I compared the concentrations of N, P, and S in both litter and mineral soil (0–15 cm depth) from three old-growth, tropical moist forests on Barro Colorado Island (BCI), Panama. Each site was on a different substrate (i.e., parent material), but otherwise had similar climate, vegetation, and topography. There were no site differences in concentrations of N and S for either litter or soil. Concentrations of litter P and soil-extractable P were greater for the andesite (igneous rock) site than for two sites on different sedimentary rocks; however, concentrations of several other litter and soil P fractions did not differ among sites. Patterns in soil P fractions suggested advanced soil development to the point that parent material has little control of P dynamics. Litter samples from each site, leached in the laboratory, released similar amounts of N, P, and S to the soil, indicating no differences in rates of turnover in the litter and in fluxes from litter into the mineral soil among sites. I expected more site differences in soil nutrient dynamics given vastly different parent materials and soil types (i.e., Oxisol vs. Alfisol) and very shallow soil on BCI that brings the parent material close to the plant root zone. Erosion and soil mixing may explain the uniformity in soil nutrient dynamics across the sites.

As a part of the surveillance effort to monitor the ecological status of Omo Biosphere Reserve in the southwestern region of Nigeria, the aerobic heterotrophic bacterial and fungal communities of the topsoil were investigated in March 1995 and April 1996, before the onset of the rainy season. Four distinct wood-tree plantations, a core strict nature reserve (SNR) area, and a buffer zone were sampled. The topsoil samples (7.5 cm depth), including the litter, were taken with an auger (8 cm diameter) and transported to the laboratory in polyethylene bags. One-gram dry weight equivalent of sample was suspended in 10 ml sterile water, and serial dilutions from it were used for the estimation of bacterial and fungal densities. The bacterial and fungal densities ranged in the order of 10⁶ and 10³ cfu/g, respectively. Out of the 18 bacterial and 16 fungal species that were obtained, 13 and 12, respectively, were isolated from the core SNR. About 46 to 69 percent of the bacteria and 50 to 83 percent of the fungi species found in the SNR were absent in different combinations in the plantations and the buffer zone; these variations were significant among the sites monitored. The bacterial and fungal species compositions were significantly different between the SNR and each of the other sites. Proportional distributions within the sites were significant only for the bacterial communities. It would appear that plantation and human activities have caused significant changes in the distribution and species richness of the heterotrophic bacterial and fungal communities relative to the undisturbed SNR area of the Omo Biosphere Reserve.

Genetic diversity and population genetic structure of Shorea leprosula was investigated using seven natural populations distributed throughout Peninsular Malaysia and one natural population from Borneo. The mean population and species level genetic diversity were exceptionally high (H_e = 0.369 ± 0.025 and 0.406 ± 0.070, respectively). Heterozygosity varied among populations, ranging from 0.326 to 0.400, with the highest values found in the populations from central Peninsular Malaysia. Correlations among ecological factors (longitude, latitude, and annual rainfall) were not significant (P > 0.05), indicating that these ecological variables were not responsible for the observed genetic differences among populations. The Bangi adult population exhibited a higher level of observed heterozygosity but lower fixation indices in comparison to its seedling population. All other seedling populations also showed positive fixation indices (F), indicating a general excess of homozygotes. This also may suggest selection against homozygotes between the seedling and adult stages. A low level of population differentiation was detected (G_ST = 0.117 with the Lambir population and G_ST = 0.085 without the Lambir population). Furthermore, gene flow (N_m) between populations was not significantly correlated with geographical distances for the populations within Peninsular Malaysia. Cluster analysis also did not reflect geographical proximity and gave little insight into the genetic relatedness of the populations. This may indicate that the populations sampled are part of a continuous population with fragmentation having occurred in the recent past.

