Fragment-size determination by workers of the grass-cutting ant Atta vollenweideri and the extent of size-matching between ant size and fragment size were investigated as a function of the distance from the nest. Foragers and their loads were sampled in four different sectors along a 30-m foraging trail in the field: directly on the harvested source, in the patch, on the trail and close to the nest. Ants sampled on the source immediately after cutting were significantly larger than carrying ants on any trail sector, whereas body mass did not differ among other samples. Thus, foraging was partitioned into at least two stages, with large ants cutting the source and smaller ants carrying the fragments to the nest. Fragments collected directly after cutting were significantly larger than those carried on the trail, indicating that the fragments were cut once again on their way to the nest. Size-matching depended on the trail sector considered. It was stronger in ants sampled closer to the nest, suggesting that carriers either collect dropped fragments according to their body mass, or cut fragments out of the dropped fragments in sizes corresponding to their body mass prior to transport.

We examined the distribution, germination, growth and photosynthetic characteristics of two co-existing morphotypes of the pioneer tree Trema micrantha at the Barro Colorado Nature Monument (BCNM), Panama. Morphotypes differed significantly in distribution and in seed characteristics. A ‘large’-seeded morphotype (endocarp mass=3.83 mg) was associated with treefall gaps in the forest interior, whereas a ‘small’-seeded morphotype (endocarp mass=1.38 mg) was found predominantly on landslides on the margins of Lake Gatun. Seeds of the small-seeded morphotype germinated faster than seeds of the large-seeded morphotype, with seedlings of the small-seeded morphotype showing both a higher Unit Leaf Rate (ULR) and a lower Specific Leaf Area (SLA). Differences in photosynthetic rates reflected differences in SLA; the small-seeded morphotype had a higher rate on a leaf area basis, while the large-seeded morphotype had a higher rate on a leaf mass basis. Although allocation patterns between morphotypes varied in a way consistent with known interspecific differences between ‘sun’ and ‘shade’ plants, relative growth rates (RGR) of the morphotypes were similar across different light conditions suggesting that factors other than light, such as water uptake efficiency, soil nutrient requirements, and perhaps seed dispersal characters may explain the habitat partitioning of morphotypes.

Microhabitat use and habitat-abundance relationships of four freshwater shrimps, Atyopsis moluccensis, Macrobrachium pilimanus, Macrobrachium trompii and Macrobrachium neglectum, were surveyed in tropical streams running through primary and secondary forests in Borneo, East Malaysia. Underwater observations revealed that A. moluccensis preferred relatively high water velocity and a boulder substrate. Macrobrachium pilimanus also preferred high water velocity and a cobble substrate, whereas M. trompii occupied stream margins with slow current and fine substrates (from POM (particulate organic matter) to pebbles). In contrast, M. neglectum was distributed relatively evenly through the stream channel. The abundance of A. moluccensis, M. pilimanus and M. trompii in the stream reaches was best explained by the abundance of boulders, cobbles and POM, respectively, suggesting that the amount of preferred microhabitat is an important factor affecting shrimp abundances in the tropical rain-forest streams. The primary-forest reaches were dominated by coarse substrates, such as cobbles and boulders, while a great proportion of the streambeds in the secondary-forest reaches were covered with sand. Owing probably to such habitat differences, the abundance of both A. moluccensis and M. pilimanus, which preferred coarse substrates, was less in the secondary- than in the primary-forest reaches. These suggested that loss of preferred habitat, namely decreased coarse substrate availability, by sedimentation resulting from riparian deforestation had altered the shrimp assemblage structures.

