Tropical rain forests harbour the most diverse plant and animal assemblages known to science, but our understanding of assemblage structure and species interactions is limited. Bats, as the only flying mammals, have the potential to exploit resources from all strata in forest communities. Thus, fruit-eating phyllostomid bats often have been categorized into canopy-, subcanopy- and understorey-foraging species, based largely upon the height at which they were most frequently captured. Here we challenge this classification and use stable carbon isotopes to assess foraging height of bat species at an Amazonian rain-forest site in Ecuador and at a Caribbean lowland rain-forest site in Costa Rica for comparison with data from mist-net captures. The proportion of the heavy stable carbon isotope 13C in relation to the lighter 12C isotope increases in plants from ground level to the canopy (0.12‰ m−1–0.18‰ m−1), and these differences in stable carbon isotope signatures are reflected in the body tissue of phytophagous bats. We used the stable carbon isotope ratio (δ13C) of wing tissue to estimate the foraging heights of 54 phyllostomid species in two Neotropical bat assemblages. Based on stable isotope data, phyllostomid species exploit food resources at all vertical strata of the forest. Capture height was not a reliable predictor of foraging height and suggests that bats most likely use lower strata to commute between foraging sites to avoid predators. Vertical stratification is likely to be a key factor promoting niche partitioning, thus promoting high local species richness in many tropical animal assemblages.

To assess the impact of habitat fragmentation on tropical avian communities, we sampled lowland forest birds on six land-bridge islands and two mainland forest sites in Lake Kenyir, Peninsular Malaysia using timed point counts, hypothesizing that insectivorous birds are the worst affected guild. We used an information-theoretic approach to evaluate the effects of area, isolation, primary dietary guild (omnivore, frugivore and insectivore) and their interactions in predicting species richness, abundance and diversity. Our analysis showed that a model that considered the effects of area, dietary guild and their interaction best explained observed patterns of species richness. But a model considering both area and dietary guild best explained the variation in abundance. Notably, insectivorous birds were singled out as the dietary guild most sensitive to fragmentation, followed by frugivorous and omnivorous birds and hence provide support for our hypothesis. Assemblages of insectivorous birds were clearly depauperate on anthropogenic forest islands in Lake Kenyir and are consistent with forest fragmentation studies in the Neotropics. Given their specialized foraging ecology and diversity, conservation of intact communities of insectivorous bird guilds in Malaysia will be critical for maintaining predator–prey interactions in lowland tropical forests.

Many tropical mammals reproduce seasonally, although the circum-equatorial climate is more stable and less seasonal than that of temperate zones. The mechanisms underlying seasonal reproduction in the tropics remain enigmatic. Female reproduction and its relation to environmental factors were investigated in the Neotropical bat species Saccopteryx bilineata. Colonies consist of year-round stable groups of several females that are defended each by an adult male. Females give birth to a single offspring each year and it is suggested that mating is restricted to November and December. In this study, it was asked whether females of a Costa Rican colony come into oestrus around the same time each year and whether oestrus times are synchronized. Oestrogen and progesterone metabolites were monitored from faeces between October and January in four years. Oestrus was identified in 32 females. In addition, climatic factors such as rainfall and temperature were monitored at the study site. Results indicate that (1) females exhibit monoestry, (2) oestrus dates cluster around the first half of December, (3) reproduction is strongly seasonal and highly predictable and (4) oestrus times are possibly influenced by long-term cues like photoperiod and short-term cues like sudden changes in rainfall and temperature.

Climate is one of the most important factors determining variation in forest structure, but whether soils have independent effects is less clear. We evaluate how climate and soil independently affect forest structure, using 89 200 stems ≥ 10 cm dbh from 220 1-ha permanent plots distributed along environmental gradients in lowland Bolivia. Fifteen forest structural variables, related to vertical structure (forest height and layering), horizontal structure (basal area, median and the 99th percentile of the stem diameter and size-class distribution) and density of life forms (tree, palm and liana), were evaluated. Environmental variables were summarized in four multivariate axes, related to rainfall, temperature, soil fertility and soil texture. Multiple regression indicates that all structural variables were affected by one or more of the environmental axes, but the explained variation was generally low (median R2 = 0.15). Rainfall and soil texture affected most forest structural variables (respectively 87% and 80%) and had qualitatively similar effects. This suggests that plant water availability, as determined by rainfall and soil water retention capacity, is the strongest driver of forest structure, whereas soil fertility was a weaker driver of forest structure, affecting 53% of the variables. Maximum forest height, palm density, total basal area and liana infestation showed the strongest responses to environmental variation (with R2 ranging from 0.31–0.82). Forest height, palm density and total basal area increased with plant water availability, while liana infestation decreased with plant water availability. Therefore, multiple rather than single environmental factors must be used to explain the structure of tropical forests.

