The types and causes of death of canopy and subcanopy trees were characterized for the tropical montane cloud forest of Tamaulipas, Mexico. Field work was carried out at the Biosphere Reserve ‘El Cielo’, located on the eastern slope of the Sierra Madre Oriental. Three study plots, each of 2 ha, were selected within the forest differing in their degree of exposure to wind. Results indicate that the tropical montane cloud forest presents a differential treefall regime derived from the great hurricane activity to which this mountain range is exposed. This effect is illustrated by the great number of fallen logs (545 trees in 6 ha) and the high percentage of uprooted (48.3%) and snapped off (37.6%) trees. Other treefall factors included the rocky and shallow soils and weakness of the dominant trees from insect attack. A differential disturbance level was obtained between zones. The disturbance was higher for the exposed zone, where the crown cover of emergent trees was not closed. Species composition of the dead material was related to the abundance of the overstorey species as standing vegetation. Ten of the 20 recorded species were able to sprout and just 7.5% of the 545 fallen trees were sprouting. Quercus germana, Q. sartorii, Clethra pringlei, Liquidambar styraciflua and Ceris canadensis, accounted for 78.5% of the gap-forming species. The type of treefall differed between species of the same genera: Q. germana was generally found uprooted, while Q. sartorii died standing probably because of previous weakening from dipterous insects.

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.

Seed-dispersal ecology in tropical montane forests (TMF) differs in some predictable ways from tropical lowland forests (TLF). Environmental, biogeographic and biotic factors together shape dispersal syndromes which in turn influence forest structure and community composition. Data on diaspore traits along five elevational gradients from forests in Thailand, the Philippines, Tanzania, Malawi and Nigeria showed that diaspore size decreases with increasing altitude, fleshy fruits remain the most common fruit type but the relative proportion of wind-dispersed diaspores increases with altitude. Probably corresponding to diaspore size decreasing with increasing elevation, we also provide evidence that avian body size and gape width decrease with increasing altitude. Among other notable changes in the frugivorous fauna across elevational gradients, we found quantitative evidence illustrating that the proportion of bird versus mammalian frugivores increases with altitude, while TMF primates decrease in diversity and density, and switch diets to include less fruit and more leaf proportionately. A paucity of studies on dispersal distance and seed shadows, the dispersal/predation balance and density-dependent mortality thwart much-needed conclusive comparisons of seed dispersal ecology between TMF and TLF, especially from understudied Asian forests. We examine the available evidence, reveal knowledge gaps and recommend research to enhance our understanding of seed dispersal ecology in tropical forests. This review demonstrates that seed dispersal is a more deterministic and important process in tropical montane forests than has been previously appreciated.

Temporal niche partitioning can be a viable mechanism for coexistence, but has received less attention than other niche axes. We characterized and compared patterns of activity, and overlap of temporal activity among the five common rodent species from a tropical semideciduous forest (TSF) and between the two common rodent species from cloud forest (CF) at El Cielo Biosphere Reserve in Mexico. Capture frequencies over 2-h intervals, obtained via live trapping (6850 trap-nights) in chosen months over 3 y formed the empirical basis for analyses. Trap transects were set from 19h00 to 07h00 and checked every 2 h. Analyses of 484 captures evinced two distinct assemblages. The TSF assemblage was diverse and with non-random temporal niche segregation, whereas the CF assemblage was depauperate with its two dominant species evincing the same activity pattern. Predator avoidance between open- and closed-microhabitat species, as well as niche complementarity may explain temporal segregation at TSF. This is the first documentation of assemblage-wide non-random temporal segregation of neotropical rodents. Time of activity may be a largely under-appreciated mechanism in other species-rich tropical rodent assemblages as well as in other species-rich biotas.

