The objective of this study was to determine the effects of gall induction on leaf phenolic compounds and their indirect effects on the subsequent attack of folivorous insects in Achatocarpus gracilis, Cordia alliodora, Guapira macrocarpa, Guettarda elliptica and Ruprechtia fusca that occur in both hillside and riparian sites at Chamela-Cuixmala tropical dry forest in western Mexico. There are differences in soil water content between riparian and hillside sites where trees in the first are mainly evergreen and deciduous in the second. A few tree species occur in both sites and their intraspecific phenological response is also different between sites. In this case, trees of a given species that occur in riparian sites will be evergreen whereas trees on hillsides of the same species will be deciduous. Four plant species had significantly greater total phenol concentrations in galled than ungalled leaves in both deciduous hillside and riparian sites. In three plant species associated with galls, host total phenol concentrations were significantly greater in short than in tall plants. The frequency and amount of folivore damage were greater in leaves without galls than leaves with galls in these four plant species. These results indicate that galling insect species may directly affect leaf phenolic concentrations and indirectly may affect the incidence and consumption of folivorous insects in tropical plant species. This may have important consequences on the preference of leaves by folivorous insects that might be excluded by galling insect species in this dry tropical system.

Leafing, flowering and fruiting of the 24 most representative species of trees and shrubs in Sudanian savannas and fallows near Bondoukuy (Burkina Faso) were studied for 2 y (May 1992 to June 1994). The period of defoliation was generally brief, and leaf flushing, flowering and fruiting occurred mostly in the dry season. The phenological behaviour varied nonetheless across species and individuals. A classification taking into account synchrony/asynchrony and duration in foliation, flowering, fruit ripening and seed dispersal phases allowed comparison of species' phenological plasticity. Species were distributed along a scale of increasing plasticity, from trees to shrubs. Along this scale, species of the Leguminosae appeared to be separated from those of the Combretaceae, the latter being generally considered as characterising fallow farming systems. These results provide new evidence about the effect of phylogenetic constraints on the phenological behaviour of the woody species of the Sudanian savannas.

RÉSUMÉ. La feuillaison, la floraison et la fructification des 24 espèces d'arbres et d'arbustes les plus représentatives des jachères et savanes soudaniennes de la région de Bondoukuy (Burkina Faso), ont été étudiées pendant 2 ans (mai 1992 à juin 1994). La défeuillaison est généralement brève et l'émergence des feuilles, la floraison et la fructification se produisent principalement pendant la saison sèche. Les comportements phénologiques varient cependant entre les individus et entre les espèces. Une classification prenant en compte le degré de synchronisme ainsi que la durée de la feuillaison, de la floraison, de la maturation des fruits et de la dispersion des graines a permis de comparer la plasticité phénologique des espèces. Celles-ci sont distribuées selon un gradient de plasticité croissante des arbres aux arbustes. Sur ce gradient les Légumineuses se dissocient des Combrétacées, espèces qui sont par ailleurs généralement considérées comme caractéristiques des jachères. Ces résultats sont une nouvelle illustration de l'effet des contraintes phylogénétiques sur le comportement phénologique des espèces ligneuses des savanes soudaniennes.

Plants often modify microsite conditions important for seedling establishment. In tropical moist forest, advance regeneration in the form of shade-suppressed seedlings is a major component of regrowth in new gaps. Tree seedlings may be filtered by the composition and structure of the forest understorey. In a lowland forest in Central Panama, we examined light availability, litter accumulation and the seedling community (abundance, diversity and composition) under and away from the canopies of a common subcanopy, clonal palm, Oenocarpus mapora, and tested whether seedling abundance varies as a function of changes in understorey light conditions and leaf litter depth. We found evidence that leaf litter was five times deeper and light availability 27% lower under O. mapora canopy than where it was absent. The probability of finding a seedling in plots under O. mapora canopies was 33% lower than in plots without the palm. Plots under O. mapora canopies also had 49% fewer species of seedlings and those seedlings came from significantly larger seeds than seedlings in plots not under the palm. Overall, seedling density was positively correlated with light availability and negatively correlated with leaf litter depth. A transplant experiment confirmed that for at least one large-seeded species of dicot, seedlings are negatively affected by the reduction in light availability and increase in leaf litter. Under O. mapora, seedlings of Gustavia superba (Lecythidaceae) had lower survivorship and biomass accumulation than away from the palm. Thus, site conditions produced by large, clonal, multi-layered palms such as O. mapora may reduce establishment of small-seeded and shade-intolerant species, potentially affecting species composition in the understorey seedling pool.