The application of stable-hydrogen isotope (δD) measurements to the study of animal movement, resource use and physiology depends on understanding factors driving variation in δD in animal tissues. The source of micro-scale variation in δD is poorly known, yet understanding micro-scale patterns of δD could shed light on important ecological processes and improve our abilities to track animal movements. Using linear and additive models, we explored the influence of micro-scale habitat use, moisture and diet on tissue δD values of Nicaraguan cloud-forest birds. Using mist nets, we captured 211 individuals of 22 resident Neotropical species at 500–1390 m asl and collected feather and claw samples. Based on three years of data from year-round sampling, our results suggest that microhabitat, seasonal shifts in moisture δD, and diet all influence bird tissue δD values. Our model results reveal a previously undescribed microgeographical effect on δD, where foraging level (understorey versus overstorey) and foraging location (forest interior versus adjacent coffee plantation) were significant predictors of δD values in bird claws and feathers. Mean claw and feather δD values among species varied from −83‰ to −19‰. Top models for claws and feathers explained 57% and 52% of variation in δD respectively. Direct comparisons of understorey (mean ± SD of −30‰ ± 15‰) versus overstorey (−50‰ ± 15‰) claw values suggest that δD may be useful in tracking vertical, micro-scale movement. Higher δD values in forest understorey birds reveal a heavy reliance upon recycled, fog moisture. Fragmentation and climate change may result in increasingly desiccated cloud forest that may exert a more negative influence on the food webs of understorey species that seem to be supported by recycled sources of moisture in the dry season.

Mbau forest covers much of the Congo, and shifts in its composition could have a large impact on the African tropics. The Ituri forest in east Congo is near a boundary between the monodominant mbau type and non-mbau mixed forest, and two 20-ha censuses of trees ≥ 1 cm diameter were carried out over 12 y to monitor forest change. Based on published diameter allometry, mbau forest had 535 Mg ha−1 biomass above ground and gained 1.1 Mg ha−1 y−1. Mixed forest had 399 Mg ha−1 and gained 3 Mg ha−1 y−1. The mbau tree (Gilbertiodendron dewevrei) increased its share of biomass from 4.1% to 4.4% in mixed forest; other common species also increased. Sapling density declined at both sites, likely because increased biomass meant shadier understorey, but the mbau tree increased in sapling density, suggesting it will become more important in the future. Tree mortality and growth rates were low relative to other tropical forests, especially in the mbau plots. Shifting toward G. dewevrei would represent a large gain in carbon in the mixed forest, but mbau is presently more important as a high-carbon stock: biomass lost during forest harvest could not recuperate for centuries due to slow community dynamics.

Food webs of streams draining tropical rain forests on Costa Rica's Pacific and Caribbean coasts were examined in the 1980s via dietary analyses, and the same streams were surveyed again in 2004 to compare trophic structure based on analysis of stable isotope ratios of fish, macro-invertebrate and plant tissues. Estimates of species’ trophic positions (TP) were calculated from stomach-contents data (51 species; 5420 specimens) and compared with TP estimates derived from analysis of nitrogen isotope ratios (82 taxa; 240 samples). Coefficients of determination for TP based on dietary versus isotopic analysis ranged from 0.18 (Quebrada Camaronal, Corcovado) to 0.73 (Quebrada Estacion, Tortuguero). Consumer taxa within all four streams spanned a broad range of carbon isotope values, indicating assimilation of variable proportions of carbon from periphyton and terrestrial vegetation that in all but one of the streams had similar δ13C values. Approximately half the consumers in all four streams had carbon ratios heavier than any of the in situ production sources examined. This pattern could be explained by consumption of other production sources that were not sampled, including periphyton taxa with variable carbon isotope signatures, or migration of prey and/or consumers between these freshwater and coastal marine habitats.

