Solar irradiance and precipitation are the most likely drivers of the seasonal variation of net primary productivity (NPP) in tropical forests. Since their roles remain poorly understood, we use litter traps, dendrometer bands and census data collected from one hectare permanent plots to quantify the seasonality of above-ground NPP components and weather parameters in 13 sites distributed along a 2800-m altitudinal gradient ranging from lowland Amazonia to the high Andes. We combine canopy leaf area index and litterfall data to describe the seasonality of canopy production. We hypothesize that solar irradiance is the primary driver of canopy phenology in wetter sites, whereas precipitation drives phenology in drier systems. The seasonal rhythm of canopy NPP components is in synchrony with solar irradiance at all altitudes. Leaf litterfall peaks in the late dry season, both in lowland (averaging 0.54 ± 0.08 Mg C ha y−1, n = 5) and montane forests (averaging 0.29 ± 0.04 Mg C ha y−1, n = 8). Peaks in above-ground coarse woody NPP appears to be triggered by the onset of rainfall in seasonal lowland rain forests (averaging 0.26 ± 0.04 Mg C ha y−1, n = 5, in November), but not in montane cloud forests.

Avian diet selection is hypothesized to be sensitive to seasonal changes in breeding status, but few tests exist for frugivorous tropical birds. Frugivorous birds provide an interesting test case because fruits are relatively deficient in minerals critical for reproduction. Here, we quantify annual patterns of fruit availability and diet for two frugivorous hornbill (Bucerotidae) species over a 5.5-y period to test for patterns of diet selection. Data from the lowland tropical rain forest of the Dja Reserve, Cameroon, are used to generate two nutritional indices. One index estimates the nutrient concentration of the diet chosen by Ceratogymna atrata and Bycanistes albotibialis on a monthly basis using 3165 feeding observations combined with fruit pulp sample data. The second index is an estimate of nutrient concentration of a non-selective or neutral diet across the study area based on tree fruiting phenology, vegetation survey and fruit-pulp sample data. Fifty-nine fruit pulp samples representing 40 species were analysed for 16 nutrient categories to contribute to both indices. Pulp samples accounted for approximately 75% of the observed diets. The results support expected patterns of nutrient selection. The two hornbill species selected a diet rich in calcium during the early breeding season (significantly so for B. albotibialis in July and August). Through the brooding and fledging periods, they switched from a calcium-rich diet to one rich in iron and caloric content as well as supplemental protein in the form of invertebrates. Calcium, the calcium to phosphorus ratio and fat concentration were the strongest predictors of breeding success (significant for calcium and Ca:P for B. albotibialis in June). We conclude that hornbills actively select fruit based on nutritional concentration and mineral concentration and that the indices developed here are useful for assessing frugivore diet over time.

Insectivorous birds and bats often protect plants through density- and trait-mediated cascades, but the degree to which insectivores reduce herbivorous arthropods and leaf damage varies among systems. Top-down interaction strength may be influenced by the biotic and abiotic context, including the presence of vegetation-disturbing animals. We tested two hypotheses: (1) insectivorous birds and bats initiate trophic cascades in tropical rain-forest understorey; and (2) the native, omnivorous collared peccary (Pecari tajacu) negates these cascades via non-trophic effects. We studied the top-down effects of birds and bats on understorey plants in north-eastern Costa Rica using 60 netted exclosures within and outside existing peccary exclosures. Excluding birds and bats increased total arthropod densities by half, both with and without peccaries. Bird/bat exclosures increased Diptera density by 28% and leaf damage by 24% without peccaries, consistent with a trophic cascade. However, bird/bat exclosures decreased Diptera density by 32% and leaf damage by 34% with peccaries, a negation of the trophic cascade. Excluding peccaries increased leaf damage by 43% on plants without birds and bats. This is the first study, to our knowledge, to demonstrate that the non-trophic activity of an omnivorous ungulate can reverse a trophic cascade.

