The intra-crop variation in the concentrations of major nutrients in the embryo-cum-endosperm (EE) fraction of seeds of 12 tree species of SE Asian lowland rain forest was investigated. Increased seed size was associated with a significant decrease in the EE concentration of N in 10 species, P and K in seven, Mg in six, and Ca in four. In contrast, the EE nutrient content increased with seed size, typically by about 1.6-fold where seed mass was 2-fold greater. In three species the allocation to protective tissues decreased with increased seed size. The results are potentially relevant to competition among young seedlings, optimal foraging by predators, allocation to defence by plants, and understanding the controls on filling of seeds.

ABSTRACT. Factors contributing to the resistance of gallery forests in savannas to the entry of fire were investigated using field observations and manipulation experiments. Mass of savanna fuels did not decrease close to forest boundaries, and in some instances increased, while savanna fuels adjacent to forests were moister than in the savanna beyond for only 1 d after rainfall. A fuel drying experiment conducted in both forest and savanna microclimates indicated that both fuel type and microclimate contributed to the resistance of forests to fire entry, although the former played a larger role. While savanna fuels in a savanna microclimate became ignitable in c. 1 d after rain, forest fuels in a forest microclimate required 4 wk to achieve ignitability. A further experiment juxtaposing forest fuels to burning savanna indicated that fire entry into forests was faciliated by deep root mats and the presence of a superficial litter layer, both of which become attenuated at the forest/savanna contact. It is concluded that fuels in these forests can reach an ignitable state late in the dry season, but that frequent fire entry is probably precluded by the tendency of savanna fires to occur earlier in the dry season and by discontinuities in fuels at the savanna/forest contact.

RESUMEN. Se investigaron los factores que contribuyen a la resistencia de la entrada del fuego a los bosques en galería de las savanas mediante observaciones de campo y experimentos de manipulación. La masa de los combustibles de la savana no decrecieron cerca de los bordes del bosque; por el contrario, en algunos casos se incrementaron, mientras que los combustibles de la savana cercanos al bosque eran mas húmedos que en la savana restante, solamente por un día despues de un evento de lluvia. Un experimento de secado de combustible, que fue conducido en los microclimas de bosque y de savana, indica que tanto el tipo de combustible como el microclima contribuyen a la resistencia de los bosques a la entrada del fuego, aunque el primer factor tuvo un papel de mayor importancia. Mientras que los combustibles de la savana en el microclima de la savana se volvieron ‘encendibles’ cerca de un día despues de lluvia, los combustibles del bosque dentro del microlima del bosque tomaron cuatro semanas para llegar a dicho estado. Un experimento que sobrepuso combustibles del bosque a la savana ardiente indicó que la entrada del fuego a los bosques se facilitó por la presencia de mantillos orgánicos profundos y de una capa superficial de hojarasca, presencia que se ve atenuada en la zona de contacto bosque/savana. Se concluye que los combustibles en estos bosques pueden alcanzar un estado de ignición tarde en la estación de sequía, pero que la entrada frecuente del fuego es probablemente evitada por la tendencia que tienen los fuegos de savana a presentarse temprano en la estación seca y por las discontinuidades de combustibles en la zona de contacto savana/bosque.

To explain very low fruit production in the tropical tree Ipomoea wolcottiana (Convolvulaceae), experiments and observations are presented regarding hypotheses that fecundity is limited by foliar herbivory, low soil phosphorus, inadequate cross-pollination, low rates of pollinator visitors, and parasitism of flower buds. A 2 × 2 factorial experiment was undertaken in two consecutive years which included control of herbivores with contact and systemic insecticides and addition of phosphorus. Phosphorus application in the second year increased the number of inflorescences and flowers by 200%, and the initiation of fruits by 400%. No response was observed for the number of mature fruits. Herbivore control and the addition of phosphorus increased seed weight by 22% in the first year, but could not be measured in the second year. Hand-pollination experiments increased the fruit set by 58% in the first year and by 75% in the second. Observations on visits to the flowers showed a six-fold difference between years in rates of pollinator visits. Parasitism of flower buds was 18% in the first year and 33% in the second. Several of the differences between years probably resulted from lesser and desynchronized flowering, due to heavy unseasonal rains in the second year. The differences in floral visitation and predation of floral buds were reflected in open fruit production: 16% in the first year and 6% in the second. All the factors investigated have important effects on the fecundity of I. wolcottiana. From the evolutionary point of view, the results suggest a complex interaction among selective forces which interact with the reproductive system. The theories of sexual selection and bet-hedging are the most plausible alternatives for explaining high levels of floral abortion in this species.