The pitchers of Nepenthes rajah, a montane carnivorous plant species from Borneo, are large enough to capture small vertebrates such as rats or lizards, which occasionally drown therein. The interactions of N. rajah with vertebrates, however, are poorly understood, and the potential mechanisms that lure vertebrates to the pitchers are largely unknown. We observed frequent visits (average: one visit per 4.2 h) of both the diurnal tree shrew Tupaia montana and the nocturnal rat Rattus baluensis to pitchers by infrared sensor camera and video recording. Both mammalian species often licked the inner surface of the pitcher lid, which harbours numerous exudate-producing glands. Analysis of volatiles extracted from the secretions of the pitcher lids by gas chromatography coupled to mass spectrometry (GC/MS) revealed 44 volatile compounds, including hydrocarbons, alcohols, esters, ketones and sulphur-containing compounds, which are commonly present in sweet fruit and flower odours. The faeces of small mammals were repeatedly observed inside the pitcher, whereas we found the body of only one Tupaia montana drowned in the 42, vital and reasonably large, surveyed pitchers. Our findings suggest that the N. rajah pitcher makes use of the perceptual biases of rats and tree shrews by emitting volatiles known from fruits. The profits that the plant obtains from the repeated visits of two small mammals, together with the provision of exudates for the mammals, comprise an exceptional case of plant–vertebrate interaction.

The species composition of vascular epiphytes and phorophytes (trees and lianas) was studied in ten 0.1-ha forest plots distributed over three landscape units (floodplains, swamps and well-drained uplands) in Colombian Amazonia. The aim was to analyse how host-preferences contributed to the patterns in epiphyte assemblages among the landscape units. In the plots 82 species (3310 plants) were holo-epiphytes, 11 species were primary hemi-epiphytes (179 plants) and 61 were secondary hemi-epiphytes (2337 plants). A total of 411 species of tree and liana were recorded as phorophytes. Detrended Correspondence Analysis and Mantel tests showed that the species composition of holo-epiphytes and secondary hemi-epiphytes differed among the landscape units. For both groups the effect of landscape unit on species composition strongly decreased after controlling for the phorophyte composition in the plots. The phorophyte composition significantly explained epiphyte composition and this effect was not removed after accounting for the effect of landscape unit. At the level of individual species, randomization tests yielded only few significant epiphyte–phorophyte associations. For 84% of the epiphyte species the average indicator of patchiness was below 1.5 demonstrating that most epiphyte individuals occurred scattered over different phorophytes. This probably hampered the analyses of host preferences for individual epiphyte species.

The Bornean climber, Nepenthes bicalcarata, is unique among plants because it is both carnivorous and myrmecophytic, bearing pitcher-shaped leaves and the ant Camponotus schmitzi within tendrils. We explored, in the peat swamp forests of Brunei, the hypothesis that these ants contribute to plant nutrition by catching and digesting its prey. We first tested whether ants increased plant's capture rate. We found that unlike most plant-ants, C. schmitzi do not exhibit dissuasive leaf-patrolling behaviour (zero patrol on 67 pitchers of 10 plants) but lie concealed under pitcher rim (13 ± 6 ants per pitcher) allowing numerous insect visits. However, 47 out of 50 individuals of the largest visitor dropped into the pitchers of five plants were attacked by ants and the capture rate of the same pitchers deprived of their ambush hunting ants decreased three-fold. We then tested whether ants participated in plant's digestion. We showed in a 15-d long experiment that ants fed on prey and returned it in pieces in seven out of eight pitchers. The 40 prey deposited in ant-deprived pitchers remained intact indicating a weak digestive power of the fluid confirmed to be only weakly acidic (pH ~5, n = 67). The analysis of 10 pitcher contents revealed that prey, mainly ants and termites, was very numerous (~400 per pitcher per plant) and highly fragmented. Altogether, these data suggest a positive effect of C. schmitzi on both prey intake and breakdown. This ant–plant interaction could thus be a nutritional mutualism involving the unusual association of carnivory and myrmecotrophy.

