All individuals of Cedrela lilloi (a valuable timber species of Meliaceae) > 4 cm DBH were sampled in three rectangular plots (c. 15 000 m2 each) in northwestern Argentina subtropical montane forests between 850 and 1350 m. Regeneration dynamics of C. lilloi was characterized by testing two hypotheses: (i) C. lilloi has a gap-phase regeneration mode, which implies a negative exponential size/age distribution at forest stand scale, growth releases due to canopy openings, and juvenile trees clumped at spatial scales of treefall gaps. (ii) There is a negative spatial association between adults and juveniles of C. lilloi which could be caused by density dependent mortality. Despite differences in age structure, diameter distribution approximated a negative exponential curve at all sites. Radial growth, measured as tree-ring widths, averaged between 2 and 4 mm y−1 which is comparable to fast growing tropical species in mature forests. Aggregation of juveniles peaked at radial distances of 8 to 15 m. Size structure, growth and spatial patterns of juveniles support the first hypothesis. In support of the second hypothesis, juveniles showed a negative spatial association with conspecific mature trees at spatial scales of less than 20 m. The Janzen–Connell model of density-dependence is one potential explanation for this pattern. Although regeneration and growth patterns of C. lilloi suggest a potential for sustainable management based on small-scale gaps, spatial dynamics at stand-scale need to be considered to ensure the regeneration of mature trees.

Epiphyte diversity as well as distribution and composition of epiphytic biomass was investigated in two lowland and two montane rain forests in Ecuador. Species numbers of epiphytes per tree were slightly higher in the montane (22–41 in Los Cedros, 33–54 in Otonga) than in the lowland forests (9–43 in Yasuni, 19–32 in Tiputini), however differences were not significant. In contrast, some epiphyte families did show significant altitudinal differences. The total epiphytic biomass per branch surface decreased from the centre of the crown to the periphery, and was generally higher in the montane (6.0 kg m−2 on central branches in Los Cedros, 1.8 kg m−2 in Otonga) than in the lowland forests (1.3 kg m−2 in Yasuni and 1.8 kg m−2 in Tiputini). Especially, dry weight of bryophytes and dead organic matter was higher in the mountains. In contrast, the biomass of green parts of vascular epiphytes on central branches was about the same in all four forests (0.4–0.6 kg m−2). A comparison with literature data from other study sites of tropical moist forests supports the observation that biomass of vascular epiphytes does not significantly change with altitude. It is discussed, that the high bryophytic biomass in montane compared to lowland forests is a major reason for differences in humus biomass between these forest types.

During the flooding of primary lowland rain forest by the filling of a hydroelectric reservoir in French Guiana, live-trapping was conducted on 175 subsequently formed islands. Different factors affecting the mammal trap efficiency were analysed. The main results show the real complexity of the sampling problem and the influence of the habitat disturbance due to the flooding. The location and/or the type of traps influenced captures of Dasypus novemcinctus, Proechimys spp., Metachirus nudicaudatus and Philander opossum in relation to their foraging and/or locomotion behaviour, and to to their size. The conformation of the islands (height and size) had no influence on the number of captures, nor on the species richness. The number of captures increased with the number of trapnights during the first water inflow and the level stretch. The best trapping success was observed during the wet season, when the strong habitat modification and the forest fragmentation became more significant. It resulted from the reduction of available land area and trophic resources. One year after the beginning of the water inflow, the decrease of the number of captures with the increasing number of trapnights reflected a real decrease of the number of mammals on the islands. The species richness increased with the number of trapnights in all periods, and was also the highest during the wet season.

Regeneration processes associated with gap-building-phase sites are described for the tropical montane cloud forest of the El Cielo Biosphere Reserve in Tamaulipas, Mexico. Twelve gap-building-phase sites c. 20 y old were selected within the forest according to their gap size and severity of disturbance (i.e. number of fallen logs). All flowering plants were recorded within the sites and specimens were identified to species level. Species were related to genera geographical affinity and growth form. Abundance, individual height and aerial crown cover were also recorded for all species. The composition for these sites was 121 vascular plant species grouped in 96 genera and 60 families. Results suggest that the advanced regeneration of shrub and tree species of the medium canopy, such as Eugenia capuli, Ternstroemia sylvatica, Turpinia occidentalis, Meliosma oaxacana, Myrsine coriacea and Podocarpus reichei, was favoured by the disturbance regime prevailing in this forest community. All of these woody species belong to tropical genera and are structurally dominant in the gap-building-phase sites. A differential response of the vegetation was obtained in these sites as a function of the severity of disturbance and gap size. This response was revealed by the dominance-diversity relationships. Small gaps (45–95 m2) or gaps created by two treefalls showed a higher diversity and density compared to larger gaps (95–150 m2) or those created by four fallen logs, which had a lower species diversity and significantly lower tree and vine densities. These results imply that gap size and the severity of disturbance have a significant effect on the regeneration of these disturbed sites.