Influence of habitat on physiological and structural characteristics was investigated for broad-leaved tropical monocotyledons in the genus Heliconia (Heliconiaceae). Seven species were selected from three different light regimes, enabling an analysis of the extent to which this genus has adapted its photosynthetic strategies and morphological characteristics to different daily photon flux densities (PFD). Predictably, light response curves showed a clear gradient with respect to light saturation and rates of maximum net assimilation (Amax). Heliconia latispatha, an open site species, showed saturation at higher PFD (1400 μmol m−2 s−1) and higher Amax (14–16 μmol m−2 s−1) than H. mathiasiae, a forest edge species (PFD 1000 μmol m−2 s−1; Amax 7.5–8.5 μmol m−2 s−1) and H. irrasa of deep-shade forest understorey (PFD 250 μmol m−2 s−1; Amax 3.5 mol m−2 s−1). Leaf blade areas were largest in open sites, and leaf specific mass was also significantly higher, but leaf support efficiency was highest in understorey species. Species in open sites had thicker leaves with more chlorenchyma, whereas deep-shade species had very thin leaves and low stomatal densities. These rapidly growing herbaceous perennials appear to allocate much of their above-ground biomass to leaf tissues and have a relatively low investment in support tissues. This contrasts with understorey palms, in which leaf form and structural investment has been interpreted as a trade-off between economy and protection against tissue loss from falling branches. Presence of below-ground rhizomes in Heliconia may provide the key to this difference.

The effect of the change in vegetation structure from closed rain forest to tall open forest on the small mammal assemblage was studied by live trapping at three sites where the ecotone was very narrow (> 20 m) near the southern end of the Wet Tropics World Heritage Area of Australia. Habitat heterogeneity was significantly higher in the mixed open forest/ecotone area than in the adjacent rain forest. There was a large change in the struture of the small mammal assemblage coincident with the vegetation discontinuity. Although the species richness of small mammals was relatively constant across the gradient, the evenness and diversity of the assemblage declined across the transition from open forest into rain forest and biomass increased, largely due to the high abundance of Rattus fuscipes in the rain forest. The results suggest that the species richness of the small mammal assemblage was not determined by the spatial heterogeneity of the vegetation struture. The species composition of the rain forest is probably related to the historical biogeography of the area whereas the species richness of the wet sclerophyll forest is probably due to a mass-area effect from the adjcant large areas of rain forest and dry sclerophyll forest. However, the evenness, and therefore the diversity of the assemblage, was strongly affected by habitat heterogeneity.

The contribution of sprouts to the long-term survival of Chlorocardium rodiei, following logging was investigated. Differently aged sprout populations on cut stumps were surveyed to determine the numbers of sprouts produced, probability of survival and growth rates. Fifty-five to 70% of the stumps produced sprouts. The proportion of stumps that sprouted was lower for the hollow stumps than for the intact ones and the number of sprouts produced per stump was reduced. No significant correlation was found between stump diameter and the number of sprouts. Some sprouts survived for at least 15 y (usually one per stump), and these had a significantly larger average diameter relative to the sprouts at younger sites. The number of sprouts and the diameter of the largest sprout showed a significant positive correlation, suggesting that the maintenance of the inherited stump root system is shared among the sprouts so that the costs per sprout are reduced. We conclude that sprouting on cut stumps may have the potential to contribute to the maintenance of C. rodiei in the canopy at logged sites.

