The extent, sequence, synchrony and correlates of diel displacements by animals can provide powerful insights into the ecological and social factors that shape an organism's day-to-day activities, but detailed data on spatial ecology are available for very few tropical taxa. Radiotelemetric monitoring of 25 slatey-grey snakes (Stegonotus cucullatus) on a floodplain in the Australian wet-dry tropics for periods of 40 to 355 d (mean=195 d, 136 locations per snake) provided extensive information on habitat use, movement patterns and home range size of these large slender-bodied colubrids. All radio-tracked animals were nocturnal, sheltering by day in soil cracks and beneath tree roots and debris. Snakes did not appear to move between 61% of successive locations and timing of movements was not synchronized among snakes. Most displacements were small (<50m), with males moving further and more often than did females. However, nesting females made occasional long-distance movements, travelling 100–400m to forested areas to oviposit but then returning to their usual home ranges. Snakes of both sexes moved further and more often during the wet-season than the dry-season. Snakes typically moved on a few successive nights then remained sedentary for the next few days, apparently reflecting cessation of activity as soon as a meal was obtained. Home ranges were small (2.9–7.4 ha) and most snakes remained in the same area throughout the year, providing a strong contrast in this respect to the large and seasonally dynamic home ranges of sympatric acrochordid and pythonid snakes.

A trapping study of five mammal species in wet sclerophyll forest adjacent to rain forest in the Australian Wet Tropics was used to examine the seasonal diversity, abundance and dung-specificity of dung beetles associated with mammal dung. A total of 542 dung beetles from 11 species within three genera was recovered from beneath the traps of 1104 mammal captures. The diversity of beetles associated with the dung of the northern bettong (Bettongia tropica), a mycophagous marsupial, differed significantly from the diversity predicted by a null model. Numbers of beetles varied significantly with type of dung, indicating preference by beetles. Beetle numbers were related positively to a 1-mo lag in monthly mean minimum temperature and less strongly to maximum temperature and rainfall. Significantly more beetles per mammal capture were detected in the wet season than in the dry season. Dung beetles showed a strong preference for either the Eucalyptus woodland (six species) or the adjacent Allocasuarina forest (four species), with only one species occurring in both habitat types. Beetle species from the Eucalyptus woodland were typically only detected in the late wet and early dry seasons, while species in the wetter Allocasuarina forest were generally collected during the late dry and early wet seasons. A significant ‘checkerboard’ species effect was detected in both time and space in both habitat types, suggesting that competition for dung was strong.

Breeding systems and dispersal mechanisms of plants (≥ 2.5 cm dbh) were examined in fragments of tropical dry forest in Central America to identify life-history characteristics associated with rarity. In particular, the richness and abundance of dioecious and mammal-dispersed trees and shrubs were examined to identify potential associations with precipitation, anthropogenic disturbance, and area. Plots totalling 1000 m2 per site were established in seven nature reserves in Costa Rica (two sites) and Nicaragua (five sites). Overall, tropical dry forests of Central America have a similar proportion of dioecious species to other lowland neotropical forests and a similar proportion of wind-dispersed plants to other tropical dry forests around the world. However, the number of dioecious and mammal-dispersed species declined with decreasing forest cover within each reserve. Although dioecious species were rare in smaller forest fragments, some of these species will not be threatened with regional extinction because they are early successional plants, they have large geographic ranges, and they are not restricted to the tropical dry forest life zone. Mammal-dispersed plants were rare in small fragments, but it is not clear whether this was due to the loss of dispersal vectors or other life-history characteristics.

Habitat fragmentation affects the ecology of tropical rain forests in many ways, such as reducing species diversity of many taxa (Laurance et al. 2002, Lovejoy et al. 1986) and increasing rates of tree mortality and canopy-gap formation near forest edges (Laurance et al. 1997, 1998, 2001). Such obvious alterations have been documented in many fragmented forests, but more subtle changes, such as those affecting plant phenology (the timing and frequency of flower, fruit and leaf production), have received far less attention. Adler & Kiepinski (2000) showed that different populations of the successional tree Spondias mombin on small man-made islands in Panama had highly synchronous flowering and fruiting. In montane forests in Colombia, Restrepo et al. (1999) demonstrated that under-storey fruit abundance was consistently increased over time near forest edges relative to forest interiors. Beyond these and a few other studies (Ackerly et al. 1990, Nason & Hamrick 1997), however, the effects of fragmentation on plant phenology have been inadequately assessed, especially in the tropics.

