Understanding the ecological function of large frugivores in tropical forests is important considering their central position in seed-dispersal networks. This study illustrates the seed-dispersal effectiveness of a western lowland gorilla (Gorilla gorilla gorilla) population in Cameroon by looking at the interaction of seed quantity, species regeneration traits and the environmental characteristics of the deposition sites (sleeping sites vs. feeding trails) in order to discuss the role of gorillas within the ecosystem. The analysis of 1030 faecal samples collected over a 3-y period (September 2009–August 2012) showed that gorillas dispersed a diverse array of species (58 spp. identified), with an average of 289 intact seeds (>5 mm) d−1 per individual. Seed damage during mouth and gut processing occurred for 12 spp., generally accounting for <5% of specific seed load. Germination trials suggested that germination success of passed seeds remained unchanged (N = 6 spp.) or improved (N = 5 spp.) after gut passage compared with hand-cleaned seeds, with one exception. Indicator species identification (IndVal method) and General Discriminant Analysis suggested that sleeping sites (N = 36), as a main type of seed deposition site, correspond to regenerating microhabitats and correspondingly degree of canopy openness is one variable discriminating significantly this type of deposition site with random locations in the forest (N = 36) used as a proxy for non-directed seed deposition on feeding trails. With a mean canopy openness of 19–27%, sleeping sites offer optimal light conditions for the seedling recruitment of non-pioneer light-demanding species, the seeds of which encompass 47.4% of the total seed load dispersed over the 3-y period. From this study, it is hypothesized that the gorilla performed directed-dispersal of some light-demanding plant species at sleeping sites, and hence, by being actively implicated in gap-phase dynamics, fulfils a valuable ecological function within its ecosystem.

Large animal species are prone to local extirpation, but ecologists cannot yet predict how the loss of megaherbivores affects ecosystem processes such as seed dispersal. Few studies have compared the quantity and quality of seed dispersal by megaherbivores versus alternative frugivores in the wild, particularly for plant species with fruit easily consumed by many frugivorous species. In a disturbed tropical moist forest in India, we examine whether megaherbivores are a major frugivore of two tree species with easily edible, mammal-dispersed fruit. We quantify the relative fruit removal rates of Artocarpus chaplasha and Careya arborea, by the Asian elephant (Elephas maximus) and alternative dispersers. Through focal watches and camera trapping, we found the elephant to be amongst the top three frugivores for each tree species. Furthermore, seed transects under A. chaplasha show that arboreal frugivores discard seeds only a short distance from the parental tree, underscoring the elephant's role as a long-distance disperser. Our data provide unprecedented support for an old notion: megaherbivores may be key dispersers for a broad set of mammal-dispersed fruiting species, and not just fruit inaccessible to smaller frugivores. As such, the elephant may be particularly important for the functional ecology of the disturbed forests it still inhabits across tropical Asia.

Although seed dispersal plays a critical role for plant regeneration, the long-term benefits of frugivores for recruitment success have been poorly explored. We evaluated the relative importance of tree species abundance and of frugivore-related factors for successful plant recruitment on 18 tree species in a lowland Colombian rain forest. We combined census data from four 1-ha plots of trees (>10 cm dbh), saplings (1–5 cm dbh) and seedlings (<1.7 m) with a dataset describing tree–frugivore interactions. Seedling abundance was higher for large-seeded species dispersed by the spider monkey, while sapling abundance was higher for large-seeded species dispersed by birds. The identity of the dispersal agent and its interaction with seed size explained 20–30% of the total variance in seedling and sapling abundance across scales. Seed size consistently influenced the species abundance of seedlings and saplings across scales, but in opposite ways. These developmental changes suggest that what is beneficial to seedlings is not necessarily beneficial to saplings. Species identity explained 10–50% of the total variance in seedling and sapling abundance among and within 1-ha plots. Overall, our findings suggest that recruitment success is context-dependent as the relative importance of the different variables addressed may shift along spatial and temporal scales.