Tepuis are uplifted mountains surrounded by forest or savannas in the Guayana Shield in northern South America. Large areas of the summits of these mountains are characterized by bare rock with vegetation isolates (‘islands’) that vary in size from less than 1 m2 to over 100 m2. The species composition, species–area relationships and the factors involved in the formation of these vegetation islands were quantified on Roraima Tepui, Canaima National Park, Venezuela. A total of 40 species were present on the islands, 35 of which were angiosperms. Orchidaceae was the family with the highest diversity, but Bonnetiaceae, Poaceae and Rapateaceae achieved higher coverage. A positive correlation between island size and species richness was found. This relationship was not linear and the best fit model between island size and richness was exponential. Of the environmental variables studied, the highest correlation was found between the log of soil volume and species richness. Soil nutrient availability, and not water, might be the limiting factor determining the species composition and diversity of these communities.

Long-term data on flower and fruit production of the forest community in two lowland tropical rain forests in north-western Sumatra are presented. The proportion of years with mast fruiting was found to be similar to that found elsewhere in Malesia. However, masting at the two sites, 70 km apart, did not coincide, and showed no correlation with the El Niño–Southern Oscillation phenomenon (ENSO). Comparisons with other sites in Malesia suggest a general waning of ENSO's impact toward western Malesia. Spatial variation at various scales in the timing of masting events was noted in Sumatra and elsewhere. This suggests that additional factors to ENSO play a role in determining forest-level mast fruiting, and we hypothesize that frugivorous animals have the opportunity to track mast fruiting. It is hypothesized that asynchrony between nearby areas in masting increases toward the western edge of Malesia.

Forest edges bordering on pasture were created by cutting and burning the surrounding Amazonian lowland rain forest in the dry season (June) of 1990. Litterfall was measured for 3.5 y along transects 10, 50, 100 and 250-m into the forest from the forest edge. Litterfall along the 10-m transects increased by up to 2.5 times that on spatial controls (250-m transects) in the dry season in which the edge was created. In the second dry season after edge creation litterfall at 10-m was lower than on controls, after which it returned to control rates in the second wet season, 1.5 y after edge creation. Litterfall 50-m into the forest was less affected; there was a smaller rise in the dry season in which the edge was cut, and no significant effects after that. At 100-m there was no effect of edge creation on litterfall. Phosphorus concentrations in litterfall were elevated along 10-m transects, 10–12 wk after edge creation, possibly because of reduced retranslocation prior to abscission. The changes in litterfall described here are inextricably linked to the biomass collapse, which occurs near forest-fragment edges in the same area.

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.

A study was conducted on the effect of extended land-use on secondary forest biomass accumulation in the Amazon. Structural measurements were made in a series of secondary forest stands, from 4–30 y old, in Brazil and Bolivia. Half of the stands were forest regrowth following clearance and only 1 y of cultivation; the other half were regrowth following 4 y or more of continuous pasture in Brazil and three or more rotations of medium-fallow agriculture in Bolivia. Above-ground live biomass was estimated using published allometric equations. Total biomass ranged from 17 to 207 Mg ha−1. Biomass of pioneer trees was poorly related to stand age, while that of later-successional trees increased linearly with age. Total biomass accumulation in Bolivia averaged 5.4 Mg ha−1 y−1 over the entire age sequence. Biomass accumulation for regrowth following short-term use was not greater than that for regrowth following medium-fallow agriculture. In Brazil, biomass accumulation averaged 9.1 Mg ha−1 y−1 over the first 12 y of regrowth and 5.9 Mg ha−1 y−1 over the entire age sequence. Biomass accumulation was significantly slower, around 5.0 Mg ha−1 y−1, for regrowth following continuous pasture than for regrowth following 1 y of cultivation.

All stems ≥ 1 cm dbh were measured, tagged, mapped and identified on a 1-ha plot of rain forest at Gambubal State Forest, south-east Queensland, Australia. The spatial patterns and size class distributions of 11 common tree species on the plot were assessed to search for mechanisms determining their distribution and abundance. The forest was species-poor in comparison to many lowland tropical forests and the common species are therefore present at relatively high densities. Despite this, only limited evidence was found for the operation of density-dependent processes at Gambubal. Daphnandra micrantha saplings were clumped towards randomly spaced adults, indicating a shift of distribution over time caused by differential mortality of saplings in these adult associated clumps. Ordination of the species composition in 25-m × 25-m subplots revealed vegetation gradients at that scale, which corresponded to slope across the plot. Adult basal area was dominated by a few large individuals of Sloanea woollsii but the comparative size class distributions and replacement probabilities of the 11 common species suggest that the forest will undergo a transition to a more mixed composition if current conditions persist. The current cohort of large S. woollsii individuals probably established after a large-scale disturbance event and the forest has not attained an equilibrium species composition.

