Responses of plants to herbivory are dependent on the type of damage and the ontogenetic stage of the plant. We compared the effects of stem pruning and defoliation on seedlings of Colophospermum mopane, an ecologically important tree species widely distributed in southern Africa. The growth of 160 greenhouse-grown juveniles were measured for 6-mo after germination and then 6-mo after treatments including 50% defoliation, 100% defoliation, 50% stem pruning and controls. Pruning resulted in 30% reductions in total leaf area, height and biomass. Partial defoliation resulted in 30% reductions in total leaf area and plant biomass. However, complete defoliation resulted in a 30% increase in biomass production, a doubling in leaf and lateral branch number, a 45% reduction in leaf size, and no change in total leaf area. Thus, completely defoliated seedlings showed greater performance than those that were only partially defoliated, indicating that C. mopane has become adapted to the chronic and severe defoliation inflicted by Imbrasia belina caterpillars. Comparison of our results with other studies indicates that C. mopane seedlings are less herbivory-tolerant than adults and that pruning has more negative effects than defoliation. Thus, seedling browsers may constrain recruitment in C. mopane, influencing its population dynamics and abundance.

Variation in understorey irradiance is both a cause and consequence of the structure and dynamics of closed-canopy forests, which are also influenced by soil nutrients and water availability. We tested the hypothesis that understorey light regimes differ among four mixed dipterocarp forest types that share the same rainfall, but grow on different soils along an edaphic gradient at one site in Borneo. Based on data from photosynthetically active radiation sensors deployed at 1-m height at 36 locations for 351 sensor-days, we found significant soil-related variation in irradiance. The more productive forest types on clay and fine loam had lower daily photosynthetic photon flux density (PPFD) than those on the more nutrient-depleted and better-drained sandy loam and loam. They also had fewer moderate to high-intensity sunflecks, and a greater proportion of their daily PPFD came from low-intensity light. Understorey irradiance did not, however, monotonically decline with increasing soil resources. Forests on intermediate soils had greater irradiance than those with more and less soil resources, due to steeper slopes. Plant communities arrayed on resource gradients are commonly used to test hypotheses of environmental factors driving their assembly. Our results indicate that consideration of multiple resource dimensions in such tests is critical.

This study examined whether high nutrient concentrations associated with leaf-cutting ant nests influence plant growth and plant water relations in Amazon rain forests. Three nests of Atta cephalotes were selected along with 31 Amaioua guianensis and Protium sp. trees that were grouped into trees near and distant (>10 m) from nests. A 15N leaf-labelling experiment confirmed that trees located near nests accessed nutrients from nests. Trees near nests exhibited higher relative growth rates (based on stem diameter increases) on average compared with trees further away; however this was significant for A. guianensis (near nest 0.224 y−1 and far from nest 0.036 y−1) but not so for Protium sp. (0.146 y−1 and 0.114 y−1 respectively). Water relations were similarly species-specific; for A. guianensis, near-nest individuals showed significantly higher sap flow rates (16 vs. 5 cm h−1), higher predawn/midday water potentials (−0.66 vs. −0.98 MPa) and lower foliar δ13C than trees further away indicating greater water uptake in proximity to the nests while the Protium sp. showed no significant difference except for carbon isotopes. This study thus shows that plant response to high nutrient concentrations in an oligotrophic ecosystem varies with species. Lower seedling abundance and species richness on nests as compared with further away suggests that while adult plants access subterranean nutrient pools, the nest surfaces themselves do not encourage plant establishment and growth.

Little is known about the relationship involving arbuscular mycorrhizal (AM) fungi and functional groups of plants that characterize different phases of tropical succession. We appraised the AM infection intensity of root cortex and spore density in the soil in sites over tropical successional gradients (grassland, secondary forest and mature forest) for several years in Araucaria, Atlantic and Pantanal ecosystems in Brazil. The intensity of AM infection decreased with advancing successional stages in all ecosystems and it was around 60–80% in early stages of succession, 37–56% in secondary forests and 19–29% in mature forests. Similarly, the AM spore number also decreased with advancing succession and was the highest in early stages (73–123 g−1), intermediate in secondary forests (32–54 g−1) and lowest in the mature forests (10–23 g−1). To verify whether such reductions influenced the potential of AM inoculum in soil, seedlings of Heliocarpus popayanensis (Malvaceae) were grown as test plants in soils obtained from five grasslands, five young secondary forests, and five mature forests in the Atlantic ecosystem. The soil inocula from the grasslands and secondary forests were 7.6 and 5.7 times more effective in stimulating seedling growth than inocula from the mature forests, respectively. Our results show that plant species in grasslands and young secondary forests stimulate the multiplication of AM fungi, leading to a higher potential of the AM inoculum. In later-successional stages, plant investment in AM fungi decreases and the potential of the AM inoculum is also reduced.