Relations between crown width and tree height and trunk diameter and tree height were measured for trees of all sizes of six large emergent species (Balizia elegans, Dipteryx panamensis, Hieronyma alchorneoides, Hymenolobium mesoamericanum, Lecythis ampla and Terminalia oblonga) in the lowland tropical wet forest at La Selva, Costa Rica. Thirty to 45 individuals were chosen per species to provide a relatively even height distribution from 1 m up to the maximum encountered height. Among small juveniles, the crown widths of the two species with the smallest leaves and horizontally spreading branches were greater than those of the four large-leaved species. On reaching and exceeding the canopy, all but one species showed pronounced increases in crown width with increasing height. These patterns are linked to leaf size and first branching height in small juveniles and low densities of emergent trees at La Selva and other tropical forests, where crown expansion in all directions enhances light interception under the equatorial sun in trees that are not crowded by neighbours of similar height.

Seasonal changes in resource use and reproduction within a guild of nectar-feeding phyllostomid bats were studied between 2002 and 2004 in a west Mexican dry forest using mist-net captures and diet analysis over 82 nights during both dry and wet seasons. The local guild consisted of three resident species, Glossophaga soricina, Leptonycteris yerbabuenae, Musonycteris harrisoni, and of Anoura geoffroyi, which was mostly captured during the wet season. The proportion of L. yerbabuenae females more than doubled during the dry season. Diet was accessed by identifying the pollen collected from the bats' bodies and by analysis of faecal material. Bats used at least 28 different nectar plants over the year. In the dry season, 91.9% of all bats carried pollen while only 54.6% did so in the wet season. Bat individuals used the highest number of plant species during the dry season, mainly the cactus Pachycereus pecten-aboriginum, various Bombacaceae and the Capparaceae Cleome spinosa. Niche overlap was higher in the dry season than in the wet season. Musonycteris harrisoni and Leptonycteris yerbabuenae showed the largest overlap and both relied heavily on cactus flowers, whereas G. soricina supplemented its nectar diet with fruits. While reproductive females of most species were found only in the flower-rich dry season, G. soricina showed a bimodal reproductive pattern.

The spread of many invasive plants is facilitated through seed dispersal by frugivorous animals. The effectiveness of various frugivores as dispersers of the seeds of Melia azedarach, a highly invasive alien tree species, was evaluated in South Africa in savanna and bushveld vegetation. During 264 h of observation, seven bird species and one bat species were recorded foraging on fruiting trees of M. azedarach. The most common visitors were the dark-capped bulbul (Pycnonotus barbatus) followed by Wahlberg's epauletted fruit bat (Epomophorus wahlbergi), but both species dropped nearly as many seeds as they dispersed. Knysna turaco (Tauraco corythaix) dispersed the highest number of fruits per minute, but occurred in low abundance in our study sites. Seed germination differed significantly between de-pulped fruits and untreated fruits after 2 mo, but was similar after 4 mo. Germination success did not differ between animal-handled and hand-depulped fruits. In contrast to the high germination success in the greenhouse, seedlings showed very low recruitment in the field. Thus, M. azedarach seems likely to benefit from frugivores (particularly dark-capped bulbul and Wahlberg's epauletted fruit bat) dispersing seeds to suitable microsites.

Concurrent nutrient and radiation limitation in forests may engender trade-offs between P-use and radiation-use efficiency in tree species. To quantify these trade-offs, structural and physiological traits were examined among five rain-forest species subjected to four levels of fertilization and two levels of radiation in a glasshouse experiment. Schist specialists, Cryptocarya lividula and Ceratopetalum virchowii, occur only on P-poor schist soils, whereas soil generalists, Cryptocarya mackinnoniana, Franciscodendron laurifolium and Myristica insipida, occur on both P-poor schist and P-rich basalt soils. Wild seedlings less than 20 cm tall and 1 y old were collected from field sites, treated with fungicide, sorted into treatments (48 plants per species), and grown for 11 mo. We hypothesized that soil specialists would possess mainly non-plastic traits conferring high P-use efficiency, whereas soil generalists would possess markedly plastic traits conferring high radiation capture and use, enabling them to outcompete specialists on P-rich soils. Only generalist C. mackinnoniana and specialist C. virchowii supported these hypotheses. Cryptocarya mackinnoniana had more plastic root mass fraction, leaf area ratio, P uptake, and higher C assimilation than C. virchowii, which resulted in greater relative growth rates in high P treatments, but lower P-use efficiency in low P treatments. In contrast, specialist C. lividula demonstrated similar trait plasticity as C. mackinnoniana, suggesting that plasticity in these traits may be poor indicators of fitness on P-poor soils.