Understanding exotic–native plant relationships within the context of landscape-scale environmental factors such as rainfall, topography, disturbance and forest structure, is important for distinguishing the role of invasions in native plant community change. In this study, the relationship between Lantana camara invasion and native forest understorey vegetation was described after accounting for environmental influences. Rainfall, terrain slope, altitude and fire frequency were measured from GIS layers, and tree density, dry above-ground L. camara biomass, and native plant abundance and species richness were measured in nested plots (four 1 × 1-m plots for grass, tree seedlings and L. camara within one 5-m-radius plot for tree saplings and herbs and shrubs within one 10-m-radius plot for trees) at 80 locations distributed across Bandipur Tiger Reserve, a tropical deciduous forest in southern India. Relationships between environmental factors and L. camara abundance were described using a multiple regression. Further, the role of L. camara abundance in explaining residual variation in native vegetation, after accounting for environmental influences, was described using linear models. Lantana camara abundance decreased with increasing tree density, supporting the notion that it thrives in disturbed forest with fewer trees. Whereas native tree seedling and herb and shrub density showed no relationship to L. camara abundance, both tree sapling density and grass volume decreased under L. camara biomass exceeding 2 kg m−2. These data suggest that, by association with grass decline and decreased recruitment of tree saplings, L. camara may be linked to the gradual transition of these tropical deciduous forests into exotic-dominated shrubland, and overall native biodiversity loss.

The present study showed that the density of germinable seeds in the soil of a humid tropical forest ecosystem at Nelliampathy, in the Western Ghats of Kerala in India, fluctuates sharply at different times of the year. There were more germinable seeds in the soil during the monsoon season. Regeneration was activated after canopy gap formation and thus a decline in the soil seed density in 1-year-old gaps was recorded. However, soil seed density increased with gap age. Differences were found between primary, late secondary and early secondary categories of species with respect to their soil seed bank. There were more seeds of primary species in the soil of natural gaps than in selection felled ones, while the seeds of early secondary shrubs and herbs were more in the soil of selection felled gaps than in natural gaps. The soil seed banks of the primary forest species, being transient, are not an important conservation tool for these species.

An important challenge of invasion biology is to understand how interactions between species traits and ecosystem properties enable alien species to become invasive at particular locations. We investigated how gap dynamics in a tropical rain forest on the island of Réunion affected the invasiveness of alien plants. In the 12 000-m2 study area, alien plants occupied 24.9% of the area of gaps, which represented 5.62% of the forest area, but only 0.8% of the understorey area. The most abundant invasive species was Rubus alceifolius, which formed dense, monospecific stands in the largest gaps (> 25 m2). Although plants could persist in the shade, a germination experiment revealed that canopy openings were essential for seedling establishment. A cyclone that struck the study area in 2002 caused a temporary thinning of the canopy, increasing light levels to above the threshold needed for germination of R. alceifolius and also stimulating the growth of established plants. We conclude that the ability of this and other alien species to colonize intact lowland tropical rain forest is strongly influenced by the prevailing gap dynamics. Because gaps are also essential for the regeneration of many native trees in our study area, there is a real danger of the forest being progressively degraded by alien plants. There are no simple solutions to controlling species such as Rubus alceifolius, but efforts should be focused mainly upon the larger gaps where the species are most invasive.

Primates are among the major groups of frugivores in the tropics, but little is known about their role in natural regeneration of African savanna ecosystems. In the savanna-forest mosaic of north-eastern Ivory Coast the olive baboon (Papio anubis Lesson) harvests fruit from at least 79 plant species. Over a 24-mo study period, 396 faecal samples from 10 groups of baboons were analysed in terms of quantitative and qualitative aspects of seed handling and dispersal (excluding grasses and sedges). Seventy-three per cent of seeds in faecal samples were undamaged. Intact seeds were from 65 species. On average, defecations contained intact seeds from 2.0 species (range = 0–10). Seed size varied between 1 and 27 mm, and 77% of the species had medium-sized to large seeds. No linear correlation between mean seed size and seed damage was found. Eighteen out of 19 species tested were viable after ingestion, but effects of gut passage upon germination varied widely. The baboon population in the study area (145 km2) dispersed an estimated 1483 intact seeds d−1 km−2 (129 seeds > 2 mm in size). The results suggest that the olive baboon is an important seed disperser in the savanna-forest ecosystem of West Africa.

Comparisons of sympatric reptile species were used to assess the variation in niche overlap for food between potential competitors at different trophic levels. Omnivorous tortoises and carnivorous vipers inhabiting the rain-forest region of West Africa were used as study models. Food niche overlap between species increased with habitat alteration in both the independent study systems: tortoises (Kinixys homeana and Kinixys erosa) and vipers (Bitis gabonica and Bitis nasicornis) showed lower values of Pianka's niche overlap index in the pristine habitat than in the altered habitat, and these differences in overlap values did not depend on chance after Monte Carlo simulations. There were higher inter-habitat food niche overlaps within-species than between-species. Permutation tests (assessed after 5000 iterations) revealed that, for both study systems, the P-values became significantly smaller with fewer resource states, thus showing the niche overlap between species really increases after habitat alteration. The observed increases in food niche overlap between species accomplished with rain-forest habitat degradation in turn may be predicted to have cumulative effects on reducing the level of forest biodiversity.