Habitat heterogeneity is an important ecological determinant of species richness. We evaluated the diversity within bird feeding guilds as related to habitat heterogeneity and land-use cover in a human-modified tropical landscape. To quantify this process, fine-scale bird census and habitat heterogeneity data were collected for a bird community in a 22.5-km2 fragmented landscape in southern Mexico. Land-use cover data derived from field surveys were used to calculate habitat heterogeneity index values and the extent of each land-use cover type in 239 grid cells of 300 × 300 m. Bird diversity values were obtained based on 1195 point-counts in these cells. Product-moment correlations and linear regression analyses were used to evaluate the relationship between bird-guild diversity values and habitat heterogeneity. A total of 109 resident bird species grouped in six feeding guilds were recorded: insectivores (42%), frugivores (21%), granivores (17%), nectarivores (9%), omnivores (8%) and carnivores (3%). Diversity values for the entire bird community were significantly positively related to habitat heterogeneity, but feeding guilds showed contrasting responses to habitat heterogeneity and the amount of land-use cover: insectivores and frugivores were more diverse and abundant in secondary forests than in any other land-cover. Our findings illustrate the importance of small landscape fragments as potential key refuges for the most diverse and specialized feeding guilds, such as granivores and insectivores.

Natural tree canopy gaps allow sunlight to penetrate to the forest floor, a major environmental component and resource for many tropical rain-forest species. We compare here how butterflies use sunny areas created by the natural gaps in canopies in comparison with adjacent closed-canopy areas. We chose butterflies as our focal organisms as they are taxonomically tractable and mobile, yet habitat sensitive. Previous studies have shown that butterfly diversity in tropical forests responds to varying degrees of canopy openness. Here we assess butterfly behavioural responses to gaps and equivalent sized closed-canopy patches. Butterfly occupancy time and behaviour were simultaneously observed 61 times in gaps and 61 times in equivalent sized closed-canopy patches across four sites in a tropical rain forest in northern Borneo. Out of the 20 most frequently recorded species, 12 were more frequently recorded or spent more time in gaps, four occurred more frequently in closed-canopy areas, and four showed no significant differences. Overall agonistic, basking, patrolling and resting were more common in gaps compared with the closed canopy. Many butterfly species have complex behavioural requirements for both gaps and closed canopies, with some species using these different areas for different behaviours. Each butterfly species had particular habitat requirements, and needed both canopy gaps and closed canopy areas for ecological and behavioural reasons, emphasizing the need for natural light heterogeneity within these systems.

A number of studies have pointed out that abiotic factors and recolonization dynamics appear to be more important than biotic interactions in structuring river–fish assemblages. In this paper, we studied the fish assemblages in 27 floodplain lakes, with perennial connection to the river, in the middle section of the Magdalena River (Colombia), to examine spatial pattern in freshwater fish diversity in relation to some environmental parameters. Our objective was to examine relationships between floodplain-lake fish communities and environmental variables associated with lake morphology, water chemistry and river–floodplain connectivity in a large river–floodplain ecosystem. During the study, a total of 18 237 fish were caught from 50 species (regional richness; 17 were migrants and 33 residents). In the present study, the most diverse order was Characiformes with 20 species, followed by Siluriformes, with 19 species. Characidae and Loricaridae were the richest families. The range of species richness (local richness) varied between five and 39 species. Similarity of local assemblages (using the presence–absence data) depends on the distance between lakes. A positive relationship was observed between the Ln of the total abundance of each species and the number of lakes where they were found. Out of all the environmental parameters taken in the lakes, only the size (Log Area) and relative perimeter length are significantly related to local assemblage species richness. It has not been possible to demonstrate that the connectivity (distance) from lakes to the main river can be considered a predictor of the local richness.

The competition for resources increases size inequality in trees, particularly under low abiotic stress. Because mangrove communities are subject to site-specific salinity (and therefore abiotic stress) gradients, these habitats should differ in height–diameter allometry and size inequality. The size inequality (by the Gini Coefficient, G) and maximum potential height (Hmax from a height–diameter asymptotic model) were determined within the mangrove forest of a coastal lagoon in Veracruz, Mexico in 20 0.25-ha plots, 10 in interdistributary basins (IBs, lower salinity) having Avicennia germinans, Laguncularia racemosa and Rhizophora mangle and 10 in mudflats (MFs, higher salinity) dominated by A. germinans. Size inequality was significantly higher in IBs (G = 0.59 ± 0.02 vs. 0.39 ± 0.03). Due to their significant intercorrelation G, total basal area and density were synthesized in one PCA axis accounting for 67% of total variance and inversely correlated with salinity (R = −0.65, P = 0.003). The height–diameter scaling model reached a stable asymptote (Hmax range: 16–21 m; coefficient of variation CV: 7.7) in IBs, suggesting that trees can still increase their diameter after achieving maximum height. In MFs, no stable asymptote was reached (Hmax range: 11–26 m; CV: 32.5), suggesting a lower growth rate of diameter in the MF trees when compared with IB trees.

