The standing live biomass of papyrus swamps is high compared with most other communities dominated by herbaceous vegetation. For the first time, measurements of below-ground rhizome and root biomass are reported here, in addition to measurements of shoot biomass, for a papyrus swamp located on Lake Naivasha, Kenya. The ratio of rhizome and root to shoot dry weight was 1.38: 1 and the total standing live biomass was 7.8 kg(dry weight) m−2. Detritus, formed from slowly decomposing papyrus, is normally the most important sink for carbon in this ecosystem. The amount of detritus retained by swamps varies but at L. Naivasha it contained almost 20 times the carbon found in living biomass. The net primary production of the swamp was calculated to be 6.28 kg(dry weight) m−2 y−1, which is amongst the highest recorded productivities for natural ecosystems. Model estimates of carbon input into the system from photosynthesis are incorporated into a carbon flow diagram for the swamp and this emphasises the importance of detritus as a sink for carbon in this ecosystem.

Frugivorous butterflies were studied in a set of forested islands (0.1 to 1.15 ha) in a reservoir in eastern Venezuela to investigate the effects of fragmentation and the resulting isolation on their abundance, diversity and species composition. While some islands showed reduced abundance and species diversity in comparison to unfragmented (or control) sites, others did not. Isolation status affected both butterfly abundance and diversity. Islands located close to their colonizing sources (0.1–1 km) tended to support similar densities of butterflies but lower numbers of species in comparison to control sites. Far fragments (1–3 km from their colonizing sources) tended to harbour lower butterfly densities in comparison to control sites but undiminished numbers of species. Species composition varied significantly between control sites and islands and amongst control sites, near islands and far islands. Interspecific differences were observed in species' responses to fragmentation. Charaxines, medium-sized satyrines, morphines and brassolines may be vulnerable to extinction after habitat fragmentation while small-sized satyrines may be relatively resistant. Observations during the dry season indicate that butterfly species may exist as mainland-island metapopulations in Lago Guri, in which small habitat fragments require recolonization every year from source populations in large islands and mainland habitat.

Frugivorous birds play an important role in parasitic mistletoe transmission among host species. The foraging behaviour and host selection of Psittacanthus schiedeanus mistletoe fruit-eating birds was studied in cloud forest remnants in Central Veracruz, México. Cedar waxwings, Bombycilla cedrorum, gray silky-flycatchers, Ptilogonys cinereus, and social flycatchers, Myiozetetes similis, eating ripe whole mistletoe fruits and defecating or regurgitating the seeds were observed. That variation in host species selection by frugivorous birds had a direct effect on mistletoe dispersion was hypothesized. To test whether mistletoe host species are visited in proportion to their abundance (total number of trees), or infested-tree abundance (total number of infested trees), or abundance of mistletoes (total number of individual mistletoe plants in each host species), the number of trees, infested trees and individual mistletoe plants were recorded. Tree abundance, infested-tree abundance and mistletoe abundance were different among host species. Frequency of birds' visits to host species varied depending on the tree, infested-tree, or mistletoe abundance. Liquidambar styraciflua was the most abundant host species with the highest number of infested trees and individual mistletoe plants, but Persea americana and Crataegus mexicana host species had the highest number of individual mistletoe plants in each tree. Our data suggest that L. styraciflua was the most selected host species by the three frugivorous birds. Host selection by birds is one behavioural factor in explaining differences in prevalence among host species, and variation in infection levels among individuals of one host species.

RESUMEN. Las aves frugívoras juegan un importante papel en la transmisión de muérdagos parásitos hacia sus árboles hospederos. La conducta de forrajeo y selección de hospedero de aves consumidoras de frutos del muérdago Psittacanthus schiedeanus fueron estudiados en remanentes de bosque mesófilo de montaña del Centro de Veracruz, México. Bombycilla cedrorum, Ptilogonys cinereus and Myiozetetes similis fueron observados consumiendo frutos maduros enteros y defecando o regurgitando las semillas. Hipotetizamos que la variación en la selección de la especie hospedera por las aves frugívoras tiene un efecto directo en la dispersión del muérdago. Para probar si las especies de árboles hospederos son visitadas en proporción a su abundancia (número total de árboles), o la abundancia de hospederos infectados (número de árboles infectados) o la abundancia de muérdagos (número total de plantas individuales de muérdago en una sola especie de árbol); el número de árboles, árboles infestados y plantas de muérdago fueron registradas. La abundancia de árboles, de árboles infestados y de plantas de muérdago fue diferente entre las especies de árboles estudiadas. La frecuencia de visitas de las aves a las especies hospederas varió de acuerdo a la abundancia de árboles, de árboles infestados y de plantas de muérdago. Liquidambar styraciflua fue la especie hospedera más abundante con el mayor número de árboles infestados y abundancia de muérdagos, pero las especies Persea americana y Crataegus mexicana presentaron el mayor número de plantas de múerdago por árbol. Nuestros datos sugieren que L. styraciflua fue la especie hospedera más seleccionada por las aves frugívoras. La selección de hospederos por las aves es uno de los factores conductuales que explican las diferencias en prevalencia entre especies de hospedero, y la variación en los niveles de infestación entre individuos de una de las especies de hospedero.

