The suitability of several methods for estimating light conditions in the understorey of tropical forests, and of different sampling schedules was evaluated. Light conditions at 16 understorey sites in a Panamanian lowland forest were continuously measured for 9 mo with quantum sensors and photodiodes. Light conditions at the sites were also assessed indirectly with hemispherical fisheye photographs, plant canopy analysis, 38-mm photographs, 24-mm photographs and a spherical densiometer. Estimates from all indirect methods, except the spherical densiometer, were highly correlated with the direct measurements. Short-term direct light measurements for a day or a week also correlated with long-term light conditions. The indirect measures differed by up to c. 70% from the direct measures relative to single site measurements. Hence, the indirect methods are inadequate where single site light conditions have to be assessed accurately. However, because light conditions encountered in the understorey varied up to 13-fold, the indirect methods were found to be well suited to rank understorey light conditions among a large number of sites. The results from frequent and infrequent sampling schedules differed only slightly, suggesting that taking indirect measures at the beginning and the end of a study offers a reasonable compromise between accuracy and sampling effort.

A permanent plot of 28 ha was established in a dense wet evergreen forest in the Western Ghats of India to study the functioning of the ecosystem. Since April 1990, 1981 trees of ≥30 cm gbh have been enumerated in a systematic sampling of five strips totalling 3.12 ha. This paper describes the main structural and floristic characteristics of the plot.

The density (635 trees ≥30 cm gbh per hectare) and basal area (39.7 m2 ha−1) are high. Despite the high diversity (Simpson's D = 0.92 and Shannon's H' = 4.56), four species are distinctly, dominant in terms of an importance value index (relative density + relative basal area). Each of these four species occupies a different layer in the ecosystem: Humboldtia brunonis Wall. (Fabaceae) dominates the undergrowth, Myristica dactyloïdes Gaertn. (Myristicaceae) the intermediate strata, Valeria indica L. (Dipterocarpaceae) the higher canopy level and Dipterocarpus indicus Bedd. (Dipterocarpaceae) the emergents. This pronounced species hierarchy is one of the most important characteristics of the evergreen forests of the Western Ghats. The two dipterocarps account for 20.1% of the total number of trees and contribute 40.9% to the total basal area. This formation can, therefore, be considered as the westernmost lowland dipterocarp forest of Asia.

Analysis of the spatial variations in the floristic composition and in the structure of the main species populations shows that two kinds of mature phases can be identified: where the topography is raised and gently sloping, the vertical structure of the stand is discontinuous, with Dipterocarpus indicus and Vateria indica forming an emergent layer above a dense undergrowth; on slopes, the stand is lower, vertically continuous and saturated with Valeria indica and Myristica dactyloïdes. The link between the structure of the stand and dynamic processes is discussed.

Drought strongly influences plant phenology, growth and mortality in tropical forests, thereby shaping plant performance, population dynamics and community structure (Bunker & Carson 2005, Condit et al. 1995). Microbial symbionts of plants profoundly influence host water relations (Lösch & Gansert 2002), but are rarely considered in studies of tropical plant physiology. In particular, plant–fungus associations, which are ubiquitous in plant communities and especially common in tropical forests, play important and varied roles in plant water status. Fungal pathogens associated with roots, vascular tissue and foliage may interfere with water uptake and transport, increase rates of foliar transpiration, and induce xylem embolism and tissue death (Agrios 1997). In contrast, rhizosphere mutualists such as ecto- and arbuscular mycorrhizal fungi may benefit hosts by increasing surface area for water uptake, enhancing stomatal regulation of water loss, and increasing root hydraulic conductivity (Auge 2001, Lösch & Gansert 2002).