The fruit-tracking hypothesis predicts a positive association between frugivores and fruit abundance over space and time. We documented hornbill diets and examined the relationship between fruit abundance and abundance of three hornbill species (Buceros bicornis, Rhyticeros undulatus and Aceros nipalensis) in the Eastern Himalaya from 2009–2012. The study was carried out at three scales: at the largest scale of the study area (15 km2), at the intermediate scale – eight 3-ha patches within the study area and at the smallest scale of individual fruiting trees. Ninety-one per cent of the 64 foraging sightings of the great hornbill were on figs while more than 50% of the foraging sightings of the wreathed (83) and rufous-necked hornbills (87) were on non-fig fruits. At the largest scale, wreathed hornbill abundance and ripe fruit abundance peaked in the non-breeding season. At the intermediate scale, wreathed hornbill abundance was positively associated with non-fig fruit availability while rufous-necked hornbill abundance was negatively associated with non-fig fruit availability. At the smallest scale, great and rufous-necked hornbill abundances were correlated with fig and non-fig fruit crop sizes, respectively. The three hornbill species track fruit availability at different scales based on diet, which has implications for their role in seed dispersal.

We report on a 2009 survey of Acacia woodlands in the Seronera area of central Serengeti, Tanzania, and compare the results to previous surveys from the 1970s–1980s. We document a substantial change in woodland structure and composition. From 1978 to 2009, woody plant density (mean ± SD) declined from 255 ± 35 trees ha−1 to 147 ± 14 trees ha−1. Canopy cover declined from 15.1 ± 1.9% to 5.7 ± 0.6%. Canopy volume declined from 1810 ± 207 m3 ha−1 to 1410 ± 121 m3 ha−1. A 19% increase in the relative canopy volume of Acacia robusta, a species avoided by the giraffe, represents a sharp decline in the quality of the giraffe's food supply, which was last comprehensively assessed in 1978. We examine these changes in the context of Pellew's 1983 woodland dynamics model describing the impacts of elephant, giraffe and fire on an Acacia tortilis tree population and determine that our data are qualitatively consistent with this model. We hypothesize that selective elephant and giraffe browsing has contributed to an increase in the relative dominance of unpalatable species, consistent with Pellew's predictions.

Epiphytic orchids are very diverse in montane forests, but fragmentation modifies this diversity. Twenty fragments were quantified to evaluate the effects of fragmentation on the alpha and beta diversities of epiphytic orchids in a montane forest located in southern Mexico. The following factors were evaluated: area, core area, shape, edge density, Euclidean nearest-neighbour distance fragment and contrast index. In each fragment, two transects of 2 × 50 m were drawn, and the trees with a diameter at breast height ≥ 20 cm were recorded. In each tree, the orchid species present were identified and quantified. Twenty-three species of epiphytic orchid in 234 phorophytes corresponding to 20 species were recorded. The epiphytic orchid richness per tree and species turnover was different between the phorophytes. The edge density and the contrast index had significant effects on the alpha diversity, while the isolation of the fragments significantly affected the beta diversity. The edge density positively affected the alpha diversity of the epiphytic orchids, likely through microclimatic changes caused by fragmentation. Drought-tolerant species were common on the edges of the fragments, and shade-tolerant species established on the core area of the fragments. This pattern most likely depends on the ecological range of the taxa, which is crucial to their development and persistence in fragmented habitats.

