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.     

Influence of Edge Exposure on Tree Seedling Species Recruitment in Tropical Rain Forest Fragments1

Edge creation has a pronounced influence on the understory vegetation, but the effects of edges on seedling species recruitment are still poorly understood. In Central Amazonia, 9–19 years after fragmentation, we recorded species richness and net seedling recruitment rate in 1 ha blocks exposed to none, one, or multiple edges within forest fragments. One‐hectare blocks were located in the center (no edge), the edge (one edge), the corners (two edges) of 10 and 100 ha fragments, and in a 1 ha fragment (four edges). In 1991, we counted all tree seedlings 5–100 cm tall found within permanent 1 m² plots located within the 1 ha blocks. In May 1993, we manually removed all seedlings that were smaller than 1 m tall from the permanent plots. Six years and five months later (October 1999), all new seedlings recruited into the plots were counted and classified into distinct morphospecies. Species richness of recruited seedlings, scaled by total seedling density, declined from the center to the edge, the corner blocks, and then to the 1 ha fragment. Overall, the four‐edged, 1 ha fragment had the poorest species richness and the non‐edged 100 ha central block the highest. The total number of recruited individuals was 40 percent less than that previously present, with the 100 ha corner having the lowest recruitment. Pairwise comparisons showed that species similarity was related to edge number for the 100 and 1 ha fragments. Species rank/abundance curves showed that a subset of species was common in all blocks within the fragments, and that the 100 ha center held more rare species than any other 1 ha block. This study demonstrated that, in a given fragment patch, the number of tree seedling species recruited varied inversely with the number of edges.     

Effects of Land Crabs on Leaf Litter Distributions and Accumulations in a Mainland Tropical Rain Forest1

The effect of the fossorial land crab Gecarcinus quadratus (Gecarcinidae) on patterns of accumulation and distribution of leaf litter was studied for two years in the coastal primary forests of Costa Rica's Corcovado National Park. Within this mainland forest, G, quadratus achieve densities up to 6 crabs/m² in populations extending along the Park's Pacific coastline and inland for ca 600 m. Crabs selectively forage for fallen leaf litter and relocate what they collect to burrow chambers that extend from 15 to 150 cm deep (N= 44), averaging (±SE) 48.9 ± 3.0 cm. Preference trials suggested that leaf choice by crabs may be species‐specific. Excavated crab burrows revealed maximum leaf collections of 11.75 g dry mass– 2.5 times more leaf litter than collected by square‐meter leaf fall traps over several seven‐day sampling periods. Additionally, experimental crab exclosures (25 m²) were established using a repeated measures randomized block design to test for changes in leaf litter as a function of reduced crab density. Exclosures accumulated significantly more (5.6 ± 3.9 times) leaf litter than did control treatments during the wet, but not the dry, seasons over this two‐year study. Such extensive litter relocation by land crabs may affect profiles of soil organic carbon, rooting, and seedling distributions.     

Species-specific Seedling Responses to Hurricane Disturbance in a Puerto Rican Rain Forest1

Seedling dynamics were followed in a Puerto Rican forest for 20 months following a severe hurricane to study the interactive effects of hurricane debris, nutrients, and light on seedling diversity, density, growth, and mortality. Three treatments (debris removal, an unaltered control with hurricane debris, and chemical fertilization added to hurricane debris) altered levels of forest debris and soil nutrients. Canopy openness was measured twice using hemispherical photographs of the canopy. We examined the demographic responses of six common species to treatments over time. Seedling densities increased for all six species but the only significant treatment effects were increased densities of the pioneer tree Cecropia and the shrub Palicourea in the debris removal treatment. Seedling growth declined with declining light levels for four species but not for the pioneer tree Alchornea or the non‐pioneer tree Dacryodes. Only Cecropia and the non‐pioneer tree Chionanthus had treatment effects on growth. Mortality also differed among species and tended to be highest in the fertilized plots for all but Cecropia and Dacryodes. We found only some of the expected differences between pioneer and non‐pioneer plants, as each species had a unique response to the patchy distributions of organic debris, nutrients, and light following the hurricane. High local species diversity was maintained through the individualistic responses of seedlings after a disturbance.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Roosting Requirements of Two Frugivorous Bats (Sturnira lilium and Arbiteus intermedius) in Fragmented Neotropical Forest1