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.

Species composition and turnover that have occurred in a series of permanent sample plots established during the 1930s and 1940s in Budongo, a semi-deciduous Ugandan forest, are reported. The plots were established as part of a sequence first used to describe forest succession, five of which have been maintained and which were last measured in 1992-1993. One plot (plot 7) provides 53 y of data from old-growth pristine forest. Plot 15 was established in wooded grassland at the forest edge and is now closed high forest. Evaluation of the remaining three plots is complicated by silvicultural interventions carried out in the 1950s. Forty species have been added since the first evaluations and a total of 188 tree species (over 80% of Budongo's forest tree flora, and including two exotics) has now been recorded from within the plots. The pattern of shade-tolerance in the original plot series conforms to patterns expected for succession with an increasing proportion of shade-tolerant species with development, and large stems appearing to ‘lag behind’ smaller stems in this respect. The time series data are less consistent, and while plot 7 increased in the proportion of shade-tolerant stems through time, the proportion of shade-tolerant species actually declines. Stem-turnover (the mean of mortality and recruitment) slowed with implied successional stage. Most species have a higher recruitment than mortality rate and stem numbers have thus increased in all plots. This is most pronounced in the putatively ‘early successional’ plot. Stem size structure has changed within the plots, with an increased proportion of smaller stems. Species show different rates of turnover and these vary from plot to plot and period to period. In plot 7, the overall mortality rate decreased with initial stem size. Estimates imply that some tree species may easily live longer than 500 y after reaching 10 cm DBH, and that 1000 y is possible. The importance of large trees in determining forest dynamics is illustrated by the finding that death of only seven stems in plot 7 contributed over 60% of net basal area losses recorded over the 53-y observation period. Many of the observed patterns were not predicted and could only have been found by long-term studies.

Changes in floristic composition and structure in a cerrado sensu stricto area were studied over a 9-y period. A sample of 19 (20-m × 50-m) permanent plots was assessed at 3-y intervals from 1985 to 1994, when all stems ≥ 5 cm at 0.30 m above the ground level were measured. There was little change in species composition over the period. Changes in density and basal area of the whole community were in the range of 5% over the 9 y. These parameters fluctuated over the years with the greatest differences found in the comparisons of 1991, 2 y after an accidental fire reached the site, with other years. The changes in density and basal area were greater than in several tropical forests, probably due to disturbances such as fire occurring at 3 to 5-y intervals. The nearly constant density and basal area suggested a highly resilient community.

The feeding behaviour of free-ranging black-handed tamarins, Saguinus midas niger, was monitored over 6 mo in a forest fragment in eastern Amazonia. The tamarins' diet included arthropods and plant exudates, but was predominantly frugivorous (87.5% of feeding records, 18 plant species) in all months. The seeds of at least six species were ingested whole and defecated intact. Ingested seeds were relatively large, but those with diameter > 1 cm or length > 2 cm were discarded. Seeds took 2–4 h to pass through the digestive tract, and more than half (50.6%) were dispersed to a habitat different from that in which they were ingested. A fifth (19.3%) of all seeds were dispersed from primary to secondary forest, indicating that S. m. niger may play an important rôle in forest regeneration, a rôle that will become increasingly important in the fragmented landscape of eastern Amazonia.

Data from a hilly forest study site at Batang Ule, Sumatra, are organized into 30 100-m × 10-m subplots lying perpendicular to the line of maximal topographic gradient, from the valley to the plateau/ridge. The following methodological question is addressed: what species diversity measures are best used in order to reveal the ecologically distinct regions in the site. The main tool used to answer this question is the α-diversity curve (Hα). Graphical examination of tree and species densities, and α-diversity curves identifies an anomalous species diversity behaviour of the ‘ridge above the slope’ subplots which may have implications on land-facet class definitions. Factor analysis of the α-diversity curves indicates that the diversity space is two-dimensional: i.e. two diversity measures are sufficient to characterize the site; the species density (H0), and the Berger-Parker index (H∞). In the

two-dimensional diversity-space three distinct species diversity groups are found which relate to the topographic gradient at the Batang Ule site. The results are compared with those for a flat homogeneous site at Pasirmayang, Sumatra. The implications of the results on land-classifications in species-diversity mapping and conservation strategy are discussed.