Anthropogenic disturbance often results in the proliferation of native species of particular groups that leads to biotic homogenization. Leaf-cutting ants are an example of such winner organisms in tropical rain forests, but their response to disturbance in dry forests is poorly known. We investigated Atta colony density in areas of tropical dry forest in Brazil with different distance to roads and vegetation cover. Atta colonies were surveyed in 59 belt transects of 300 × 20 m, covering a total area of 35.4 ha. We found 224 Atta colonies, 131 of which were active and belonged to Atta opaciceps (87 colonies, 2.45 ha−1), A. sexdens (35 colonies, 0.98 ha−1) and A. laevigata (9 colonies, 0.25 ha−1). The density of active colonies sharply decreased from 15 ± 2.92 ha−1 in the 50-m zone along roads to only 2.55 ± 1.65 ha−1 at distances up to 300 m. The reverse pattern was observed for inactive colonies. Active Atta colonies preferentially occur in areas with low vegetation cover, while inactive colonies prefer areas with high vegetation cover. We demonstrate for the first time that anthropogenic disturbances promote the proliferation of leaf-cutting ants in dry forest in Brazil, which may affect plant regeneration via herbivory and ecosystem engineering as demonstrated for rain forests.

The spatial heterogeneity of tropical forest epiphytes has rarely been quantified in terms of biomass. In particular, the effect of topographic variation on epiphyte biomass is poorly known, although forests on ridges and ravines can differ drastically in stature and exposure. In an Ecuadorian lower montane forest we quantified epiphytic biomass along two gradients: (1) the twig–branch–trunk trajectory, and (2) the ridge–ravine gradient. Twenty-one trees were sampled in each of three forest types (ridge, slope, ravine positions). Their epiphytic biomass was extrapolated to stand level based on basal area–epiphyte load relationships, with tree basal areas taken from six plots of 400 m2 each per forest type. Our results document the successional addition and partial replacement of lichens by bryophytes, angiosperms and finally dead organic matter along the twig–branch–trunk trajectory. Despite having the highest tree basal area, total epiphytic biomass (mean ± SD) of ravine forest was significantly lower (2.6 ± 0.7 Mg ha−1) than in mid-slope forest (6.3 ± 1.1 Mg ha−1) and ridge forest (4.4 ± 1.6 Mg ha−1), whereas maximum bryophyte water storage capacity was significantly higher. We attribute this pattern to differences in forest dynamics, stand structure and microclimate. Although our study could not differentiate between direct effects of slope position (nutrient availability, mesoclimate) and indirect effects (stand structure and dynamics), it provides evidence that fine-scale topography needs to be taken into account when extrapolating epiphytic biomass and related matter fluxes from stand-level data to the regional scale.

The role of regeneration strategies in structuring tropical tree communities has been a recent focus of attention of tropical ecologists. Canopy discontinuities created by tree-falls are considered to be critical to many rain forest trees for establishing and completing their life cycles. It was investigated whether it is advantageous for two common large-seeded and shadetolerant Amazonian tree species, the palm Astrocaryum murumuru and the legume, Dipteryx micrantha, to have their seeds dispersed to gaps. The dispersal process was simulated by placing experimental seeds and seedlings in gaps and in the shaded understorey, and then following their performance. Seeds and seedlings from the two species were used. The Cox proportional hazard model was used to estimate the survival functions and compare survivorship curves. Survival of Astrocaryum seeds, and of Dipteryx seeds and seedlings, was higher in two gap zones (crown and bole) than in the shaded understorey (both near and far away from a conspecific adult). A higher proportion of seedlings of both plant species survived within gaps than in the shaded understorey over an 18-wk period. One year after the seedlings had been transplanted into gaps these patterns still held. Naturally occurring Astrocaryum seeds survived better beneath conspecific adult trees located within 30 m of gaps than beneath trees located 100 m from gaps. Variation in gap size was significant for Dipteryx seeds, but not for Dipteryx seedlings and Astrocaryum seeds and seedlings. Dipteryx seeds survived longer in smaller than in larger gaps. By colonizing tree-fall gaps, these two species improved their chances of completing their life cycles.