The trophic ecology of Konaino Creek, a small mountain headwater stream draining rainforest in the aseasonal tropics on Bougainville Island, Papua New Guinea, was examined and a food web was constructed. The major source of energy in Konaiano Creek was allochthonous detritus, most of which had been terrestrially degraded to fine particulate organic matter rather than entering the stream as leaf litter. This fine detritus was collected by the filter-feeders (mostly Simuliidae and also Hydropsychidae) which formed the dominant functional feeding group (64.4% of the fauna). Thus filterers processed most of the allochthonous detritus and made the energy available to other trophic levels, rather than shredders (1.7% of the fauna) which perform this role in temperate headwater streams. Collector-gatherers made up 22.7% of the fauna, carnivorses, mostly Odonata, Decapoda (crabs) and Hydrobiosidae, comprised 2.8% of the fauna and grazer-scrapers made up 7.4%. The latter were inhibited by low instream production owing to heavy shading and the instability and abrasion of the substrate due to frequent spates. In comparison, the trophic ecology of the nearby, coastal, Bovo River (with a catchment mainly in rainforest but mostly cleared with introduced species at the study site) was quite different and it was dominated by collector-gatherers (74%) and grazer-scrapers (15%).

Many studies suggest that edaphic variables are major determinants of frog distributions. However, leaf-litter depth and soil characteristics are influenced by distance from streams, so the apparent relationship between edaphic characteristics and species distributions could be an artefact of the dependence of species on free water for reproduction. Therefore, we investigated the effect of edaphic variables on the mesoscale distribution of frog species not dependent on free water for reproduction. We evaluated the effects of soil texture, pH, slope, number of trees and leaf-litter volume on the distribution of nine terrestrially reproducing anuran species in the Reserva Ducke, a 100-km2 terra firme forest preserve in central Amazonia. Diurnal and nocturnal assemblages of anuran species were sampled in 72 plots systematically distributed across the reserve. We sampled the diurnal anuran assemblage by visual encounter in 250 × 1-m plots and the nocturnal assemblage in 250 × 20-m plots using both auditory and visual surveys. The majority of terrestrially breeding anuran species were influenced by topographic and/or edaphic variables, such as slope, soil clay content and pH. However, responses to environmental predictors differed among species. Most species occurred throughout all environmental gradients and relationships with soil characteristics were subtle, indicating that these species occur in the majority of habitats in Reserva Ducke. The results of this study indicate that terrestrially breeding frogs are habitat generalists that show little mesoscale beta diversity associated with habitat variation.

The Monteverde Cloud Forest Reserve protects a variety of primary montane forest communities on volcanic parent materials. We describe the structure and composition of the forest to provide background information for epiphyte and nutrient cycling studies and for comparison with other tropical montane forests. In a 4-ha study plot in leeward cloud forest, density of stems (2062 individuals ha−1 for stems >2 cm dbh, 555 individuals ha−1 for stems > 10cm dbh) and stem basal area (73.8 m2 ha−1 for stems >2 cm, 62.0 m2 ha−1 for stems >10 cm dbh) were high relative to other montane forests. Stems in a subset of the plot (c. 1/3 of the area) were identified and assigned to 47 families, 83 genera and 114 species, which is rich compared with other montane forests. Large stems had a higher spatial variability of structural and floristic characteristics than small stems.

Fine roots and their turnover represent a dynamic aspect of below-ground biomass (BGB) and nutrient capital in forest ecosystems, and account for a significant fraction of net primary productivity (NPP) (Cuevas 1995, Vogt et al. 1990). On a weight basis, coarse roots contribute more to total ecosystem biomass than fine roots, but they account for only a small portion of annual root production (Eamus et al. 2002). Despite the fact that fine roots may compose less than 2% of total ecosystem biomass, they may contribute up to 40% of total ecosystem production (Vogt et al. 1990). Therefore, estimates of root production, like estimates of root biomass, should differentiate between coarse- and fine-root production.