Tropical forest insects are vertically stratified between the canopy and understorey. Using 60 traps set at two heights above the forest floor (30 at 15 cm and 30 at 1 m) we compared abundances in capture height, persistence in traps, and sex of three co-occurring understorey butterflies (Cithaerias pireta, Dulcedo polita and Pierella helvina) in Costa Rica. We captured, marked and released 283 individual butterflies (65 C. pireta, 79 D. polita, 139 P. helvina) and showed all three species were captured more often in low traps, and P. helvina was captured only in low traps. The probability of remaining in traps for 24 h did not differ significantly for D. polita and P. helvina, but was significantly lower for C. pireta. The odds of trapping either sex did not differ significantly for P. helvina and C. pireta, but they were significantly lower for D. polita males. We experimentally demonstrate that these co-occurring species fly and feed just above the forest floor, but differ with respect to their persistence in traps and attraction to traps by sex. Our study implies that closely related species can exhibit behavioural differences that may influence population abundance estimates in multi-species studies.

The degree to which geographical location rather than environment affects the maintenance of high tropical forest beta diversity on altitudinal gradients is not well understood. Forest composition and its relationship to climate, soil, altitude and geographical distance were determined across an 1114-km2 landscape in south Pacific Costa Rica spanning an altitudinal gradient (0–1500 m asl). In 37 0.25-ha plots, > 200 species of dicot trees (≥ 30 cm dbh) and canopy palms (≥ 10 cm dbh) were found. Ordination analysis showed strong species composition patterns related to altitude; plot coordinates on the main axis showed negative correlations to the abundance of lowland-forest species Iriartea deltoidea (r = −0.54) and Brosimum utile (r = −0.65), and positive correlations to higher-altitude species Alchornea glandulosa (r = 0.63), Quercus sp. (r = 0.50) and Ocotea sp. 2 (r = 0.48). Mantel correlations, correlograms and variation partitioning analysis of relationships between floristic composition and spatial and environmental factors indicated that spatial location of the plots – potentially, dispersal limitation – was the single most important (R2adj = 0.149) driver of beta diversity, but that environmental heterogeneity also plays an important role. In particular, palm species turnover was strongly related to soil chemical properties. The effects of dispersal limitation on floristic assembly could determine the future distribution of plant communities as a result of climate change.

Neotropical fruit-eating bats play a crucial role in forest regeneration by dispersing seeds of pioneer plants from forests into deforested areas. However, later in succession bats may carry seeds in both directions. We used an isotopic approach to reveal the direction of seed transfer mediated by three co-existing short-tailed fruit bats (Carollia castanea, C. sowelli and C. perspicillata) between a forest and an adjacent mid-successional site (>15 y since deforestation); two habitats where individuals of the genus Piper differed in stable carbon isotope ratios by ~2.5‰. In a feeding experiment, we confirmed that δ13C of seeds is not altered by digestive processes. We then collected seeds defecated by bats of the genus Carollia and found that δ13C of these seeds is higher than those of Piper individuals growing in the forest, irrespective of whether bats were captured in or outside the forest. We conclude that bats of the genus Carollia were more likely to carry seeds from successional areas into the forest than in the opposite direction.

We examined stand dynamics and biomass along an altitudinal gradient in a tropical montane forest (TMF) in the disturbance-prone Cordillera Central, Dominican Republic. We tested the general hypothesis that chronic disturbance by fire, wind, floods and landslides results in a landscape of relatively low above-ground biomass with high rates of mortality, recruitment and growth as compared with other TMFs. We also expected above-ground biomass to decrease with altitude in part due to declines in growth and increased biomass losses from mortality with increasing altitude. We resurveyed 75 0.1-ha plots distributed across the altitudinal gradient (1100–3100 m asl) 8 y after they were established. Our observations provided mixed evidence on these hypotheses. Turnover rates were high (> 2% y−1) and significantly greater on windward slopes. Above-ground biomass (mean = 306 Mg ha−1, 95% CI = 193–456 Mg ha−1) was highly variable but comparable to other TMFs. Altitudinal patterns of declining biomass and above-ground growth matched observations for other TMFs, whereas mortality and recruitment exhibited no altitudinal trends. More quantitative studies in a variety of TMF settings are needed to better understand how natural disturbance, complex environmental gradients and species dynamics interact to regulate carbon storage, sequestration and turnover across altitudinal gradients in TMFs.