Patterns of species composition in a neotropical termite assemblage were studied in relation to early effects of forest fragmentation as well as other sources of environmental and spatial heterogeneity. Termite diversity surveys were carried out at three mainland forest sites, and 10 islands of varying size, in an area of lowland tropical forest that had been flooded 4 y earlier, during the creation of the Petit Saut hydroelectric project in French Guiana. The ‘ghost forest’ of dead emergent trees in the flooded zone was also studied for its influence on island termite assemblages. Results suggested that the effects of forest fragmentation upon the total assemblage, and upon soil-feeders in particular, were subordinate to the influence of forest understorey palm density, and the closely associated gradients of soil humus depth and soil pH. Moreover, gradients for these three variables were uncorrelated with forest fragmentation and probably reflected spatial environmental heterogeneity pre-dating inundation events. Nevertheless, factors associated with forest fragmentation appeared to have had a significant effect on changes in termite species composition across the study site, primarily in structuring the wood- and leaf-litter-feeder assemblage. Effects upon the latter were not apparently a result of influx of species from the ghost forest. Purely spatial variation also influenced β-diversity changes in species composition across the site. In conclusion, the effects of forest fragmentation upon termites appear to have been relatively mild compared with other faunal groups, 4 y after flooding. Nevertheless, we predict that the effects of fragmentation on termite assemblages will ultimately be negative. This study also points to the importance of Amazonian understorey palms in structuring a tropical forest termite assemblage.

We investigated the effects of rainfall on the distribution and abundance of the satyrine butterfly Ragadia makuta in selectively logged and unlogged forest on Borneo. In 1997-98, there was a severe El Niño-Southern Oscillation (ENSO) drought, and annual surveys over a 4-y period showed that abundance of R. makuta was greatly reduced during the drought, but that populations quickly recovered after it. Monthly surveys over a 12-mo period of typical rainfall showed that high rainfall in the month preceding surveys significantly reduced butterfly abundance. Butterfly abundance and distribution did not differ between selectively logged and unlogged areas in either monthly or annual surveys and there was no difference between selectively logged and unlogged areas in the pattern of post-drought recovery. These results indicate that the abundance of R. makuta was significantly reduced both after high rainfall and during severe drought, but that these impacts were short-lived and were not affected by habitat disturbance. ENSO droughts on Borneo naturally often lead to widespread forest fires and thus impacts of ENSO events for butterflies are more likely to be due to indirect effects of habitat loss, rather than direct effects of drought on butterfly population dynamics.

Shadehouse design may have a significant influence on the results of shadehouse studies investigating the light responses of tropical seedlings. Shadehouse effects were investigated by examining the seedling growth and morphological properties of four shade-tolerant tree species from north-east Queensland in the forest and in neutral-density shadehouses for up to 1 y. Understorey plots were compared with 0.8% shadehouses, and single-treefall gaps were compared with 10% shadehouses. Seedlings of all species showed higher growth rates in the shadehouses compared with the equivalent forest treatments. Nonetheless, trends found between high- and low-light growth rates in the shadehouse seedlings were mirrored in forest-grown seedlings. Specific leaf area (SLA) values in the shadehouses provided a good approximation of SLA in the forest for species in both light treatments. In contrast, biomass allocation and leaf area ratios were only comparable for forest and shadehouse seedlings grown in the high-light treatment. Furthermore, seedlings grown in shadehouses differed significantly from forest-grown seedlings in terms of their root morphology. Despite the differences in absolute values of plant characters in the shadehouse and forest seedlings, 65% of significant differences detected between high- and low-light plant characters in shadehouse-grown seedlings were also found in the forest-grown plants.

Nodulated legume trees comprised 43% of the stand basal area in the low, most frequently flooded microsites, and 23% in higher, drier microsites in a tropical freshwater swamp forest in French Guiana. Dinitrogen fixation in Pterocarpus officinalis, Hydrochorea corymbosa and Inga pilosula was confirmed by acetylene reduction assay (ARA), presence of leghaemoglobin in nodules and the 15N natural abundance method. The results for Zygia cataractae were inconclusive but suggested N2 fixation in drier microsites. Nodulated Inga disticha had a 15N-to-14N ratio similar to non-N2-fixing trees, but ARA indicated nitrogenase activity and leghaemoglobin was present in nodules. All bacterial strains were identified as Bradyrhizobium spp. according to the partial 16S rDNA sequences, and they were infective in vitro in the model species Macroptilium atropurpureum. About 35-50% of N in the leaves of P. officinalis, H. corymbosa and I. pilosula was fixed from the atmosphere. Dinitrogen fixation was estimated to contribute at least 8-13% and 17-28% to whole-canopy N in high and low microsites, respectively. Symbiotic N2 fixation appears to provide both a competitive advantage to legume trees under N-limited, flooded conditions and an important N input to neotropical freshwater swamp forests.