The abundance, richness and species composition of frog, reptile, bird and mammal faunas varied along an extensive gradient from 470 to 1406 mm annual rainfall, and between three contrasting soil types (clay, loam and sand) in northern Australia. Patterns varied between and within vertebrate classes. In general, the most fertile soils in the highest rainfall sites supported the greatest species richness and abundance, but this association with fertility broke down at lower rainfall sites. Frogs were richest and most abundant at high rainfall sites, especially on clay soils, presumably because these had greatest water availability. Clay soils supported few reptile species, but these were often at relatively high abundance. High rainfall sites supported the richest reptile faunas. On sand and loam soils, bird species richness varied little along the rainfall gradient, but richness declined very substantially on clay soils. This was probably largely due to the far more marked vegetation structural change on clay soils than on other substrates. Few mammals were reported, and no clear trends were associated with either rainfall gradient or soil texture. Turnover in species composition along the rainfall gradient was gradual and limited on sand and loam soils, but far more marked on clay soils. There were few cases of replacements of ecologically comparable species along the gradient. These patterns reflect the disparate history, fragmentation and landscape positioning of clay soil environments, relative to the far more homogeneous eucalypt-dominated vegetation on sand and loam soils. Although comparable studies are lacking on other continents, patterns revealed here may be idiosyncratic due to the virtual monopolisation by eucalypts of the environment across the very extensive rainfall gradient.

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.

The dependent flora was surveyed on 20 trees at a 1.5-ha site in montane rain forest at 2600 m altitude in western Venezuela. Vascular species were recorded over the whole site and totalled 120 epiphytes, 21 climbers, 3 hemiepiphytes, 5 nomadic vines and 6 mistletoes. Non-vascular species were recorded within 95 sample plots and totalled 22 mosses, 66 liverworts and 46 macrolichens. The angiosperm species were restricted in geographical range to the Neotropics; 22.1% were endemic to Venezuela. Pteridophyte and bryophyte species were largely restricted to the Neotropics but few were endemic. Macrolichen species were mostly pantropical or cosmopolitan; only 9.6% were restricted to the Neotropics and none was endemic. Canonical Correspondence Analysis found the environmental variables most closely correlated with variation in community composition to be height above ground and a horizontal gradient reflecting differences in forest structure. The epiphytic vegetation was classified using Two-way Indicator Species Analysis into a Syrrhopodon gaudichaudii–Elaphoglossum hoffmannii group of lower trunks, an Omphalanthus filiformis–Maxillaria miniata group of intermediate levels and an Usnea–Parmotrema group of upper crowns. Diversity increased with height above ground; non-vascular diversity was greatest in upper crowns whereas vascular diversity was greatest at intermediate levels. Similarity levels were low among sample plots of the same community, whereas between-tree and between-stand similarities were relatively high.

This study demonstrates that elephant seed-dispersal is vital for Balanites wilsoniana (Zygophyllaceae), a forest canopy tree with no other effective dispersers. Non-dispersed seeds suffered high mortality (84%) and low germination (3%). Elephants were reliable at visiting fruiting trees (46%), ingesting available seeds (55%) and dispersing the fruit crop (26%). Compared with unpassed seeds, seeds passed through an elephant's gut had improved germination (54.9% vs. 2.9%) and reduced time to germination (82 vs. 132 d). Given predation rates under parent trees, elephant gut passage is expected to improve absolute germination by 66%. When this change in germination is divided by germination of unpassed seeds, elephant gut passage is expect to improve relative germination by as much as 4000%. Although post-dispersal predation occurred in dung containing B. wilsoniana seeds, dispersed seedlings were abundant. Compared with seedlings under parent trees, dispersed seedlings had greater survival and height. Dispersed seedlings survived in a diversity of environments, thrived under high-light conditions, and were negligibly affected by post-dispersal density. Although elephants are not essential for B. wilsoniana germination, their actions greatly increase seedling establishment and move a large proportion of seeds. This high degree of effectiveness, together with very low non-dispersed seed survival, provides strong evidence that B. wilsoniana is dependent on elephants for its long-term persistence.