The responses of plants to shade and foliar herbivory jointly affect growth rates and community assembly. We grew 600 seedlings of ten species of the economically important Dipterocarpaceae in experimental gradients of shading (0.3–47.0% of full sunlight) and defoliation (0, 25%, 50% or 75% of leaf area removed). We assessed stem diameters initially, after 2 and 4 mo, and calculated relative growth rates (RGR) with a linear model. Shading interacted with defoliation, reducing RGR by 21.6% in shaded conditions and 8.9% in well-lit conditions. We tested three hypotheses for interspecific trade-offs in growth responses to shading and defoliation. They could be positively related, because both reduce a plant's access to carbon, or inversely related because of trade-offs between herbivore resistance and tolerance. We observed, however, that species varied in their response to shading, but not defoliation, precluding an interspecific trade-off and suggesting that plants tolerate shade and herbivory with differing strategies. Shading most strongly reduced the growth of species with less-dense wood and larger seeds. The strong and variable growth responses to shade, contrasted with the weak and uniform responses to defoliation, suggest that variation in light availability more strongly affects the growth of tropical tree seedlings, and thus community assembly, than does variation in herbivory.

Fires can significantly affect the structure, floristic composition and biomass content of tropical forests, which are not adapted to this disturbance. To assess the impact of understorey fires on above-ground biomass, this study was conducted in the northern Brazilian Amazon (Roraima state), where uncontrolled forest fires are recurrent. Fifty plots (0.25 ha each) distributed across five fire disturbance classes were inventoried. Losses in biomass stocks were significant (−57% and −63%) for forests that suffered from recurrent fires and progressively occurred until some point between 3 and 7 y after the last fire, as deduced from previous studies. Twelve years after a fire event, biomass stocks were reconstituted, although differences in the floristic composition were associated with greater fire severity, particularly driven by the dominance of the Cecropia spp. In thrice-burned forests, live biomass reached levels of secondary forests of the same region. For large trees (> 50 cm dbh), the reduction (−54%) in biomass was significant in thrice-burned areas. These findings highlight the threat that fires represent for tropical rain forests and emphasise the need for long-term surveys in order to distinguish between the effects of fire severity, fire frequency and time-since-fire artefact.

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.

Spatial association patterns reflect underlying mechanisms of coexistence, community structure of plant species in tropical forests. We hypothesized that if spatial associations between two species shift toward segregation patterns during the course of growth, deterministic mechanisms, such as interspecific competition and habitat differentiation, would prevail, whereas if no directed change in spatial associations between two species is observed and, consequently, the initial association pattern is retained through growth, the two species would experience weak interspecific competition and show no habitat differentiation. To assess the underlying mechanisms operating between confamilial species, we analysed spatial associations among 11 dipterocarp species in terms of three growth stages distinguished on the basis of dbh in the Pasoh 50-ha plot in Peninsular Malaysia. We analysed the spatial associations of all possible combinations among identical stages (165 pairs) and among different stages (330 pairs) for each pair of 11 species, except between identical species. Our previous study revealed that the 11 species could be characterized into two classes: seven fast-growing species exhibited high growth and mortality rates, spatial aggregation on a small scale, and positive habitat associations, while four slow-growing species exhibited low growth and mortality rates, spatial aggregation on a large scale, and no habitat associations except one. Spatial segregation was observed between fast-growing species (32 pairs, 17%) and between species of different classes (35 pairs, 14%), but not between slow-growing species. Throughout the growth stages, positive associations with other species were maintained for slow-growing species versus fast-growing species. In contrast, changes in initial associations toward segregation were observed more in fast-growing species. These results indicated that interspecific competition or habitat differentiation dominated for fast-growing species, while non-directed random processes dominated for slow-growing species.

Spatial distribution of resources is known to govern animal distribution and behaviour. However, few empirical studies have formally evaluated this relationship. Unlike previous studies in which a patch or gap of floral resources is defined a priori by the observer at a subjective perception scale, we used the Spatial Analysis by Distance IndicEs (SADIE) to assess the location, length and spatial co-occurrence of patches and gaps of Palicourea padifolia inflorescences and hummingbird activity (feeding, perching, vocalizing, flying past and agonistic behaviour) in a tropical montane cloud forest of central Veracruz, Mexico. Along a 1010-m transect, both resource and hummingbird activity had a distribution approximately 200% more aggregated than expected by chance, at a scale of tens to hundreds of metres in length. In addition, aggregation patterns of resource and overall and agonistic hummingbird activity were found to be positively associated in 2009 but negatively in 2010. Campylopterus curvipennis and Amazilia cyanocephala were the most frequent species involved in vocal and agonistic activity. The difference observed between the two years may be due to changes in the composition and dominance of hummingbird species with different foraging strategies. In both years, hummingbird overall activity was positively correlated to size of resource patches.