Seed-dispersal ecology in tropical montane forests (TMF) differs in some predictable ways from tropical lowland forests (TLF). Environmental, biogeographic and biotic factors together shape dispersal syndromes which in turn influence forest structure and community composition. Data on diaspore traits along five elevational gradients from forests in Thailand, the Philippines, Tanzania, Malawi and Nigeria showed that diaspore size decreases with increasing altitude, fleshy fruits remain the most common fruit type but the relative proportion of wind-dispersed diaspores increases with altitude. Probably corresponding to diaspore size decreasing with increasing elevation, we also provide evidence that avian body size and gape width decrease with increasing altitude. Among other notable changes in the frugivorous fauna across elevational gradients, we found quantitative evidence illustrating that the proportion of bird versus mammalian frugivores increases with altitude, while TMF primates decrease in diversity and density, and switch diets to include less fruit and more leaf proportionately. A paucity of studies on dispersal distance and seed shadows, the dispersal/predation balance and density-dependent mortality thwart much-needed conclusive comparisons of seed dispersal ecology between TMF and TLF, especially from understudied Asian forests. We examine the available evidence, reveal knowledge gaps and recommend research to enhance our understanding of seed dispersal ecology in tropical forests. This review demonstrates that seed dispersal is a more deterministic and important process in tropical montane forests than has been previously appreciated.

An important challenge of invasion biology is to understand how interactions between species traits and ecosystem properties enable alien species to become invasive at particular locations. We investigated how gap dynamics in a tropical rain forest on the island of Réunion affected the invasiveness of alien plants. In the 12 000-m2 study area, alien plants occupied 24.9% of the area of gaps, which represented 5.62% of the forest area, but only 0.8% of the understorey area. The most abundant invasive species was Rubus alceifolius, which formed dense, monospecific stands in the largest gaps (> 25 m2). Although plants could persist in the shade, a germination experiment revealed that canopy openings were essential for seedling establishment. A cyclone that struck the study area in 2002 caused a temporary thinning of the canopy, increasing light levels to above the threshold needed for germination of R. alceifolius and also stimulating the growth of established plants. We conclude that the ability of this and other alien species to colonize intact lowland tropical rain forest is strongly influenced by the prevailing gap dynamics. Because gaps are also essential for the regeneration of many native trees in our study area, there is a real danger of the forest being progressively degraded by alien plants. There are no simple solutions to controlling species such as Rubus alceifolius, but efforts should be focused mainly upon the larger gaps where the species are most invasive.

Primates are among the major groups of frugivores in the tropics, but little is known about their role in natural regeneration of African savanna ecosystems. In the savanna-forest mosaic of north-eastern Ivory Coast the olive baboon (Papio anubis Lesson) harvests fruit from at least 79 plant species. Over a 24-mo study period, 396 faecal samples from 10 groups of baboons were analysed in terms of quantitative and qualitative aspects of seed handling and dispersal (excluding grasses and sedges). Seventy-three per cent of seeds in faecal samples were undamaged. Intact seeds were from 65 species. On average, defecations contained intact seeds from 2.0 species (range = 0–10). Seed size varied between 1 and 27 mm, and 77% of the species had medium-sized to large seeds. No linear correlation between mean seed size and seed damage was found. Eighteen out of 19 species tested were viable after ingestion, but effects of gut passage upon germination varied widely. The baboon population in the study area (145 km2) dispersed an estimated 1483 intact seeds d−1 km−2 (129 seeds > 2 mm in size). The results suggest that the olive baboon is an important seed disperser in the savanna-forest ecosystem of West Africa.

Understanding exotic–native plant relationships within the context of landscape-scale environmental factors such as rainfall, topography, disturbance and forest structure, is important for distinguishing the role of invasions in native plant community change. In this study, the relationship between Lantana camara invasion and native forest understorey vegetation was described after accounting for environmental influences. Rainfall, terrain slope, altitude and fire frequency were measured from GIS layers, and tree density, dry above-ground L. camara biomass, and native plant abundance and species richness were measured in nested plots (four 1 × 1-m plots for grass, tree seedlings and L. camara within one 5-m-radius plot for tree saplings and herbs and shrubs within one 10-m-radius plot for trees) at 80 locations distributed across Bandipur Tiger Reserve, a tropical deciduous forest in southern India. Relationships between environmental factors and L. camara abundance were described using a multiple regression. Further, the role of L. camara abundance in explaining residual variation in native vegetation, after accounting for environmental influences, was described using linear models. Lantana camara abundance decreased with increasing tree density, supporting the notion that it thrives in disturbed forest with fewer trees. Whereas native tree seedling and herb and shrub density showed no relationship to L. camara abundance, both tree sapling density and grass volume decreased under L. camara biomass exceeding 2 kg m−2. These data suggest that, by association with grass decline and decreased recruitment of tree saplings, L. camara may be linked to the gradual transition of these tropical deciduous forests into exotic-dominated shrubland, and overall native biodiversity loss.