Despite the important functional role of deep roots in withdrawing water during drought, direct measurements of root distribution are very rare in tropical rain forests. The aim of this study was to investigate the root distribution of Entandrophragma cylindricum, a common tree species in the Central African semi-deciduous rain forest, in Ferralsols and Arenosols. We dug two pits to a depth of 6 m in Ferralsols and two pits to a depth of 3 m in Arenosols, close to E. cylindricum trees. The vertical soil profiles were divided into 10 × 10-cm grid cells and the roots counted were distributed in three diameter classes. We fitted a root distribution model to our dataset. We found that vertical root distribution was shallower in Arenosols than in Ferralsols. Root penetration was not stopped even by a Ferralsol with high gravel content in its subsoil. Overall, our measurements showed that 95% of all roots were distributed to depths of between 258 and 564 cm from the soil surface, which is much deeper than the 95 cm depth previously reported in the literature for tropical rain forests. As sampling depth could explain this discrepancy, we recommend a sampling depth of at least 3–5 m to accurately estimate root distribution. The drier the dry season, the deeper the sampling depth should be. Our results are consistent with global models of root distribution in forest ecosystems, which are driven by climate variables. We thus suggest that deep rooting could be common in rain forests with a marked dry season.

Repeated use of sleeping trees (STs) by frugivores promotes the deposition and aggregation of copious amounts of seed, thus having key implications for seed dispersal and forest regeneration. Seed-rain patterns produced by this behaviour likely depend on the frequency of use of these sites, yet this hypothesis has been poorly tested. We evaluated community-level seed-rain patterns produced by the spider monkey (Ateles geoffroyi) over 13 mo in latrines located beneath 60 STs in the Lacandona rain forest, Mexico. Because this primate is increasingly ‘forced’ to inhabit fragmented landscapes, we tested whether sleeping-tree fidelity (STF) differed among sites and between continuous and fragmented forests. We also tested whether seed-rain patterns were associated with STF within each site and forest type. STF was highly variable among STs (average = 7 mo, range = 1–12 mo), but did not differ among study sites or forest types. STF was positively associated with seed abundance, species diversity and species turnover. Nevertheless, STF tended to be negatively related to seed community evenness. These results are likely due to the most frequently used STs being in areas with greater food density. Our results demonstrate that site fidelity shapes community-level seed-rain patterns and thus has key ecological implications.

The identification of seed dispersers and predators is essential to understand the effect of anthropogenic disturbances, and the associated defaunation process, on tropical forest dynamics in Central Africa. In this study, the animals involved in seed predation and dispersal of Dacryodes normandii (Burseraceae), an endozoochorously dispersed tree species endemic to Gabonese forests, were identified in a site in south-east Gabon using two complementary methods: direct observation and camera-trap monitoring of fruit piles. The combined sampling effort (172 h of direct observations and 796 d of camera trapping) led to the identification of six disperser and eight predator species of D. normandii seeds. With high frequency of consumption (88% and 57% of their visits, respectively) and long visit duration (83 and 23 min, respectively), the western lowland gorilla and central chimpanzee were identified as the main dispersers of this species. Seeds passed through the gorilla gut exhibited high germination success (68%). Rodents were identified as predators of D. normandii seeds, potentially displaying rare secondary dispersal through scatter-hoarding. The results of this study highlight the importance of great apes in the seed dispersal of this tree species.

The spatial distribution of adult trees is typically not expected to reflect the spatial patterns of primary seed dispersal, due to many factors influencing post-dispersal modification of the seed shadow, such as seed predation, secondary seed dispersal and density-dependent survival. Here, we test the hypothesis that spatial distributions of primary seed shadows and adult trees are concordant by analysing the spatial distributions of adult Genipa americana trees and the seed shadow produced by its key primary disperser, the capuchin monkey (Cebus capucinus) in a tropical dry forest in Costa Rica. We mapped the dispersal of G. americana seeds by the capuchins during focal animal follows (mean = 463 min, n = 50) of all adults in one free-ranging group over two early wet seasons (May–July, 2005 and 2006). We mapped the locations of all G. americana trees within a 60-ha plot that lay within the home range of the capuchin group. We conducted multiple spatial point pattern analyses comparing degrees of clustering of capuchin defecations and G. americana trees. We found that adult tree distributions and primary dispersal patterns are similarly aggregated at multiple spatial scales, despite the modification of the primary dispersal patterns and long dispersal distances.