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 red crab Gecarcoidea natalis is the most abundant land crab on Christmas Island, Indian Ocean, and is found wherever native rain forest is still intact. The largest crabs grow to > 120 mm carapace width and reach > 500 g in mass. Red crabs are diurnally active, and moisture appears to be the most important factor governing surface activity – activity was nil below 77% RH, relatively low at 87% RH, and high above 95% RH. The crabs retreat to the interior of their burrows during dry periods, but emerge rapidly in response to rain. Because of their dependence on moisture for surface activity, red crabs show greater levels of activity during the wet season. Annual breeding migrations and moulting activity decreased the density of surface-active crabs at the main study sites for several weeks at the start of the wet season, and fruit and leaf fall caused dramatic local increases in the density of active red crabs. Density and biomass at the main study sites averaged 1.3 crabs m-2 and 1454 kg ha-1, respectively, and 1.2 crabs m-2 and 1137 kg ha-1 at a total of five sites across the island. Rough calculations suggest that the island-wide population is at least 100 million individuals. Red crabs are the dominant consumer on the forest floor, and consume seeds, seedlings and leaf litter. Because they are both abundant and widespread, these animals have the potential to be the most important determinant of pattern and process in rain forest on Christmas Island.

Along eight forest–savanna transects at seven semi-deciduous forest islands in the southern Comoé National Park data on spatial distribution of tree-size classes and environmental parameters (fuel load, shading by upper tree layers, and soil depth) were collected. For dominant tree species, a sequential series was observed from the forest border into the forest interior. At the forest border, Anogeissus leiocarpus was the most abundant tree with juveniles (<1 cm dbh) reaching highest density values (mean of 502 individuals ha−1) at the outer periphery of the forests. Regression analysis of juveniles of dominant tree species and environmental parameters resulted in a separation of forest and savanna species. Forest tree species regenerated well at forest sites, but also in the shade of A. leiocarpus stands. We conclude that (1) the studied forest islands advance against savanna by sequential succession, and (2) A. leiocarpus has a high potential to regenerate at savanna–forest boundaries under moderate fire impact and on rather shallow soils. The potential of A. leiocarpus to act as an important pioneer in the replacement of savanna by forest due to its effective regeneration at savanna sites and subsequent modification of site conditions, especially fire intensity by shading out savanna grasses, is discussed.

Patterns of population fluctuation, reproductive activity and age structure were studied in populations of the marsupial Micoureus demerarae occupying two small (7.0 and 8.8 ha) fragments of Atlantic Coastal Forest in southeastern Brazil, from 1995 to 1998. Males, but not females, were observed to move between populations. Estimated sizes of the populations in each fragment were very small, usually below 20 individuals. Breeding usually occurred from September to April. Population peaks came mostly by the end of this season, the delay reflecting the time required for the young to become trappable. In August 1997, the area was hit by a fire severely affecting the smaller fragment. Populations were synchronous before the fire, although they became asynchronous after it, possibly in the short term only. Small population sizes, synchrony and presumable male-biased migration are all likely to make the set of populations more vulnerable to extinction than expected for a metapopulation.

In the Central Amazonian floodplains, several hundred tree species grow in areas that are periodically flooded by nutrient-rich white-water rivers (várzea) and by nutrient poor black-water rivers (igapó). Seed masses of 31 species from várzea and 27 species from igapó were compared taking into consideration their taxonomic relatedness. Overall average seed mass was higher (mean = 7.08 g) in nutrient-poor igapó than in nutrient-rich várzea (mean = 1.16 g). In igapó, the species growing at high elevations on the flooding gradient had significantly higher seed masses than the species growing at low elevations. In várzea, no difference was found between species growing at high and low elevations. Four large-seeded species from igapó occurring at high elevations on the flooding gradient were responsible for most of the difference in average seed mass between forest types. These data suggest that at low positions in the flooding gradient in igapó, selection pressure on seed size is probably the same as in várzea. At sites with short periods of flooding in igapó forests, on high levels in the flooding gradient, the need for rapid height growth may have selected for species with larger seeds which enable seedlings to be less dependent on soil nutrients.

Long-term growth rates of 32 palms (Prestoea montana) were studied using height and diameter measurements taken four times between 1946 and 1982. All leaf scars were counted to estimate leaf production rate, and the distance between scars was measured to establish relationships between leaf production and height growth. Height and diameter growth and leaf production of these palms were not constant during this 36-year period. A greater distance between leaf scars was indicative of rapid height growth but not of high leaf production rate. Dominant palms always grew fast (> 20 cm/yr) in height when they were small, but height growth slowed down when they reached the canopy. Small palms with slow height growth remained suppressed and lost the capacity to grow rapidly in height as adults. Diameter growth decreased with age from 0.07 cm/yr (SE = 0.02) between 1946–51 to 0.01 cm/yr (SE = 0.003) between 1951–82. Annual leaf production averaged 4 leaves/yr over the 36 years of study, was significantly different (P = 0.001) between 1946–51 (5.6 leaves/yr) and 1951–82 (3.8 leaves/yr), and was 4.6 leaves/yr for palms that grew >5 m since 1946 and 3.7 leaves/yr for those that remained suppressed. The best estimate for the mean age of all palms was 61.1 yr (SE = 2.4), obtained from long-term population leaf production rates.