Leaf quality (nutrient concentration) and herbivory were studied in a set of dominant tree species in regenerating, secondary tropical dry forests growing on limestone in the Yucatán Peninsula. The study's objective was to assess how variation in nutrient availability affects leaf nitrogen and phosphorus concentrations and herbivory. Our study included an area of young forest (10 y old), with phosphorus-poor soils, and an area of old forest (∼60 y old), in which soil P was comparatively less limiting. The foliage of representative species of each site was studied, including Acacia gaumeri, Leucaena leucocephala and Lysiloma latisiliquum at the young forest site, and A. gaumeri, Bursera simaruba and Pithecellobium dulce at the old forest. Four independent plots (12 × 12 m) at each forest were either left intact (controls) or fertilized with N, with P, or with N plus P for three consecutive years. Relaxation of nutrient limitations at the young forest resulted in an increase in leaf phosphorus and herbivory in all species. In contrast, N and P fertilization at the old forest site did not consistently affect leaf nutrient concentration and leaf damage. In this site the main response to fertilization was an increase in leaf nitrogen only in B. simaruba and an increase in herbivory in A. gaumeri; leaf phosphorus was unaffected by soil fertilization. Overall, we observed that, subsequent to nutrient addition, leaf phosphorus concentration and herbivory in the leguminous species increased at the young site; leaf nitrogen concentration in the non-leguminous species increased at the old site while herbivory only increased in A. gaumeri, following a pattern similar to that observed in the young site. We conclude that the regulatory mechanisms of leaf quality and damage by herbivores will vary, depending on the details of the site's nutrient limitations and the identity of species.

The effects of defoliation on leaf and propagule production, and leaf chemistry of the mangrove Kandelia candel (L.) Druce were evaluated in a manipulative experiment in Hong Kong. Artificial defoliation of leaf lamina at 50% of the length of midrib resulted in significant reduction in leaf, twig and propagule production, and size of the latter. Through the negative effects on propagule number and size, severe herbivory may influence fitness of the mangrove, and, thus, community structure. In contrast, no apparent adverse effects on growth and production were observable at 25% defoliation. Defoliation also significantly affected leaf chemistry of the trees, particularly those suffering 50% defoliation. Concentrations of soluble tannins and carbohydrates in leaves were significantly lower at 50% defoliation compared with the control. Total nitrogen also decreased significantly with increased per cent defoliation. Plants surviving in stressful habitats, such as mangroves, are probably more affected by loss of leaf biomass than those surviving in favourable environments.

Fleshy diaspores (fruits, seeds) comprise a large portion of the litter on the floor of tropical forests, and interactions involving litter-foraging ants and diaspores are common in these areas. In this study, the interactions between ants and non-myrmecochorous diaspores (i.e. not adapted to dispersal by ants) were surveyed along a 1.4-km transect in a restinga forest (sandy soil) on the coast of south-eastern Brazil. During 2 y of monthly samplings, 562 interactions involving 48 ant species and 44 species of diaspore (0.02–11.10 g) were recorded. Ant–diaspore associations involved a considerable part of the ground-dwelling ant community. Large ponerine ants individually removed the diaspores up to 13 m, whereas small ants (myrmicines) normally recruited workers and consumed the diaspore on the spot. Ant-derived benefits to diaspores of non-myrmecochorous plants included secondary dispersal (small to medium-sized diaspores), and increased germination success after seed cleaning by ants. Large ponerine ants such as Odontomachus chelifer and Pachycondyla striata were the main seed vectors. Seedlings of three species were associated with nests of O. chelifer. The results indicate that ants play an important role in fruit/seed biology in the restinga forest.

