We investigated the association between lianas and Bertholletia excelsa (Brazil nut), a long-lived, emergent tree of significant ecological and economic importance in Amazonia. Our objectives were: (1) to determine the relationship between crown liana load and liana number, basal area, and origin in relation to the B. excelsa host; and (2) to determine the relationship between liana load and B. excelsa fruit and nut production, diameter growth, and crown form, position and area. One hundred and forty trees (≥50 cm dbh) were selected with representatives of 10 diameter classes and four liana load categories. To quantify fruit and nut production, fruit counts and nut fresh weights per tree were measured in 2002 and 2003, and annual diameter growth was quantified using dendrometer bands. Trees with lianas produced significantly fewer fruits and had reduced nut fresh weights than liana-free trees. Trees with the most extensive liana loads (>75% crown coverage) were 10.2 times more likely to have crown forms categorized as less than half-crowns or few branches than trees with reduced liana loads. No statistically significant relationship was found between liana load and tree diameter growth. Results suggest that liana cutting might increase B. excelsa fecundity and commercial nut yields.

The diets of the acouchy Myoprocta exilis, agouti Dasyprocta leporina and paca Agouti paca, the three largest terrestrial rodents of French Guiana forests, were investigated from stomachs collected over several years. These species are primarily frugivores. The percentage of fruit in their diet decreased, and seasonal variation increased from the smallest species, the acouchy, to the largest, the paca (contents in dry mass: respectively, 98.5% ± 0.6%, 87.4% ± 3.5%, and 83.9% ± 7.6%). They differed from each other in the proportions of pulp and secondary foods (leaves, fibre, insects) consumed annually as well as seasonally. Dietary diversity in the acouchy was half that of the two other species. The seasonal variation in consumption of different fruit parts and of secondary foods by these species corresponded well with the fruit production in the forest, and was linked to their reproductive events. Overall, the diets of these three large rodents appeared intermediate between the small rodent species and the artiodactyls (peccaries and brocket deer) inhabiting the same areas, which also concords with their body mass and their place within the guild of terrestrial mammals.

We used tree-rings to reconstruct long-term patterns of suppression, release and growth among five sympatric canopy tree species representing the full range of shade tolerance in a seasonal tropical forest in western Thailand. We expected that the frequency and duration of suppression and release events would be positively correlated with shade tolerance. All five species showed evidence of major and moderate growth releases. As expected, Melia azederach, an extreme heliophile, had the fewest releases. However, among the other species the number of major releases was consistent across the range of shade tolerance. The most significant difference among the species was the number of moderate releases recorded. There was a general positive correlation between the number of moderate releases and shade tolerance; however, Chukrasia tabularis, a relatively shade-intolerant species, had an anomalously high number of moderate releases. The study species also showed considerable variation in canopy accession strategies. The least common canopy accession strategy was establishment in the understorey and growth into the canopy in the absence of any gaps. However, with the exception of Melia, all four study species had one or more individuals that successfully reached the canopy using each of the four canopy accession strategies. These results highlight the importance of periodic or episodic bouts of gap formation on canopy tree recruitment and the utility of tree-rings for reconstructing long-term growth patterns in tropical trees.

Investigations of bats in naturally fragmented ecosystems may help refine assumptions about bat responses to fragmentation of their habitats by human activity. Bat species assemblages were studied for a 3.5-y period in a naturally fragmented landscape in the north-west Yucatan Peninsula, Mexico. Bats were systematically sampled using mist nets in a total of 16 forest islands of four categories (four sites each): large (>20 ha), small (<5 ha), far and near (sites located >10 km or <1 km away from the nearest forest island, respectively). We captured a total of 1134 bats representing 17 species. Bat diversity and species richness were similar among categories of forest island. Fruit-eating bats (78–93% of captures) were significantly more abundant in large and in small than in forest islands in the far or near categories. Differences in density of Manilkara zapota trees in the forest islands might underlie the observed variations in the abundance of frugivorous bats. Distances traversed by bats (0.65–38 km) between forest islands (typically 100–300 m away), facilitates the mobility of bats across the landscape. However, the moderately rich bat species assemblage detected, suggests that other species may not be able to persist in such naturally fragmented ecosystems.

