A seed-removal experiment was conducted to assess the role of Proechimys semispinosus (Central American spiny rat) as potential predator and disperser of Astrocaryum standleyanum (black palm) seeds. One hundred fresh ripe A. standleyanum fruits were placed in semipermeable cages on each of 14 small islands in Panama, and seed disappearance rates were calculated for each island. Spiny rat density and biomass were determined by live-trapping on each island for 5 consecutive days and 4 nights. Censuses of fruiting trees were conducted on each island to control for effects of food availability on removal of palm fruits. Disappearance rates were related positively to total spiny rat density, density of adult and subadult spiny rats, and spiny rat biomass, but the density of fruiting trees accounted for very little variation. Spiny rats evidently prey heavily on A. standleyanum seeds, based on the high removal rates and on feeding observations of captive individuals. Since spiny rats may scatterhoard A. standleyanum seeds, they may also function as effective seed dispersers if seeds are removed to favourable germination sites unavailable to other seed predators. Results indicate that spiny rats, because of their abundance and wide distribution, may be important but overlooked predators and dispersers of A. standleyanum seeds and of other large-seeded tree species.

Bukit Belalong, Brunei, is a small mountain (913 m) of uniform shale lithology with continuous primary rain forest from its base to its summit. Three 0.25 ha plots were established at each of three altitudes (200, 500 and 850 m) to investigate the altitudinal zonation of the soils and the vegetation. One soil profile from each altitude is described and chemical analyses of the soils indicate that they are similar at all altitudes. In each plot all trees ≥10 cm dbh were measured and identified as far as possible. Maximum tree height was greatest at 200 m (60 m) and least at 850 m (33 m). Tree density (number of individuals ha−1) increased with altitude. The forest at 850 m was the most species- and family-rich, but since the ratio of species to individuals did not vary significantly among altitudes, the higher species richness is attributed partly to the larger number of trees sampled. The Dipterocarpaceae was the most important family in terms of basal area at all altitudes, but its proportion of basal area was much smaller at 850 m than at the other altitudes. The Dipterocarpaceae was the most diverse family at 200 m and 500 m, but at 850 m its diversity was exceeded by the Lauraceae, Myrtaceae and Euphorbiaceae and equalled by the Clusiaceae. The forests at 200 m and 500 m are classified as lowland rain forest and that at 850 m is classified as a lower montane forest.

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.

One of the major uncertainties concerning the role of tropical forests in the global carbon cycle is the lack of adequate data on the carbon content of all their components. The goal of this study was to contribute to filling this data gap by estimating the quantity of carbon in the biomass, soil and necromass for 23 long-term permanent forest plots in five life zones of Venezuela to determine how C was partitioned among these components across a range of environments. Aboveground biomass C ranged from 70 to 179 Mg ha−1 and soil C from 125 to 257 Mg ha−1, and they represented the two largest C components in all plots. The C in fine litter (2.4 to 5.2 Mg ha−1), dead wood (2.4 to 21.2 Mg ha−1) and roots (23.6 to 38.0 Mg ha−1) accounted for less than 13% of the total C. The total amount of C among life zones ranged from 302 to 488 Mg ha−1, and showed no clear trend with life zone. In three of the five life zones, more C was found in the dead (soil, litter, dead wood) than in the live (biomass) components (dead to live ratios of 1.3 to 2.3); the lowland moist and moist transition to dry life zones had dead to live ratios of less than one. Results from this research suggest that for most life zones, an amount equivalent to between 20 and 58% of the aboveground biomass is located in necromass and roots. These percentages coupled with reliable estimates of aboveground biomass from forest inventories enable a more complete estimation of the C content of tropical forests to be made.

Patterns of trap response in the Central American spiny rat (Proechimys semispinosus), a frugivorous forest rodent, were examined in relation to age, sex, density and resource abundance. Populations on four small islands (designated numbers 4, 8, 14, and 52) in Gatun Lake (central Panama) were sampled by live-trapping for four nights every month for four years. Trappability was calculated as the proportion of all individuals known to be alive that were actually captured. Mean body mass at first capture was within the range of subadult body masses and differed among islands but not between sexes. Trappabilities summed over the entire study were generally low and differed among islands and age classes (juveniles, subadults and adults) and between sexes. In general, adults were more trappable than juveniles and subadults, and females (particularly adults) were more trappable than males. Trappability and the number of captures per individual varied seasonally and were generally greatest during December and January. Monthly estimates of these two variables were examined for autocorrelation, and the order of the autoregressive error model was determined separately for each island population. The arcsine of trappability and the number of captures per individual were regressed on log10 densities of spiny rats and of fruiting trees and lianas after controlling for autocorrelation. Both trappability indices were negatively related to the density of fruiting trees and lianas on islands 8 and 14, indicating that spiny rats may be seasonally food-stressed. The number of captures per individual was related positively to the density of spiny rats on island 52 and negatively on island 8. Results indicate that spiny rat populations must be sampled with greater intensity and duration than temperate forest rodents.