Small-scale spatial association of the distribution for 55 abundant tree species with two environmental factors (humus depth and surface microtopography) was examined in two 1-ha plots of a heath (kerangas) forest in Central Kalimantan, Indonesia. More than 80% of the 55 species showed a significant habitat preference in humus depth and/or relative elevation in at least one plot. In particular, ten species occurring in both plots showed a consistent significant preference for humus depth or relative elevation in the two plots. Using randomization tests, however, only five species significantly associated with humus depth and no species with relative elevation. These results suggest that edaphic and topographic factors, especially humus depth, contribute to determining local spatial distribution and floristic composition of abundant tree species in the forest.

Effects of litter cover on insect herbivory, leaf-fungal damage, and seedling performance (survival, growth in height and leaf number), were assessed on established Nectandra ambigens seedlings in three sites of a Mexican tropical rain forest. Naturally occurring seedlings were measured for height, leaf number and standing levels of leaf damage by herbivores and pathogens. Thereafter, seedlings were exposed for 1 y to three litter treatments: (1) litter addition, (2) control and (3) continuous litter removal. There was a significant effect of site on the proportion of surviving seedlings and on the relative growth rates in terms of leaf number. Overall, seedlings with higher initial leaf number showed higher survival and produced more leaves after 12 mo than seedlings with lower initial leaf number. Litter addition resulted in seedlings with lower relative growth rates in height and higher levels of damage by herbivores and pathogens. On average, leaf area damaged by pathogens was < 6% and herbivory was < 27%. Seedlings with higher levels of herbivory showed higher levels of pathogen attack. Herbivory and pathogen attack had no effect on seedling performance. This study reveals the importance of litter-cover on the interactions between seedlings, herbivores and pathogens, as well as on the performance of tropical seedlings.

We investigated the spatial distribution of fruiting in Dryobalanops aromatica (Dipterocarpaceae), a mast-seeding emergent tree that is found in the tropical rain forests of Borneo. Of 393 adults (diameter at breast height (dbh) > 30 cm) in a 52-ha area, 143 (36.4%) individuals fruited. A second-order spatial analysis, based on Ripley's K-function, revealed that fruiting trees were significantly (P < 0.05) aggregated with respect to the total population over distances of 15–115 m. Thus, the spatial distribution of fruiting trees was more aggregated than expected from the adult tree distribution, which itself was aggregated within the study plot. Logistic regression analysis showed that fruiting trees had a significantly greater dbh and were surrounded by more conspecific adults per ha than non-fruiting individuals. Moreover, fruiting trees were found at higher elevations, on steeper slopes, and on sandy rather than clayey soils. In contrast, annual diameter growth, adult density over areas < 1 ha, and slope convexity did not significantly affect fruiting probability. The ratio of seedling (20 cm ≤ h < 60 cm) and sapling (60 cm ≤ h < 300 cm) density to adult density was higher in sites at higher elevations, on steeper slopes, and on sandy soils than that for lower elevations, flat slopes and clayey soils. This suggests that the aggregation of fruiting trees occurred not only at the time of the study, but that it had occurred repeatedly in the past, at the high-elevation, steep-sloped, sandy sites. Thus, site conditions probably affect the dynamics and spatial structure of local populations through differences in fruiting frequency among trees experiencing different site conditions.