How important is ecotypic differentiation along elevational gradients in the tropics? Reciprocal transplants of two shrubs, Clibadium erosum (Asteraceae) and Psychotria berteriana (Rubiaceae), and a palm, Prestoea acuminata var. montana (Palmaceae), were used to test for the effect of environment and population origin on growth and physiology in the Luquillo Experimental Forest of Puerto Rico. Two sites were used, one at Pico del Este (1000 m in cloud forest) and one at El Verde (350 m in lower montane rain forest). At the cloud forest site, plastic barriers were erected around a subset of the plants to examine if protection from wind affected survival or biomass accumulation. Survival of C. erosum and P. berteriana was not affected by site, population origin, or the presence of barriers. For P. acuminata var. montana, survival was higher for plants with barriers, but not affected by site and population origin. Plants of C. erosum and P. berteriana at El Verde grew larger than at Pico del Este, but there was no effect of population origin or barrier treatment on biomass accumulation for these species. For P. acuminata var. montana, there was no effect of environment, population origin, or barrier treatment on biomass accumulation. Light-saturated photosynthetic rate (A_max) of C. erosum, P. berteriana, and P. acuminata var. montana, as well as leaf anatomical characteristics of C. erosum, were unaffected by environment, population origin, and barrier treatment. On balance, there seems to be little evidence of ecotypic differentiation in these species along the gradient.

Cacti growing in forests potentially experience growth limitation due to reduced light availability. To test this hypothesis, we studied the population structure of Opuntia echios var. gigantea at 15 sites on the south side of Isla Santa Cruz, Galápagos Islands, Ecuador. Populations were located in communities ranging from arid scrub at low elevations to closed-canopy tropical dry forest at higher elevations. Ordination confirmed the existence of a strong elevation–vegetation gradient. Opuntia abundance peaked at lower elevations (<ca 30 m), with lower densities in closed-canopy sites. For populations in scrub vegetation, density declined fairly regularly with plant height. Populations in forested sites had few plants of intermediate height, suggesting periodic recruitment. Scrub populations had random dispersion, while those in forests were aggregated. The change in spatial pattern may be related to a change in primary reproductive mode from asexual propagation via fallen fruits to propagation via fallen cladodes. Height was significantly correlated with stem diameter. Intercepts of these relationships increased toward higher elevations, probably in response to the increasing height of the surrounding canopy.

After Hurricane Andrew crossed southern Florida (U.S.A.) on 24 August 1992, native and exotic pioneer species in subtropical hardwood forests (hammocks) regenerated from seed banks. Regeneration occurred in hammocks of metropolitan Dade County and the Long Pine Key region of Everglades National Park. The density of the native pioneer Trema micrantha was significantly higher in hammocks of Long Pine Key than in those of metropolitan Dade County. In contrast, the basal area of the exotic pioneer Carica papaya was greater in Dade County hammocks than Long Pine Key hammocks. Although T. micrantha tended to be restricted to areas of soil disturbance (tip-up pits) formed by trees uprooted during Hurricane Andrew, especially in Long Pine Key, C. papaya was located throughout hammocks. These results suggest differences in the regeneration niches in which the native T. micrantha required more specific disruptions (i.e., both canopy and soil) than C. papaya (only removal of canopy) for establishment. A broad regeneration niche could in part account for the capability of an exotic species with a dormant seed bank to invade native subtropical forests following natural large-scale disturbances.

This study tested three hypotheses regarding how plants respond to the spatial heterogeneity in light availability in the rain forest understory: (1) understory plants occur preferentially in the lighter parts of the understory; (2) understory palms are more shade tolerant than other understory plants; (3) rain forest plants differ in their ontogenetic response to understory light conditions. The study was carried out in old-growth rain forest in the Yasuní National Park, Amazonian Ecuador. The hypotheses were tested by comparing the distributions of 20 plant species (1454 individuals) over microsites with differing degrees of exposure to canopy gaps to the background distribution of these microsites in the forest. The gap exposure of a given microsite was described by an index based on the number and size of gaps in the canopy to which the site was exposed. Two plant height classes were studied: 0.80–2.49 and 2.50–5 m. The first and third hypotheses were accepted, while the second hypothesis was rejected. The results for the individual species corresponded well with what is known from earlier studies about the ecology of these species or close relatives, suggesting that the patterns observed can be generalized for Neotropical rain forests. Notably, the most abundant species in the study represent several different life history strategies. Thus, abundance in the rain forest understory can be achieved by several different strategies. This suggests that niche differentiation in terms of response to small changes in understory light conditions may be an important factor in the maintenance of the high local plant species richness of tropical rain forests.

