Addressing what affects predation among caterpillars, we conducted an experiment in a Bornean rain forest on 212 clay models of Tortricidae caterpillars (the herbivore) and 53 trees of Kopsia pauciflora (the host), located either in the open or under closed canopies. We predicted that the frequency of predatory attacks towards caterpillars increases (1) in canopy gaps and (2) on leaves damaged by herbivory, but (3) decreases among caterpillars that wrap their body in leaves. Each plant with caterpillar models was consecutively allocated to one of four treatments: caterpillars artificially rolled in leaves vs caterpillars on unrolled leaves, and caterpillars on artificially damaged vs undamaged leaves. Each time, caterpillar models were placed on the plants for 48 h, and then replaced with new models that were subjected to a new treatment. On average, our caterpillar models had a 21% chance of being attacked per 24 h. More attacks were performed by insects (81.6%) than birds. The attack frequency did not depend on the canopy cover or on leaf damage, but decreased among models rolled in leaves. This pattern was mainly attributable to insect attacks, which were also more frequent than bird attacks. Overall, the Tortricidae caterpillars seem to suffer comparable predation rates in the open and under closed canopies irrespective of the herbivory damage of leaves, but their leaf-rolling behaviour might reduce predation.

Leaf-cutting ants are dominant herbivores in Neotropical rain forests, and their colony densities increase in disturbed habitats such as forest edges. However, while it is well-established that leaf-cutting ants profit from changes to the food-plant community, the phylogenetic dimension of this ant–plant interaction remains poorly understood in fragmented forests. We studied diet composition of Atta cephalotes in the edge and interior of Atlantic forest in north-east Brazil (8°30′S, 35°50′W). We applied phylogenetic signal analysis to investigate the diet across plant lineages and performed phylogenetic generalized linear models to analyse the diet in both habitats. We found a phylogenetic signal in diet and in leaf mechanical resistance, which means that A. cephalotes selects closely related food plants with less resistant leaves. Most preferred species belong to Malpighiales, Rubiaceae and Melastomataceae. We also found that irrespective of phylogeny, ants select food plants with less resistant leaves, both in edge and interior. However, ants choose more abundant plants only in edges. High abundance of optimal diet facilitates foraging in forest edges and explains why colony densities increase in disturbed habitats. Finally, by favouring or disfavouring specific clades, leaf-cutting ants contribute to changes in the phylogenetic structure of tropical rain forests, e.g. phylogenetic impoverishment.

We monitored native forest regeneration over 11 y in a eucalyptus plantation and compared it with the neighbouring primary forest. For the plantation forest, we hypothesized that species richness, density, basal area and densities of old-growth species would increase over time, and compared to the primary forest, plantation forest would have higher species richness and density, but lower densities of old-growth species. In 2016, we repeated the protocol of a study that sampled the plantation forest in 2005, with thirty 10 × 10-m plots and enumerating trees (≥10 cm diameter), saplings (>1 to <10 cm diameter) and seedlings (<1 cm diameter). In the plantation forest, for trees, the species richness, density of gap, bird-dispersed and mammal-dispersed species increased by 67%, 156%, 116% and 238% respectively; whereas for saplings, density of gap, bird-dispersed and small-seeded species declined by 45.2%, 51% and 18.2% respectively over time; and seedling densities did not change across functional groups. Stand basal area increased by 80.1% in the plantation forest. The primary forest had 446% greater density of closed-canopy trees compared with plantation forest. Contrary to our prediction, the plantation forest did not accumulate significant densities of old-growth species over time, probably due to demographic filters that prevent them from attaining maturity.

We compared exotic pasture grass cover near the edges of 20–25-y-old secondary forests (N = 8) with those of mature forests (N = 8), bordering actively grazed pastures on the Pacific Coast of Ecuador. We estimated grass cover in 224 1 × 3-m plots along transects that ran from the pasture edge into forest interiors (11–44 m). Using a spline regression, we divided the transects into three segments: exterior (in the pasture), edge and interior (in the forest). With a stepwise regression, we tested the effect of transect section, forest type and distance from edge on grass cover. Forest type, distance from edge, interior transect section and the combined effect of distance from edge in both the interior and exterior sections explained variation in grass cover. Grass abundance was higher and penetrated further into secondary than mature forests. Edge proximity and differences in forest canopy openness likely favours recruitment and persistence of pasture grasses.