As tropical forest fragmentation accelerates, scientists are concerned with the loss of species, particularly those that play important ecological roles. Because bats play a vital role as the primary seed dispersers in cleared areas, maintaining healthy bat populations is critical to natural forest regeneration. Observations of foraging bats suggest that many Neotropical fruit‐eating species have fairly general habitat requirements and can forage in many different kinds of disturbed vegetation; however, their roosting requirements may be quite different. To test whether or not general foraging requirements are matched by equally broad roosting requirements, we used radiotelemetry to locate roost sites of two common frugivorous bat species (Sturnira lilium and Artibeus intermedius) in a fragmented forest in southeastern Mexico. Sturnira lilium roosted inside tree cavities and selected large‐diameter roost trees in remnant patches of mature forest. Fewer than 2 percent of trees surveyed had a mean diameter equal to or greater than roost trees used by . S. lilium, Artibeus intermedius roosted externally on branches and vines and under palm leaves and selected roost trees of much smaller diameter. Compared to random trees, roost trees chosen by A. intermedius were closer to neighboring taller trees and also closer in height to these trees. Such trees likely provide cryptic roosts beneath multiple overlapping crowns, with sufficient shelter from predators and the elements. While males of A. intermedius generally roosted alone in small trees within secondary forest, females roosted in small groups in larger trees within mature forest and commuted more than three times farther than males to reach their roost sites. Loss of mature forest could impair the ability of frugivorous bats to locate suitable roost sites. This could have a negative impact on bat populations, which in turn could decrease forest regeneration in impacted areas.     

Pleistocene Plant Fossils in and near La Selva Biological Station, Costa Rica1

Radiocarbon dating and ⁴⁰Ar/³⁹Ar analysis of overlying tephra indicate that plant fossil assemblages exposed by stream erosion and well construction in and near La Selva Biological Station in eastern lowland Costa Rica are Pleistocene in age. We identified plant taxa based on wood, leaves, fruits, seeds, pollen, and spores examined from three sites at ca 30 m elevation. Extrapolating from modern ranges and surface temperature lapse rates suggests paleotemperatures 2.5–3.1°C cooler than at present     

Fragmentation Effects on Diversity of Wasp Community and Its Impact on Fig／Fig Wasp Interaction in Ficus racemosa L.

Habitat fragmentation usually results in alteration of species composition or biological communities. However, little is known about the effect of habitat fragmentation on the fig/fig wasp system.In this study, we compared the structure of a fig wasp community and the interaction between figs and fig wasps of Ficus racemosa L. in a primary forest, a locally fragmented forest and a highly fragmented forest.Our results show that, in the highly fragmented forest, the proportion of pollinator wasps is lower and the proportion of non-pollinator wasps is higher compared with the primary forest and locally fragmented forest. The proportion of fruits without pollinator wasps in mature fruits is also greatly increased in the highly fragmented forest. The proportion of galls in all female flowers increases in the highly fragmented forest, whereas the proportion of viable seeds does not change considerably. The disruption of groups of fig trees results in a decrease in pollinator wasps and even might result in the extinction of pollinator wasps in some extreme cases, which may transform the reciprocal interaction between figs and fig wasps into a parasite/host system. Such an effect may lead to the local extinction of this keystone plant resource of rain forests in the process of evolution, and thereby, may change the structure and function of the tropical rain forest.     

Density, Distribution, and Attributes of Tree Cavities in an Old-Growth Tropical Rain Forest

Tree cavities are a critical resource for many animals, especially as nesting sites for birds. Patterns of cavity distribution in temperate forests are well studied, yet little is known of cavities in tropical forests, despite a hypothesized decrease in cavity availability with decreasing latitude. We studied cavity density and distribution in a wet lowland tropical forest in Costa Rica and compared our results with estimates from forests around the world. Cavities at our site were common, occurred frequently in living trees, and were often formed by damage or decay rather than by woodpeckers. Most cavities had small openings, and woodpecker-created cavities were nonrandomly oriented. Contrary to prediction, cavity density appears to increase from the poles to the tropics. We suggest potential mechanisms to explain these patterns.     

