We examined the effects of the 1997–98 El Niño drought on nine rain forests of Mount Kinabalu, Borneo, at four altitudes (700, 1700, 2700 and 3100 m) on contrasting geological substrata (ultrabasic versus non-ultrabasic). Measurements of rainfall and atmospheric aridity indicated that the departure from normal conditions during the drought became greater with increasing altitude. During 1997–99 (drought period) compared to 1995–97 (pre-drought period), median growth rates of stem diameter of trees decreased for both smaller (4.8–10 cm) and larger (≥ 10 cm) diameter classes in the six upland forests (≥ 2700 m on ultrabasic substrata and ≥ 1700 m on non-ultrabasic substrata), but for neither diameter class in the other forests. The majority of species decreased or did not change growth rates during 1997–99, whereas some did increase. Tree mortality increased during 1997–99, at the larger diameter class in the two lowland forests (700 m) on both substrata, and at least at the smaller diameter class in the four upland forests (≥ 1700 m) on non-ultrabasic substrata. In two of these upland forests, mortality was restricted to particular understorey species. Mortality did not significantly increase in the three upland forests (≥ 1700 m) on ultrabasic substrata; this suggests that the adaptation to nutrient-poor soils might have provided the resistance to drought.

The land crab, Gecarcinus quadratus (Gecarcinidae), affects plant diversity in a mainland, neotropical rain forest through selective seedling consumption. In Costa Rica's Corcovado National Park, G. quadratus lives in fossorial populations with up to six crabs m−2 in coastal forest extending c. 600 m inland from the Pacific Ocean. A narrow transition zone (c. 30 m) separates the crab zone from the crabless zone that comprises the remainder of the 55 000-ha park. The composition of dicotyledonous plants from the adjacent zones were distinct: crabless zone transects (240 m2) contained 97 species representing 76 genera and 41 families; crab zone transects contained 46 species representing 38 genera and 25 families. Despite the narrow zonal separation, the crabless zone shared only 27.4, 29.9 and 52.8% of its species, genera and families, respectively, with the crab zone; in contrast, crab zone transects shared 56.5, 60.5 and 88% of their species, genera and families with the crabless zone. This abrupt diversity gradient suggests a powerful selection pressure in the crab zone preventing the establishment of certain dicotyledonous species. Two years of experimental crab exclusion suggested that G. quadratus reduces seedling density and diversity. Densities of seedlings (3–25 cm tall) within exclosures increased 144% over baseline values while control densities decreased. In contrast, seedlings at the cotyledon stage and taller (26–50 cm) were unaffected by crab exclusion. Preference tests conducted in the crab zone revealed a five-fold ratio of mortality rates for seedlings transplanted from the crabless zone and crab zone respectively. However, when protected from crabs, transplanted crabless and crab zone seedlings survived similarly well over 6 mo (83 and 70% respectively). Together, data from transects and experiments suggest that G. quadratus influences plant community composition.

Lianas of Yasuní National Park and the Huaorani Ethnic Reserve, Ecuador were sampled using 0.2-ha subsamples from 12 1-ha plots. Using surveys and herbarium collections, the total species richness of lianas in the park and reserve is estimated to be close to 500 species. Terra firme habitats are significantly more species rich than floodplain habitats, but density of stems is not significantly different between habitats. The most abundant liana species is Machaerium cuspidatum Kuhlm. & Hoehne (Fabaceae) in both floodplain and terra firme habitats, representing more than 10% of all stems in census plots. Analysis of rarity indicates that none of the liana species encountered on the 12 plots is rare in terms of geographic range, and many species are not rare in terms of population size or habitat specificity. The rank order of abundance, combined with the coefficient of variability in species distribution among plots, is used to identify the 38 species (an oligarchy) that dominate forests in Yasuní.

Remnant trees in fields may facilitate regeneration in fallows by attracting seed dispersers and creating favourable sites for plant establishment. Ntumu shifting cultivators in Cameroon clearing rain forest spare some trees to provide shade for crops. This study examines their effects on regeneration by comparing vegetation in quadrats beneath and away from their crowns, in fallows of varying age (3-20 y). For fallows of all age classes, plant diversity was not significantly different between positions. Trees accounted for larger proportions of species and stems beneath remnant trees. Herbs, principally large monocots, accounted for larger proportions away from trees. Basal area was much higher in quadrats beneath trees. In all quadrats, the majority of individuals belonged to species with animal-dispersed seeds. These accounted for a larger proportion of individuals beneath trees (75%) than away (64%). Wind-dispersed species accounted for a larger proportion of stems away from trees (23.6%) than beneath (11.7%). These data suggest that enhanced seed rain by attraction of perching animals, demonstrated in a companion study, nfluenced regeneration patterns. Effects on regeneration were similar beneath trees of an animal-dispersed (Pycnanthus angolensis, Myristicaceae) and a wind-dispersed (Triplochiton scleroxylon, Sterculiaceae) species. Remnant trees in fallows may facilitate succession by nucleation, resulting in more rapid restoration of forest and conservation of nutrients in plant biomass.