Rain forests at Los Tuxtlas are islands surrounded by anthropogenic vegetation containing invasive species. Epizoochory is an important dispersal syndrome among invasive species but ecological studies are scant. In order to evaluate the movement of epizoochore diaspores into the mature forest, we selected two abundant secondary herbs with contrasting diaspore morphology, and five dispersers with different pelage type. We used stuffed mammals as a dummy and passed them through patches of these two plant species, counting the number of adhered diaspores. For dispersal into the forest, diaspores were manually adhered to dispersers, these were pulled along 100-m forest transects, counting the number of diaspores remaining at distance intervals. We observed the highest amount of detachment in the first 20 m of the forest edge. For Desmodium incanum diaspores with sticky hairs, adhesion to dispersers showed no differences among pelage types, therefore dispersal is determined mainly by diaspore detachment. Dispersers with long pelage (Didelphis marsupialis, Nasua narica and Procyon lotor) move diaspores for shorter distances, while dispersers with short pelage (Silvilagus floridanus and Philander opossum) are more likely to move diaspores for longer distances. In Pavonia schiedeana more diaspores (seeds with spines and hooks) were attached and remained attached on dispersers with long and thick pelage. Mean distance travelled for P. schiedeana diaspores varied between 11 and 2700 m. In D. incanum the distance travelled did not exceed 500 m. Based on comparisons between dispersal patterns and on ecological data of these two plant species and their potential dispersers, P. schiedeana has a higher probability of invading the forest and finding suitable habitats for establishment than D. incanum.

The effect of tillage, clipping, precipitation and temperature on above-ground grass production was investigated in permanent plots at a Zambian savanna site for 5 y (1996^2001) by the harvest method. Mean species richness was 4.6 species m-2 with a total of 15 species at the study site. Mean end-of-season grass phytomass was 464 g m-2 with no statistically significant differences among years in control quadrats. Grass phytomass recovered within two wet seasons after tillage and a similar trend was observed after cessation of a 2-y monthly harvesting regime. However, in experimental quadrats, plot, treatment and year had significant effects on grass production. Previous- and current-season precipitation had no significant effect on end-of-wet-season grass phytomass but phytomass of the previous season explained 27-53% of the variation in end-of-wet-season phytomass. Grass production peaked in the wet season and declined sharply as the dry season progressed. The interaction between precipitation, temperature, harvesting and duration of the dry season explained 81-91% of the variation in daily grass production but the significance of the interactions varied with season and duration of harvesting. Precipitation-use efficiency (PUE) of grasses declined from 2.25 g m-2 mm-1 at the start of the wet season in December to 0.25 g m-2 mm-1 at the end of the season in March. Monthly harvesting for 1 y reduced PUE to less than 25%. The results indicate very complex relationships between above-ground grass production (dependent variable) and climate and land-use (independent) factors that makes the prediction of grass production in central southern African savannas difficult.

The aims of this study were to investigate the roosting ecology and social organization of the spotted-winged fruit bat, Balionycteris maculata (Megachiroptera), within an old-growth Malaysian dipterocarp forest, and test the hypothesis that males spend a significantly greater proportion of the night in the immediate vicinity of their day roosts than females. Balionycteris maculata roosted in cavities and formed small harem groups that consisted of one adult male accompanied by up to nine adult females and their dependent young. Both male and female bats foraged locally, within approximately 1 km of their roost sites. However, in contrast to females, which remained away from the roost site throughout the night, both harem and solitary adult males divided their nightly activity between foraging and returning to their roost sites. Individual females were also found roosting sequentially with up to three different males. Hence, these results suggest that B. maculata has a resource-defence polygynous mating system, in which the roost cavity may represent a critical and defendable resource by which males recruit and gain improved reproductive access to females. Roost cavities occupied by B. maculata were found within a number of different forest structures, including ant nests, termite nests and the root masses of epiphytes. The consistent shape and position of the cavities, along with a single observation of cavity enlargement, provide the first evidence of roost-cavity excavation, by a megachiropteran bat, within the nests of social insects. Although the cues by which females select their roost sites are unclear, some aspect of roost cavity size, or condition, may provide a possible means by which females can assess the reproducÍtive fitness of potential mates.