(Accepted 31st July 1999)

In the ‘elfin’ cloud forest of Serrania de Macuira, exchangeable ammonium and nitrate, and the rates of soil nitrogen mineralization and nitrification were measured in soil samples under different water, temperature and mineral nutrient additions. The effects of nitrogen, phosphorus and nitrogen plus phosphorus fertilization on radial trunk growth were measured in three tree species. In the cloud forest soils, concentrations of ammonium were much higher than those of nitrate. Nitrate was higher in samples collected during the afternoon than during the morning, probably as a result of leaching during the night or nitrification during the day. When samples were incubated under different water and temperature treatments, rates of nitrogen mineralization and nitrification increased more with changes in soil water content than with changes in temperature. Nitrification was significantly increased in soils amended with ammonium or with ammonium plus phosphorus, suggesting that nitrification is substrate-limited. Fertilization with nitrogen and phosphorus resulted in significantly increased girth increments in Guapira fragrans (Dum. -Cours.) Little and Rapanea guianensis Aublet. Myrcianthes fragrans (Sw.) D.C. did not respond to the fertilization. The results of this study support the hypothesis that the characteristics of montane rain forest in small and large tropical mountains (the ‘Massenerhebung’ effect) are greatly controlled by soil water conditions and related soil nitrogen availability.

For analysing field data as well as for modelling purposes it is useful to classify tree species into a few functional types. In this paper a new aggregation of tree species of the dipterocarp rain forests in Sabah (Borneo), Malaysia, is developed. The aggregation is based on the two criteria successional status and potential maximum height. Three classes of successional status (early, mid and late successional species) and five classes of potential maximum heights (≥ 5 m, 5–15, 15–25, 25–36, > 36 m) lead to a combination of 15 functional types. The criteria of the developed classification are chosen to suit for applications with process-based models, such as FORMIX3 and FORMIND, which are based on photosynthesis production as the main process determining tree growth. The concept is universal and can easily be applied to other areas. With this new method of grouping a more realistic parametrization of process-based rain forest growth models seems to be possible.

Termites were surveyed at an altitude of 1000 m in Maliau Basin, Sabah (Borneo), in two primary forest habitats: lower montane forest and stunted facies of upper montane forest. Soil, wood, litter, termite nests, and arboreal microhabitats up to a height of 2 m above ground were sampled using a belt transect protocol. Two transects were run in each forest type. The upper montane forest transects collected 11 and 13 species, compared with 15 and 19 in the lower montane forest transects. With the addition of species records from casual sampling (conducted with roughly equal effort in each forest type), totals of 18 species were found in the upper montane forest, and 34 species in the lower montane forest. Similar species richness and relative abundance of wood-feeding termites were found in both forest types. However, the lower montane forest had greater richness and relative abundance of species that feed on soil and extremely decayed soil-like wood. This difference is attributed mainly to (i) the well developed, organic-rich soils of the lower montane forest being more suitable for soil-dwelling and mound-building termites compared with the upper montane forest which has sandy soils with a thick covering of peat, and (ii) the low-stature and open canopy of the upper montane forest creates microclimatic conditions which are adverse for soil-feeding termites compared with the high, closed canopy in the lower montane forest. As expected, comparisons with similar studies in lowland forests in Southeast Asia confirmed that the upland forests are relatively depauperate.

Observations were made of foraging frequency and food selection by species of Macrotermitinae in a savanna habitat in the distal reaches of the Okavango Delta, Botswana. A removal experiment in the field was also conducted. Observable foraging showed seasonal peaks in frequency, related to temperature in Macrotermes michaelseni and to moisture in sympatric soil nesting Termitidae. Foraging was mostly on woody litter of the dominant canopy trees and mammalian dung. There was considerable overlap in the selection of food items between species but M. michaelseni were seen more frequently on items of woody litter greater than 6 mm in diameter. In Petri-dish trials Microtermes spp. workers were aggressive towards M. michaelseni individuals and in the field they foraged more frequently and effectively when M. michaelseni numbers were experimentally reduced. Macrotermes michaelseni appeared to avoid competition with other fungus-growing species by spatial separation within the habitat and some limited separation in food selection, but intense competition is probably rare as space and food resources may rarely be limiting.

Tropical Cyclone ‘Rona’ crossed the coast of the Daintree lowlands of northeastern Australia in 1999. This study reports on its impact on forest canopy openness at six lowland rain forest sites with contrasting management histories (old-growth, selectively logged and regrowth). Percentage canopy openness was calculated from individual hemispherical photographs taken from marked points below the forest canopy at nine plots per site 3–4 mo before the cyclone, and at the same points a month afterwards. Before the cyclone, when nine sites were visited, canopy openness in old-growth and logged sites was similar, but significantly higher in regrowth forest. After the cyclone, all six revisited sites showed an increase in canopy openness, but the increase was very patchy amongst plots and sites and varied from insignificant to severe. The most severely impacted site was an old-growth one, the least impacted a logged one. Although proneness to impact was apparently related to forest management history (old-growth being the most impacted), underlying local topography may have had an equally strong influence in this case. It was concluded that the likelihood of severe impact may be determined at the landscape-scale by the interaction of anthropogenic with meteorological, physiographic and biotic factors. In the long term, such interactions may caution against pursuing forest management in cyclone-prone areas.