The relative importance of different bottom-up-mediated effects in shaping insect communities in tropical secondary forests are poorly understood. Here, we explore the roles of vegetation structure, forest age, local topography (valley vs. hill top) and soil variables in predicting fruit-feeding butterfly and tree community composition, and tree community composition in predicting fruit-feeding butterfly community composition, in different-aged naturally regenerating and primary forests of Kibale National Park, Uganda. We also examine which variables are best predictors of fruit-feeding butterfly species richness or diversity. Butterflies (88 species) were sampled with a banana-baited trap and trees (98 taxa) with a 40 × 20-m sampling plot at 80 sampling sites. The environmental variables explained 31% of the variation in the tree community composition, the best predictors being local topography, forest age and cover of Acanthus pubescens (a shrub possibly arresting succession). The fruit-feeding butterfly community composition was better predicted by tree community composition (explaining 10% of the variation) rather than vegetation structure, local topography or soil factors. Environmental variables and tree species richness (or diversity) were poor predictors of butterfly species richness (or diversity). Our results emphasize the importance of tree community to recovery of herbivorous insect communities in tropical secondary forests.

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.

Secondary forests occupy a growing portion of the tropical landscape mosaic due to regeneration on abandoned pastures and other disturbed sites (Asner et al. 2009). Tropical secondary forests and degraded old-growth forests now account for more than half of the world's tropical forests (Chazdon 2003), and provide critical ecosystem services (Brown & Lugo 1990, Guariguata & Ostertag 2001).

Little is known about how land-use changes affect interspecific interactions such as fruit–frugivore mutualisms. Forest age could affect both fruit sugar concentrations via differences in light availability or disperser abundance, and fruit removal rates via differences in bird and plant community composition. We examined how these two factors are affected by forest age in a Costa Rican rain forest. We compared seven young-secondary forest species, seven old-growth forest species, and Miconia nervosa growing in both forests. We measured sugar concentrations in fruits and manipulated the location of paired fruiting branches, measuring subsequent fruit removal. Sugar concentration means were on average 2.1 percentage points higher in young-secondary forest species than in old-growth forest species, but did not differ among Miconia nervosa fruits from the two forests. Fruit removal rates were higher in young-secondary forest for 86% of young-secondary forest species, 71% of old-growth forest species, and on average for both young-secondary and old-growth forest Miconia nervosa individuals. Higher sugar concentrations in young-secondary forest plants could reflect stronger competition for dispersers, while experimental fruit removal results suggests the opposite patterns of competition; fruits are more likely to be removed by dispersers in young-secondary forest independent of fruit nutrient concentration.

We mapped the occurrence of canopy gaps periodically between 1978 and 1990 in a 1.5 ha study plot within a 70-year-old (in 1978) Neotropical forest on Barro Colorado Island (BCI), Republic of Panama. The total area of the forest under canopy gaps in the plot averaged 4.3% (3.1% to 5.7%, 95% CI). There was high year-to-year variability in the rate of new gap formation. On the basis of repeated observations for four yearly intervals, the annual rate of new gap formation ranged from 0.45% y−1 to 6.5% y−1. Most gaps were small. The mean size of individual gaps originally was 79 m2 (range: 8-604 m2). However, large gaps (≥150 m2) occurred more frequently than expected for a secondary forest on BCI. Gaps closed rapidly the first year after formation but the rate of closure slowed thereafter. Despite the absence of any obvious environmental gradients, gaps were spatially clustered. Even in this relatively small plot, there seemed to be distinct gap-prone and gap-free areas.

The soil of remnant primary rainforest on granite in Singapore is very acidic (pH mostly 3.5–4.2 at 0–10 cm) and has unusually low total concentrations of nitrogen and phosphorus, but the mean concentrations of N and P in the living leaves and freshly fallen leaves of the more shade-tolerant species are within the ranges found for other lowland forests on infertile oxisols and ultisols. The concentration of Ca in freshly fallen leaves is very low. The soil under secondary forest (bselukar) on sandstone dominated by Adinandra dumosa (Theaceae) was degraded during use for agriculture. It has the same pH range but even lower values of total N and P. The mean concentrations of N, P, K, Ca and Mg in the living leaves are slightly higher than in leaves of the more shade-tolerant trees of primary forest, but lower than in the leaves of the species which require canopy gaps for establishment or early onward growth. The few species sampled in both primary forest and belukar show no consistent trend in foliar concentrations. The degraded soil has selected species with inherently lower foliar concentrations.