The present study showed that the density of germinable seeds in the soil of a humid tropical forest ecosystem at Nelliampathy, in the Western Ghats of Kerala in India, fluctuates sharply at different times of the year. There were more germinable seeds in the soil during the monsoon season. Regeneration was activated after canopy gap formation and thus a decline in the soil seed density in 1-year-old gaps was recorded. However, soil seed density increased with gap age. Differences were found between primary, late secondary and early secondary categories of species with respect to their soil seed bank. There were more seeds of primary species in the soil of natural gaps than in selection felled ones, while the seeds of early secondary shrubs and herbs were more in the soil of selection felled gaps than in natural gaps. The soil seed banks of the primary forest species, being transient, are not an important conservation tool for these species.

To address how seasonality affects the richness and abundance of tropical insects, we compared the canopy and understorey communities of fruit-feeding butterflies in a seasonal Atlantic forest in south-eastern Brazil. Butterflies were sampled over 1 y using a standardized design with baited traps. A total of 2047 individuals in 69 species were recorded (1415 in the canopy, 632 in the understorey). Clear differences were found between canopy and understorey, with significantly higher butterfly abundances in the canopy. We observed two marked peaks of abundance and richness in both strata; one at the transition from dry to the wet seasons, and the other at the transition from wet to dry seasons. We found lower species turnover throughout the year in the canopy. We interpret this as evidence that temperature is more important than rainfall in explaining the yearly variation of abundance in vertical strata. The higher temperatures found in the canopy may allow butterflies to maintain activity in this stratum all year round, whereas the understorey is subject to colder temperatures, thus presenting a higher species turnover. These results improve our understanding of diversity gradients between evergreen and seasonal tropical forests, allowing insights into how climate and beta diversity gradients interact.

A representative sample of waterways in Liberia was surveyed for crocodiles, and all three species of African crocodiles were observed. Contrary to previous reports, only Nile crocodiles inhabit the Monrovia mangrove swamps and brackish-water mouths of rivers; there are no slendersnouted or dwarf crocodiles in these habitats. There is partitioning of habitats among the three species in Liberia: Nile crocodiles in mangrove swamps and river mouths (brackish water); slendersnouted crocodiles in rivers through rain forest (freshwater); and dwarf crocodiles in small streams through rain forest (freshwater), some entering the adjacent river. Dwarf crocodiles utilize burrows in the stream banks.

Nowhere are crocodiles abundant, and there is an apparent absence of adults. Although 40% of Liberia is forested, deforestation is occurring rapidly by foreign timber companies and slash-and-burn agriculture practised by the largely rural population. Hunting with firearms, although illegal, is widespread, both for subsistence and commercially, without concern for depletion. Mangrove ecosystems near human population centres are being destroyed. The combination of the above factors has apparently depleted the crocodile populations in Liberia.

Survival of Afrotropical primary forests depends not only on habitat protection but also on the protection of animal species such as frugivorous primates, recognized as the most important seed dispersers for many plants. Here we investigate seed-dispersal services by the bonobo (Pan paniscus) in an evergreen lowland tropical rain forest of the Congo Basin. In the long-term research site of LuiKotale, we investigated food habits and seed processing based on 22 mo of behavioural observation, seed trial experiment and long-term daily GPS tracking of a habituated ape community. Bonobos were mainly frugivores (66% of all feeding sessions), spending about 3.5 h d−1 swallowing seeds that were transported for an average of 24 h in the gut. On average, an individual bonobo dispersed 172 kg y−1 of seeds (or 220000 seeds y−1) of more than 91 plant species by endozoochory over an average distance of 1.2 km from the parent tree. Passed seeds germinated more rapidly, more successfully and had greater post-dispersal survival than unpassed seeds. Bonobo-dispersed plants accounted for 40% of tree species and 65% of individual trees in the study site (12 1-ha plots census). Since bonobos show little functional overlap with other frugivores, the loss of their seed-dispersal services is likely to affect forest structure and dynamics. Our results justify description of the threatened bonobo as a gardener of the Congo forest.

Comparisons of sympatric reptile species were used to assess the variation in niche overlap for food between potential competitors at different trophic levels. Omnivorous tortoises and carnivorous vipers inhabiting the rain-forest region of West Africa were used as study models. Food niche overlap between species increased with habitat alteration in both the independent study systems: tortoises (Kinixys homeana and Kinixys erosa) and vipers (Bitis gabonica and Bitis nasicornis) showed lower values of Pianka's niche overlap index in the pristine habitat than in the altered habitat, and these differences in overlap values did not depend on chance after Monte Carlo simulations. There were higher inter-habitat food niche overlaps within-species than between-species. Permutation tests (assessed after 5000 iterations) revealed that, for both study systems, the P-values became significantly smaller with fewer resource states, thus showing the niche overlap between species really increases after habitat alteration. The observed increases in food niche overlap between species accomplished with rain-forest habitat degradation in turn may be predicted to have cumulative effects on reducing the level of forest biodiversity.

