxa found in the seed bank. We discuss these results in the context of the main factors that have been postulated to influence the abundance, life form, and species composition of tropical forest seed banks, and explore the role of the latter during intermediate phases of tropical forest succession and regeneration.     

Seed size in tropical trees: a comparative study of factors affecting seed size in Peruvian angiosperms

Evolutionary comparative methods taking into account the potential effects of relatedness reveal that, among 202 species of animal-dispersed tropical woody angiosperms from Peru, large seeds may be an adaptation to: (1) dispersal by mammals rather than by birds and (2) greater plant height. Using the most powerful techniques currently available, appropriate comparisons were made between related species or species groups at all taxonomic levels, thus allowing use of all species in the data set. Even so, the results show no evidence of adaptation (i.e., a functional relationship) of seed mass to successional syndrome or to growth form (trees or vines). In previous cross-species analyses, seed mass has been linked to both of these ecological variables. The lack of relationship between seed mass and habitat use or growth form may be due to targeting the wrong trait to predict these two variables, or because seed mass is subject to exaptation, i.e., a particular seed mass has arisen for other reasons, but allows invasion into a habitat and currently serves to maintain that species in the habitat. Differentiation between these two explanations for the lack of correlation between seed mass and successional syndrome or growth form would entail species-by-species examination of the function of seed mass under natural conditions. The effect of relatedness on the ecological correlates of seed mass and plant height is determined for each of three taxonomic schemes. It is concluded that the three schemes do not differ greatly in the results obtained.     

Characteristics of bird species using forest and agricultural land covers in southern Costa Rica

Our understanding of why tropical forest species differ in their ability to inhabit agroecosystems is limited, despite the link between this ability and the likelihood of population decline for species inhabiting regions undergoing widespread conversion of forest to agriculture. We used logistic regression and data from southern Costa Rica to develop a model based on natural history characteristics to distinguish between forest species that did or did not use agricultural land covers. We sampled birds along 15 3.0-km routes, seven in forest and eight in agriculture, five times over three years. Each species was classified as an F-species, detected only in forest, a G-species, detected in both forest and agriculture, or an A-species, detected only in agriculture. Thirty percent of species were F-species, 42% were G-species, and 28% were A-species. Based on the logistic regression model, the likelihood of being a G-species, as opposed to an F-species, was low for species that were dependent on forest interior, had a stenophagous diet, and a small elevational range. Weight, resident versus migrant status, and whether or not a species was insectivorous, were not significant predictors of being a G-species. For all F–G species pairs, the model correctly predicted the G-species 70% of the time. The model provides a first step in identifying those characteristics that predispose forest species to use agricultural land. In addition, our results indicate that the structurally simple agricultural systems of the study region are of limited value for a large proportion of the regional species pool.     

Fruit removal efficiency and success: influence of crop size in a neotropical treelet

The reproductive success of animal-dispersed plants is closely linked to the number of seeds that they are able to disperse. The fruit crop size hypothesis states that a plant with large fruit crop size will attract more dispersers than a plant with a smaller fruit crop, which may result in more seeds being dispersed from the foremost. In this study, we experimentally examined the effect of crop size and other factors on primary seed dispersal in a neotropical shrub/tree, Casearia corymbosa (Flacourtiaceae). We used two predictive variables of reproductive success, which produce an accurate picture of seed dispersal ratio: fruit removal efficiency (proportion of a fruit crop removed by frugivores) and fruit removal success (relative contribution of each individual tree to the number of fruits removed in the population). We established two levels of fruit crop size at the C. corymbosa individuals, using plants with large (150 fruits) and small crops (50 fruits). We found that individual plants with larger crops had significantly higher values of fruit removal efficiency (92.6%) and success (5%) than plants with smaller crops (69.3% and 1.3%, respectively). Fruit removal efficiency was related to vegetation type, plant height and fruit width, but the variance explained by these variables was low ( < 8%). Fruit removal success was significantly related to crop size ( > 90% of the variance explained). These results suggest that fruit removal efficiency and success are strongly related to fruit crop size of C. corymbosa plants.     

Interaction between ants and seeds of a nonmyrmecochorous neotropical tree, Cabralea canjerana (Meliaceae), in the Atlantic forest of southeast Brazil

On the forest floor of two Atlantic forest sites in southeast Brazil, we recorded 26 ant species (12 genera) interacting with the seeds of Cabralea canjerana (Meliaceae), a typical ornithochorous tree whose seeds are covered by a lipid-rich aril. The ants treat the arillate seeds in three different ways: (1) the large ponerine ants Pachycondyla striata and Odontomachus chelifer individually remove the seeds to their nests, (2) many species (Pheidole spp.) recruit workers to remove the aril on the spot, or (3) Solenopsis spp. recruit nestmates and cover the seeds with soil before removing the aril on the spot. The ants remove the aril exceptionally rapidly, and removal greatly facilitates seed germination. Seed predation by insects below fruiting trees is severe, and field experiments using vertebrate exclosures showed that rodents also prey heavily upon seeds found near parent trees. Ponerine ants actively remove seeds from this predation-prone zone. By removing bird-manipulated and naturally fallen seeds, ants can play a key role in the fate of medium-sized seeds like those of C. canjerana.