Bioassays of nutrient limitation were carried out for Melastoma malabathricum growing on soil from primary lowland dipterocarp rainforest in Singapore, and for Dillenia suffruticosa on soil from adinandra belukar, a nutrient-poor secondary forest type dominated by Adinandra dumosa. Three questions were addressed. 1. What is the nutrient most limiting to growth in primary forest? 2. What is the nature of nutrient limitation under conditions of adequate P supply? 3. Is there a qualitative difference in the nature of nutrient limitation under primary forest and adinandra belukar? Results showed that there was a strong limitation by P availability in both primary forest and adinandra belukar under the experimental conditions used. Once plants had an adequate P supply, all other nutrients became limiting to growth in primary forest soil. These findings are interpreted as support for the hypothesis that P availability would limit the productivity of moist tropical forests in general in the absence of mycorrhizas; tentative conclusions are drawn on the assumption that most woody tropical plants are mycorrhizal. It is argued that limitation by major cations may be common on old, highly leached tropical rainforest soils.

Hong Kong is on the northern margin of the Asian tropics. The original forest cover was cleared centuries ago but secondary forest has developed since 1945 at many sites protected from fire and cutting. There are also older forest patches maintained behind villages for reasons of ‘feng shui’, the Chinese system of geomancy. All plants >2 cm dbh were identified and measured in forty-four 400-m2 plots. Detrended correspondence analysis showed a floristic continuum, with the montane sites (>500 m) most distinct and some overlap between lowland post-1945 secondary forest and the feng shui woods. The 30–40 year-old secondary forest is dominated by Persea spp. Montane forest is similar but lacks several common lowland taxa of tropical genera and includes more subtropical taxa. The feng shui woods have the most complex structure and contain some tree species not found in other forest types. Their origin and history is obscure but we suggest that both planting and selective harvesting have had a role in their current species composition

Root biomass, root lengths, and mycorrhizal associations were compared in a series of primary and Acacia-dominated secondary rainforest stands on nutrient-poor, red podzolic soils developed from low grade Palaeozoic metasediments. Five soil cores to 200 mm depth were collected at random locations from each of 20 sites. Ten of these sites were in 20–25 m high closed secondary forest (30–40 y old) dominated by Acacia aulacocarpa and ten sites were located in primary, selectively-logged, rainforest (28–32 m tall). Arbuscular mycorrhizas were the only form of association found in the primary forest sites. Ectomycorrhizas dominated the secondary forest sites although arbuscular mycorrhizas were also present. The primary forest sites had significantly higher root biomass (34.4 ± 17.8 t ha-1) and root length (33,400 ± 3,200 km ha-1) than the secondary forests (11.6 ± 4.6 t ha-1 and 25,200 ± 4,800 km ha-1 respectively), and this was interpreted as a reflection of the greater allocation of biomass to roots necessary to support the greater above ground biomass. The specific root length in the secondary forest (340 ± 119 cm g-1) was twice that of the primary forest (154 ± 65 cm g-1) indicating that the trees in the secondary forests achieved a degree of soil exploration which was comparable to that in the primary forest with less than half the biomass allocation to roots. The dominance of ectomycorrhizas in the secondary forest was associated with the prevalence of Acacia aulacocarpa, and the results cannot be extended to other secondary forests in the region. The implications that the dominant ectomycorrhizal associations have for the patterns of successional development and the patterns of species colonization in these Acaria-dominated secondary forests are discussed.

Seed dispersal is the first stage of colonization, and potentially affects recruitment. This process deserves more attention in tropical montane cloud forests (TMCF), since secondary succession is common owing to episodic disturbances. We studied annual seed rain in 10 nearby forest stands, ≈7 to ≈100 y following shifting agriculture, and one primary forest stand in southern Mexico to test the hypothesis that seed rain is limited at the scale of neighbouring fragments and that such limitation differs among species with different dispersal modes and successional origin. Annual seed rain was heterogeneous among forest fragments probably due to the prevalence of local seed dispersal, differences in stand age and the proportion of zoochory, and may help explain the patchy distribution of species observed in TMCF. Seed rain abundance and species diversity per unit trap area increased with the age of the stand. Biotically dispersed seeds increased towards older stands relative to abiotically dispersed seeds. Late-successional seeds were rarer in early successional stands than pioneer seeds in late-successional stands, suggesting that long-distance dispersal is generally more common for pioneer plants. Seed dispersal appears to constrain forest regeneration and to influence fragment species composition as a function of the distance from the source forests.

Litterfall quantity and quality may respond to alterations in resource availability expected with ongoing land-use and climate changes. Here, we quantify the effects of altered resource availability on non-woody litterfall quantity and quality (nitrogen and phosphorus concentrations) in eastern Amazonian forest regrowth (Brazil) through two multi-year experimental manipulations: (1) daily irrigation (5 mm d−1) during the dry season; and (2) fortnightly litter removal. Consistent with other tropical forest data litterfall exhibited seasonal patterns, increasing with the onset of the dry season and declining with the onset of the rainy season. Irrigation did not affect litterfall mass and had little impact on nitrogen (N) or phosphorus (P) concentrations and return, except for decreasing litter P concentration at the end of two irrigation periods. Litter removal did not alter litterfall mass or P concentration, but progressively reduced litterfall N during the course of the experiment. Overall, these results suggest significant resistance to altered resource availability within the bounds of our experimental treatments; our findings may help to constrain carbon and nutrient cycling predictions for tropical forests in response to land-use and climate changes.