Nutrient subsidies from one ecosystem to another serve a critical link among ecosystems. The transfer of materials across the terrestrial-to-marine boundary is considered to be driven by hydrological connectivity, but animal movement can provide another pathway for nutrient transfers. In two separate studies we assessed the role mammals (bats and kangaroos) play in alleviating nutrient limitation in mangrove forests in Australia. At Lizard Island, we measured tree growth and foliar elemental and isotopic composition of trees growing within and outside a large flying fox roost. In Western Australia, we measured foliar elemental and isotopic composition of trees within two forests frequented by kangaroos that feed in spinifex grasslands and shelter in the shade of the mangroves. We compared those with mangroves from adjacent forests that are not frequented by kangaroos. We show that at both locations, the mangrove forest receives terrestrial nutrient subsidies through animal movement. At Lizard Island dominant mangrove species were significantly enriched in nitrogen within the bat roost, as evidenced by higher foliar N concentrations (by up to 150%), N:P and N:C ratios in trees within the roost compared with trees outside the roost. The isotopic signature of foliar N was significantly enriched in 15N by 1–3‰ within the roost, further suggesting that the source of the N enrichment was the bat roost. Growth rates of mangroves within the roost were nearly six times higher than trees outside the roost. In the arid coast of Western Australia, we show elevated foliar 15N abundance of up to 3‰ in mangroves where kangaroos shelter relative to trees where they do not. Thus, this study presents two examples for mammalian herbivore mediated localized transport of nutrients from terrestrial to marine ecosystems, consequently affecting mangrove tree growth, productivity and forest structure.

Bats are important components in tropical mammal assemblages. Unravelling the mechanisms allowing multiple syntopic bat species to coexist can provide insights into community ecology. However, dietary information on component species of these assemblages is often difficult to obtain. Here we measured stable carbon and nitrogen isotopes in hair samples clipped from the backs of 94 specimens to indirectly examine whether trophic niche differentiation and microhabitat segregation explain the coexistence of 16 bat species at Ankarana, northern Madagascar. The assemblage ranged over 4.4‰ in δ15N and was structured into two trophic levels with phytophagous Pteropodidae as primary consumers (c. 3‰ enriched over plants) and different insectivorous bats as secondary consumers (c. 4‰ enriched over primary consumers). Bat species utilizing different microhabitats formed distinct isotopic clusters (metric analyses of δ13C–δ15N bi-plots), but taxa foraging in the same microhabitat did not show more pronounced trophic differentiation than those occupying different microhabitats. As revealed by multivariate analyses, no discernible feeding competition was found in the local assemblage amongst congeneric species as compared with non-congeners. In contrast to ecological niche theory, but in accordance with studies on New and Old World bat assemblages, competitive interactions appear to be relaxed at Ankarana and not a prevailing structuring force.

Constructing community fruit–frugivore networks has proved challenging in tropical forests to date, particularly in lowland Amazonia, which hosts the most diverse spectrum of frugivorous vertebrates and morphological fruit types worldwide. We assessed data on fruit resource production, frugivore assemblages and corresponding fruit–frugivore networks in two contrasting forest types along the Rio Juruá of western Brazilian Amazonia: seasonally flooded várzea (VZ) and unflooded terra firme forest (TF). Over 2 y we conducted monthly surveys of fruit patches and medium- to large-bodied vertebrate frugivores within three 100-ha plots (two TF, one VZ), supplemented by fruit surveys along 67 5-km transects distributed across two contiguous forest reserves (41 TF, 26 VZ). Observations of trophic interactions were supplemented by semi-structured interviews with experienced hunters and fishermen from 16 local communities. The resultant binary networks contained low proportions of all potential interactions (TF: 25.7%, VZ: 19.4%) between 36 functional groups of frugivores and 152 plant genera and, while we report significant heterogeneity in fruit resource use among broad frugivore guilds within each forest type, recursive partitioning analysis failed to clearly match differences in fruit selection to fruit traits. The annual flood pulse in várzea forests had an overriding influence on the species turnover of both fruit resources and frugivores between the two forest types, with higher-order effects on network structure.