The vast array of species found in the humid lowland tropical forests of the world are frequently divided into two general groups, light-demanders and shade-tolerators (Denslow 1980, Swaine & Whitmore 1988). In this dichotomy the characteristics of the gap-demanders are often given to include small seed size, dormancy mechanisms, potentially rapid growth, and requirement of direct sunlight for germination and establishment. Conversely, shade-tolerators are considered to have large, short-lived seeds, the potential to germinate in canopy shade and an ability to grow slowly in deep shade (Swaine & Whitmore 1988). Failing to conform to either general pattern, small-seeded shade-tolerant species have been documented from both the Central American and South-east Asian tropics (Ellison at al., Metcalfe & Grubb 1995). Some of these small-seeded species have been shown to be able to germinate in filtered light approximating to canopy shade (Metcalfe 1996), and seedlings of other small-seeded species have been recorded growing under a closed canopy (Grubb 1996, Kiew 1988, Raich & Gong 1990). Several taxa shown by Metcalfe (1996) and Metcalfe & Grubb (1997) to be small-seeded and shade-tolerant were found in considerable numbers in the soil seed bank in Sabah by Kennedy (1991), and one (Urophyllum glabrum) by Putz & Appanah (1987) in Peninsular Malaysia. Species with small seeds (less than about 1-10 mg) do not have the resources to emerge from underneath a covering of leaf litter, and require litter-free sites for successful establishment (Guzmán-Grajales & Walker 1991, Molofsky & Augspurger 1992, Putz 1983).

Spatial vegetation utilization of elephants was investigated within mixed woodland savanna along the Chobe River in northern Botswana in the dry season of 1998. Using multiple linear regression, accumulated stem breakage by elephants was predicted by a terrain index, distance to water, stand density, number of trees > 4 m tall, tree height, density of Combretum apiculatum, C. elaeagnoides, C. mossambicense and the density of other (accumulated) tree species. Within mixed woodland at 2–7 km distance from the river fine-grained terrain ruggedness was the most important factor contributing to 55% of observed differences in use by elephants, while distance to water and the density of C. apiculatum contributed an additional 20% and 4%, respectively to the multiple linear regression model. Stem breakage was, on average, almost twice as high in rugged terrain compared with flat terrain at similar distance to water within the same vegetation type. Rugged terrain had 2–3-fold higher proportion of plots with very high Combretum shrub densities. These results suggest that the terrain index may be useful in management, predicting the areas most sensitive to vegetation change in a woodland system with increasing elephant densities.

The small size of many African protected areas makes adjacent rangelands potentially important in the local survival of wild animals. In order to assess the importance of pastoral areas to wild ungulates, we studied density and habitat choice of wild ungulates and cattle in Lake Mburo National Park, Uganda, the adjacent exclusively pastoral Nshara Dairy Ranch and on private land consisting of a mixture of ranching and subsistence farms. Transects, in the three land-use zones, were walked during the wet season and the data were analysed by DISTANCE sampling technique. We found significantly higher total density of wild ungulates on the dairy ranch compared with the National Park and private land. There was no significant difference in total wild animal density between the National Park and private land. Impala (Aepyceros melampus), zebra (Equus quagga), bushbuck (Tragelaphus scriptus) and waterbuck (Kobus ellipsiprymnus) had significantly higher densities on the dairy ranch compared to the National Park. Only eland (Taurotragus oryx) density was higher in the National Park compared to private land. Wild ungulates and cattle showed a high degree of habitat overlap, generally preferring open grassland. Our study shows that high densities of wild ungulates are not necessarily associated with protected areas. Pastoral areas may be important for populations of wild herbivores during the growing season despite a pronounced presence of livestock.