Competition and predation have often been referred to as key forces regulating community structure. However, in the last decade great attention has been paid to the structuring role of positive interactions between plants, which occur when the presence of one species enhances fitness, survival or growth of another (Bruno et al. 2003, Callaway et al. 2002). Positive and negative interactions are unlikely to occur separately in nature (Brooker & Callaghan 1998, Holmgren et al. 1997), and the overall importance of positive interactions on community structure tends to be higher in resource-poor environments (Callaway et al. 2002). Moreover, this balance can be subjected to spatial and temporal shifts (Callaway 1998, Morris & Wood 1989, Tielbörger & Kadmon 1997) linked with plant ontogenetic development and/or changes on resource availability (see Callaway & Walker 1997).

Herbivory is important in tropical woody plants, but the few data available suggest that rates of herbivory are mostly low in epiphytes. We quantified herbivory at the community level in five bromeliad, three orchid and five fern species of a Mexican humid montane forest. Leaf area loss was <1.5% in bromeliads and orchids, but much higher (7–20%) in ferns. Damage was positively correlated with leaf nitrogen content but not with leaf life span. In contrast to low leaf damage, many bromeliads were infested by curculionid larvae feeding on the meristematic tissue at the ramet base, and we estimate that this accounts for 18 and 31% of ramet and shoot death of large individuals of Tillandsia punctulata and T. deppeana, respectively. Herbivory in flowers, capsules or inflorescence stalks reduced fecundity by c. 14–18% in three of the five bromeliads and by 90% in the orchid Lycaste aromatica, but had little effect on the other species. These data show that even if the leaf area consumed is indeed low in epiphytic orchids and bromeliads, the less conspicuous damage done to reproductive organs and meristematic tissue can have a strong effect on fecundity and survival.

A representative sample of waterways in Liberia was surveyed for crocodiles, and all three species of African crocodiles were observed. Contrary to previous reports, only Nile crocodiles inhabit the Monrovia mangrove swamps and brackish-water mouths of rivers; there are no slendersnouted or dwarf crocodiles in these habitats. There is partitioning of habitats among the three species in Liberia: Nile crocodiles in mangrove swamps and river mouths (brackish water); slendersnouted crocodiles in rivers through rain forest (freshwater); and dwarf crocodiles in small streams through rain forest (freshwater), some entering the adjacent river. Dwarf crocodiles utilize burrows in the stream banks.

Nowhere are crocodiles abundant, and there is an apparent absence of adults. Although 40% of Liberia is forested, deforestation is occurring rapidly by foreign timber companies and slash-and-burn agriculture practised by the largely rural population. Hunting with firearms, although illegal, is widespread, both for subsistence and commercially, without concern for depletion. Mangrove ecosystems near human population centres are being destroyed. The combination of the above factors has apparently depleted the crocodile populations in Liberia.

Using data from forest sites in Sarawak, East Malaysia, a demographic comparison is made of pioneer species in the genus Macaranga with understorey trees of the primary forest in the closely related genus Mallotus. In primary forest, saplings and trees of these genera arc uncommon and have low growth rates. For the six years following disturbance by logging, a Macaranga population was characterized by high rates of diameter growth and recruitment. Eleven years after disturbance, seedling and sapling abundance had declined sharply. In contrast, the Mallotus population was stable in size class distribution and increased slightly in total population size during the 11 years after logging. Growth rates of Mallotus trees increased after logging but were still much less than Macaranga trees.

A morphological comparison shows that most Macaranga species, in contrast with Mallotus species, have large hollow twigs that are occupied by ants and have larger, hairier, thicker, more toothed leaves with longer petioles, axillary inflorescences with more flowers, infructescences with more fruit, fruits that are smaller in size and smaller seeds.