For a total of 17 mo, we investigated the impact of a road constructed through pristine tropical lowland rain forest, on the composition and structure of the resident anuran community in the Ulu Temburong National Park (Brunei Darussalam). One year after road construction, eight new anuran species had immigrated into the impacted area. Encounter surveys were conducted and radio-transmitters attached to a total of 16 frogs to identify habitat use and distribution of the largest (>130 mm SVL) immigrant species (Limnonectes ingeri) in comparison with a similar sized resident congener. A strong preference for roadside habitat was found for the immigrant. However, several incursions of more than 500 m beyond the road into the pristine forest were also recorded. Over 200 stomach content samples as well as stable nitrogen isotope ratios of 76 individuals were used to assess the trophic ecology of the two species. Invertebrate diet composition did not differ. However, the immigrant species ate over 400% more frogs than its resident congener and had a significantly higher δ15N ratio, indicating a higher position in the food web. This suggests that L. ingeri acts as a new top frog predator in the system and indicates that it may have a significant negative impact on the local anuran community. Overall, road construction in the absence of logging has led to the immigration of new frog species with impacts far beyond the linear disturbance of the road itself.

Although what drives the abundance and habitat selection of ungulates is a long-standing question, coherent datasets investigating the influences of rainfall, competition and fire on ungulates are unusual. Over 4 y we carried out extensive monthly road transects in Ithala Game Reserve, South Africa, to determine the demographics and habitat occupancy of the region's prevalent grazer (wildebeest) and mixed-feeder (impala). Habitat occupancy was determined using a GIS-based approach. We obtained 8742 sighting records, encompassing 8400 wildebeest and 10071 impala. Annual rainfall did not significantly correlate with population sizes of either species. Fecundity of wildebeest, but not of impala, showed a significant positive relationship with rainfall specifically over the perinatal period (November–December), whilst no significant relationships were found for either species between fecundity and rainfall over the previous year, 2 y, rut (February–April) or height of the dry season (June–August). Impala unexpectedly favoured browse habitats to grassland year round, probably consequent on competition for grass with wildebeest. Dry-season grass flushes attracted both wildebeest and impala. The study emphasized how rainfall, competition and fire regimes may affect differently grazers compared with mixed-feeders.

The keystone plant resources (KPR) concept describes certain plant species in tropical forests as vital to community stability and diversity because they provide food resources to vertebrate consumers during the season of scarcity. Here, we use an 8-y, continuous record of fruit fall from a 1.44-ha mature forest stand to identify potential KPRs in a lowland western Amazonian rain forest. KPRs were identified based on four criteria: temporal non-redundancy; year-to-year reliability; abundance of reproductive-size individuals and inferred fruit crop size; and the variety of vertebrate consumers utilizing their fruit. Overall, seven species were considered excellent KPRs: two of these belong to the genus Ficus, confirming that this taxon is a KPR as previously suggested. Celtis iguanaea (Cannabaceae) – a canopy liana – has also been previously classified as a KPR; in addition, Pseudomalmea diclina (Annonaceae), Cissus ulmifolia (Vitaceae), Allophylus glabratus (Sapindaceae) and Trichilia elegans (Meliaceae) are newly identified KPRs. Our results confirm that a very small fraction (<5%) of the plant community consistently provides fruit for a broad set of consumers during the period of resource scarcity, which has significant implications for the conservation and management of Amazonian forests.

To what extent tropical forest persisted in the Malay-Thai Peninsula during the Last Glacial Maximum, or contracted southwards with subsequent post-glacial expansion, has long been debated. These competing scenarios might be expected to have left contrasting broad spatial patterns of diversity of forest-dependent taxa. To test for a post-glacial northward spread of forest, we examined latitudinal clines of forest-dependent bat species at 15 forest sites across Peninsular Malaysia. From captures of 3776 insectivorous forest bats, we found that low richness characterized the north of the study area: predicted richness of 9–16 species, compared with 21–23 in the south. Predicted species richness decreased significantly with increasing latitude, but showed no relationship with either seasonality or peninsula width. Analyses of beta-diversity showed that differences between communities were not related to geographical distance, although there was evidence of greater differences in species numbers between the most distant sites. Assemblages were consistently dominated by six cave-roosting species from the families Rhinolophidae and Hipposideridae, while another 16 species were consistently rare. We suggest that these observed patterns are consistent with the hypothesized northward expansion of tropical rain forest since the Last Glacial Maximum, but emphasize that more surveys in the extreme north and south of the peninsula are required to support this assertion.