Secondary fallow vegetation in parts of the Papua New Guinea lowlands is dominated by the shrub Piper aduncum L. that originates from South America. Here we report on its seed bank, growth rate and biomass accumulation. P. aduncum accounted for 69 % (408 m−2) of the seed bank in the forest and 53 % (1559 m−2) of the seed bank under fallow. About 90 % of the tree seed bank at the fallow site was dominated by P. aduncum whereas this was 78 % in the forest soil. Two-year-old P. aduncum had grown to 4.5 m height and had accumulated 48 Mg dry matter (DM) per ha of above ground biomass. The rate of biomass accumulation increased from 10 Mg DM ha−1 y−1 in the first year to 40 Mg DM ha−1 y−1 in the second year when 76 % of the biomass consisted of mainstems. The highest growth rate of 134 kg DM ha−1 d−1 occurred when P. aduncum was 17-mo-old. Aggressive invasion and monospecific stands of P. aduncum are explained by its dominance in the seed bank, fast growth, and high rates of biomass accumulation. P. aduncum is a major competitor to indigenous tree species and presents a threat to Papua New Guinea's rich biodiversity.

One manner in which elephants utilize trees is by removing their bark. This type of utilization is concentrated on the largest trees in the landscape. The role of bark removal in increasing the vulnerability of large trees to fire and the mechanism through which fire damage is mediated were investigated in Kruger National Park, South Africa, by experimentally removing bark and burning Acacia nigrescens stems with diameters ranging between 30 and 68 mm. Also, field surveys were conducted subsequent to natural fires in order to investigate mortality patterns of large trees with dbh greater than 15 cm with bark removed by elephants. An increasing probability of mortality was associated with increasing amounts of bark removal but only if trees were burned. When trees had bark removed but were not burnt, simulating damage only to cambium and phloem, none of the 12 treated stems died in the 4-mo period over which the experiment ran. Moreover, low levels of cambium damage were detected in large burned stems. This suggests that bark removal increases fire-induced xylem damage and that this damage contributes towards stem mortality. In a survey of 437 large trees, bark removal by elephants was frequent on large stems (44%) and larger trees have greater amounts of bark removed. Post-fire mortality of large trees was significantly associated with increasing bark removal and stem diameter. These results indicate that bark removal by elephants increases the vulnerability of stems to fire, resulting in mortality of large stems otherwise protected from fire.

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.

Leaf litter production by lianes (=lianas) and trees was compared in an evergreen rain forest in subtropical Australia. Several successional stages were represented at the main site. Lianes contributed 2.2% of total basal area (69.6 m2 ha-1) of this site, but 24% of leaf litterfall (5.9–6.5 tha-1 y-2 over two years. Minor year-to-year variation in litterfall was attributed to incidence of severe storms, and drought. Lianes were responsible for about 17% of leaf litterfall in spring, 21% in the summerpeak, and 40% in autumn, more lianes than trees being deciduous. Leaf litter production by 23 individual species of liane, in relation to their basal area within the main site, was, on average, 15 times as great as that by 34 tree species, but declined more steeply between the species-groups of early and later succession. Tendrillar lianes, unlike twiners and scramblers, were confined to the ‘early’ successional group, and their foliage was spread across canopy surfaces, maximizing light interception. It was concluded that this contributed to greater production of leaf litter, per unit basal area, by tendrillar than by other climbers, and to the successional decline in leaf litterfall from lianes.

Although fire is occurring at greater frequencies and spatial scales in the moist tropics, few studies have examined the ecological impacts of fire in tropical montane cloud forest (TMCF). This study, conducted in the Chimalapas region of Oaxaca, Mexico, documents changes in live tree biomass, live fine-root biomass, and fallen and standing dead wood 4 y following deep ground fires occurring in TMCF during the 1997–98 El Niño Southern Oscillation event. Forests growing on two different substrates (metamorphic and sedimentary) and having three different statures (mean canopy heights: 20–30 m, 15–20 m and 4–6 m) were assessed within six paired plots established on adjacent burned and unburned forest sites. Total live tree biomass was 82% and 88% lower for burned TMCF growing on metamorphic and sedimentary substrates, respectively, compared with unburned TMCF. Nearly 100% of the living biomass was killed in elfin TMCF located on exposed sedimentary limestone at the highest elevations. Live fine-root biomass in the upper organic soil horizon of burned TMCF sites was 49% lower on metamorphic substrates and 77% lower on sedimentary substrates compared with unburned sites. The amount of total dead wood was 3- to 14-fold greater in burned forests compared with unburned forests. These results suggest that first-time fires in relatively undisturbed TMCF can cause dramatic changes in live above- and below-ground biomass at levels greatly exceeding values reported for most lowland tropical rain forests. These patterns may be attributed to the slower decomposition rates and thick organic soils typical of TMCF, combined with the relatively fast drainage associated with steep topography and, in some locations, sedimentary limestone-derived substrates.