Hong Kong is on the northern margin of the Asian tropics. The original forest cover was cleared centuries ago but secondary forest has developed since 1945 at many sites protected from fire and cutting. There are also older forest patches maintained behind villages for reasons of ‘feng shui’, the Chinese system of geomancy. All plants >2 cm dbh were identified and measured in forty-four 400-m2 plots. Detrended correspondence analysis showed a floristic continuum, with the montane sites (>500 m) most distinct and some overlap between lowland post-1945 secondary forest and the feng shui woods. The 30–40 year-old secondary forest is dominated by Persea spp. Montane forest is similar but lacks several common lowland taxa of tropical genera and includes more subtropical taxa. The feng shui woods have the most complex structure and contain some tree species not found in other forest types. Their origin and history is obscure but we suggest that both planting and selective harvesting have had a role in their current species composition

Damage to Hawaiian Acacia koa forest by Hurricane Iniki was assessed by comparison with our previous measures of stand structure and leaf area index (LAI) at sites along a precipitation/elevation gradient on western Kauai. Reductions in LAI ranged from 29 to 80% and were correlated with pre-hurricane LAI and canopy height. The canopy damage resulted in a large pulse of litter, ranging from 4 to 19 t ha-1 across our study sites. In the first year post-hurricane, LAI recovery and relative increase in stand basal area were negatively correlated with the fraction of canopy LAI lost. The two major overstorey species, A. koa and Metrosiderous polymorpha, were damaged less than the sub-canopy species, Dodonaea viscosa and Psidium guajava. Although D. viscosa and P. guajava were both severely damaged in the hurricane, the alien species P. guajava had much higher survival than the native D. viscosa, both as adults and as seedlings. However, seedling density of P. guajava was limited by low recruitment. At these sites, there was no drastic change in species composition following hurricane disturbance, and forest structure and productivity had recovered to a great degree within 2 y.

The amount of leaf litter fall produced by different tree species in tropical forests varies in space and time. Falling litter may cover seeds and thereby enhance their survival by making their detection by seed predators more difficult. Tests were made to determine whether Astrocaryum murumuru and Dipteryx micrantha seeds survive better in microsites covered by leaf litter. Seed numbers and litter cover on the forest floor were experimentally manipulated. How natural variation in leaf litter cover and thickness affects seed and seedling survival of these two plant species was also examined. Seed survivorship was significantly higher for both plant species in microsites with leaf litter than in those with bare soil. Results from an experiment in which the litter was not disturbed showed that Astrocaryum seed survival was positively correlated with litter thickness (defined as the number of overlying dead leaves). Astrocaryum seedling survival was also significantly affected by leaf litter; more seedlings survived in shallow litter. Leaf litter had no effect on Dipteryx seedling survival. The results of the study suggest that early recruitment of both Astrocaryum and Dipteryx is influenced by the spatial distribution and amount of forest leaf litter.

Similarities and differences among eight upland rain forest sites in Peruvian Amazonia were measured separately by using Melastomataceae, pteridophyte and tree species compositions and edaphic characteristics of the sites. All three plant groups showed a similar pattern among the sites, and this pattern could be explained by edaphic differences but not by geographical distances among the sites. The explicability of site-specific edaphic characteristics on the basis of geological history is discussed. The results suggest that both pteridophytes and Melastomataceae can be used as indicators of floristically different rain forest types that are edaphically defined. Distribution patterns of these plant groups can be studied much more rapidly than the patterns of trees and therefore both Melastomataceae and pteridophytes may be used in large scale phytogeographical studies that are urgently needed in the face of rapidly advancing deforestation.