Ontogenetic and seasonal variation in diet was examined for 11 species of insectivorous forest-floor frogs and lizards from a lowland wet forest in north-eastern Costa Rica. Specimens were collected systematically over an entire seasonal cycle and represented individuals of all sizes. Individual prey items were removed from stomachs of preserved specimens, measured and identified. Ontogenetic shifts in prey size were pervasive. Ontogenetic shifts in prey composition were limited to four species; these were not the species with greatest range in body size, nor the species with the broadest diets. Small prey types (ants, mites, collembolans) decreased in representation and large prey types (roaches, orthopterans, millipedes) increased in importance over ontogeny; this could be because prey selection is based primarily on prey size or because of different prey preferences among age classes. There is little evidence for size-structure in this assemblage. There is no evidence that total availability of arthropod prey varies among seasons, but some evidence that preferred prey are less common in the wet season. Diet was similar between lizards and frogs. Lizards were more likely to have empty stomachs, but also greater stomach volume, than frogs; this indicates a difference in food-gathering strategies. Our study indicates strong similarity between frogs and lizards in diet despite enormous differences in physiology and behaviour.

Termite foraging was monitored with cellulose baits in both the wet and dry season in the semi-arid Okavango Delta region of northern Botswana. Seven wood-feeding species were found. Fungus-growing termites dominated the assemblage in all plots, and four species (Macrotermes michaelseni, Odontotermes sp. A, Allodontermes sp. and Microtermes sp.) were found in most. Foraging by smaller-bodied Allodontermes sp. and Microtermes sp. was largely confined to the wet season. Foraging by M. michaelseni and Odontotermes sp. A was more constant, but showed a general peak in the early dry season (May and June), likely due to nutrient and energy demands associated with alate production. Despite extensive overlap in distributions of these species, co-occurrences on baits were significantly less frequent than expected by chance. In addition, incidence in a plot of Allodontermes sp. and Microtermes sp. was negatively correlated in the wet season, as was that of M. michaelseni and Odontotermes sp. A in the dry season. Differences in body size, life history, and nest structure produce temporal and spatial heterogeneity in the foraging and distribution patterns of termites, and therefore suggest that consideration of community composition can improve prediction of the influence of termite assemblages on decomposition.

This paper addresses the effects of habitat fragmentation on the phyllostomid bats of the Atlantic rain forest in Brazil, by comparing community structure (species richness and capture frequency) and the frequency of three bat species sampled along 36 transects encompassing six habitat categories: interiors and edges of large (>1000 ha) and small fragments (<100 ha), and the surrounding matrix of second-growth forests and areas of shade cocoa plantation. Species composition, richness and total captures were not directly affected by forest size per se, although the frequency of one dominant forest species (Artibeus obscurus) was significantly lower in small fragments compared with larger ones. The high connectivity among forest patches in the study area and the ability of some species to use the surrounding matrix of secondary forests and shade cocoa plantations possibly precludes the insularization effect. Qualitative habitat changes induced by fragmentation, such as edge formation and forest regrowth affected bat community structure; both modified habitats comprised a limited subset of the species assemblage found in the interiors of mature forests. The results presented here provide evidence of impoverished bat assemblages in man-modified habitats linked with deforestation and overall disturbances related with forest fragmentation.