The effect of tree canopy cover on soil properties was studied over three periods: middry season (January), mid-growth period (August) and peak-growth period (October) in three 1 ha plots in savanna of north-west Nigeria. The objective was to find out whether tree canopies change the nutrient status of the soil under them relative to adjacent grasslands. Soils under tree canopies were found to have significantly higher levels of organic matter, calcium, magnesium, potassium, total exchangeable bases, cation exchange capacity and pH than those in open grasslands. Nitrogen and phosphorus were slightly higher in soils under tree canopies than those in the open grasslands. Trees 7 m and above had more influence on soil properties than smaller trees. Differences in soil properties among the study plots were due to differences in their soil texture. Seasonal trends were observed in organic matter, carbon: nitrogen ratio, cation exchange capacity, phosphorus, calcium, sodium, total exchangeable cations and percentage base saturation.

A study of the benthic leaf litter community of the Tarumazinho, a central Amazonian stream (03° S, 60° W) is presented. This small tributary of the Rio Negro is of the blackwater type with a very low nutrient content. The sampling stations were towards the mouth of the tributary where it flowed through annually inundated forest and the work was carried out during the low water season. Previous studies concluded that these acid waters were of very low productivity and held impoverished faunas. However, this study showed that leaf litter banks held a diverse aquatic community of unexpectedly high biomass. This was because the banks supplied food and shelter and were available during the low water period when other habitats had dried out. The macrofauna was dominated by fish and shrimps. From an examination of gut contents a food web was produced. Primary consumers were found to take mainly detritus and fungi and it was concluded that allochthonous inputs were the main source of energy and nutrients. No feeding specialists were found and all the predators took prey from more than one trophic level.

The natural 15N abundance method for estimating symbiotic biological N2-fixation was tested on legume trees from two rain forests on contrasting soils (oxisols and spodosols) in French Guiana. When possible, the significance of N2-fixing species in the plant community was evaluated in terms of density, biomass and contribution of N2-fixation to the building up of the total nitrogen mass in the leaves. Of the two sites, the rain forest on spodosols was the less favourable for application of the δ15N method: the available soil nitrogen was isotopically similar to fixed-N2. Hence, the results showed that a reliable estimate of N2-fixation could not be obtained. A substantial contribution of fixed-N2 to the nitrogen nutrition of legumes was found on oxisols, with an average value of 54 % Ndfa (Nitrogen derived from the atmosphere). The contribution of the N2-fixing legumes to the biomass of the stand was estimated to be 2 t ha−1 for the leaf biomass and 136 t ha−1 for the total above-ground plant biomass. With 7.5 % of trees in the stand able to fix N2 (462 out of 6156), N2-fixation was estimated to be 7 kg ha−1 y−1. These results are the first use of the δ15N method to estimate nitrogen input by N2-fixing legumes to a natural rain forest. The inter-site variability observed in the δ15N of the non-fixing plants suggested different nitrogen-cycling processes in the two soils. The δ15N of the non-N2-fixing plants could be related to the soil nitrogen availability and be used as an indicator of efficient or non-efficient nitrogen-cycling rain forests. The spatial variability of the δ15N in the plant-available soil nitrogen pool and the nitrogen balance in tropical rain forests are discussed.

Germination and seedling resprouting capacity of the very large-seeded tree species Prioria copaifera Griseb. (Fabaceae) was studied in the seasonally moist forest of Barro Colorado Island, Panamá. Seeds with 60% of their cotyledonary mass removed did not suffer a reduced probability of germination compared to unmanipulated seeds and seeds infested with up to eight insect larvae germinated as well as uninfested seeds. Unmanipulated seeds were able to consecutively produce up to four functional resprout shoots after excision of the initial, and subsequent, fully expanded shoots. Even seeds with up to 60% of their reserves removed showed some capacity to resprout. Less than 10% of seeds we encountered in the field 2 mo after the end of the fruiting season were in a viable state, with the majority (55%) of mortality attributable to insect or pathogen damage. Of the 46 seeds that were viable, 30% had suffered partial removal of seed reserves similar to our manipulation treatments. These results indicate that P. copaifera seeds are capable of tolerating severe seed and shoot damage. The selective advantage conferred by damage tolerance may in part have contributed to the evolution of large seed size in this species.

Campos de murundus (earthmound fields) are common throughout the cerrado (savanna) in Brazil. They have a clearly defined distribution particularly where they are associated with ground water. There is a distinctive woody vegetation on the discrete mounds contrasting with surrounding grass-covered depressions.