This study addresses patterns of nutrient dynamics on a precipitation gradient (1800–3500 mm y−1) in lowland tropical forest with heterogeneous soil parent material, high plant species diversity and large changes in species composition. Mean foliar concentrations of phosphorus, potassium, calcium and magnesium decreased with increasing precipitation, whereas foliar carbon:nitrogen increased with increasing precipitation. Mean foliar nitrogen:phosphorus varied from 16.4–23.8 suggesting that plant productivity at these sites is limited by phosphorus. Total soil nitrogen increased as a function of foliar litter lignin:nitrogen, whereas net nitrogen mineralization rates decreased with increasing lignin:N indicating that as litter quality decreases, more soil nitrogen is held in soil organic matter and the mineralization of that nitrogen is slower. Extractable phosphorus in soil was negatively correlated with foliar litter lignin:phosphorus, illustrating effects of litter quality on soil phosphorus availability. Overall, the results suggest that variation in plant community composition along this precipitation gradient is tightly coupled with soil nutrient cycling. Much of our understanding of effects of precipitation on nutrient cycling in tropical forest is based on precipitation gradients across montane forest in Hawaii, where species composition and soil parent material are constant. Our results suggest that variation in parent material or species composition may confound predictions developed in model island systems.

Resumen: Este estudio trata sobre los patrones de dinámica de nutrientes en un gradiente de precipitación (1800–3500 mm y−1) en un bosque tropical de tierras bajas con material parental heterogéneo, alta diversidad de especies de plantas, y un gran cambio en la composición de especies. Las concentraciones promedio de fósforo, potasio, calcio y magnesio disminuyen con un aumento en la precipitación, mientras que la proporción carbono:nitrógeno aumenta con un aumento en la precipitación. La proporción promedio de nitrógeno foliar:fósforo varía de 16.4 a 23.8, lo cual sugiere que estos sitios tienen limitaciones de fósforo. El nitrógeno total del suelo aumenta como una función de la proporción de lignina foliar de la hojarasca:nitrógeno, mientras que la tasa de mineralización del nitrógeno neto disminuye con un aumento de la proporción de lignina:nitrógeno, lo cual indica que mientras la calidad de la hojarasca disminuye, más nitrógeno del suelo es retenido en la materia orgánica y la mineralización de ese nitrógeno es más lenta. El fósforo extraíble está correlacionado negativamente con la proporción de lignina foliar de la hojarasca:fósforo, lo cual ilustra el efecto de la calidad de hojarasca en la disponibilidad de fósforo del suelo. En resumen, estos resultados sugieren que la variación en la composición de plantas de una comunidad a lo largo de este gradiente de precipitación tiene un efecto considerable en la retroalimentación de los ciclos nutricionales del suelo. Gran parte de nuestros conocimientos sobre el efecto de los gradientes de lluvia en ciclos nutricionales del suelo han sido basados en estudios de gradientes de precipitación de bosques de montaña en Hawaii, en donde tanto la composición de especies y el material parental del suelo son constantes. Nuestros resultados sugieren que la variación del material parental del suelo y la composición de especies pueden confundir las predicciones desarrolladas en sistemas modelos de islas.

The study of seed fate is crucial for understanding fruit-frugivore interactions. One factor that can greatly influence the fate of seeds dispersed through mammal defecation, is the dung that accompanies the seeds. Dung attracts dung beetles and rodents; the former eat and bury dung, the latter feed on seeds. In this study the fate of Micropholis guyanensis subsp. guyanensis seeds surrounded by no dung and by 5, 10 and 25 g of howler monkey dung was followed until seedling establishment. The depths at which dung beetles bury the seeds were measured, and the effect of burial depth on seedling emergence was determined. Although initial seed fates differed among treatments, the same percentage of seedlings established from seeds without faecal material, and from seeds with 5, 10 and 25 g of dung. However, a significantly higher proportion of seedlings established from buried seeds when compared to seeds that remained on the surface. The percentage of seedlings establishing in a controlled germination experiment decreased significantly with increasing burial depth. The effect of dung beetle activity should be taken into consideration when assessing the role that mammal endozoochory plays on seed dispersal ecology of tropical plants. These results provide further evidence for the emerging realization that the study of secondary seed dispersal and post-dispersal events is crucial for a more complete understanding of plant regeneration.