Ecologists agree on the importance of hurricanes in community dynamics in tropical forests but little is known of the responses of herbaceous species to large disturbances. The passage of Hurricane Georges over the island of Puerto Rico provided a unique opportunity to study the post-hurricane recovery of Heliconia caribaea, a large understorey herb. Five weeks after Georges, the types of vegetative damage, mortality and recovery were recorded and their relationship with plant size was established. The relationship between recovery strategy and leaf or stem damage severity was also determined. The abundance of H. caribaea seedlings in sites with severe or little forest canopy damage was recorded to determine the effects of canopy cover on seedling establishment. Most individuals showed severe leaf and stem damage. This was not related to plant size. Mortality was low and caused by fallen trees. Recovery was not associated with severity of leaf damage but more individuals with no signs of regeneration appeared among plants that had severe stem damage. Seedling abundance was higher in areas with severe canopy damage relative to those with closed canopy suggesting that hurricanes may benefit H. caribaea by creating suitable habitats for seedling establishment. We argue that recovery strategies like those exhibited by H. caribaea may be common among understorey plants, particularly in areas regularly subjected to large-scale disturbances.

Our study examines the effect of seasonal rains on soil organic C dynamics in a tropical deciduous forest ecosystem in Western Mexico. At the end of the wet season, an accumulation of labile nutrient forms developed and was maintained during the dry season. This accumulation enhances microbial activity in the first rains of the wet season. For example, the litter samples of the dry season had a higher C and N mineralization than those of the wet season. Similarly, the January soil samples had higher C mineralization than October soil samples (55 and 34 μg C g-1 d-1, respectively). These results suggest that the quality of C is strongly affected by the seasonality of rains, which in turn influences microbial activity. This seasonality also influences nutrient redistribution between soil aggregate fractions. Chemical changes across seasons suggest that soil organic matter associated with macro-aggregates represents the main source of energy for microbial activity at the beginning of the wet season, while micro-aggregates protect the labile nutrient forms during the growing season.

One hundred and nine soil samples were collected in 69 different localities along two transects, one North-South, about 900 km in length, and the other East-West (750 km in length), across Senegal and Gambia. The first transect followed a rainfall gradient and the second a human density gradient. The relationship between carbon content, C/N ratio and soil type on the abundance and species distribution of the nematodes along the transects was studied. Results showed that short-term fallows did not influence the specific structure of the communities, when compared with the nematode communities of fields located in the immediate vicinity, where Scutellonema cavenessi and Tylenchorhynchus gladiolatus were the dominant species. The expected negative influence of human disturbance on nematode occurrence seemed to be compensated by greater crop diversity, mainly near the towns. Less-disturbed areas also maintained a high diversity, but were characterized by the presence of particular species such as Xiphinema spp. Soil type was the most important factor affecting the species composition of the nematode community. As a result, nematode communities followed a distribution in areas, corresponding to the successive soil types, but did not change in relation to the human or climatic gradients. At a large scale, the study of plant-parasitic nematodes can give both different and complementary information on the ecological trends of an area to that of free-living nematodes.

Relative densities of mahogany species vary across tropical Africa and correspond with changes in soil fertility and moisture status. Seedling growth of four co-occurring African mahoganies (Entandrophragma spp.) was studied in relation to soil nutrient and moisture status in a shade-house experiment. On naturally occurring forest soils, E. cylindricum and E. utile exhibited increased relative growth rate (RGR) and decreased root mass ratio (RMR) with an increase in soil fertility while E. angolense and E. candollei did not. Changes in leaf morphology with fertility did not correspond to changes in species performance. On moist, fertile soils, E. angolense outperformed congeners but E. candollei performed equally well on moist infertile soils. Entandrophragma cylindricum performed as well as E. angolense and E. candollei on two of three soil moisture stress treatments but E. utile consistently performed poorly. Comparative seedling performance fitted well with limited available data on the distribution of Entandrophragma spp. in relation to soil fertility and moisture gradients and suggests that within-forest microsite heterogeneity may help explain the distribution of Entandrophragma species within the Dzanga-Sangha Dense Forest Reserve, Central African Republic.