A community of frugivorous weevils was studied by quantitative rearing of 57 weevil species represented by 10485 individuals from 326 woody plant species in lowland rain forest in Papua New Guinea. Only fruits from 35% of plant species were attacked by weevils. On average, weevils were reared from only 1 in 33 fruits and 1 kg of fruit was attacked by 2.51 individuals. Weevil host specificity was relatively high: 42% of weevil species fed on a single plant genus, 19% on a single plant family and only 16% were reared from more than one family. However, monophagous specialists represented only 23% of all reared individuals. The average 1 kg of fruits was infested by 1.84 individuals of generalist weevils (feeding on allogeneric or allofamilial host species), 0.52 individual of specialists (feeding on a single or several congeneric species), and 0.15 individual of unknown host specificity. Large-seeded fruits with thin mesocarp tended to host specialist species whereas those with thick, fleshy mesocarp were often infested with both specialists and generalists. Weevils tended to avoid small-seeded, fleshy fruits. The low incidence of seed damage (3% of seeds) suggests that weevils are unlikely to play a major role in regulating plant populations via density-dependent mortality processes outlined by the Janzen–Connell hypothesis.

Mass-flowering plant species are often pollinated by social bees that are able to use the abundant resource by recruiting workers from their colonies. In this study we surveyed pollinators on the mass-flowering perennial crop coffee (Coffea arabica) in its native range in Ethiopia. Previous studies in areas where coffee is introduced often find the social honeybee, Apis mellifera, to be the dominant pollinator. In those areas, the bee-species composition visiting coffee varies with a higher bee diversity closer to forest or in less modified habitats. We surveyed pollinators of coffee under different shade-tree structures, by collecting hoverflies and bees landing on coffee flowers in 19 sites in south-west Ethiopia. We found the native honeybee (A. mellifera) to be the dominant visitor of coffee flowers in all sites. Honeybee abundance was not affected by the local shade-tree structure, but was positively affected by the amount of coffee flower resources. Other pollinators were positively affected by complex shade-tree structures. To conclude, the honeybee is clearly the dominant pollinator of coffee in Ethiopia along the whole shade-tree structure gradient. Its high abundance could be a consequence of the provision of traditional bee hives in the landscape, which are colonized by wild swarming honeybees.

Dung beetles are known to perform important ecological functions, such as secondary seed dispersal of vertebrate-defecated seeds. We found that dung beetles also move buried seeds upwards, with positive consequences for seedling establishment. In the Lacandon rain forest of southern Mexico we conducted field experiments to address three questions: (1) What proportions of different-sized seeds buried by dung beetles are exhumed by them? (2) Does upward relocation of seeds caused by dung beetle activity promote seedling establishment? (3) Does recurrent beetle activity increase seedling establishment? Using 4-mm, 8-mm and 12-mm beads as seed mimics, embedded in howler-monkey dung, we found that 2–6% of buried beads were later exhumed by beetles, with smaller beads exhumed more often. In small plots (N = 100) where beetles were allowed to bury dung and seed rain was excluded, seedling establishment was over three times higher compared with plots without beetle activity. In plots (N = 8) where we placed dung on four occasions in 1 mo, seedling establishment was more than twice as high compared with plots with single-time dung placement. We believe that our findings open up interesting research opportunities to help further elucidate this newly discovered ecological function of dung beetles.