Studies in western Amazonian forests have found that similarities in soil cation concentration and texture explain floristic similarities between sites, when these are measured using trees, pteridophytes or Melastomataceae. However, it is not known to what extent the three plant groups react to the same soil characteristics, because tree studies have almost always been conducted in different areas than studies on the understorey plant groups. We made inventories in 23 sites representing non-inundated rain forests on clayey to loamy soil in three regions of western Amazonia. Significant Mantel correlations between the floristic patterns of trees and pteridophytes were found in all three regions when floristic differences were measured with species presence–absence data. When species abundance data were used, and when the floristic patterns of trees and Melastomataceae were compared, significant correlations were found in one or two regions. Mantel correlations between plant groups were highest in the two regions where the observed variation in soil characteristics was largest. In all regions, the same soil variables emerged with significant Mantel correlations with trees, pteridophytes and Melastomataceae. Soil calcium and magnesium were most frequently retained in the models of multiple regression on distance matrices. On average, soil differences explained 50% of the variation in floristic differences (range = 14–84%), and geographical distances explained 16% (range = 0–64%). Our results demonstrate that beta diversities of the three plant groups are highly correlated, and that much of this congruence is explained by similar reactions to soil variation. These results support the idea that pteridophytes, and to a lesser degree Melastomataceae, can be used as indicators of general floristic and edaphic patterns in Amazonian rain forests. Since understorey plants are much quicker to inventory than trees, this would make it possible to recognize and map floristic patterns over huge areas of lowland Amazonia within a reasonable time.

We developed allometric relationships between tree size parameters (stem diameter at breast height (dbh), at ground surface (D0) and tree height) and leaf, stem, small-root (diameter <5 mm) and total root biomass in various tropical secondary-forest trees in Sarawak, Malaysia. In total, 136 individuals from 23 species were harvested to measure above-ground parts. Root systems of 77 individuals of 16 species were also excavated. The coefficients of correlation for the obtained allometric relationships between tree diameter and plant-part biomass showed high values, ranging from 0.83 to 0.99. In addition, there were few interspecific differences in relationships for all biomass parts, except for leaves. We also found relatively high coefficients of allometric relationships between tree height and plant-part biomass ranging from 0.83 to 0.94. Comparison of above- and below-ground biomass equations for various tropical rainforests implies that our allometric equations differ largely from the equations for tropical primary forests. Thus, choosing both above- and below-ground allometric equations for biomass estimation in tropical secondary forests of South-East Asia requires careful consideration of their suitability.

In tropical moist forests, length of the dry period may have a profound influence on leaf dynamics, plant growth and survival. To evaluate the role of light and water availability on seedling performance, a 1-y experiment was carried out in a tropical moist forest in the Bolivian Amazon in which seedlings of three tree species (Brosimum lactescens, Cedrela odorata and Schizolobium amazonicum) were planted in gaps and the understorey. Variation in length of the dry period was simulated by subjecting part of the seedlings to a water treatment at the end of the dry period. Gaps and understorey had a similar soil moisture content, which varied between 39% in the wet season and 16% in the dry season. Height and leaf growth rates were higher in gap compared to understorey plants, and in the wet compared to the dry season. A high growth during the wet season provided gap plants with a decisive size advantage over understorey plants during the dry season. Their larger root system allowed gap plants to explore a larger surface area and deeper soil layers for water. Consequently, gap plants of Cedrela experienced a shorter deciduous period (22 d) compared to understorey plants (61 d). Watering at the end of the dry season cued the flushing of new leaves by Cedrela, although it did not lead to a higher plant growth.