After a reconnaissance ground and air-photograph survey of the nature and distribution of mounds within the Federal District, four one-hectare plots were chosen for detailed study within the ecological reserve of the University of Brasília. Measurements were made of the size, shape and frequency of mounds. Numbers varied from 26 to 61 ha-1 with a uniform distribution occupying 10–50% of the plots. Heights ranged from 0.05 to over 2 m. The generally semi-elliptical shape in ground plan had an average size of 7 by 5.5 m with no evidence of preferred orientation although water scouring from runoff partly influences the morphology. Volumes varied from 0.01 m3 to 141.5 m3. The soils of the murundus differed from those of surrounding depressions by being better drained, with bright colours and strong cohesive structures; they were more argillic with lower base saturation and pH values. Such sites favour colonization both by cerrado plants and by termites. A classification of the murundus is postulated. Although this paper does not consider the origins of murundus, they appear to relate more closely to drainage and differential erosion than to termite activity.

Diameter increments, recruitment and mortality were studied over a six-year period in a 3.02 ha sample in an area of 64 ha of undisturbed gallery forest alongside the Gama stream. The study site is located in Fazenda Água Limpa in Brasília, Federal District, Brazil. Trees ≥ 31 cm gbh (10 cm dbh) were measured every three years from 1985 in 151 (10 m × 20 m) permanent plots in Gama forest.

The mean annual diameter increment for trees ≥ 10 cm dbh was 0.25 cm y-1. Variability was high with coefficients of variation c. 100% or more. Light-demanding and large upper storey species tended to grow faster than shade-tolerant and lower storey species. Mean increments for the community increased with increasing diameter classes.

The mean annual mortality rate for the community was 3.5% y−1 while the recruitment was 2.7%. The community was dynamic with 86% of all species either suffering mortality or gaining recruits or both during the six years. The net difference between recruitment and mortality was zero for 55 species. The recruitment compensated for the mortality of most of the abundant species.

Although the pollination biology of many individual plant species has been investigated in the Oriental region, there have been very few community-level studies. The two most comprehensive of these were in the primary mixed dipterocarp forest of Lambir Hills National Park, Sarawak (4°20′N: Momose et al. 1998, Sakai et al. 1999) and in the warm temperate evergreen broad-leaved forest and cool temperate mixed forest on Yakushima Island (30°N: Yumoto 1987, 1988). Hong Kong (22°17′N) lies midway between these sites, at the northern margin of the tropics, where winter temperatures fall below 10 °C at sea-level for a few days every year and there are occasional frosts above 400 m (Dudgeon & Corlett 1994). Latitudinal effects, however, are compounded in comparisons with other well-studied East Asian sites, by centuries of massive human impact, leaving a degraded landscape of steep, eroded hillsides, covered in fire-maintained grassland, secondary shrublands and, locally, secondary forests (Zhuang & Corlett 1997). This history has left a relatively impoverished fauna but a surprisingly diverse flora, including 400 native tree species (Corlett & Turner 1997). In these circumstances, failures of pollination and dispersal mutualisms might be expected to accelerate the loss of plant species from the landscape (Bond 1994, Kearns & Inouye 1997). Previous studies have shown that most woody vegetation in Hong Kong is dominated by species whose seeds can be dispersed by the commonest avian frugivores, the light-vented and red-whiskered bulbuls (Pycnonotus sinensis (Gmelin) and P. jocosus (Linn.)) and the Japanese white-eye (Zosterops japonicus Swinhoe) (Corlett 1996, 1998), but there is no equivalent information available on pollination biology.

Miombo woodland in Zambia is divided into wetter and drier types by the 1100 mm mean rainfall isohyet. In the present study two wetter (Northern, Northwestern) and three drier (Central, Eastern, Western) miombo subtypes were recognized.

A survey of the woody flora of 94 old-growth and 58 coppice stands in the five miombo subtypes revealed some significant differences in species diversity among them. The floristic differentiantion of miombo subtypes was best achieved by the use of the Shannon species diversity index (H′) and species density (number of species 0.1 ha−1 or 0.4 ha−1). H′ in miombo decreased from 2.247 in Northwestern to 2.136 in Northern, 2.052 in Eastern, 1.918 in Central and 1.824 in Western subtypes. Although species density was not significantly different between wetter miombo subtypes, this decreased in a similar fashion from 18.78 0.1 ha−1 in Northwestern to 18.30 in Northern, 16.65 in Eastern, 14.79 in Central and 12.29 in Western subtypes. The low species diversity in Western drier miombo was attributed to the combined effects of low soil fertility and rainfall.

Although the mean number of species per 0.1 ha plot was higher in coppice samples, species richness (d) and index of diversity (H′) were lower than in old-growth samples. These results indicate lower species diversity in regenerating miombo. The dynamic aspects of species structure should therefore be considered in the future conservation of biological diversity in miombo woodland.