The effects of herbivory by the leaf-cutting ant Atta laevigata Fr. Smith on the re-establishment of forest trees in an abandoned farm near Manaus, central Amazonia, were investigated. Experimental seedling transplants and observations on seedlings which emerged naturally in the study area showed that damage by leaf-cutting ants negatively affected tree seedling survival and growth. However, excluding leaf-cutting ants from experimental plots for 20 mo did not significantly increase tree seedling densities. The number of seedlings emerging varied considerably between plots and this obscured any effect the ants may have had on seedling survivorship. Taller seedlings, and seedlings attacked only once, suffered less mortality than smaller seedlings and seedlings attacked twice or more. In general, mortality was greater for those species preferred by the ants, indicating that selective herbivory by leaf-cutting ants affects tree species composition. The number of seedlings attacked by A. laevigata remained approximately constant throughout the period of this study in spite of the fact that the number available for attack increased. Thus, the chance of any individual seedling being attacked declined with time, suggesting that the effect of Atta herbivory on tree establishment is stronger during the first few years of forest regeneration.

The effects of escape distance to parental trees and tree clusters on the removal of Brazil-nut seeds (Bertholletia excelsa, Lecythidaceae) by vertebrate seed predators were examined in an entirely undisturbed stand of Brazil-nut trees of eastern Amazonia. Population density estimates, based on line-transect censuses, are also presented for Bertholletia trees and agoutis (Dasyprocta leporina), the most important scatterhoarder and seed predator of Brazil-nuts at this site. Seed removal experiments were conducted within and outside a natural Bertholletia tree cluster (castanhal) during both the wet and dry seasons. While there were no within-cluster effects of escape distance from parent trees on seed removal rates, overall seed removal within the cluster was significantly greater than that well outside the cluster. Moreover, removal rates in the wet season were consistently higher than those in the dry season both within and outside the tree cluster. Results suggest that the probability of early seed survival for Bertholletia, in relation to distance to seed sources, operates on different spatial scales, and that seed predators allocate greater foraging effort to scattered seeds during the fruitfall (wet) season, when buried seed stocks are being cached by agoutis.

As forest fragmentation and intentional burning of grasslands increase, the frequency of fires penetrating the dry and subhumid tropical forests of Bolivia is also likely to increase. To expand our understanding of the role of fire in tropical dry forest, the physical and thermal properties of barks of tree species were studied to determine their relative resistances to cambial damage by fire. For 16 tree species found in the dry forest of the Lomerío region of eastern Bolivia, bark thickness, moisture content, and specific gravity were measured. Insulating capabilities of bark were measured by obtaining cambial and surface temperatures during experimental wick fires. Bark thickness on trees 5-100 cm dbh (diameter at 1.4 m) ranged from 2–51 mm and both thick- and thin-barked species were represented. For all species, bark thickness increased as stem diameter increased. Bark thickness explained more (63%) of the variation in peak cambial temperatures during fires than did bark moisture content (4%) or specific gravity (1%). A threshold bark thickness of 18 mm was associated with the ability to withstand lethal cambial temperatures during the experimental, low intensity fires. For 13 of the 16 species included in this study, trees ≤20 cm dbh have bark thickness below the threshold 18 mm and, therefore, are likely to experience cambial injury from low intensity fires. Our results suggest that the forest presently characteristic of the Lomerío region did not develop with frequent fires and that species composition is likely to be substantially affected by an increase in fire frequency.

Two synthetic DNA probes composed of tandemly repeated ‘core’ sequences (GACA and GATA) were used to examine genetic diversity at multiple variable-number-tandem-repeat (VNTR) loci within and among four spatially isolated Philippine populations of Rubus moluccanus L. (Rosaceae), an uncultivated bramble widely distributed throughout southeast Asia – Malesia. Central goals were to determine whether apomictic propagation was detectable in R. moluccanus and to examine whether populations isolated on separate islands, or on mountain tops on a single island, were genetically differentiated. Sampling 22 individuals per population, the findings include: (1) no two individuals shared identical VNTR band profiles and thus apomictic propagation was not detected; (2) the mean number of bands scored per individual was 24.3 (SD = 3.91) and the proportion of polymorphic loci within populations ranged from 0.69 to 1.00 (mean = 0.86 ± SD = 0.085); (3) the average proportion of bands shared between individuals within populations ranged from 0.39 to 0.67 (0.50 ± 0.067), while average interpopulation similarity ranged from 0.21 to 0.50 (0.32 ± 0.092); and (4) estimated heterozygosity within populations ranged from 0.42 to 0.79 (0.62 ± 0.083), while interpopulation heterozygosity ran from 0.62 to 0.81 (0.74 ± 0.062). Permutation tests were used to estimate the statistical significance of differences in similarity between populations. A Luzon population isolated by 1200 km was always significantly different in similarity tests when compared with each of the other three populations, all located on Mindanao (and separated by at least 100 km but less than 250 km). Of the latter three, only one population differed significantly from the other two in the degree of VNTR markers shared, possibly reflecting biogeographic partitioning suggested for the island. Across all populations, average estimated Fst was 0.154, although mean inter-island Fst (0.224) was significantly higher than mean intra-island Fst (0.085). These results suggest that, while intra-island gene flow is relatively high, significant differentiation of tropical species may occur even over short distances on individual islands. Preservation of only a limited number of populations may result in a significant loss of genetic diversity in such species.