A high-diversity primate community consisting of 13 species was examined at an entirely undisturbed and remote terra firme forest near the headwaters of the Urucu River, Amazonas, Brazil. Twelve of these species used a 900-ha study plot, located 4 km or more inland from the river, and comprised a total group density of 16 groups km−2, an overall density of 146 individuals km−2, a crude community biomass of 381 kg km−2, and a metabolic biomass of 278 kg0.75 km−2. These species were occasionally found together within small (≥1 km2) areas of forest, so long as they included the full complement of available forest types. More frequently, however, only half the species co-occurred in a single habitat type at any one time, reflecting a relatively consistent pattern of subcommunity spatial organization. Habitat generalist species making extensive, yearround use of the predominant habitat type (high forest), were common, whereas habitat specialists were uncommon to very rare, and confined to minor habitats. Those species coexisting throughout the year in high forest were found to segregate vertically from one another. These and other modal patterns of community structure were disrupted seasonally by long-distance, inland movements by two ‘igapó’ forest species living in large groups – squirrel monkeys and white-face capuchins – to high forest areas, which apparently resulted from a food bottleneck in their habitat. Another species found in large groups – woolly monkeys – also exercised seasonally vagrant movements within the high forest matrix, which coincided with periods of greatest scarcity of mature fleshy fruits.

Maximum height was assessed for tree species from seven temperate deciduous forests, one subtropical forest and one tropical forest and combined with published tree heights for three other tropical forests. The temperate deciduous forests showed a strong concentration of canopy species and a dearth of subcanopy species. In contrast, the four tropical forests showed more uniform distributions of maximum heights, while the subtropical forest had an intermediate distribution. The tropical and subtropical sites had greater densities of small trees than did the temperate sites and most of these small trees were members of small-to medium-sized species. Sapling recruitment per unit stem basal area increased with declining maximum height in Panama, which is consistent with the criterion for coexistence of species of differing stature derived from Kohyama's forest architecture hypothesis. Greater penetration of light into gaps and favourable conditions for growth over most of the year may allow more smaller-statured species to coexist with canopy trees in tropical vs. temperate forests.

Fig pollinators (Agaonidae, Chalcioidea) lay their eggs in fig inflorescences (Ficus, Moraceae). Reproductive success for both partners is thus largely dependent on the dispersal of these tiny wasps. Some are known to cover substantial distances (> 10 km) using wind above the canopy. However, fig ecology is extremely varied, and hence one might also expect a diversity of pollinator dispersal strategies. We studied fig pollinator dispersal in Sarawak (2001 and 2004) and Peninsular Malaysia (2003). The results indicate substantial differences in dispersal ecology between the pollinators of monoecious and dioecious figs. Monoecious-fig pollinators were common, and species composition and rank abundances were similar between years despite short sampling periods. Substantial temporal and spatial variation in their production is thus smoothed out by long-distance dispersal. Some species whose hosts do not occur at our Sarawak site and are rare throughout Borneo were caught, suggesting exceptionally long-distance dispersal in these species. Conversely, few dioecious-fig pollinators were caught and species overlap between years was low. Dispersal range in many dioecious-fig pollinators may be more restricted. At a finer scale, among genera pollinating monoecious figs we found marked differences in flight behaviour (height and time-of-dispersal). We relate these findings to the ecology of their hosts, and discuss the implications for fig–fig-pollinator coevolution.

Floristic and structural comparisons and a numerical classification were performed to identify the priority areas for conservation of genetic resources in central Brazil. The main type of woody vegetation, cerrado sensu stricto (savanna woodland with 10–60% tree cover), was sampled under a uniform methodology in six selected sites (total sample of 6 ha) scattered in a physiographic unit called the Pratinha Plateau located between 15–20° S and 46–49° W. A total of 6971 trees belonging to 139 species and 42 families was measured. Each site had a combination of 50–80 species and approximately 30 families but only 22 species from 14 families were common to all sites. Similarity decreased with distance, indicating the existence of geographical gradients as the physical characteristics were relatively uniform and the soils were dystrophic at all sites. The most dissimilar sites, in the localities of Patrocínio-MG and Paracatu-MG were recommended for the establishment of conservation units. The already existent conservation units arc all clumped in the Federal District and are not enough to protect the genetic heritage of the plateau. Floristics and phytosociological surveys based on land system zoning were recommended as useful tools in planning conservation areas at a regional level to preserve the maximum biodiversity.