Fragmented populations are usually exposed to the negative effects of reduced gene flow, genetic drift and population differentiation. These effects result in the collective loss of genetic variation, thereby reducing the probability of population adaptation to new environmental conditions and increasing the risk of extinction. Forest fragments commonly exhibit suboptimal site conditions, which can result in enhanced clonal reproduction, and a potential reduction in clonal diversity due to increased selfing and inbreeding depression. The clonal diversity, genetic diversity and structure of Heliconia angusta (Heliconiaceae) were assessed using AFLP-markers. We analysed six patches in the continuous forest (Atlantic rain forest, State of Rio de Janeiro) and eight patches (155 leaf samples in total) in five nearby forest fragments (age of oldest fragment: c. 50 y; size range: < 5–100 ha). Clonal diversity (Pd) of patches was slightly, yet significantly, lower in forest fragments compared with continuous forest. Measures of genetic diversity of patches in forest fragments did not differ from those in the continuous forest. A STRUCTURE analysis did not show any clear clustering of patches in the continuous forest and forest fragments. Our results suggest that H. angusta has not yet suffered from the anticipated negative effects of fragmentation.

Nucleation, leading to the formation of tropical forest patches in open areas, has occurred in many parts of the world. This study examined the role of Ficus natalensis as a nucleus and estimated seed dispersers in the nucleation process. Seed rain and post-dispersal fate of Syzygium guineense ssp. afromontanum (a common forest tree species) under eight F. natalensis crowns were compared with those in other woodland microsites. In addition, focal observations of frugivore were conducted at F. natalensis in the woodlands and S. guineense ssp. afromontanum in the forest, when both species were fruiting. Most dispersed seeds (85%) were found under F. natalensis, and the number of dispersed seeds was significantly greater under F. natalensis than at other microsites. Germination (n = 600) and survival percentages (n = 384) were greater under F. natalensis compared with those in treeless open microsites. These results suggest that F. natalensis act as efficient nuclei in nucleation process. Results of focal observations at S. guineense ssp. afromontanum (100 h) showed that Schalow's turaco consumed 63% of fruit and swallowed all pecked fruit. Schalow's turaco also visited fruiting F. natalensis frequently (0.4 h−1) during 80 h of observations, indicating that Schalow's turaco is an important seed disperser during nucleation.

Positive plant interactions have strong effects on plant diversity at several spatial scales, expanding species distribution under stressful conditions. We evaluated the joint effect of climate and grazing on the nurse effect of Croton wagneri, by monitoring several community attributes at two spatial scales: microhabitat and plant community. Two very close locations that only differed in grazing intensity were surveyed in an Ecuadorian dry scrub ecosystem. At each location, two 30 × 30-m plots were established at four altitudinal levels (1500, 2630, 1959 and 2100 m asl) and 40 microsites were surveyed in each plot. Croton wagneri acted as community hubs, increasing species richness and plant cover at both scales. Beneath nurses mean richness and cover values were 3.4 and 21.9%, and in open areas 2.3 and 4.5%, respectively. Magnitude of nurse effect was dependent on climate and grazing conditions. In ungrazed locations, cover increased and diversity reduced with altitude, while grazed locations showed the opposite trend. In ungrazed plots the interactions shifted from positive to negative with altitude, in grazed locations interactions remained positive. We conclude that the nurse effect is a key mechanism regulating community properties not only at microsite but also at the entire community scale.

Amphibians are known to be sensitive to environmental change but their responses at the level of metacommunities to short-term environmental variability are poorly understood. We used field data from two protected areas, La Pera and Nahá (Chiapas, Mexico) to test for variation in metacommunity properties for two consecutive years (2009 and 2010). Amphibians and accompanying environmental data were recorded to a standardized protocol within each landscape, in four or five transects of 50 × 2-m per patch, for 30 and 31 patches, respectively. We found 23 species in La Pera and 30 in Nahá (21 species shared). Metacommunity structure was analysed using reciprocal averaging (RA) ordination by means of metrics for coherence, turnover and boundary clumping, with Spearman rank correlation used to examine relationships with environmental variables. The metacommunity structure varied differentially among the landscapes between years, being classed as quasi-Gleasonian in La Pera in both years, but Clementsian for Nahá in 2009 and Gleasonian for Nahá in 2010. In further illustration of variation between years, in 2009 the principal community gradient (RA axis 1) for La Pera was significantly positively correlated with altitude (r = 0.36), forest disturbance status (r = 0.78), mean canopy cover (r = 0.79) and mean litter depth (r = 0.67), while in 2010 it was correlated with latitude (r = 0.38), mean grass-layer height (r = 0.38), incidence of rainfall prior to sampling (r = 0.35) and presence of temporary ponds (r = 0.45). Our findings support the notion that amphibians respond to short-term environmental changes by individualistic movement within the landscape as well as via population dynamic responses.