Secondary forests account for 40% of all tropical forests yet little is known regarding their suitability as habitat for diurnal large mammals and game birds. This is especially so for second-growth that develops on large areas of degraded land. We address this by investigating assemblages of large-bodied birds and mammals in extensive patches of secondary forest in the Jarí region of the north-eastern Brazilian Amazon, comparing species richness and abundance against that of adjacent undisturbed primary forests. We conducted 184 km of line-transect censuses over a period of 3 mo, and found that although primary and secondary forests held a similar abundance of large vertebrates, the species composition was very different. Secondary forests supported a high abundance of ungulate browsers (0.85 vs 0.44 indiv. per 10 km) and smaller-bodied primates (15.6 vs 4.6 indiv. per 10 km) compared with primary forests. However, large prehensile-tailed primates were absent (black spider monkey Ateles paniscus) or at very low abundance (Guyanan red howler monkey Alouatta macconelli) in secondary forest. The abundance of large frugivorous/granivorous birds was also low in secondary forests compared with primary forests (22.6 vs 37.1 individuals per 10 km, respectively). Faunal assemblages appear to reflect food resource availability. Concurrent vegetation surveys indicated that secondary forests had high levels of terrestrial and understorey browse. Fruit production was largely restricted to pioneer trees such as Bellucia and Inga spp. Although these regenerating forests were an important habitat for large mammals and birds, they were limited in terms of faunal richness, particularly dispersers of large-seeded plants.

The species richness, number of plants, biomass, and species composition of holo- and hemi-epiphytes were recorded in fifty-six 0.04-ha plots, distributed over forest fallows of 2–30 y old and mature forests in lowland Amazonia (Amacayacu National Park, Colombia). A total of 9190 epiphytic plants representing 162 species were recorded on 4277 phorophytes. Seventy species were classified as holo-epiphyte and 85 as hemi-epiphyte. Aroids were most diverse (58 species) and represented 76% of the total recorded biomass. Anemochory was more dominant among holo-epiphytes and zoochory among hemi-epiphytes. The species richness, density and biomass of both holo- and hemi-epiphytes increased significantly from young fallows to old fallows and mature forests. Hemi-epiphytes had greater density and biomass than holo-epiphytes. In canonical ordination, forest age did not relate to the species composition of holo-epiphytes. However, for hemi-epiphytes, the age effect was significant, suggesting that species turnover takes place in the ageing fallows.

The lowland rain-forest ecosystem in Sundaland (Borneo, Sumatra, Java, the Malay Peninsula south of 10°N, and associated islands) has been recognized as a biodiversity hotspot (Myers et al. 2000). However, it is suffering from huge amounts of disturbance, and it is predicted that South-East Asia will lose three-quarters of its rain forest by the turn of next century (Sodhi et al. 2004).

Tropical pioneer tree species are considered as a functional group characterized by a suite of ecological characteristics such as high light demand and high photosynthetic capacities. This study compared the photosynthetic characteristics of eight co-existing pioneer tree species in 3–4-y-old and about 6-m-tall secondary forest stands in Sulawesi, Indonesia. Its objectives were (1) to determine the range and interspecific variation in six photosynthetic parameters, and (2) to identify morphological and chemical leaf traits that can predict light-saturated net photosynthetic rates (on a leaf area or leaf mass basis, Amax-area or Amax-mass). Species averages of Amax-area in sun leaves ranged between 14.2 and 20.3 μmol m−2 s−1 (mean 17.5) which is high compared with literature data. Among the co-existing species, average leaf size (56–896 cm2) differed by a factor of 16, specific leaf area (SLA, 10.7–21.4 m2 kg−1) and leaf nitrogen content (19.6–33.9 g kg−1) twofold. At the species level, Amax-area was not correlated with leaf N content but decreased significantly with leaf size. Amax-mass showed a higher interspecific variation than Amax-area, and was positively correlated with SLA and leaf N content (slope: 13.4 nmol CO2 g N−1 s−1). Both, Amax-area and Amax-mass were more closely related to leaf morphological attributes than to leaf N. We conclude that the tropical pioneer tree species studied do not form a homogeneous functional group in terms of photosynthetic performance. Rather, a considerable variation in leaf morphology and nitrogen content exists, which also shows up in a substantial variation in Amax-mass and, to a lesser extent, in Amax-area.