Estimates of insectivorous bat diversity in the Palaeotropics have largely been hampered by the lack of long-term studies employing appropriate capture techniques. Using a variety of trapping methods, 45 insectivorous bat species were captured in approximately 3 km2 of primary dipterocarp rain forest in Malaysia over 8 y. The cumulative site list for Kuala Lompat Research Station, Krau Wildlife Reserve, now stands at 51 insectivorous species. Although this is likely still not a complete list, it is already one of the most species-rich in the world. We attribute much of our success in recording this diversity to the extensive use of harp traps. Of the 45 species, 38 were captured in an intensive harp-trapping programme (> 1030 harp-trap nights) of the forest interior (22 species exclusively so). Insectivorous bats of the forest interior are thus a key component of Old World bat diversity, particularly in South-East Asia, and are dominated by taxa capable of detecting and capturing prey in cluttered environments (Hipposideridae, Rhinolophidae, Kerivoulinae and Murininae).

The relationship between seedling recruitment (under and away from parent trees) and the behaviour of seed dispersers and predators, was explored in a three-year study in Kibale National Park, Uganda. On the basis of 1261 hours of observation, the foraging activity of the diurnal frugivores which fed on fruit from six tree species was quantified. The fate of dispersed and non-dispersed seeds and seedlings was examined experimentally. The findings suggest that a trade-off exists between factors that promote seedling growth in areas with high seedling density and factors that promote dispersal by frugivores. For example, dispersal of Mimusops bagshawei increases both seed and seedling survival; seeds placed away from adult conspecifics had a 8% lower probability of disappearing than seeds placed under adults and seedlings away from adults had a 30% greater probability of surviving than seedlings grown under adults. In contrast, for Uvariopsis congensis, dispersed seeds had a 56% greater probability of disappearing than seeds directly under a parent tree, while the survival of dispersed and non-dispersed seedlings was similar. Non-dispersed seed and seedling disappearance were correlated with the percentage of the fruit crop removed from focal trees, suggesting that the ability to survive under an adult maybe related to other aspects of the tree's life history.

To examine the potential contribution of herbarium material to the description and analysis of tropical tree phenology, flowering times and geographical distribution were graphed for 1673 flowering collections from 18 species native to Neotropical dry forests and phenological differences between species were analysed. These include the timing and duration of flowering as well as morphological differences such as flowering on leafless twigs vs flowering on shoots bearing old or new foliage. Species-specific flowering periods of herbarium collections are similar to those observed in phenological field studies, but are often longer because of the larger geographical and temporal sampling range. Conspecific collections of different geographical origin show distinct differences in flowering periodicity, which are correlated with differences in the timing and intensity of the dry season. Interspecific differences in the timing of phenology relative to the dry season indicate differences in the control of phenology by seasonal drought. Herbaria thus represent a large potential source of phenological information which can either supplement and extend phenological field studies or provide phenological information for dry forest species not studied in the field but well represented in herbarium collections.

Current literature on the response of tropical tree seedlings to fertilization was analysed. Fertilization effects on biomass accumulation, relative growth rate and tissue nitrogen (N) and phosphorus (P) were examined for seedlings grown under relatively high light. Higher light conditions in gaps are expected as human disturbance expands in tropical landscapes. Two questions were asked: (1) Does nutrient addition enhance growth or result in nutrient accumulation in the tissues of tropical tree seedlings grown in pots under high light? (2) Are these responses conditional on their adaptations to light availability, seed mass or growth form? Most of the species (73% of light-demanding and 60% of shade-tolerant) demonstrated a positive growth response to fertilization. The magnitude of the response of light-demanding species was more than twice that of shade-tolerant species. A higher proportion of shade-tolerant species accumulated nutrients in their tissues (up to 95% for P), but over half the light-demanding species did so as well. For both types of species, a higher percentage accumulated P than N. Responses in growth and in P accumulation were negatively correlated with seed mass. Shrubs responded more than trees in biomass accumulation, but there was a trend toward greater response by trees in relative growth rate and N and P accumulation. The results suggest that variation in nutrient availability could affect the structure of tropical forests regenerating from large-scale disturbance. In fertile sites, competitive exclusion may occur within the light-demanding species, resulting in a decline in local tree diversity. Depending on the effects of human disturbance on nutrient availability, loss of tree diversity at a local scale may compound diversity loss due to habitat destruction at the landscape scale.