Secondary succession in a 19-y old tropical rainforest of French Guiana was investigated through the distribution pattern of primary forest species. The experimental plot of 25 ha was logged to simulate conditions of a paper pulp cut. Enumeration of all plants of height ≥ 50 cm in 52 subplots 10 m × 10 m indicated the number of species and individuals in five species categories; short-lived pioneer species, pioneer species that persist in mature forest, primary forest species capable of germinating in shade, primary forest species that indicate a preference for clearings (small gaps) in mature forest and understorey primary forest species. Change in proportion of the most abundant species (≥ 1 cm dbh) over the last 15 y indicated a decrease in short-lived pioneer species and an increase in pioneers that can persist in the mature forest. Proportion of species categories per height class indicated that primary forest species were mainly found in the lower height class in this forest which is largely dominated by secondary forest tree species forming a canopy that rarely exceeds 20 m. Individuals of primary forest species were encountered on ARBOCEL as sprouts from large stumps, sprouts from juveniles or as untraumatised saplings or juveniles. The distribution pattern in a 4-ha area of 11 selected primary forest canopy tree species revealed an overall trend in recolonisation of this species category in the secondary forest. Considered individually, the clustered distribution of stems of Eperua grandiflora showed the importance of regeneration through sprouting or remaining individuals. The gradient distribution, oriented from the edge to the centre of the studied plot, and displayed by the stem densities of Virola michelii, suggested a recent progressive dispersal of seeds. A more dynamic recolonisation front was observed for the primary forest understorey species Cassipourea guianensis and was probably linked to both the initial logging conditions and the fact that this species fruits when relatively young.

The relationship between mycorrhizal colonisation and phosphorus acquired by seedlings of the arbuscular mycorrhizal tree Oubanguia alata Bak f. (Scytopetalaceae) and the ectomycorrhizal tree Tetraberlinia moreliana Aubr. (Caesalpiniodeae) was evaluated at low and high inorganic phosphorus availability. AM colonisation was positively correlated with phosphorus uptake by O. alata at low, but not at high phosphorus availability. Seedlings growth was positively related to arbuscular mycorrhizal colonisation at both low and high phosphorus availability, suggesting that growth promotion by arbuscular mycorrhizas is not simply related to an increase of phosphorus uptake. In contrast, phosphorus uptake by T. moreliana was correlated with EM colonisation at both low and high phosphorus availability, but there was no relationship between growth and ectomycorrhizal colonisation. Promotion of phosphorus uptake by arbuscular mycorrhizas and ectomycorrhizas at low phosphorus availability is consistent with the co-occurrence of the two types of mycorrhiza in tropical rain forests where available soil phosphorus is low. However, ectomycorrhizal colonisation may also be of advantage where inputs of phosphorus rich litter raise the phosphorus status of the soil, as seen in the groves of ectomycorrhizal trees in Korup National Park, and may be one of the factors reinforcing local dominance by these trees.

This paper presents a method for estimating monkey numbers in a large area of forest where there is a gradient of monkey densities. The method is illustrated using data collected in the northeastern forests of Gabon during an earlier project. These forests are sparsely populated and there are few roads. The density of Cercopithecus nictitans increases with distance from the nearest road. A geographic information system (GIS) divided the forest into bands of increasing distance from the nearest road. The number of monkeys in each band is the product of the monkey density in that band and the area of the band. Summing across bands gives the population estimate; the standard error can be estimated by bootstrapping. The optimum sample size can be estimated by simulation. Combining estimates of the density gradient with a GIS is a cost-effective method of censusing primates in extensive forests.

Sympatric lizards in a transitional forest of Roraima, Brazil, dividing Amazon rain forest from savanna, contained a lizard assemblage of 16 species representing six families. Lizards varied in size, activity (diurnal versus nocturnal), microhabitats used, exposure to various conditions of light availability, prey types, and prey sizes. Overlaps in microhabitat occurrence varied from 0 (no overlap) to 1.0 (total overlap) whereas overlaps in prey types varied from 0.005 to 0.607. Microhabitat overlaps were higher overall than dietary overlaps. Pseudocommunity analyses on microhabitat data indicate that the community is not randomly assembled and that two distinct guilds exist, a leaf-litter guild and an arboreal guild, each with four species. Similar analyses on diet data revealed no apparent guild structure ast the first rank (nearest neighbour). Lizard diets did not differ from a random assortment based on prey type. At lower levels, the assemblage was structured with respect to food. Variation in prey use among lizard species was tied more closely to the effect of lizard body size on prey size (lizards ate different-sized prey items). Although exploitative competition among species may maintain structure within this assemblage it does not necessarily cause the observed differences. Several species are nearly identical ecologically to sister taxa in other environments and within different lizard assemblages suggesting that composition of the local assemblage limits the species that can enter the assemblage. Finally, lack of structure at the lowest (most similar) neighbour ranks may reflect the impact of a transitional habitat on stability of species interactions at the local level.