Our aim was to evaluate the role of sucrose and the role of smell, taste and touch in the selection and consumption of fruit in wild spider monkeys. We recorded the feeding bouts of 14 adults for 9 mo in the Otoch Ma’ax Yetel Kooh Reserve, Punta Laguna, Yucatan, Mexico. For each of 2346 inspections on fruits of six species the consumption or rejection and the use of touch, smell and taste was recorded. Ten fruit samples (five ripe and five unripe) from each species were collected and the sucrose concentration was determined with a refractometer. As expected, sucrose concentrations were higher in ripe than unripe fruits. The difference in sucrose concentration between ripe and unripe fruits was positively associated with the proportion of inspections on ripe fruits and the proportion of consumed ripe fruits. Furthermore, the senses of touch and taste were used more often when fruits were ripe, whereas the sense of smell was used more often when fruits were unripe. The results suggest that sensory cues and sucrose concentration play important roles in fruit selection in spider monkeys.

To understand the variation in community dynamics of tropical montane forests along gradients of soil fertility, death, recruitment and growth of trees (≥5 cm diameter) were monitored over 14 y (1997–2011) in nine plots placed in a matrix of three geological substrate types (Quaternary sediments, Tertiary sedimentary rocks and ultrabasic rocks) and three topographical units (ridge, middle and lower slopes) on Mount Kinabalu, Borneo. The plot area was 0.05 ha for ridge, 0.1 ha for middle slope and 0.2 ha (on ultrabasic rocks) and 1 ha (on the other substrates) for lower slope. Recruitment rates did not show a consistent pattern across geological substrates or topographies. Mortality rates were relatively high in almost all plots during the 1997–1999 period, including the El Niño drought, and in three plots on ultrabasic rocks during 2001–2005. Binomial logistic regression analyses showed that mortality during 1997–1999 increased with soil fertility (soluble phosphorus). Background mortality, excluding these periods, did not differ across geological substrates or topographies. The average growth rate during 1997–2011 was higher on more fertile soils and positively correlated with mortality during 1997–1999. We suggest that a high mortality rate during the drought period was related to high species diversity on more fertile soils, whereas a lower growth rate was related to stunted structures on poorer soils.

We studied the removal of seeds of three species of large-seeded tree (Astrocaryum standleyanum, Attalea butyracea and Dipteryx oleifera) from three different heights within six study plots in a lowland forest in central Panama. Fresh fruits with intact seeds fitted with industrial sewing bobbins were placed within semi-permeable exclosures. Removed seeds were tracked to deposition sites, and seed fate was determined. Removals were likely perpetrated by two small rodents, the strictly terrestrial Proechimys semispinosus and the scansorial Sciurus granatensis, because they were the most abundant small rodents in the study site during the study period and were of sufficient size to remove large seeds. Rodent abundance and fruit availability were estimated by conducting censuses. Nine microhabitat variables were measured at each deposition site to determine if these two rodents were preferentially depositing seeds in sites with certain characteristics or were randomly depositing seeds. During the study, rodents handled 98 seeds, 85 of which were not predated upon and could potentially germinate. Removal rates were not influenced by rodent abundance or fruit availability. Seeds were most frequently moved <3 m and deposited with the fruit eaten and the seed intact. However, some seeds did experience relatively long-distance dispersal (>10 m). Rodents preferentially deposited seeds in locations with large logs (>10 cm diameter), dense herbaceous cover, and an intact canopy. The number of large logs was different from random locations. Despite not being able to determine long-term fate (greater than c. 1 y), we show that these small rodents are not primarily seed predators and may in fact be important mutualists by dispersing seeds relatively long distances to favourable germination sites.