Leaf phenology was studied in individuals of a canopy species, Elateriospermum tapos (Euphorbiaceae), at various ontogenetic stages, in a Malaysian rain forest. The timing of leaf emergence was not synchronized among sapling individuals, and was not correlated with any meteorological factors of the preceding month. The timing of leaf fall in saplings was positively correlated with net radiation, and maximum and minimum temperature, but negatively correlated with relative humidity the preceding month, although these correlations were weak. The leaf production rate was larger under higher light, but the leaf fall rate was not related to the light regime of the saplings. Thus, leaf production was enhanced by the light availability for each individual, while leaf fall may have been enhanced by drought stress. Non-synchronous leaf production appears to be important for sapling growth, allowing saplings to occupy better-lit space quickly. On the other hand, tall trees showed a clear synchronous leaf-fall pattern, with an annual cycle, and no meteorological factors were correlated with the timing of leaf fall. Mature trees of this species produced flowers simultaneously with new leaves, after shedding their leaves. This suggests that the need to synchronize flowering might be the primary determinant of leaf production phenology in mature individuals.

Throughfall volume and interception of bulk precipitation events were measured during individual rain events of differing magnitudes in a primary wet tropical forest at La Selva Biological Station, Costa Rica. The relationship between canopy structure and throughfall were examined to identify key sources of spatial variation. Geostatistical analyses were also used to examine the spatial variation in throughfall, spatial autocorrelation and to determine minimum distances for independence of collectors. Throughfall volume was collected from 56 ground-based (funnel-style) collectors. Throughfall was collected for 26 separate precipitation events during July and August 1998. Per cent cover, distance to nearest tree, distance to nearest leaf were also estimated for each collection point. A weak relationship was found with per cent cover (r2 = 0.11). No relationship was found between throughfall and distance to the nearest leaf above the collector. Estimated interception was 1.88 mm (r2 = 0.94) with increased variance as bulk precipitation increased. A range distance of 45 m was estimated from variograms, strongly suggesting that large tree canopies and gaps are the source of much of the spatial variance in throughfall volume. Interception was reduced by 19% if only spatially independent collectors were used.

Amomum villosum Lour. (Zingiberaceae) is an obligate shade plant. Its leaves naturally roll up upon exposure to bright sun. This paper characterizes the effects of its leaf rolling and light on photoinhibition of photosynthesis and thermal dissipation activity under field conditions with sufficient soil moisture. Between the plants from bright and shaded parts of a canopy gap, both maximum photosynthetic rate (mean ≈ 6.5 μmol m-2 s-1) and stomatal conductance (148 vs. 131 mmol m-2 s-1) were not significantly different, but apparent quantum yield (0.021 vs. 0.025) and dark respiration rate (0.49 vs. 0.30 μmol m-2 s-1) were. In the foggy morning, when the incident irradiance was low, photoinhibition had already occurred in plants in the bright part of the canopy gap and also in plants in the shaded part of the gap but to a much lesser degree. Photoinhibition accelerated with increase of incident irradiance and relaxed in the afternoon. Thermal dissipation as indicated by non-photochemical fluorescence quenching (NPQ) increased rapidly in the morning and continued at a high rate in the afternoon. Prevention of leaf rolling resulted in acceleration of photoinhibition. Artificial inhibition of the xanthophyll cycle led to acceleration of both photoinhibition and inactivation of PSII reaction centres, and decrease of NPQ. Photoinhibited leaves in either control or treatment plants recovered overnight. These results show that A. villosum is able effectively to prevent photodamage through the mechanisms of leaf rolling, thermal dissipation and reversible inactivation of PSII reaction centres.