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N-fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N-fixing symbionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer-term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.

Phenological characteristics of 453 individuals representing 39 tree species were investigated in two dry forests of the Lomerío region, Department of Santa Cruz, Bolivia. The leaf, flower, and fruit production of canopy and sub-canopy forest tree species were recorded monthly over a two-year period. Most canopy species lost their leaves during the dry season, whereas nearly all sub-canopy species retained their leaves. Peak leaf fall for canopy trees coincided with the peak of the dry season in July and August. Flushing of new leaves was complete by November in the early rainy season. Flowering and fruiting were bimodal, with a major peak occurring at the end of the dry season (August–October) and a minor peak during the rainy season (January). Fruit development was sufficiently long in this forest that fruiting peaks actually tended to precede flowering peaks by one month. A scarcity of fruit was observed in May, corresponding to the end of the rainy season. With the exception of figs (Ficus), most species had fairly synchronous fruit production. Most canopy trees had small, wind dispersed seeds or fruits that matured during the latter part of the dry season, whereas many sub-canopy tree species produced larger animal- or gravity-dispersed fruits that matured during the peak of the rainy season. Most species produced fruit annually. Lomerío received less rainfall than other tropical dry forests in which phenological studies have been conducted, but rainfall can be plentiful during the dry season in association with the passage of Antarctic cold fronts. Still, phenological patterns in Bolivian dry forests appear to be similar to those of other Neotropical dry forests.

Most orchids studied thus far show long-term resource adjustments to increases in fruit production within a flowering season, but none of these offers rewards to their potential pollinators. If nectar production is energetically expensive, then resources utilized to produce fruits and seeds may be even more limited in pollinator-rewarding orchids than in non-rewarding ones. Thus, resource adjustments may be more dramatic or entirely different in nectar producing plants. In this study, we performed artificial hand-pollinations for two consecutive flowering seasons in the nectar producing orchid Comparettia falcata, and tested whether or not fruit set, seed set, and seed viability were limited by the quantity of pollinations or by resources. In addition, we compared mechanisms of short-term (fruit abortion within seasons) and long-term consequences (percent change in leaf length and change in flower number per plant between seasons, probability of shoot and inflorescence production, and mortality) between hand-pollinated and unmanipulated plants. The relationships among plant traits related to vegetative size and reproduction also were examined. Hand-pollinations showed some negative effects. Fruit set was higher in hand-pollinated plants in the first season but was similar to the controls in the second. Seed set was significantly lower and abortions were higher than in unmanipulated plants. On the other hand, some of our measurements were unaffected by the hand-pollination treatment. Unexpectedly, there were no significant differences between groups in percent change in leaf length, change in flower number per plant between seasons, or the probability of shoot and inflorescence production. Although there was a strong correlation between leaf size and the number of flowers produced within a season, associations between leaf size and traits related to current or future reproduction were not consistent. Like other epiphytic orchids, pollination limitation occurred within a single season in C. falcata, but increases in fruit production also resulted in reduced lifetime fitness as estimated by a compounded fitness index. Contrary to all other epiphytic orchids studied, long-term adjustments to increased fruit production in C. falcata through reduction in future growth or flower and inflorescence production were either minor or lacking. Our results suggest that the nature of plant strategies associated with resource constraints during sexual reproduction may be dependent on whether or not plants have evolved traits that are costly.