Understanding the habitat preferences of native and non-native species may offer valuable insights into the mechanisms favouring invasion of disturbed habitats. This study investigated the determinants of trap-site detection probability of three native (Maxomys surifer, Maxomys whiteheadi and Leopoldamys sabanus) and one invasive (Rattus rattus) species of terrestrial murid (Muridae) in logged and unlogged forests of northern Borneo. We established four and two trapping grids in repeatedly logged and unlogged forest, respectively, for a total of 500 sampled trap sites. From these, we obtained 504 detections of the four species over 3420 trap nights. For each species, probability of detection was modelled as a function of both the structural components and disturbance level of the forest patch measured around each trap site. Each of the four species showed contrasting microhabitat preferences: M. surifer favoured increased canopy closure and intermediate ground and understorey vegetation cover; M. whiteheadi preferred increased ground vegetation cover and canopy height; L. sabanus favoured sites with larger amounts of coarse woody debris and less leaf litter; and R. rattus was associated with increased ground vegetation cover. Within logged forest, detection probabilities of the three native species did not vary significantly with level of patch disturbance, whereas that of the invasive R. rattus increased markedly in more degraded sites. This latter finding will have increasingly important implications when considering the rapid degradation of forests in the region, and the resulting expansion of suitable habitat for this competitive species.

South-East Asian tropical rain forests experience sporadic, but profuse, seed production after general flowering, leading to the synchronous emergence of various seedlings and subsequent seedling dynamics, which play a crucial role in determining species distribution and coexistence. We examined the relative importance of both biotic (initial height, conspecific seedling density) and abiotic (canopy openness, per cent sand, soil water content) drivers using survival data for 1842 seedlings of 12 dipterocarp species for 1.5 y following mast fruiting in an old-growth Bornean tropical rain forest. More than 30% of all dipterocarp seedlings survived 1.5 y after mast fruiting. When all species were analysed together, we found that initial seedling height, canopy openness and conspecific seedling density affected dipterocarp seedling survival. Negative density dependence indicated that predators were not satiated, but dipterocarp seedlings rather suffered from host-specific natural enemies or intraspecific competition. Species-level analyses of seven dipterocarp species showed large variation in response to biotic and abiotic factors. These results suggest that interspecific differences in the relative importance of biotic and abiotic effects on seedling survival might contribute to species coexistence.

In order to understand the stoichiometric balance between foliar nitrogen (N) and phosphorus (P) on P-poor soils, we investigated how foliar N and P attributes (i.e. N and P concentrations in green and senesced leaves, N and P resorption efficiencies) of 30 tropical tree species co-vary along a gradient of soil P availability across three forests on Mount Kinabalu, Borneo. We found strong and positive correlations between foliar N and P in the concentrations and resorption efficiencies within each forest and across the three forests. Slopes of standardized major axis between foliar N and P concentrations for both green and senesced leaves were not different among the three forests, although the values of the scaling exponent in the relationships of foliar N to P across the three forests were significantly lower than 1. We suggest that down-regulation of N concentrations in green leaves on P-poor soils is one of several possible mechanisms explaining why N concentrations decrease with decreasing P concentrations in both green and senesced leaves toward a lower P availability in soils. On the other hand, the physiological and ecological reasons why N and P resorption efficiencies are positively correlated with each other across tree species remain unclear.

Body size in animals is an important trait affecting species niche differentiation and restricting similarity. Using camera-trap data over 2008–2010, we used photo-captures from 50 cameras spread throughout Mudumalai Tiger Reserve (Western Ghats, India) to assess the activity budgets of 21 mammal species ranging in body size from 1 kg to 2088 kg. Large carnivores were mostly cathemeral whereas small cat and civet species were purely nocturnal. Mongoose species were mainly diurnal possibly due to their terrestrial feeding habits and reduce competition with other sympatric small carnivores. All large and small-bodied herbivores were cathemeral and nocturnal respectively, whereas medium-sized herbivores were active during the day. Overall, small mammals tended to be mostly nocturnal, whereas large mammals were cathemeral mainly due to energy requirements and other ecological constraints. Body size showed significant negative relationship with mean vector length (clustering of activity in time) thus implying that the daily amount of time being active increased with body size. The shorter activity time (12 h) in small mammals resulted in higher mean vector length probably to utilize the available time to fulfil energy needs. The observed cathemeral activity in large mammals may be associated with travel over larger areas to acquire large quantities of food therefore they are active for a longer duration. Our results clearly support the allometric relationship between body size and activity budgets in mammals and its association with niche differentiation.