Striga asiatica (Scrophulariaceae) is an obligate root hemiparasite of mainly C4 grasses (including cereals). It is the most widespread of the 42 Striga species occurring in many semi-tropical, semi-arid regions of mainly the Old World. Examination of herbaria specimens revealed that S. asiatica has a wider geographical distribution, is present at higher altitudes and occurs in a more diverse range of habitats than previously reported. The host range is also larger than previously reported and is likely to include a large number of C3 plants. Morphology of examined specimens revealed variation in size and corolla colour suggesting the existence of ecotypes. Climate may exert a significant influence on the distribution of S. asiatica given the diversity of potential host plants and their distribution beyond the current recorded range of S. asiatica.

The role of regeneration strategies in structuring tropical tree communities has been a recent focus of attention of tropical ecologists. Canopy discontinuities created by tree-falls are considered to be critical to many rain forest trees for establishing and completing their life cycles. It was investigated whether it is advantageous for two common large-seeded and shadetolerant Amazonian tree species, the palm Astrocaryum murumuru and the legume, Dipteryx micrantha, to have their seeds dispersed to gaps. The dispersal process was simulated by placing experimental seeds and seedlings in gaps and in the shaded understorey, and then following their performance. Seeds and seedlings from the two species were used. The Cox proportional hazard model was used to estimate the survival functions and compare survivorship curves. Survival of Astrocaryum seeds, and of Dipteryx seeds and seedlings, was higher in two gap zones (crown and bole) than in the shaded understorey (both near and far away from a conspecific adult). A higher proportion of seedlings of both plant species survived within gaps than in the shaded understorey over an 18-wk period. One year after the seedlings had been transplanted into gaps these patterns still held. Naturally occurring Astrocaryum seeds survived better beneath conspecific adult trees located within 30 m of gaps than beneath trees located 100 m from gaps. Variation in gap size was significant for Dipteryx seeds, but not for Dipteryx seedlings and Astrocaryum seeds and seedlings. Dipteryx seeds survived longer in smaller than in larger gaps. By colonizing tree-fall gaps, these two species improved their chances of completing their life cycles.

The seed dispersal system of a neotropical tree, Cabralea canjerana (Meliaceae), was studied in two forested areas in southeastern Brazil. The first study site, Parque Estadual Intervales (PEI), is a 49,000-ha reserve composed mostly of old-growth Atlantic rain forest. The second site, Mata de Santa Genebra (MSG), is a 250-ha fragment of old-secondary semideciduous forest whose present bird fauna differs markedly from the original, in part as a consequence of forest fragmentation. At PEI 35 bird species ate the diaspores of C. canjerana. Black-tailed tityra (Tityra cayana, Tyrannidae) was the main seed disperser, but several other species were also important seed dispersers. In contrast, at MSG C. canjerana diaspores were eaten by 14 bird species. At this area, the red-eyed vireo (Vireo olivaceus, Vireonidae) was the most important seed disperser, but it was also a ‘waster’ which dropped seeds beneath parent plants, or carried them to sites unsuitable for germination. At PEI, exposed seeds on the forest floor were heavily preyed upon by rodents and insects. Insects destroyed mainly seeds deposited near to parent plants. Insect predation was less intense at MSG than at PEI. The rodent density at MSG was unusually small, and part of the post-dispersal seed predation may be done by terrestrial birds, such as doves and tinamous, which are especially common at MSG. Some of the differences recorded between the seed dispersal systems of C. canjerana at PEI and MSG may have been the result of the fragmentation and isolation of the latter area.