Dioecy in the Solanaceae is rare, occurring in <1 percent of the species worldwide and known in only two species from South America. We report the occurrence of cryptic dioecy in the Neotropical genus Deprea. Studying herbarium material of the Venezuelan endemic D. paneroi revealed morphological distinctions that tentatively served to divide collections into male and female morphs. This discovery prompted investigation of population structure, crossing studies, and morphometric analyses of 95 individuals from the largest known wild population and a small sample of greenhouse-grown plants. About 60 percent of the population was reproductive, and of these, nearly equal numbers exhibited female or male characteristics. Plants that bore fruit and small sterile anthers were characterized as female; those that lacked fruit but produced pollen in large anthers were characterized as male. The morphology, receptivity, and developmental phenology of pistils were identical in both forms. Crossing studies revealed significant enlargement of the ovaries in open-pollinated female flowers and female × male hand-pollinations. The ovaries in open-pollinated male flowers and male × male hand-pollinations never enlarged. These results suggest that the only legitimate combination is female × male. Dioecy in the family and possible mechanisms for the evolution and maintenance of dioecy in D. paneroi are discussed.

A high frequency of dieback and mortality was found in Rhizophora mangle (red mangrove) in coastal mangrove forests of southwest Puerto Rico. Eight plots were established in a variety of mangrove environments to assess the extent and severity of the dieback and mortality. The imperfect fungus Cytospora rhizophorae was isolated consistently from stem dieback and associated canker tissues. No other pathogenic microorganisms were isolated. Pathogenicity was demonstrated in greenhouse and field inoculation experiments with C. rhizophorae using young R. mangle seedlings. In field studies of seedlings inoculated, mortality was 33 percent, and greenhouse mortality was as high as 50 percent. Cytospora rhizophorae was isolated consistently from diseased mangrove tissue; the fungus was grown in pure culture and then inoculated into healthy red mangroves, causing cankers. Koch's postulates were fulfilled by infecting R. mangle with C. rhizophorae, producing disease symptoms, and successfully re-isolating C. rhizophorae from the diseased tissue. This is the first documentation of the pathogenicity of C. rhizophorae associated with mortality in red mangrove. In the coastal mangrove forests of southwest Puerto Rico, C. rhizophorae may play an important role in the frequency of dieback and mortality in R. mangle, producing slow stem diameter growth and causing frequent stem and root wounds.

The Janzen–Connell model of tropical forest tree diversity predicts that seedlings and young trees growing close to conspecific adults should experience higher levels of damage and mortality from herbivorous insects, with the adult trees acting as either an attractant or source of the herbivores. Previous research in a seasonal forest showed that this pattern of distance-dependent herbivory occurred in the early wet season during the peak of new leaf production. I hypothesized that distance-dependent herbivory may occur at this time because the new foliage in the canopy attracts high numbers of herbivores that are limited to feeding on young leaves. As a consequence, seedlings and saplings growing close to these adults are more likely to be discovered and damaged by these herbivores. In the late wet season, when there is little leaf production in the canopy, leaf damage is spread more evenly throughout the forest and distance dependence disappears. I tested three predictions based on this hypothesis: (1) the same species of insect herbivores attack young and adult trees of a given plant species; (2) herbivore densities increase on adult trees during leaf production; and (3) herbivore densities in the understory rise during the course of the wet season. Censuses were conducted on adults and saplings of two tree species, Quararibea asterolepis and Alseis blackiana. Adults and saplings of both species had largely the same suite of chewing herbivore species. On adults of Q. asterolepis, the density of chewing herbivores increased 6–10 times during leaf production, but there was no increase in herbivore density on adults of A. blackiana. Herbivore densities increased 4.5 times on A. blackiana saplings and 8.9 times on Q. asterolepis saplings during the wet season, but there were no clear trends on the adults of either species. These results suggest that the potential of adult trees as a source of herbivores on saplings depends on the value of new leaves to a tree species' herbivores, which may differ across tree species.