How do animals respond to an unpredictably heterogeneous environment? Ants foraging in the leaf litter of tropical wet forests experience unpredictably fluctuating food resources. To study how an ant species responds to these changes, foragers were tracked to determine home ranges of 51 colonies of Aphaenogaster araneoides, in three sites in a Costa Rican tropical wet forest. Of these colonies 16 were excavated to measure colony size, colony growth, and reproductive investment. These demographic variables were compared with two measures of home range quality: leaf litter dry weight and mass of arthropods. Home range areas of colonies were highly correlated with colony size, and moderately correlated with resource abundance. Colony growth was independent of colony size, as is found in other ants in unpredictable environments. The growth of colonies was closely associated with resource abundance. Production of the male reproductive caste was closely tied to the size of a colony rather than growth, but male production in slow-growing colonies was limited. Colonies foraging within high-quality environments grew at a faster rate, but reproduction was mainly correlated with colony size. Furthermore, it was found that the frequency of foragers in long-term treatment plots with supplemental food and reduced leaf-litter quality was not significantly different from the frequency of foragers in control plots. This rain-forest ant does not modify its home range areas in response to poor environments, and as a result, small-scale environmental heterogeneity strongly determines growth and reproduction.

Infestation rates by palm bruchid beetles (Bruchidae: Pachymerini) on Attalea palm seeds are so high that most endocarps remaining near parent trees contain larvae rather than endosperm a few months after the peak fruiting season. Vertebrates feeding on palm endocarp contents may therefore be feeding on animal rather than plant matter, and the timing of use of endocarps may be influenced by the presence of larvae rather than by background fruit availability. On Maracá Island Ecological Reserve, Roraima State, Brazil, white-lipped peccaries (Tayassu pecari), agoutis (Dasyprocta leporina) and squirrels (Sciurus igniventris) all fed on bruchid beetle larvae from Attalea maripa palm endocarps. Squirrels fed on both larvae and endosperm year-round in all habitat types; agoutis fed on both larvae and seeds only at times and at sites where alternate foods were not available; and white-lipped peccaries fed almost exclusively on bruchid larvae in high-palm-density habitats. Differences in timing and location of endocarp use by these species may provide an index of mammal community dynamics in neotropical forests.

In tropical forests, severe droughts caused by El Niño events may strongly influence the water relations of tree seedlings and thereby increase their mortality. Data on known-aged seedlings of three common shade-tolerant canopy tree species (Trichilia tuberculata, Tetragastris panamensis and Quararibea asterolepis) in a Panamanian moist forest are presented. Seedling survival during a severe El Niño dry season (1997-98) was compared with prior long-term survival data, and levels of drought stress were assessed by measuring plant water potentials and gas exchange characteristics. Contrary to prediction, dry-season seedling survival was not dramatically reduced in any species compared with that expected in ‘normal’ years. In Trichilia and Quararibea, pre-dawn water potentials averaged −2 MPa and midday water potentials about −3 MPa. Stomatal conductances were very low, averaging 26 mmol m-2 s-1 for Tetragastris and 11–13 mmol m-2 s-1 for Trichilia and Quararibea. Photosynthetic rates also were very low but consistently positive, averaging 0.8–1.1 μmol m-2 s-1. The findings suggest that, once established, seedlings of common tree species in this semi-deciduous forest may be tolerant of drought events.

Pollen analysis of a 150-cm-long core from Pantano de Pecho (0°20′S, 78°37′W) at 3870 m altitude in the Ecuadorian páramo documents altitudinal migrations and the composition of the upper forest line prior to deforestation. Four successive radiocarbon dates of 293 ± 41 14C y BP, 498 ± 40 14C y BP, 626 ± 33 14C y BP, and 729 ± 44 14C y BP show that the record includes the last c. 730 radiocarbon y, corresponding to the last c. 660 calendar years (cal y BP). The natural upper forest line was at a minimum altitude of 3400-3500 m between c. AD 1290 to 1315 (zone 1), from c. AD 1315 to 1350 at 3500-3600 m (zone 2), from c. AD 1350 to 1640 at 3600-3700 m (zone 3), from c. AD 1640 to c. 1765 at 3750 m (zone 4), and from c. AD 1765 to the present at 3700-3650 m (zone 5). The most important taxa were Alnus, Hedyosmum, Miconia, other Melastomataceae, Gunnera and Solanaceae. Since c. AD 1350 Podocarpus was continuously present with low abundance, but possibly not close to the upper forest line. Rarer elements of the upper montane forest were Dodonaea, Myrsine, Weinmannia, Myrica, Myrtaceae, Sapium, Juglans, Piper, Euphorbiaceae and Rubiaceae. Human disturbance and deforestation are shown by the presence of Rumex, Spermacoce pollen and charcoal particles. We surveyed the vegetation composition from isolated forest patches located between 3650 m and 4300 m. TWINSPAN analysis indicates forest patches up to 3950 m have a similar floristic composition to closed forest below the upper forest line. We argue that this apparent similarity does not necessarily mean that the slopes between 3750 and 3950 m were covered by closed forest in the past.