Little is known about the change of species-rich tropical insect communities along habitat gradients. Diversity and species richness of geometrid moths were investigated at 15 anthropogenically disturbed sites in a montane area in the Andes of southern Ecuador representing a successional gradient. These plots were compared with six closed-forest understorey sites. We collected a total of 23,720 individuals representing 868 morphospecies. Local diversity increased with forest recovery and decreased with increasing distance to the natural forest. 18.6% of all species were found as unique singletons. The mean proportions of local singletons differed significantly between three succession classes. Forest understorey showed a higher proportion of singletons than early and late successional stages. Ordination of the moth samples showed a clear separation of geometrid ensembles at successional sites vs. the forest understorey sites. Patterns of species turnover were influenced by the degree of habitat openness, and to a lesser extent by elevation. However, faunal differences were not related to geographical distances between the sampling sites. In conclusion, geometrid moth ensembles of regenerating Andean montane forest remain diverse, but change significantly in composition relative to adjacent natural forest, whereas the diversity and composition of the geometrid fauna are far more strongly affected in non-forested habitats and abandoned pastures.

Tropical plants have been suggested to have higher levels of mechanical, chemical and biotic defences than temperate plants. However, comparisons have usually included deciduous species within the temperate group, which confounds the analysis since deciduous species are predicted to have a different strategy with respect to investment, nutrition and defence than evergreen species. In this study we examined levels of defence and nutrition in five evergreen species of Eucryphia occurring along a latitudinal gradient in eastern Australia, grown under common conditions in a glasshouse. From the resource-availability hypothesis we predicted the opposite gradient in defence investment, i.e. that lowest levels of defence will occur in tropical species with potentially high growth rates and annual productivity. However, we found an increase in cell wall content, total phenolics and tannin activity, and a decrease in protein availability, with decreasing latitude and/or increasing mean annual temperature. Hence, there was a trend of increasing defence (although not in leaf toughness) and declining nutritional quality towards the tropics. These latitudinal trends were recorded in both mature and expanding leaves. The same trends were observed in leaves of two species collected from the field, indicating that the results were not peculiar to the experimental growth regime. The latitudinal trend in defence did not support our prediction based on the resource availability hypothesis and may indicate that herbivore pressure is providing an overriding selection pressure, although there are alternative explanations.

Seedling growth under three shade levels was studied at Varanasi, India, for five tree species of tropical dry forest, which differed in life-history traits. Three of these were small-seeded pioneer (Albizia procera, Acacia nilotica and Phyllanthus emblica) and the other two were large-seeded non-pioneer (Terminalia arjuna and Terminalia chebula) species. Seedlings of all the species were subjected to three light levels (80–100%, 20–30% and 3–7% of full sunlight) comparable to sunlit gaps and shaded microsites in the dry forest. After 4 mo of shade treatment, height, basal area, biomass and other growth traits, viz. RGR (relative growth rate), NAR (net assimilation rate), and SLA (specific leaf area) were determined. Etiolation and plasticity indices were calculated. Reduction in seedling height, biomass and relative growth rates and enhancement in SLA due to shade was greater for small-seeded pioneer species. Seedlings from large-seeded non-pioneer species exhibited a stronger etiolation response to shade than seedlings from small-seeded species. Phenotypic plasticity indices for basal area, plant biomass and relative growth rate were greater for the three small-seeded early successional species (A. procera, A. nilotica and P. emblica), indicating their specialization in a more favourable light environment such as large gaps and forest peripheries. The non-pioneer and pioneer species differed only in the degree of shade tolerance, and we suggest that dry forest species cannot be strictly categorized into two distinct groups (shade tolerant vs. intolerant), rather shade preference and gap preference would be the more expressive terms. Marked environmental heterogeneity in terms of irradiance and the phenology of dry tropical trees permits coexistence of species of varying ecological traits, contributing to the maintenance of diversity in the dry forest.