The response of vascular epiphyte communities following natural or human disturbance has been little studied. Over 5 y, we evaluated the post-stripping recolonization of vascular epiphytes in cloud forest. Vascular epiphytes were experimentally removed from branch and trunk plots (1 m in length) on five trees in two secondary cloud forest fragments in southern Mexico. Similarity between colonizer and established communities was compared in each fragment using a further five trees with no stripping. All seedlings were recorded yearly. Non-vascular epiphyte cover was estimated in each plot. The recolonization rate was very high; after 5 y, epiphyte density of the colonizer community (27.4 ± 6.8 individuals per segment) reached similar values to those of the established community (26.7 ± 3.3) in nearby trees. While similarity (composition and abundance) between the colonizer community and established community was high (81%), diversity accumulation curves indicated that the colonizer community presents a lower diversity of epiphytes (5.5 equivalent species) than the established community (11.4). Colonization of xerophytic bromeliads was high, while pteridophytes and orchids presented reduced recovery. The immediately surrounding source of propagules had a strong influence on recolonization. In both the colonizer and established communities, dominance rank was bromeliads > peperomias > pteridophytes. The results show that the recovery capacity of epiphytic vegetation in secondary forest is high, if propagule sources are close by. However, at 5 y after disturbance, it is unclear whether the colonizer community would present the same species composition as the established community or if it would give rise to a different community.

The importance of deterministic and stochastic processes in structuring ecological communities is an enduring debate. Although this debate is nearly a century old, the extent to which communities are structured by species interactions or chance events is a central issue in ecology. We examined the assemblages of plants living inside 119 birds’ nest ferns (Asplenium goudeyi), which are endemic to Lord Howe Island. Specifically, we investigated whether patterns of species richness and community composition were influenced by fern size, fern isolation and plant dispersal abilities. Fern size and fern isolation significantly predicted plant community richness. At the community level, plant composition patterns did not deviate from randomized expectations. Individual species occurrences increased with increasing community richness, and no species exclusions were observed. Wind-dispersed taxa, which accounted for 29% of all species, were well represented in isolated ferns. Comparatively, poorer dispersers were confined to ferns nearest the forest at the base of the cliffs. We suggest that dispersal plays a key role in structuring plant communities living within birds’ nest ferns, and that species interactions are less important. Our study emphasizes the importance of epiphytes with a nest-like growth form as habitat for plants in a harsh environment.

Not much is known about the nitrogen (N) uptake capacity and N-form preference of tropical trees. In a replicated labelling experiment with 15N-ammonium, 15N-nitrate and dual-labelled glycine applied to saplings of six tree species from southern Ecuadorian montane forests, we tested the hypotheses that (1) the saplings of tropical trees are capable of using organic N even though they are forming arbuscular mycorrhizas, and (2) with increasing altitude, tree saplings increasingly prefer ammonium and glycine over nitrate due to reduced nitrification and growing humus accumulation. Three- to 5-y-old saplings of two species each from 1000, 2000 and 3000 m asl were grown in pots inside the forest at their origin and labelled with non-fertilizing amounts of the three N forms; 15N enrichment was detected 5 days after labelling in fine roots, coarse roots, shoots and leaves. The six species differed with respect to their N-form preference, but neither the abundance of ammonium and nitrate in the soil nor altitude (1000–3000 m asl) seemed to influence the preference. Two species (those with highest growth rate) preferred NH4+ over NO3−, while the other four species took up NO3− and NH4+ at similar rates when both N forms were equally available. After 13C-glycine addition, 13C was significantly accumulated in the biomass of three species (all species with exclusively AM symbionts) but a convincing proof of the uptake of intact glycine molecules by these tropical montane forest trees was not obtained.