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.

Multiple species of primate disperse seeds and differentially contribute to the seed rain in tropical forests. The goal of this study was to examine seed dispersal by a primate community of five monkey and two ape species in the Dja Reserve, Cameroon. The density of primates in the reserve was calculated to be 77 individuals km-2. Analysis of 5789 faecal clumps demonstrated that 40% of monkey and 74% of ape faecal clumps possessed whole seeds. Six of the seven focal species acted as seed dispersers; faecal clumps passed by the black-and-white colobus (Colobus guereza) did not contain any whole seeds during the study. Seed passage trials on four captive monkey species showed monkeys to have an average seed passage time of 22 hours and defecation rate of five times per day. From the above results, the primate community was estimated to defecate 1129 seeds km-2 d-1. Seeds passed by the primate community came from 125 species of trees, lianas and shrubs, equivalent to at least 34% of the known tree flora. Rarefaction curves indicated that additional collection effort would identify more seed species passed by primates. Germination studies demonstrated that primate-passed seeds are viable. The number of seeds and plant species dispersed suggests that the primate community plays an important role in the maintenance of forest structure.

This study demonstrates that forest elephants (Loxodonta africana) play a key role as seed dispersal agents for the upper canopy forest tree, Balanites wilsoniana. Seed that passed through elephants had a much greater probability of germinating (50.7%) than seeds from fruits eollected directly from the tree (3%). Of 515 seeds that were collected from under parent trees, 89.1% were rotten at the time of collection, 2.6% had been eaten by forest pigs, 2.8% had been killed by squirrels, and 0.7% had been killed by insect damage. The remaining 4.8% of the seeds were still alive. In 150 hours of focal observations on B. wilsoniana trees, two fruits were seen to be destroyed by squirrels, no fruits were seen to be consumed by potential dispersers, and no evidence of scatter hoarding was found. Experimental seedling stations placed out under parent trees and away from parent trees indicated no significant effect of dispersal on seedling survival. Although elephants are not essential for the successful germination of B. wilsoniana seeds, they do greatly increase probability of germination and play a vital role in their dispersal, suggesting that this is a highly interdependent plant-animal seed dispersal system.

The influence of local precipitation and temperature on long-term growth dynamics in two species of seasonally dry tropical forest trees were investigated. Growth records were extracted from tree rings in Guanacaste province, Costa Rica. These chronologies provide a long-term (c. 85-y) record of tree growth for two species with contrasting phenologies. Annual growth, in both species, was dependent on annual and/or monthly variation in local precipitation but less so on temperature. For each species, however, patterns of growth reflected unique degrees of sensitivity to monthly rainfall and rainfall during previous years. It is hypothesized that such differences were due to the rooting depth of these species. A review of the literature also indicated similar diverse cambial growth responses by tropical trees to variation in annual and monthly climate. Lastly, it was shown that variation in longer term fluctuations in the Pacific and Atlantic oceans, as measured by the El Niño Southern Oscillation (ENSO) and North Atlantic Oscillation (NAO), significantly influenced local precipitation in Guanacaste only during the wettest portion of the wet season. Such temporal sensitivity may have differentially influenced the longer-term growth of some tropical tree species but not others. Together, these results support the hypothesis that tropical tree species respond individualistically to variation in local and regional climate and that some tropical assemblages may in fact be structured by species-specific differences in soil water-use.