Discrete humus layers are common on podzols under temperate coniferous and tropical heath forests, and patchy layers also occur under some temperate broadleaved forests on non-podzolic soils. We used multiple data sets to test the reported association of humus with oligotrophic but non-podzolic soils under non-heath dipterocarp forest at Lambir, Sarawak. We examined the distribution, morphology and nutrient dynamics of necromass on soils derived from sandstone and shale. Concentrations of the main mineral nutrients were lower in fresh litter on the very oligotrophic sandstone soils than on shale. The rates of litterfall were similar, so that annual litterfall fluxes of all nutrients were lower on sandstone. The lower nutrient concentrations and fluxes in the litter on sandstone resulted in slower decomposition, longer residence times and larger standing crops of forest-floor necromass, with lower concentrations of nutrients. The necromass on sandstone sequestered significantly more N, K and Mg but less Ca and Mn than on shale, with no significant difference for P. The variations in necromass nutrient dynamics were associated with morphological differences. There were mats of densely rooted humus under the litter on sandstone, whereas litter lay directly over the mineral topsoil on shale. Spatial associations with soil nutrients were weak for necromass thickness, but clear for humus. The proportions of nutrients in the litterfall and necromass reflected the stoichiometric profiles of the soils. We attribute the differences in necromass nutrient dynamics and their association with soil reserve nutrients to lower rates of nutrient replenishment from the weathering of sandstone than from shale. Necromass characteristics are robust field indicators of multivariate edaphic differences in these and other tropical forests on Acrisols/Ultisols derived from Tertiary clastic sediments.

Competitive interactions among pioneer species may have a significant impact on the course of secondary succession in tropical rain forests. We predicted that the outcome of competitive interactions in early succession would vary with soil nutrient availability. To test this hypothesis we grew seven pioneer tree species alone and in dense competitive mixtures, with four nutrient treatments: no nutrient addition, and N, P, and N plus P addition. Performance of plants grown alone and in mixtures was strongly nutrient limited. However, contrary to expectation, the competitive hierarchy among the seven species was almost identical among the four nutrient treatments. The dominant species, Melastoma malabathricum, accounted for > 70% of total stand biomass in all nutrient treatments. Seedlings of this species had higher rates of gas exchange and initial growth, and lower root allocation than the other species. Profiles of light availability within the competitive stands indicated that light levels at ground level were well above levels at which pioneer species can successfully survive and grow, yet seedlings of species other than Melastoma remained stunted. Leaf N concentrations in all stands were 25–55% reduced by competitive interactions, and N addition increased relative competitive performance in only Melastoma suggesting that Melastoma was particularly effective at N acquisition, limiting nutrient uptake by the other species. Toward the end of the experiment individuals of Melastoma began to reproduce, suggesting that the competitive hierarchy would have changed in a longer-duration experiment.

Fire history was reconstructed from fire-scarred individuals of the endemic pine (Pinus occidentalis) along climatic gradients in the Cordillera Central, Dominican Republic. We analysed variation in fire frequency by climate, elevation and aspect (windward and leeward of the central massif). A high correspondence between known fires and fire-scar chronologies indicates that the primary rings of this species are annual. Evidence was found for 41 fire years since 1727 A.D.; 28 were landscape-scale fires, nine of which were exceptionally large and linked to El Niño-triggered droughts. Mean fire return interval (FRI; the mean of individual samples) and mean fire interval (MFI; the composite mean of a group of samples) were used to estimate the upper and lower range in fire frequency. Mean FRI of the entire study area was 31.5 y (±24.9 y SD) and MFI for landscape-scale fires was 5.6 y (±4.1 y SD). The fire regime varied significantly with elevation and aspect. Mean FRI was 42.1 y (±27.6 y SD) and MFI was 9.8 y (±16.7 y SD) on moister windward zones, and mean FRI was 16.7 y (±7.8 y SD) and MFI 4.2 y (±1.9 y SD) in drier leeward zones. On windward slopes, high-elevation mean FRI (26.4 y) was significantly shorter than at middle (44.5 y) and low elevations (58.4 y). On leeward slopes, mean FRI did not vary significantly with elevation. The strong windward elevational patterning of the fire regime is driven by the trade wind inversion which traps moist air below 2000–2300 m. Such elevational patterns may be characteristic of montane fire regimes throughout the tropical trade wind belt.