An exhaustive floristic inventory was conducted in a 400-ha block of tropical semideciduous forest in the Chiquitanía region of Santa Cruz, Bolivia. A total of 501 species were collected using both quantitative and traditional plant collecting methods. Trees represented the most diverse life form (124 spp), followed by herbs (101 spp), lianas (85), shrubs (66), herbaceous climbers epiphytes (15) and parasites (two). Floristic diversity was greatest for the forest floor community, when compared to either the understorey or canopy tree communities. Quantitative data were obtained for 336 species in 100 plots which contained a series of nested subplots to sample smaller-stature plants. Stem density for trees (dbh≥5 cm) was 914 trees ha−1, with a total density for all life forms estimated to be 135,000 plants ha−1; the total basal area for the forest was estimated to be 27.6 m2 ha−1. Species richness when measured by standard methods was found to be one of the highest for a dry forest region reported for the Neotropics with a mean of 70.8 spp 0.1 ha−1 (dbh ≥ 2.5 cm) and 50 spp ha−1 (dbh ≥ 10 cm). Comparison of life forms and vertical strata showed that the flora on the forest floor was more diverse than all other vertical strata combined. Habitat heterogeneity was studied using ordination procedures based on floristic data and to identify the characteristic species of three plant communities: granite outcrops, valley forest and upland forest. The most abundant tree species in the study area were Acosmium cardenasii, Neea hermaphrodita, Aspidosperma tomentosa and Galipea trifoliata, while the species with the greatest basal area were Anadenanthera colubrina, Acosmium cardenasii, Caesalpinia floribunda, Aspidosperma tomentosa, Piptadenia viridiflora, Chorisia speciosa, Tabebuia impetiginosa, Centrolobium microchaete, and Machaerium scleroxylon. Most canopy and understorey tree species had a population structure characterized by numerous juveniles and relatively few large trees, while emergent species tended to have a size-class distribution with relatively few juvenile individuals. Lianas and canopy trees were predominantly anemochorous, while understorey trees and shrubs were predominantly zoochorous; herbaceous species were largely autochorous, with only fern species relying on wind dispersal (sporochory). A comparison of structural attributes with other dry forest areas in the Neotropics, demonstrates the heterogeneous nature of vegetation types that are commonly assigned to this ecosystem. The Chiquitano dry forest shares many floristic elements with the semideciduous forests of the Andean piedmont of northwestern Argentina, the Misiones region of eastern Paraguay and northeastern Argentina, as well as the Caatinga region of northeastern Brazil.

Dispersal of seeds away from the parent plant may facilitate escape from density-dependent seed mortality. However, many post-dispersal events can have a profound influence on the survival of dispersed seeds. By incorporating seeds in the dung that dung beetles process for consumption and oviposition, dung beetles could enhance seed survival if they remove seeds from areas of high predation risk and place them in locations that avoid subsequent predation and that are suitable for germination. The role of dung beetles in seed survival was investigated over 15 mo in Kibale National Park, Uganda. Depths of seeds buried by beetles, levels of predation on buried and unburied seeds, and germination success of seeds buried to different depths were examined. Results suggest that by burying seeds dung beetles increase the probability that seeds will escape predation and germinate. Of seeds placed in dungpiles, 69% remained at the surface, while 25% were buried from 1–3 cm in depth. Larger seeds were buried more shallowly than smaller seeds. Buried seeds were less likely to be removed by predators than seeds at the surface. Germination of seeds buried at 1- and 3-cm depths was significantly higher than seeds buried at 10 cm. For the species tested, many seeds were buried by dung beetles between 1 and 3 cm and at this depth there was a high probability of escaping predators and germinating. This demonstrates the potential ecological importance of dung beetles in facilitating seed survival and provides data to consider the role of dung beetles in the evolution of seed attributes.