Temporal and Spatial Diversity and Distribution of Arboreal Carabidae (Coleoptera) in a Western Amazonian Rain Forest1

Diversity of arboreal carabid beetles was sampled by fumigation in 100 3 × 3 m stations within a 100 × 1000 m terra firme forest plot in Ecuadorian Amazonia. Nine sampling dates from January 1994 to October 1996 yielded 2329 individuals belonging to 318 species of which more than 50 percent were undescribed species. A high percentage of the species sampled were rare; the proportion that occurred once per sampling date (singletons) ranged from 50.0 to 62.5 percent. Estimates of species richness were from 82 to 282 species of arboreal carabids in the study plot on a given sampling date. Most richness values were greater than 173 species. Species accumulation curves attained asymptotes for all but one sampling date, indicating that an adequate level of sampling effort was used to characterize the diversity of carabid fauna. Total accumulation curves based on pooled data failed to reach asymptotes. There was a high turnover in species composition between sampling dates; less than 50 percent of the species between the majority of sampling dates were shared, suggesting that the total species pool may be extremely large. Although species composition changed seasonally, species richness varied little. Spatial autocorrelation analysis revealed that the structure of this species assemblage was significantly patterned at distances below 280 m. Taken together, the large percentage of undescribed species, die failure of the overall species accumulation curves to level off, and the high turnover in species composition indicate that the species diversity of carabid beetles is far higher than previously thought.     

Home Range, Habitat Use, and Activity of Baird's Tapir in Costa Rica1

Home range size, habitat utilization, and activity patterns of five adult Bairds tapirs (Tapirus bairdii) were studied via radiotelemetry from June 1995 to May 1996 in Corcovado National Park, Costa Rica. Estimates of 4153 animal locations were made. Home range sizes (95% minimum convex polygon) for the entire study period (wet season and dry season) averaged 125.0 ha (SD = 72.7), 94.9 ha (SD = 47.6), and 96.8 ha (SD = 51.2), respectively. Monthly home range sizes averaged 55.5 ha (SD = 32.1) and did not vary between seasons. Across the entire study, secondary forest, the most commonly used habitat type (61.3%) was utilized more than expected as predicted by availability (49.6%), while the second most used habitat, primary forest, was used (25.0%) less than expected (36.2%). The same general habitat selection pattern was observed in the dry season. Wet season habitat utilization values showed tapirs using both primary and secondary forest habitats in proportion to their availability. Diurnal and nocturnal activity levels were estimated at 20.2 and 80.4 percent, respectively. More diurnal and less nocturnal activity was observed during the wet season compared to the dry season.     

Variation in Seedling Density and Seed Predation Indicators for the Emergent Tree Dipteryx panamensis in Continuous and Fragmented Rain Forest1

Seedling density and the condition of stony endocarps of the tree Dipteryx panamensis were assessed in protected continuous forest and two forest fragments exposed to hunting and selective logging. Seedling density was higher in forest fragments than in continuous forest, while more whole endocarps and fewer chewed and half endocarps were found in fragments, indicating lower seed predation at fragment sites. These findings appear to contradict two earlier D. panamensis studies and we discuss methodological differences that could account for our disparate results. Hunting and fragmentation effects on mammal populations are suggested as a cause for the altered recruitment pattern in fragments.     

Seed Dynamics in Relation to Gaps in a Tropical Montane Rainforest of Hainan Island, South China： （I） Seed Rain

Seed dynamics is an important part of stand dynamics in forest ecosystems. In this paper, 26 gaps were randomly selected to study the influence of gaps on the spatial and temporal patterns of seed rains in a tropical montane rainforest of Hainan Island, South China. Three zones for each gap, including outside gap zone （Non-gap）, transitional gap zone （EG-CG）, and central gap zone （CG）, were designed, and fourseed traps （each lm x lm in size） were placed in each zone. Seed rains were collected by these traps every 10 days from June 2001 to May 2002. Seed rain varied greatly with season and generally exhibited a pattern of unimodal change during the study period： seed abundance and species richness were both greater in the wet season than in the dry season. Gaps significantly influenced the temporal patterns of both species richness and density of seed rains. Gaps had no significant influences on the spatial distribution patterns of seed rain species richness, but significantly affected the spatial distribution pattern of seed rain densities. Among the three different zones of gaps, the outside gap zone generally received more seeds inputs than the two other gap zones.     