The aim of this study was to investigate the spatiotemporal variation in richness, abundance, structure and composition of phyllostomid bats over a successional gradient in a tropical dry forest in south-eastern Brazil. Four successional stages (pasture, early, intermediate and late) were sampled in the northern part of the state of Minas Gerais. Bats were sampled using mist nets at three sites for each of the four successional stages (12 sites in total) during eight periods between 2007 and 2009. A total of 537 individuals were captured (29 recaptured), distributed among four families and 22 species. Bat abundance and richness varied in space, being higher in the late-successional stage, and over time, being significantly lower during the dry season. When compared between guilds, only the abundance of omnivores varied significantly during the sampled months. Our results demonstrate that areas of late-successional stages showed higher bat richness and abundance in comparison with areas undergoing secondary succession. Our results also suggest the use of early-successional areas as flying routes by bats can lead to failure to detect differences in bat composition within successional gradients. We suggest future studies should assimilate landscape-level analyses into their studies to better evaluate the effects of successional gradients on bat assemblages.

Activity patterns of animals are generally influenced by many factors. We hypothesized that the behavioural responses (i.e. activity time-budget allocated to vigilance, feeding and moving) of mountain nyala (Tragelaphus buxtoni) should vary with habitat type, season, group-size and sex-age class. We randomly established a total of 12 permanent walking transects with the aid of a GPS device across three major habitat types used by the mountain nyala (i.e. four transects in each habitat). Following each transect, we conducted focal-animal observations to quantify the time-budget allocated to vigilance, feeding and moving. A total of 119 and 116 focal-animals were assessed in the wet and dry season respectively. Moreover, along each transect, seven habitat variables were collected in systematically laid 109 circular plots each with a 5-m radius (i.e. 31, 41 and 37 plots in the cleared vegetation, plantation and natural forest respectively) in the wet and dry season. We developed behavioural models by correlating the time-budget (i.e. proportion of time vigilance, feeding and moving) of the focal-animals in accordance with habitat variables, group-size and sex-age class. In the wet season, mountain nyala devoted most of their time to vigilance, but they allocated the largest proportion of their time to moving in the dry season. Vigilance differed among the three habitats and was highest in the cleared vegetation during the dry season. Contrary to expectations, adult males were more vigilant than both adult females and sub-adults during the dry season. The behavioural models based on time-budget help to predict how the mountain nyala perceive their environment and trade-off between food acquisition and safety in the wet and dry season. The study also improves our understanding of the adaptive behavioural ecology of the endangered mountain nyala.

Understanding patterns in plant and herbivorous insect diversity across spatial and temporal scales is fundamental to ecology, but comparative multi-taxonomic studies in tropical seasonally dry forests remain scarce. In 36 sites, distributed over three forest age classes (5–10 y, 10–30 y, >100 y) and three seasonal forest types (dry, intermediate, humid), we sampled plants of different stem diameter classes while caterpillars were sampled across vertically distributed forest layers during three seasons over the year. We recorded 299 plant species and 485 caterpillar morphospecies. For large woody plants, species numbers showed a gradually increasing trend with forest age in the intermediate and humid forest types, while the main portion of explained variation in overall species turnover was accounted for by the forest type × forest age interaction (21.3–23.1% of 44.4–48.7%). Ordinations and multivariate pairwise comparisons suggested a faster but also very distinct successional development of species diversity of large plants in the driest compared with humid and intermediate forest types. In contrast, highest species numbers of small plants in the undergrowth was often found in the 5–10 y-old vegetation across forest types, whereas forest type was the major factor in overall species turnover (contributing 24.2% of 48.7% explained variation). Caterpillar species turnover was most correlated to species turnover of small plants; however, variation in caterpillar species diversity appears to be mostly regulated by seasonal cues, and to a lesser extent by patterns of regional turnover and local diversity of undergrowth plant species.