An analysis was conducted of floristic patterns contained in 48 1-ha tree plots distributed at 29 sites in seven neotropical countries, with a primary emphasis on the Amazonian region. Analyses were made with family level data, using detrended correspondence analysis and multidimensional scaling to generate two-dimensional ordinations. Dissimilarity values for all pairs of plots were then used to compare forest composition at both local (flooded vs unflooded forests) and regional scales (e.g., western vs central vs eastern Amazonia). The predominate family of trees in a large majority of Amazonian and Guianan forests (by number of stems) is either Palmae or Leguminosae (sensu latu), followed by Moraceae and Euphorbiaceae. The forests of western Amazonia are particularly rich in palms, Moraceae, and Myristicaceae, whereas those of eastern Amazonia and the Guianas are rich in Lecythidaceae and Chrysobalanaceae. Dissimilarity between sites increases with distance for both flooded and unflooded forests. The tree communities of flooded and unflooded forests within a region tended to resemble one another more closely than forests of either type resembled the homologous forests of the adjoining regions. Within Amazonia the edaphic properties of each region and its geological history are tightly interrelated. It is therefore difficult to distinguish between evolutionary and ecological interpretations of the results.

Patterns of buttress formation in tropical trees vary greatly within and among species. In Kibale National Park, Uganda, some form of a buttres was found on 23% of the 78 species (1785 trees) sampled from a variety of distantly related families. Large differences in buttress formation were documented within a single family and even within the same genus. Previous studies have suggested that buttresses are mechanical adaptations to counter asymmetric loads experienced during brief critical phases in a tree’s development and these persist after the need for a mechanical support has disappeared. As a tree grows from the understorey, up to the canopy, or emerges from the canopy, the potential number of occasions that a tree will experience directional stress increases. Many canopy level trees will probably have been in the vicinity of a treefall gap during their development, while emergent trees may experience gap exposure in addition to wind stresses associated with canopy emergence. Therefore, it is predicted that understorey trees should have fewer and less developed buttresses (after correcting for overall tree size) than canopy trees, which should have fewer and less developed buttresses than emergent trees. Detailed measurements of buttresses from 194 trees of eight species support this prediction. There was no evidence that trees thought to have experienced directional stress associated with selective logging almost 30 y ago had increased the number or size of existing buttresses. The pattern of buttressing in Kibale generally supports the idea that buttresses are mechanical adaptations to counter episodic asymmetric loads, and that buttresses persist after the need for a mechanical support has disappeared.

ABSTRACT. The spatial heterogeneity of element fluxes was quantified by measuring litterfall, throughfall and litter decomposition for 1 y in 30 randomly located sampling areas in a lowland dipterocarp rain forest. The idea tested was that turnover of elements is more variable than turnover of dry matter in a forest with extremely high tree species diversity. In spite of the low fertility of the soil (an ultisol), total litter production (leaves, trash, and wood <2 cm in diameter) was high (1105 g m−2 y−1) with inputs to the forest floor of carbon, nitrogen, phosphorus, calcium, magnesium, potassium, manganese and iron of 550, 15.3, 0.47, 6.26, 2.49, 4.75, 0.95 and 0.14 g m−2 y−1 respectively. Throughfall was 81% of the annual rainfall and transferred 22.2, 1.37, 0.14, 1.07, 0.67, 0.39, 7.92, <0.06, and <0.06 g m−2 y−1 of organic carbon, nitrogen (all forms), phosphorus, sulphur, calcium, magnesium, potassium, manganese and iron, respectively. Average turnover rates of nutrients in litter were highest for potassium and decreased in the sequence calcium, magnesium, carbon, nitrogen and phosphorus. Concentrations of phosphorus, nitrogen and potassium in litterfall, litter mass and topsoil were closely correlated with each other. Concentrations of calcium and manganese were positively correlated with each other and with elevation. Variations in leaf chemistry and total litterfall caused the spatial heterogeneity of element input to the forest floor to have a coefficient of variation of 30 — 70%, depending on the element. Due to the strong positive correlation between element fluxes and pools, the spatial variability of turnover rates (CV c. 20%) was lower than that of element input. Turnover rates for K varied by a factor of 4, and for Ca by a factor of 2.8 when the different sites were compared. The results strongly suggest that the most important factor determining spatial heterogeneity of organic matter and element dynamics on the forest floor is the site-specific amount of leaf fall, rather than spatially variable decomposition rates.