Ecology of Aglaia mackiana (Meliaceae) Seedlings in a New Guinea Rain Forest1

The large seeds of Aglaia mackiana (Meliaceae) germinate and produce vigorous seedlings under closed canopies or in large gaps. To assess seedling ecology after germination, we measured growth, herbivore damage, and survivorship of seedlings over one year. The sample included shaded seedlings from dispersed seeds, undispersed seeds under parent trees, and seedlings transplanted to gaps. We quantified the light environment using hemispherical canopy photographs taken above seedlings at the beginning and end of the one–year study. Seedlings transplanted to gaps grew faster and had more leaves, larger total leaf surface area, longer secondary roots, and greater root mass than shaded seedlings. Seedlings in gaps did not differ from shaded seedlings in survivorship or amount of herbivore– and pathogen–caused leaf damage. The canopy photographs taken one year apart suggest there is a rough equilibrium in closed canopies with slight changes occurring around an average light level. Sites with < 0.06 ISF (a unitless, relative measure of canopy openness or reflected sunlight) tended to remain the same with minor fluctuations toward brighter or darker. Sites with canopy openness > 0.06 ISF tended to close; few gaps grew larger. Seedlings under parenr trees and seedlings away from parent trees had similar amounts of leaf damage and virtually identical survivorship after 18 months, but seedlings under parent trees had slower growth rates and smaller total leaf surface areas. Dispersal did not strongly benefit seeds via escaping high levels of mortality or comperition around the parent.     

Seed Dispersal by the Sloth Bear (Melursus ursinus) in South India1

Previous studies have shown that sloth bears are mainly myrmecophagous, but in this study, 39 and 83 percent (wet and dry season, respectively) of scats surveyed in South India contained fruits. Seeds from three of six plant species collected from scats germinated faster than seeds not passing through bears; germination percentage was not higher in any of the six species. These results suggest that the sloth bear may play an important role in the population dynamics of fleshy‐fruited species of the region.     

Influence of Microclimate on the Occurrence of Cyanobacteria in the Phyllosphere in a Premontane Rain Forest of Costa Rica

Abstract: The occurrence of cyanobacteria in the phyllosphere of several understory plants was investigated during a one and a half year study in a primary premontane rain forest in Costa Rica. Altogether, seven species of epiphyllous cyanobacteria were identified and are discussed here. The ecology of the two most frequent species in the understory, Scytonema javanicum and Sc. hofmannii , was studied in further detail. Within the Understory, the relative abundance of these species, as well as of epiphyllous bryophytes, was more influenced by air humidity than by light. The humidity, characterized as potential evaporation, also had influence on the pseudosuccession of the phyllosphere vegetation. While at moist sites bryophytes and both species of Scytonema appeared nearly simultaneously on 6–9-month-old leaves, at dryer sites Sc. javanicum , and especially Sc. hofmannii , appeared 6–9 months after bryophytes had established. On 13–15-month-old leaves bryophytes and Scytoriema were usually well established. On average, 20–30% of the leaf area of 2–5-year old leaves of Spathacanthus hoffmannii (Acan-thaceae, an abundant understory shrub) were covered by bryophytes, 2–3% by Sc. javanicum , and 0.1–0.2% by Sc. hofmannhi . An influence of phorophyte species on the occurrence and abundance of these two species of Scytonema could not be detected. However, Sc. hofmannii was more frequent on leaves with higher bryophyte cover, while Sc. javanicum was independent from bryophytes. A comparison of leaves from the understory and from the canopy showed that the composition and abundance of cyanobacteria species changed with height above forest floor.     