Lianas are abundant in tropical forests around the world, yet little is known about their ecology. In this study we report the results of a survey of ten common liana species on 27 host tree species in Cooloola National Park, Queensland, Australia. We found significant associations between some liana and tree species. Our results indicate that host tree diameter was important in explaining liana presence or absence. Host tree fruit type was correlated with the presence or absence of different liana types. Tendrillar and root-climbing lianas tended to occur most often on mid-sized, fleshy-fruited trees, and on smaller, non-fleshy-fruited trees. The presence of stem twining lianas tended to decrease with increasing tree diameter for both fleshy- and non-fleshy-fruited trees, although this effect was more pronounced for fleshy-fruited trees. In general, lianas were most prevalent on trees possessing intermediate levels of bark roughness and flakiness. Although we established associative patterns between liana species and these host characteristics, the model suggested that other unidentified variables may also be important in determining presence or absence of liana species.

The allometry of crown shape and trunk diameter with tree height were analysed for four tree species in a tropical lowland rain forest, southeast Brazil. The dimensional relationships of a subcanopy species (Garcinia gardneriana) were contrasted with those of two canopy (Chrysophyllum flexuosum and Swartzia simplex) and one emergent species (Sloanea guianensis). For all trees ≥ 1 cm dbh, we recorded dbh, total height, branching height, crown area, crown width and crown length. Observed allometric relationships indicated interspecific variation in trunk diameter and crown shape with height. All species conformed to the elastic similarity model, except the emergent one that showed thicker trunks and a scaling exponent conforming to the constant stress model. The general allometric function used to describe the overall relationship (all sizes combined) did not specify exceptional variation in crown shape between species of contrasting adult stature (emergent vs. subcanopy species). However, when allometric relationships through ontogeny were considered, different strategies of growth, maintenance and expansion of crown became evident. Crown shapes were much more variable in canopy and emergent species than in the subcanopy one, suggesting that larger-statured species might be more flexible in the relative allocation of energy to height, diameter, and crown growth than smaller-statured ones. Notwithstanding, it is suggested that it is not possible adequately to predict allometric relationships only by adult stature/canopy position. Allometric variation may be also related to size-dependent changes in demographic traits and/or different responses to light availability among tree species.

To study the impact of the annual long-term flooding (flood-pulse) on seasonal tree development in Amazonian floodplains, the phenology and growth in stem diameter of various tree species with different leaf-change patterns were observed over a period of 2 y. The trees of the functional ecotypes, evergreen, brevi-deciduous, deciduous and stem-succulent showed a periodic behaviour mainly triggered by the flood-pulse. Trees have high increment during the terrestrial phase. Flooding causes a shedding of some or all leaves leading to a cambial dormancy of about 2 mo and the formation of an annual ring. Studies carried out in tropical dry forests verify a strong relationship between the phenological development and the water status of the trees, strongly affected by seasonal drought. The comparison of the phenology and the diameter growth of the corresponding ecotypes in floodplain forest and a semi-deciduous forest in Venezuela shows a displacement of at least 2 mo in the periodicity, except for stem-succulent tree species. For stem-succulent trees it remains unclear which factors influence phenology and stem diameter growth.

Laboratory and growth-chamber experiments were used to evaluate whether there was evidence for nutrient retention by tropical terrestrial ecosystems being a two-stage process involving first soil adsorption and then plant absorption. Quartz sand with and without Fe and Al oxide coatings were treated with nutrient solution, then subjected to a leaching regime that simulated early wet-season conditions at a tropical location. Nutrient cations applied were rapidly lost in the initial leaches from quartz sand without oxide coatings, but showed a more gradual loss from oxide-coated sand, indicating temporary adsorption by the latter. In a second experiment, oxide-coated sand with and without seedlings of Grevillea robusta (a non-mycorrhizal tree species) were subjected to a similar treatment and leaching losses were compared. The presence of seedlings significantly reduced the losses of all nutrient cations, with the effect being minimal for Na and greatest for K, confirming that plants can gain access to temporarily adsorbed nutrients. More typical tropical soil-vegetation systems are likely to possess properties that magnify both the adsorptive and absorptive processes that have been documented in these experiments, justifying extrapolation of the experimental results to these natural systems. The existence of a two-stage process of nutrient retention provides a plausible explanation for the resistance of most tropical ecosystems to rapid loss of nutrients following events such as fires, which provide acute nutrient loading to the system.