Seed dispersal is a process critical to the maintenance of tropical forests, yet little is known about the interactions of most dispersers with their communities. In the Dja Reserve, Cameroon, seed dispersal by the hornbills Certaogymna atrata, C. cylindricus and C. fistulator (Aves: Bucerotidae) was evaluated with respect to the taxonomic breadth of plants dispersed, location of seed deposition and effects on seed germination. Collectively, the three hornbill species consumed fruits from 59 tree and liana species, and likely provided dispersal for 56 of them. Hornbill-dispersed tree species composed 22% of the known tree flora of the site. Hornbill visit lengths, visit frequencies, and seed passage times indicated that few seeds were deposited beneath parent trees; in five hornbill/tree species pairings studied, 69–100% of the seeds ingested were deposited away from the parent trees. Germination trials showed that hornbill gut passage is gentle on seeds. Of 24 tree species tested, 23 germinated after passage by hornbills; of 17 planted with controls taken directly from trees, only four species showed evidence of inhibition of germination rate, while seven experienced unchanged germinated rates and six experienced enhanced germination rates. Results suggested that Certaogymna hornbill rank among the most important seed dispersers found in Afrotropical forests, and they deserve increased conservation attention. Ceratogymna hornbills are likely to become increasingly important in forest regeneration as populations of larger mammalian seed dispersers (such as forest elephants and primates) diminish.

The nutrient contents and accessions in litterfall over a period of 3 y are reported for undisturbed areas and at two sites disturbed by selective harvesting in tropical rain forest in North Queensland, Australia. Mean concentrations (mg g&supminus1; dry weight) of nutrients in litterfall ranged from 10 to 12 for nitrogen; 0.33 to 0.43 for phosphorus; 3.6 to 4.3 for potassium; 6.0 to 10.5 for calcium; and 1.7 to 2.6 for magnesium. These concentrations are in the middle to lower part of the spectrum of values recorded for tropical forests. Accessions of nutrients in litterfall (kg ha&supminus1; y&supminus1;) ranged from 59 to 64 N; 1.9 to 2.4 P; 20 to 24 K; 34 to 63 Ca; and 9 to 16 Mg. These rates, particularly for N and P, are among the lowest recorded for tropical forests. There were no consistent between-site differences in total nutrient accessions in small litterfall. In terms of the contribution of litterfall to the accessions of nutrients to the forest floor, this suggests that the logged sites have recovered from the effects of selective harvesting within 25 y. Nutrient accessions at each site were distinctly seasonal, with maximum accessions occurring in the late dry season to late in the wet season. Leaf-fall accounted for the largest proportion of nutrient accessions over the study period, although at certain times accessions in both reproductive material and wood were significant. A cyclone which crossed the coast near the study sites resulted in large nutrient accessions over a short period but had little effect on the total annual accession. A comparison with previous studies of litterfall in Australian tropical rainforests indicates that nutrient return in litterfall is directly related to soil fertility.

Geophagy (soil eating) by the forest elephant (Loxodonta africana cyclotis) and other mammal species in the Dzanga National Park, Central African Republic, has led to large treeless licks. The geological and physical properties of these licks were studied and the chemical characteristics and particle sizes of geophagical lick soils were compared with soils from the forest and from licks where evidence of geophagy was absent. Average lick size of 20 licks in the Hokou lick area was 12,900 m2 (SE = 3,400 m2). Licks were situated exclusively in areas of outcropping dolerite intrusions. Physico-chemical analysis of the geophagical lick soil showed significantly higher quantities of sodium, potassium, calcium, magnesium, phosphorus, manganese and clay compared with the other soils. Several samples of the geophagical lick soil had lower nutrient levels than the mean level of forest topsoil, indicating that nutrient supplementation is probably not the only reason for geophagy. The high clay content of the geophagical soil may help absorbing and combating secondary plant compounds. Pronounced geophagy in rainforests may be a sign for poor habitat quality for large mammals living in these ecosystems.

Many organisms create or alter resource flows that affect the composition and spatial arrangement of current and future organismal diversity. The phenomenon called ecosystem engineering is considered with a case study of the mound building termite Macrotermes michaelseni. It is argued that this species acts as an ecosystem engineer across a range of spatial scales, from alteration of local infiltration rates to the creation of landscape mosaics, and that its impacts accrue because of the initiation of biophysical processes that often include feedback mechanisms. These changes to resource flows are likely to persist for long periods and constrain the biological structure of the habitat. The value of ecosystem engineering is discussed as a holistic way of understanding the complexity of tropical ecology.