This paper presents data from a standardized series of line-transect censuses on the species and subspecies composition, population density, and crude biomass of western Amazonian primate communities occurring at eight flooded (= vázea) and 12 unflooded (= terra firme) forests. These were located primarily along one of the largest white-water tributaries of the Amazon (= Solimōes), the Juruá river. On average, terra firme forests contained twice as many primate species, lower population densities, and less than half of the total community biomass than did adjacent várzea forests. There was a clear habitat-dependent positive association among primate species, particularly within várzea forests, as well as marked shifts in guild structure between forest types. Species turnover between these two forest types involved primarily understorey insectivores (e.g. Saguinus sp.), which do not occur in seasonally inundated forest. These were consistently replaced by squirrel monkeys (Saimiri sp.), which are extremely abundant in annually flooded várzea forests. Similarly, large-bodied folivores such as red howler monkeys (Alouatta seniculus) were uncommon or rare in terra firme forests, but very abundant in várzea forests, even though they are hunted less intensively in the former than in the latter. This can be largely explained by the nutrient-rich alluvial soils of young floodplains, compared to the heavily weathered terra firme soils occurring even within short distances of major white-water rivers. This study clearly shows a reversed diversity/density pattern resulting from the lower species richness, but high overall community biomass of seasonally flooded Amazonian forests, which can now be generalized for a wide range of terrestrial vertebrate taxa, including amphibians, birds, and several other orders of mammals.

The fate of seeds of eight tree species was followed during 4 y of storage in the soil of an Afromontane forest at Gara Ades in the eastern highlands of Ethiopia. Seeds were enclosed in nylon mesh bags and buried at 5 cm soil depth. The bags were exhumed at intervals and the viability of the seeds was assessed by germination and cutting tests. Seeds of Bersama abyssinica and Ekebergia capensis germinated in the soil almost completely within a year after burial. The seeds of Juniperus procera, Olea europaea and Podocarpus falcatus also germinated to a substantial degree in the soil but with a distribution over several years, and some seeds of these species remained viable at the end of the 4-y period. Germination in the soil was very low in seeds of Acacia abyssinica and Croton macrostachyus throughout the whole burial period and the seeds kept their viability. In C. macrostachyus fresh seeds were highly dormant, but after 3 y or more in the soil they germinated readily in the laboratory suggesting an altered dormancy with time in the soil. Dormancy in seeds of A. abyssinica and Indigofera rothii was not altered throughout the study period as evidenced by marginal or no germination during incubation in the laboratory. The differential seed behaviour observed during storage in the soil can be an indicator of the regeneration strategy of the species studied. B. abyssinica, E. capensis, J. procera, O. europaea and P. falcatus form seedling banks on the forest floor and lack persistent soil seed reserves in contrast to A. abyssinica, C. macrostachyus and I. rothii which accumulate reserves of long-lived seeds in the soil. The generally high levels of dormancy and somewhat extended viability in the soil, even in several of the species producing seedlings in undisturbed forest, may have been selected for under a climate of seasonal drought and unreliable rainfall that characterizes this region.

Man-made grasslands dominated by Imperata cylindrica (L.) Beauv. in forested areas of lowland Nepal are commonly cut and/or burned annually. Changes in grass forage quality following different treatments of cutting and burning and axis deer (Axis axis) response to such habitat manipulations were investigated. Samples of matured grass were collected in December 1990, February and April 1991 from three experimental sites: cut, burned, cut-and-burned. Four locations on cut-and-burned grassland were repeatedly sampled at 12-d intervals from January to April 1992. Numbers of axis deer were recorded during the dry season of 1991/1992 on grassland plots receiving the following treatments: cut, cut-and-burned, and uncut/unburned (controls). Based on grass quality differences between December and February and between December and April, cut-and-burned treatments gave the greater increase in forage quality. N was significantly higher on cut-and-burned plots than on cut plots both in February and in April, while Na, K and P was significantly higher in February. On plots cut-and-burned in January, Ca concentrations were relatively low while the P content fell below required levels for domestic stock towards the end of the dry season in April. Na concentrations were below the minimum required levels for both domestic and wild ruminants during the whole period. When an entire grassland was cut, deer density increased gradually. When the same area was subsequently burned, the daily deer density increased much more rapidly. Axis deer preferred burned plots compared to plots neither cut nor burned and to cut plots. Plots burned in late February had higher densities of axis deer than plots burned 1.5 mo earlier. When nearby recently burned plots were available, deer density was reduced on plots burned earlier.