Madagascar has one of the world's highest rates of human population increase, which is coupled with an increase of resource exploitation, particularly food and firewood. Forests are cleared and converted to rice fields or plantations (mainly Eucalyptus or pine). How does deforestation affect the amphibian diversity of the original biotope, the rain forest? To answer this question, the amphibian fauna of intact rain forests was compared with that of secondary forests, Eucalyptus plantations and rice fields. The main consequence of rain forest disturbance was loss of amphibian species. Compared with an intact forest, species richness in secondary forests, Eucalyptus plantations and rice fields were 54%, 46% and 12%, respectively. Species number and individual density increased with increasing structural complexity of the habitat and the presence of water bodies. The reproductive strategy of the species could be decisive for the presence or absence of single species in different habitats. With increasing degradation the percentage of species spawning in water increased. Correspondingly, Hyperoliidae and Raninae were characteristic of degraded habitats, whereas Microhylidae and Mantellinae were representative of natural habitats.

Seed removal was evaluated at the macro- and micro-habitat level in areas differing in successional stage in the Bolivian Amazon. The successional stages consisted of secondary forests of 2, 10 and 20 years old and primary forest. Seeds of nine tree species were artificially dispersed and the number of seeds removed was evaluated over 7 weeks. Several stand characteristics were measured at the sites where seeds were dispersed. Seed removal at the end of the experiment varied from 50 to 100% depending on the species, and from 74 to 90% depending on successional stage. In general, the removal rate decreased with an increase in age of successional stage. The seed removal rate was related to liana density and not to litter thickness. Different microhabitat characteristics explained the seed removal rate of four species but microhabitat characteristics did not explain the decrease in seed removal rate with an increase in forest age. The results support the idea that post-dispersal seed removal reduces the number of seeds available for germination, consequently playing an important role in the regeneration of abandoned agricultural areas.

Bactrocera Macquart (Diptera: Tephritidae) is a genus of nearly 500 species, the majority of which are endemic to tropical South-East Asia and Australasia (Drew 1989). Nearly all Bactrocera have frugivorous larvae that feed on the fleshy fruit of tropical trees, shrubs, and vines, and some species are major horticultural pests (White & Elson-Harris 1992). In this paper,where the term ‘fruit fly’ is used, we are referring explicitly to species of Bactrocera.

The reproductive ecology of piassava palm (Attalea funifera Mart.) was investigated for 19 months in the Atlantic forests of eastern Brazil. Its breeding biology is characterized by the production of functionally female, male and bisexual inflorescences. Staminate sterility is positively correlated with number of pistillate flowers. Staminate and pistillate flowers are receptive at anthesis. Bagging tests for apomixis were negative. Although geitonogamy can occasionally occur, outcrossing predominates. Pollen is relatively dry, but wind pollination is not a significant factor in cross pollination. Although inflorescences are visited by large numbers of insects, pollination is effected mainly by small sap beetles (Mystrops sp.) and weevils (Phyllotrox tatianae), which feed and breed in fertile staminate flowers. Pistillate flowers, which offer no reward to pollinators, appear to attract insect visitors by olfactory and visual imitation of staminate flowers, that is, via Bakerian mimicry. Flower production, fruit maturation, leaf production and leaf abscission occur continuously throughout the year. Flowering exhibits a distinct peak during the warmest months (Dec–Apr). Given its ubiquity throughout the region, A. funifera may well serve as a flowering and fruiting keystone species.

Damage caused by elephants was monitored in two woodland stands of Zakouma National Park (Chad) between February and March 1998. The Acacia seyal savanna was more severely damaged (29.8% of trees damaged of which 13.2% severely) than the Combretaceae savanna (26.5% of trees damaged of which only 4.2% severely). Nearly all severely damaged trees showed resprouts (respectively 86.8% and 88.5% in Combretaceae and A. seyal savannas). Both low damage rate and low mortality rate indicate that no serious ‘elephant problem’ occurred in Zakouma National Park. Elephants selected trees to damage according to species and height. Such selection also occurred for severe damage in A. seyal savanna, but only height affected severity of damage in Combretaceae savanna. In both savannas, the plots close to a water point were the most frequently damaged. In Combretaceae savanna, the distribution of damaged trees was clumped. On the other hand, spatial auto-correlation was not significant for the severity or the frequency of damage in A. seyal savanna. Neither tree density nor diversity of woody species affected the spatial patterns of damage or severe damage.