Savannas intermingled with gallery forests are dynamic habitats typical in Africa. This study aims to determine if differences in species traits lead to non-overlapping distribution of gallery-forest and savanna species and abrupt transition between gallery forest and savanna. Tree species densities were measured in 375 plots of 1500 m2 covering a total sample area of 56.25 ha along forty 3-km transects located at right angles to a riverbed with gallery forest into surrounding savanna. Location, vegetation type, soil physical properties, erosion and fire occurrence were recorded as site factors. Data analysis included the quantification of co-occurrence patterns, threshold indicator taxa analysis and fuzzy set ordination. The gallery forest–savanna gradient predicted floristic composition of plots with a correlation of 0.595 but its accuracy was locally modified by the occurrence of fire and the physical properties of soil that covered more than 30% of the range of residuals. The distribution of gallery-forest and savanna tree species did not overlap. Along the gallery forest–savanna gradient, savanna species gradually increased in density while gallery-forest species showed a community threshold at 120 m from the river beyond the width of gallery forest. The forest species driving this trend should play an important role in the dynamics of gallery forest–savanna boundaries.

The ‘campos de murundus’ (carthmound fields) are typical landscapes of the cerrado (savanna) region of Central Brazil and are characterized by many rounded earthmounds called ‘murundus’ which are scattered over a grassland surface. As these areas are normally liable to seasonal water-logging, the earthmounds are true habitat islands capable of bearing woody plants typical of the cerrado, which are normally very intolerant of root inundation. The vegetation of an area of murundus field at Cuiaba, in the State of Mato Grosso, Central Brazil, was surveyed for the purpose of evaluating the island-effect on the plant community growing on the mounds/islands. Two main aspects were considered: the influence of the size of the mound and the differences between the types of cerrado occurring on the mounds and on the areas surrounding the murundus field. Most plant species occurring on the mounds were also present in the flora of the surrounding cerrado. However, only a fraction of these species was able to colonize the mounds successfully. The number of species on the mounds was found to be significantly correlated with mound area. Speciesarea relationships were better described with the power model S = C(Az) than with the exponential model S = C + z (logA) (fitted with linear regression). The mound size also had a strong influence on the spatial distribution of the species on the surface of the mounds. The species were organized in an increasing number of concentric topographic belts corresponding to a gradient of soil depth above the Hooding level.

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.

(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.

A trapping study of five mammal species in wet sclerophyll forest adjacent to rain forest in the Australian Wet Tropics was used to examine the seasonal diversity, abundance and dung-specificity of dung beetles associated with mammal dung. A total of 542 dung beetles from 11 species within three genera was recovered from beneath the traps of 1104 mammal captures. The diversity of beetles associated with the dung of the northern bettong (Bettongia tropica), a mycophagous marsupial, differed significantly from the diversity predicted by a null model. Numbers of beetles varied significantly with type of dung, indicating preference by beetles. Beetle numbers were related positively to a 1-mo lag in monthly mean minimum temperature and less strongly to maximum temperature and rainfall. Significantly more beetles per mammal capture were detected in the wet season than in the dry season. Dung beetles showed a strong preference for either the Eucalyptus woodland (six species) or the adjacent Allocasuarina forest (four species), with only one species occurring in both habitat types. Beetle species from the Eucalyptus woodland were typically only detected in the late wet and early dry seasons, while species in the wetter Allocasuarina forest were generally collected during the late dry and early wet seasons. A significant ‘checkerboard’ species effect was detected in both time and space in both habitat types, suggesting that competition for dung was strong.

Resolving the carbon (C) balance in the Amazonian forest depends on an improved quantification of production and losses of particulate C from forested landscapes via stream export. The main goal of this work was to quantify litterfall, the lateral movement of litter, and the export of coarse organic particulate matter (>2 mm) in four small watersheds (1–2 ha) under native forest in southern Amazonia near Juruena, Mato Grosso, Brazil (10°25′S, 58°46′W). Mean litterfall production was 11.8 Mg ha−1 y−1 (5.7 Mg C ha−1 y−1). Litterfall showed strong seasonality, with the highest deposition in the driest months of the year. About two times more C per month was deposited on the forest floor during the 6-mo dry season (0.65 Mg C ha−1 mo−1) compared with the rainy season (0.3 Mg C ha−1 mo−1). The measured C concentration of the litterfall samples was significantly greater in the dry season than in the rainy season (49% vs. 46%). The lateral movement of litter increased from the plateau (upper landscape position) towards the riparian zone. However, the trend in C concentration of laterally transported litter samples was the opposite, being highest on the plateau (44%) and lowest in the riparian zone (42%). Stream-water exports of particulate C were positively correlated with streamflow, increasing in the rainiest months. The export of particulate C in streamflow was found to be very small (less than 1%) in relation to the amount of litterfall produced.