Livestock graze Indian forests to varying extents but their impact on wild native ungulates is rarely understood. Negative impacts of sympatric livestock on chital (Axis axis) demography and food availability were assessed and compared in the Gir Forest, India, at different spatio-temporal scales. No difference in average group size (mean ± SE) (7.11 ± 0.8 indiv.) (short-term response), fawn to doe ratio (0.43 ± 0.03) (short- to medium-term response), chital density (44.8 ± 7.1 indiv. km−2) (medium- to long-term response), and rate of population increase (r = 0.07 ± 0.014) (long-term response) was found between areas sympatric and livestock-free at the larger spatial scale of Gir Forest. Instead, chital density was correlated with rainfall (r = 0.92). After controlling for confounding factors of rainfall, vegetation community, terrain and lion density, chital density was 62% higher for livestock-free compared with sympatric areas but other demographic parameters showed no statistical difference. Peak above-ground biomass was greater in livestock-free (3255 ± 209 kg ha−1) compared to sympatric areas (1438 ± 152 kg ha−1), but chital food was more abundant in moderately grazed areas compared to livestock-free areas. Overall, long-term livestock grazing has depressive effects on chital but in the short term habitat productivity and suitability overrides the depressive effects of sympatric livestock.

Vegetation and environment change mutually during secondary succession, yet the idiosyncrasies of the vegetation effect on the understorey environment are poorly understood. To test whether the successional understorey environment changes predictably and is shaped by the structure and seasonality of tropical dry forests, we estimated basal area and vegetation cover, and measured understorey temperature, light and moisture conditions, in 17 plots forming a 60-y chronosequence and a mature forest. Light and air and soil temperature decreased with time (75−15% of open-sky radiation, 31.7−29.3 °C, and +2.5 °C to −0.5 °C relative to ambient, respectively), whereas relative humidity increased (67−74%). Soil water availability increased with early-successional development (−45 to −1 kPa) but decreased afterwards (to −18 kPa). The first axis of a PCA of the rainy-season environment explained 60% of the variation and was strongly related to air temperature and relative humidity. During tropical dry-forest succession, such factors may be more important than light, the reduction in which is not extreme compared with taller and more vertically stratified wet forests. Seasonality significantly affected the successional environmental gradients, which were marked mainly during the wet season. Environmental heterogeneity was higher in the wet than in the dry season, and larger for resources (light and water) than for conditions (temperature and humidity). The wet-season increase in environmental heterogeneity potentially creates differential growing scenarios; the environmental harshness of the dry season would mostly challenge seedling survival.

We examined the consequences of habitat fragmentation on the assemblage of floral visitors and pollinators to male- and female-phase inflorescences of the understorey dominant palm Astrocaryum mexicanum at the Los Tuxtlas tropical rain forest. In six forest fragments ranging from 2 to 700 ha, we collected all floral visitors, pollinators and non-pollinators, to male- and female-phase inflorescences at the time of their greatest activity. We used multivariate and mixed-effects models to explore differences in guild composition between sexual phases of inflorescences and the effects of forest fragment size on several metrics of the assemblages of floral visitors. We detected 228 786 floral visitors, grouped into 57 species, across the six forest fragments. On average, abundance and species richness of floral visitors to female-phase inflorescences were higher than to male-phase ones. Forest fragmentation had no effect on species richness but negatively affected Shannon's diversity index. Overall, the most abundant species of floral visitors were predominantly found in inflorescences of plants from the large fragments. In contrast, most of the less common species were more abundant in the smallest fragments. The abundance of pollinators (those found on inflorescences of both phases and dusted with pollen that was carried to flower stigmas), and the ratio of pollinators to other floral visitors, increased with fragment size in both sexual phases of the inflorescences but these effects were significantly stronger on male-phase inflorescences than on female-phase inflorescences. These results show that tropical forest fragmentation correlates with changes in the composition of flower visitors to a dominant palm, with a reduction in the abundance of pollinators, but that such changes co-vary with the sexual phase of the plants.