(Accepted 31st July 1999)

In the ‘elfin’ cloud forest of Serrania de Macuira, exchangeable ammonium and nitrate, and the rates of soil nitrogen mineralization and nitrification were measured in soil samples under different water, temperature and mineral nutrient additions. The effects of nitrogen, phosphorus and nitrogen plus phosphorus fertilization on radial trunk growth were measured in three tree species. In the cloud forest soils, concentrations of ammonium were much higher than those of nitrate. Nitrate was higher in samples collected during the afternoon than during the morning, probably as a result of leaching during the night or nitrification during the day. When samples were incubated under different water and temperature treatments, rates of nitrogen mineralization and nitrification increased more with changes in soil water content than with changes in temperature. Nitrification was significantly increased in soils amended with ammonium or with ammonium plus phosphorus, suggesting that nitrification is substrate-limited. Fertilization with nitrogen and phosphorus resulted in significantly increased girth increments in Guapira fragrans (Dum. -Cours.) Little and Rapanea guianensis Aublet. Myrcianthes fragrans (Sw.) D.C. did not respond to the fertilization. The results of this study support the hypothesis that the characteristics of montane rain forest in small and large tropical mountains (the ‘Massenerhebung’ effect) are greatly controlled by soil water conditions and related soil nitrogen availability.

In tropical forests, severe droughts caused by El Niño events may strongly influence the water relations of tree seedlings and thereby increase their mortality. Data on known-aged seedlings of three common shade-tolerant canopy tree species (Trichilia tuberculata, Tetragastris panamensis and Quararibea asterolepis) in a Panamanian moist forest are presented. Seedling survival during a severe El Niño dry season (1997-98) was compared with prior long-term survival data, and levels of drought stress were assessed by measuring plant water potentials and gas exchange characteristics. Contrary to prediction, dry-season seedling survival was not dramatically reduced in any species compared with that expected in ‘normal’ years. In Trichilia and Quararibea, pre-dawn water potentials averaged −2 MPa and midday water potentials about −3 MPa. Stomatal conductances were very low, averaging 26 mmol m-2 s-1 for Tetragastris and 11–13 mmol m-2 s-1 for Trichilia and Quararibea. Photosynthetic rates also were very low but consistently positive, averaging 0.8–1.1 μmol m-2 s-1. The findings suggest that, once established, seedlings of common tree species in this semi-deciduous forest may be tolerant of drought events.

The ‘campos de murundus’ (carthmound fields) are typical landscapes of the cerrado (savanna) region of Central Brazil and are characterized by many rounded earthmounds called ‘murundus’ which are scattered over a grassland surface. As these areas are normally liable to seasonal water-logging, the earthmounds are true habitat islands capable of bearing woody plants typical of the cerrado, which are normally very intolerant of root inundation. The vegetation of an area of murundus field at Cuiaba, in the State of Mato Grosso, Central Brazil, was surveyed for the purpose of evaluating the island-effect on the plant community growing on the mounds/islands. Two main aspects were considered: the influence of the size of the mound and the differences between the types of cerrado occurring on the mounds and on the areas surrounding the murundus field. Most plant species occurring on the mounds were also present in the flora of the surrounding cerrado. However, only a fraction of these species was able to colonize the mounds successfully. The number of species on the mounds was found to be significantly correlated with mound area. Speciesarea relationships were better described with the power model S = C(Az) than with the exponential model S = C + z (logA) (fitted with linear regression). The mound size also had a strong influence on the spatial distribution of the species on the surface of the mounds. The species were organized in an increasing number of concentric topographic belts corresponding to a gradient of soil depth above the Hooding level.

The development of metacommunity theory, which suggests that the diversity and composition of communities is influenced by interactions with other communities, has produced new tools for evaluating patterns of community change along environmental gradients. These techniques were used to examine how plant communities changed along elevation gradients in montane tropical forests. Two transects of 0.1-ha vegetation plots were established every 50 m in elevation in the mountains of eastern Puerto Rico. The transects ranged from 300 m to 1000 m asl and 400 m to 900 m. In each plot, all free-standing woody stems greater than 1 cm in diameter at 130 cm in height were marked, measured and identified. Additional data on three similar transects were taken from the literature. The upper or lower boundaries of species ranges were significantly clumped along all five transects. Coherence, a measure of the number of gaps in species distributions, was also significant across all transects, and three transects showed significant, albeit low, nestedness. Four sites had significant species turnover. These results suggest that metacommunity techniques can be useful in searching for patterns of community change present in montane tropical 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.