Abandoned plantations of coffee, tea and other commercial crops offer opportunities for understanding ecological processes in modified forest ecosystems. Unlike tree plantations tea is maintained as a shrub with a continuous dense short canopy that precludes large-frugivore activity thereby limiting dispersal of forest species to such areas. In this study we determine how location and density of Grevillea robusta a shade tree in tea plantations and proximity of plantations to forests influences seed arrival from forests into the plantations. We also estimate the importance of dispersal modes in the colonization processes. We laid 10 × 10-m plots at three distance intervals from the forest edge in three different plantation types with varying shade tree densities. Within the plots we laid four 1× 1-m subplots at the corners of the plot. We estimated species richness, abundance and categorized the seeds into dispersal modes in these plots. Grevillea robusta increased species richness of seeds by three times and abundance of seeds by 3–30 times compared with plantations without them. Higher density of G. robusta increased seed input changed species composition and altered species dominance in the plantations. Distance to forests influenced seed arrival in plantations without G. robusta trees and plots 95 m from the forest did not have any seeds in them. No such effect was seen in plantations with G. robusta trees. Seeds dispersed by birds or a combination of birds and mammals contribute 30% of the seeds reaching the plantations with G. robusta and this was not influenced by distance from the forest. In plantations without G. robusta bird dispersal is restricted to 25 m from the forest edge. In general density of shade trees has a strong influence on seed arrival which can negate the forest proximity effect and enhance natural forest colonization.

We analysed the effect of a hurricane on a plant–ant network and on vegetation cover. Plant cover was sampled using linear sampling in several vegetation types: deciduous forest, a dry forest, sand dune pioneers, sand dune scrub, ecotone of freshwater marsh, deciduous forest and dune scrub, and mangrove forest. We sampled ant–plant interactions and vegetation cover before and after Hurricane Karl hitting (September 2010) the central coast of the state of Veracruz, Mexico. The pre-hurricane network consisted of 16 plant and 25 ant species in 52 associations. The post-hurricane network consisted of 17 plant and 20 ant species in 56 associations. We found a significant decrease in the total linear cover of EFN-bearing plants between October 2009 (646 m, no hurricane effect) and October 2010 (393 m, after hurricane Karl) (total sample length 2025 m). Both networks were significantly nested (0.999 and 0.973, P < 0.001), suggesting that network topology remained similar. Our results show changes in several network characteristics and species proportions. The number of plant species that contributed to nestedness vs. idiosyncratic species did not differ significantly in the pre-hurricane network, while the number of plant species that contributed to nestedness vs. idiosyncratic species did differ significantly in the post-hurricane network. The number of ant species that contributed to nestedness vs. idiosyncratic species differed significantly in the pre-hurricane network, and also in the post-hurricane network. Differences in nestedness contributions of species before and after the hurricane reflect an alteration from a generalized, highly nested, more stable pre-disturbance network, to a more low-degree or specialized network (i.e. fewer interactions among generalist species, those species with the most associations). The maintenance of important core components of the network after a huge disturbance, suggests a short-term resilience typical of mutualistic networks.

Research has increasingly established that mesoherbivores influence the regeneration of woody plants. However the relationship between mesoherbivore density and degree of impact, and the spatial component of this impact, has not been well established. Using a novel sampling design, we assessed in iMfolozi Park, South Africa, the impact of impala (Aepyceros melampus) across the full complement of woody species within the home range, evaluating its spatial component and relationship to impala density. We used four GPS collars, in separate breeding herds, and a GIS to detect zones of different density of impala in the landscape, thus defining a fine-grain browsing gradient. We assessed impact on woody recruits (≤ 0.5 m height) across this gradient by means of 1600 random 1 × 1-m quadrats. Densities of woody seedlings, and mean percentage of remaining canopy, were significantly less in areas of high impala density versus low-density areas. There was a significant correlation between increasing impala density and decreasing density of favoured woody recruits. We propose a hypothesis of impala-induced patch dynamics. It seems likely that the ubiquitous impala may create and sustain a shifting mosaic of patches, and thus function as a key determinant of landscape heterogeneity.