Local species richness and between-site similarity in species composition of parasitoid wasps (Hymenoptera: Ichneumonidae; Pimplinae and Rhyssinae) were correlated with those of four plant groups (pteridophytes, Melastomataceae, Burseraceae and Arecaceae) in a western Amazonian lowland rain forest mosaic. The mosaic structure of the forest was related to variation in soils within the non-inundated terrain. Significant matrix correlation between patterns in parasitoid wasp species composition and plant species composition was found. Most of the overall correlation was due to idiobiont parasitoids of weakly concealed hosts, which attack host larvae and pupae in exposed situations, with two of the four ecologically defined parasitoid groups showing no correlation at all. A positive correlation between the number of plant species and the number of Pimplinae and Rhyssinae species at a site was found when the latter was corrected for collecting effort. Consequently, the degree of floristic difference between sites may be indicative of the difference in species composition of ichneumonids, and the species richness of plants may serve as a predictor of the species richness of parasitoid wasps. Although these results were obtained in a mosaic including structurally and floristically clearly different types of rain forest, the correlation coefficients were relatively low, and the present results lend only weak support to the idea of using plant distributions as indicators of animal distributions.

Stand structure dynamics during early secondary forest succession were related to mortality, growth and recruitment rates, and the dependence of these demographic processes on fallow age and initial stand structure attributes was evaluated. In 11 secondary tropical rain-forest sites (1.5–19 y) in Chiapas, Mexico, one plot of 10 × 50 m was established. Diameter and height were measured for all trees ≥ 1 cm dbh, and their survival, growth and recruitment was monitored over a 2-y period. Changes in stand structure were especially fast in the first 5 y of succession, and decreased rapidly afterwards, which resulted from similar stand-level changes in relative mortality, growth and recruitment rates. Demographic processes were negatively related with initial stand basal area, but independent of initial tree density. Basal area was a better explanatory variable of the among-stand variability in these rates than fallow age. Results suggest that asymmetric competition and resulting patterns of tree-thinning are major driving forces determining secondary forest successional pathways. Fallow age per se is a compound variable reflecting community organization at a certain point along the successional axis, while community structure drives succession. Sudden mass mortality among dominant species in some stands showed that early secondary forest succession is not always a gradual and unidirectional process.

Intraspecific synchrony in the phenology of tropical forest trees has implications for satiation of browsers and seed predators, pollination, dispersal and interpreting plant responses to environmental cues and stresses. Community phenological studies have been constrained in their consideration of these issues by the lack of concise quantitative measures of intraspecific synchrony. We propose a novel application of Colwell's predictability indices to overcome this deficiency. Its application is demonstrated by evaluating the leaf and reproductive phenologies of 8–16 individuals of each of 20 monsoon forest tree species from northern Australia monthly for 30 mo. The indices yielded substantial additional information over that provided by summaries averaged amongst conspecifics. Marked intraspecific asynchronies in leaf flush or flowering occurred in five evergreen species and two deciduous fig species, and were associated with prolonged individual commitments to these states. Five partly deciduous species exhibited marked intraspecific variation in the seasonal extent of leaf loss. We interpret the observed variation in intraspecific synchrony in terms of contrasting water-use and pollination strategies. The synchronicity indices are concise and robust to small and variable sample sizes, and may thus enhance studies of species-rich assemblages.

We examined effects of experimental manipulations of vegetation cover on recruitment, mortality and density of seedlings (20–100 cm tall) and saplings (≥100 cm tall) of woody growth forms over a 2.5-y period. We created four treatments in each of three 15–20-y-old tropical forest stands in Costa Rica: a large canopy gap (270–350 m2), a small canopy gap (50–100 m2), understorey vegetation removal, and an unmanipulated control treatment. Creation of canopy gaps, especially large ones, increased first-year recruitment and density, as well as overall mortality of seedlings. Saplings experienced lower mortality and more prolonged gap-enhanced recruitment and density than seedlings. Removal of understorey vegetation had little or no effect on tree seedling and sapling dynamics. Recruitment and density of lianas responded only to large gaps, whereas understorey species responded to both gap treatments and to spatial heterogeneity within gaps. Tree species exhibited diverse regeneration requirements, whereas liana and understorey species were more specialized to the high and low ends of the light availability gradient, respectively. Canopy gaps provide a critical mechanism for regeneration of lianas, and canopy tree species that dominate during the early stages of secondary forest succession. The choice of management system for these secondary forests can determine the direction and rate of succession.