Biopedturbation, or animal-caused soil disturbance, may be important for development and maintenance of small-scale heterogeneity in ecosystems with the potential to alter seed mortality and recruitment. However, its role in tropical forests has been largely ignored. This study explored effects of vertebrate biopedturbation on seed (1) microsite heterogeneity, (2) predation and (3) germination in a West African rain forest. Exclosure experiments were used to study how biopedturbation altered burial and movements of seeds of four common canopy species. Effect of seed burial on removal by seed predators was also examined. Germination of the dominant canopy species (Raphia palma-pinus) in swamp forest was tested within artificial disturbances mimicking that of a locally common but endangered mammal, the Liberian mongoose (Liberiictis kuhni), which was estimated to turn over the entire forest floor in this habitat in c. 8 mo. Seed exposure to biopedturbation for 20 d (n = 80) led to an overall 6.5-fold increase in small-scale horizontal movement and increased probability of burial (6–52% higher), varying by species. Burial effectively eliminated seed removal for all four species (n = 160) by seed predators over 20 d. Germination of Raphia palma-pinus seeds (n = 100) was enhanced by 17.5% on average over 4 mo in simulated disturbances. Results suggest biopedturbation may be important for seedling recruitment and that loss of species with this functional role could have underappreciated yet important impacts on tropical plant communities.

Lantana camara, a shrub of Central and South American origin, has become invasive across dry forests worldwide. The effect of the thicket-forming habit of L. camara as a dispersal and recruitment barrier in a community of native woody seedlings was examined in a 50-ha permanent plot located in the seasonally dry forest of Mudumalai, southern India. Sixty 100-m2 plots were enumerated for native woody seedlings between 10–100 cm in height. Of these, 30 plots had no L. camara thickets, while the other 30 had dense thickets. The frequency of occurrence and abundance of seedlings were modelled as a function of dispersal mode (mammal, bird or mechanical) and affinities to forest habitats (dry forest, moist forest or ubiquitous) as well as presence or absence of dense L. camara thickets. Furthermore, frequency of occurrence and abundance of individual species were also compared between thickets and no L. camara. At the community level, L. camara density, dispersal mode and forest habitat affinities of species determined both frequency of occurrence and abundance of seedlings, with the abundance of dry-forest mammal-dispersed species and ubiquitous mechanically dispersed species being significantly lower under L. camara thickets. Phyllanthus emblica and Kydia calycina were found to be significantly less abundant under L. camara, whereas most other species were not affected by the presence of thickets. It was inferred that, by affecting the establishment of native tree seedlings, L. camara thickets could eventually alter the community composition of such forests.

Orchid species are well known for their highly specialized pollinator interactions. To better understand the reproductive biology of the tropical epiphytic orchid Bulbophyllum variegatum on Réunion, we investigated the floral morphology, breeding system, pollinator diversity, floral scent profile and fruiting success of about 30 individuals in three natural populations during two consecutive flowering seasons. Controlled hand-pollination experiments in two populations showed that the species is self-compatible, but requires pollinator service to achieve reproduction. Videotape pollinator observations were conducted during two flowering seasons for 56 h and revealed that B. variegatum is pollinated by a single species of fly from the Platystomatidae. This fly seems to be attracted by the unpleasant scent produced by the flowers, and does not receive any reward after achieving pollination. In addition, no egg-laying behaviour was observed. Bulbophyllum variegatum thus exhibits a typical sapromyiophilous pollination syndrome which constitutes the first proven case of sapromyiophily within the genus Bulbophyllum on the Mascarene Archipelago. Hand pollinations further showed that fruit set was not significantly higher for flowers that received outcross pollen than for those that were self-crossed (53% and 44% respectively). Fruit sets under natural conditions were significantly different among populations, ranging from 0.5% to 24.3%. This low fruit production is likely due to infrequent pollinator visits, particularly in disturbed forests where the pollinator has never been observed.