Survival, Growth, and Ecosystem Dynamics of Displaced Bromeliads in a Montane Tropical Forest1

Epiphytes generally occupy arboreal perches, which are inherently unstable environments due to periodic windstorms, branch falls, and treefalls. During high wind events, arboreal bromeliads are often knocked from the canopy and deposited on the forest floor. In this study, we used a common epiphytic tank bromeliad, Guzmania berteroniana (R. & S.) Mez, to determine if fallen bromeliads can survive, grow, and reproduce on the forest floor and evaluate the potential impact of adult dispersal on plant and soil nutrient pools. Bromeliads were transplanted to and from tree stems and the forest floor and monitored intensively for six months; survival, growth, and impacts on ecosystem nutrient pools were followed on a subset of plants for 16 months. Six months after transplanting, bromeliad mortality was low (3%), and 19 percent of study individuals had flowered and produced new juvenile shoots. Mortality on the subset of plants followed for 16 months was 14–30 percent. Although survival rates were relatively high in all habitats, bromeliads transplanted to trees grew significantly more root length (x?± SE: 189 ± 43 cm) than those moved to the forest floor (53 ±15 cm) and experienced lower rates of leaf area loss. All transplanted bromeliads rapidly altered the substrate they occupied. Individuals transplanted to and among trees rapidly decreased base cation concentrations but significantly increased P concentrations of their underlying substrate. On the ground, bromeliads increased C, N, and P concentrations within nine months of placement. Our results suggest that in this montane tropical forest, bromeliads respond rapidly to displacement, locally modify their substrates, and can access the resources needed for survival regardless of habitat.     

Reproduction of the Bat Glossophaga commissarisi (Phyllostomidae: Glossophaginae) in the Costa Rican Rain Forest During Frugivorous and Nectarivorous Periods1

I studied the reproduction of the 9 g nectarivorous bat Glossophaga commissarisi (Phyllostomidae: Glossophaginae) in relation to its food choice in a Costa Rican lowland rain forest over more than 2 yr. Reproduction was bimodal with birth peaks in April/May and October/November. The first birth peak coincided with a period of frugivory, whereas the second occurred during a period of almost exclusive nectarivory. All adult females recaptured in consecutive reproductive periods were reproductively active on both occasions, indicating that individual G. commissarisi females regularly reproduce twice per year at the study site. Throughout the annual cycle the mean testis length of the males correlated best with the proportion of females lactating, suggesting a post partum estrous pattern. Animals became reproductively active at about 1 yr of age. Fecal samples showed that females consumed significantly more insects than males.     

The Structure of Ground Beetle Assemblages (Coleoptera: Carabidae) at Fruit Falls of Melastomataceae Trees in a Brazilian Terra Firme Rain Forest1

Although most Carabidae are recognized as predacious, recent studies have shown that some tropical rain forest Carabidae use the small seeds of fallen fruit from some tree species as a key food resource (particularly figs [Moraceae] but also those from other families). Assemblages of carabid beetles (Coleoptera: Carabidae) at fruit falls of the Melastomataceae tree genera, Bellucia, Loreya, and Miconia, were studied in a terra firme rain forest near Manaus (Amazonia, Brazil) from February 1992 to August 1996. Of 5483 specimens of 14 carabid species collected at 53 fruit falls, 98.2 percent of the collected beetles represented the seed-feeding genus Notiobia. Four of the 7 Notiobia species were found to reproduce at Melastomataceae fruit falls. The most abundant species, N. glabrata, was found to be a Melastomataceae seed generalise while N. maxima seemed to prefer the seeds of Bellucia, and N. aulica and Af. umbrifera the seeds of Miconia. The presence of larvae indicated reproduction of N. glabrata at fruit falls of all three tree genera and reproduction of N. maxima at those of Bellucia. Fruit falls of different species and genera of Melastomataceae appeared to occur at different times of die year and not continuously throughout the year. The seed-feeding Notiobia species, which specialize on Melastomataceae, used fruit falls of figs as migratory “stepping stones” during periods when there were no Melastomataceae fruit falls. The comparable suite of fig seed-feeding specialist species of Notiobia did not appear to do the reverse and seemed to avoid Melastomataceae fruit falls. The carabid assemblage at fig fruit falls was more diverse than the assemblage at those of Melastomataceae. Among the Melastomataceae, carabid species richness was highest at Miconia fruit falls and lowest at Loreya fruit falls. The three most common predatory species caught at Ficus fruit falls were also caught at those of Melastomataceae and appeared to reproduce at both fruit fall types. The proportion of the carabid assemblage accounted by predatory species was much lower at Melastomataceae (1.8%) than at fig fruit falls (8%).