The small size of many African protected areas makes adjacent rangelands potentially important in the local survival of wild animals. In order to assess the importance of pastoral areas to wild ungulates, we studied density and habitat choice of wild ungulates and cattle in Lake Mburo National Park, Uganda, the adjacent exclusively pastoral Nshara Dairy Ranch and on private land consisting of a mixture of ranching and subsistence farms. Transects, in the three land-use zones, were walked during the wet season and the data were analysed by DISTANCE sampling technique. We found significantly higher total density of wild ungulates on the dairy ranch compared with the National Park and private land. There was no significant difference in total wild animal density between the National Park and private land. Impala (Aepyceros melampus), zebra (Equus quagga), bushbuck (Tragelaphus scriptus) and waterbuck (Kobus ellipsiprymnus) had significantly higher densities on the dairy ranch compared to the National Park. Only eland (Taurotragus oryx) density was higher in the National Park compared to private land. Wild ungulates and cattle showed a high degree of habitat overlap, generally preferring open grassland. Our study shows that high densities of wild ungulates are not necessarily associated with protected areas. Pastoral areas may be important for populations of wild herbivores during the growing season despite a pronounced presence of livestock.

The temporal impact of clearing trees on litter production, litter decomposition and on C, N and P release through decomposition of litter was examined in the pasture systems of a semi-arid zone of central Queensland. Paired sites for cleared pastures (developed from clearing woodlands) and uncleared (intact woodland) pastures were selected to represent three dominant tree communities of the region i.e. Eucalyptus populnea, E. melanophloia and Acacia harpophylla, with three different time-since-clearing (5, 11–13 and 33 y) treatments. Yearly litter production was greater at uncleared sites (1732–1948 kg ha−1 y−1 for eucalypt and 2596 kg ha−1 y−1 for acacia communities) compared with cleared sites (1038–1282 kg ha−1 y−1 for eucalypt and 1100 kg ha−1 y−1 for acacia communities averaged over three time-since-clearing treatments). Rates of litter decomposition and of release of C, N and P from decomposing litter were higher at cleared than uncleared sites for all three tree communities. The cleared and uncleared sites did not differ significantly in total amount of C and N released per year since the concentrations of C and N were greater in litter from uncleared sites but the rate of release was less than that at cleared sites. Slow but continuous release of nutrients in eucalypt and acacia woodlands may be an adaptation of these communities to maintain the nutrient cycle and to avoid leaching of nutrients in the nutrient-poor soils of the region.

The presence and density of two sympatric, large-sized vipers (the Gaboon viper, Bitis gabonica and the nose-horned viper, Bitis nasicornis) were studied along several transects, during both dry and wet seasons, and at different times of day, in southern Nigeria (West Africa). Three habitat types were found along the various transects (mature rain forest (MF), secondary rain forest (SF), swamp forest (SW)). The detection probabilities for these vipers were modelled with a set of competing models, and the various models were ordered by Akaike Information Criterion procedures. Two classes of models were used: the single-season model, and the multi-species model. The best models (single-season model) suggested that: for the Gaboon viper, habitat types SF and SW were particularly important in detecting this species, especially during the rainy season at 08h00–16h00. For nose-horned vipers, the best models had SW and MF as site-covariates. Application of the multi-species model revealed that there were different detection functions if both species are present at a site, with a ‘negative’ interaction of occupancy between the species. Females and males were similarly detectable in a logistic regression model, but feeding status and pregnancy slightly increased detection probability in a logistic regression model. Viper density was modelled by a DISTANCE sampling procedure. The density of one species tended to be inversely correlated to the density of the other, suggesting that (1) the rain-forest environment does not support abundant populations of both vipers when sympatric, and (2) the two Bitis species subtly partition the habitat resources.