To examine interspecific variation in the intensity of ant defense among three sympatric species of obligate myrmecophytes of Macaranga (Euphorbiaceae), we measured the ratio of ant biomass to plant biomass, ant aggressiveness to artificial damage on host plants, and increase in herbivore damage on host plants when symbiont ants were removed. Increase in herbivore damage from two- and four-week ant exclusion varied significantly among the three species. The decreasing order of vulnerability to herbivory was M. winkleri, M. trachyphylla, and M. beccariana. The ant/plant biomass ratio (= rate of the dry weight of whole ant colonies to the dry weight of whole aboveground plant parts) and ant agressiveness also varied significantly among the three species; the orders of both the ant/plant biomass ratio and ant aggressiveness were the same as in the herbivory increase. These results indicated that the intensity of ant defense differs predictably among sympatric species of obligate myrmecophytes on Macaranga. In addition to the interspecific difference in the total intensity of ant defense, when symbiont ants were excluded, both patterns of within-plant variation in the amount of herbivore damage and compositions of herbivore species that caused the damage differed among species. This suggests that the three Macaranga species have different systems of ant defense with reference to what parts of plant tissue are protected and what herbivorous species are avoided by ant defense. Thus, it is important to consider the interspecific variation in ant defense among Macaranga species to understand the herbivore community on Macaranga plants and the mechanisms that promote the coexistence of multiple Macaranga myrmecophytes.

Dispersal quality, as estimated by the cumulative effects of dispersal, germination, seed predation, and seedling survival, was examined for Beilschmiedia pendula (Lauraceae) in Monteverde, Costa Rica. I determined the pattern of dispersal by finding seeds deposited by birds, protected the seeds from seed predators with cages to assess germination and seedling survival, and examined seed predation rates with marked seeds. Seed predation, germination, and seedling survival were compared between seeds naturally dispersed by birds and seeds placed at randomly located sites.

Approximately 70 percent of seeds dispersed by birds (N = 244) were deposited <10 m from crown edges of fruiting B. pendula trees, although some seeds were dispersed at least 70 m away. Larger seeds were more likely to be dispersed under or close to the parent trees, and larger seeds produced larger seedlings. Seed size was not correlated directly with seedling survival, but larger seedlings at three months were most likely to survive one year. Seed predation by mammals and insects and seedling mortality due to fungal pathogens were concentrated beneath the crowns of parent trees. Seedlings and saplings were more abundant beneath fruiting B. pendula trees, but individuals farther away were taller on average. Thus, dispersal is beneficial for B. pendula, but such benefits appear most pronounced at a small spatial scale; seeds dispersed >30 m from the crown edges actually had a lower probability of survival than those dispersed 10–20 m. Only 10 percent of B. pendula seeds received high-quality dispersal in terms of landing in the zone with the highest per seed probability of seedling survival 10–20 m from parental crowns.

We examined characteristics of roosting sites utilized by two flying fox species (Pteropus tonganus and P. samoensis) in American Samoa. The colonial roosting sites of P. tonganus were observed over a ten-year period, including two years when severe hurricanes devastated bat populations and destroyed roost trees. Prior to the hurricanes, roosts were located on cliff faces above the ocean or steep mountainsides, locations that were either inaccessible to people or in protected areas where hunting was not allowed. In the years immediately following the hurricanes, P. tonganus colonies split into smaller groups that moved frequently to different locations. Four years after the second hurricane, colonies had coalesced and returned to many of the traditional roosting sites used before the hurricanes. Common tree species in upland and coastal forest were selected as roosts. The isolated locations selected for P. tonganus roosts were apparently the result of hunting pressure on the colonies. The solitary roosts of P. samoensis were observed during 29 months. Roosting bats were well concealed and hard to detect within the forest; even bats on exposed branches were cryptic. Mature primary forest was favored as roosting habitat. Individual bats used specific branches or trees as roosts and returned to them for up to 29 months. Unlike P. tonganus, people did not alarm roosting P. samoensis easily and some roosts were located near houses and along roads.