Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

Heterogeneity of Light Availability and its Effects on Simulated Carbon Gain of Tree Leaves in a Small Gap and the Understory in a Tropical Rain Forest1

To clarify the small-scale heterogeneity of light regimes in a rain forest, photosynthetic photon flux density (PFD) was measured at 1-min intervals during six days at 12 microsites in each of two plots, a small gap and an understory in Pasoh Forest Reserve, Peninsular Malaysia. Frequency distribution of microsite PFD was unimodal with the peak value between 16 and 32 μmol/m²/sec in the small gap, but between 8 and 16 μmol/m²/sec in the understory. In the small gap, PFD was more variable among microsites; total daily PFD and daily sunfleck PFD exceeding 10 μmol/ m²/sec tended to be higher (P <0.05; t-test) compared to those in the understory. Sunfleck PFD exceeding 50 μmol/ m²/sec, however, showed no difference between the two plots. Diffuse PFD transmittance, defined as the ratio of PFD in the forest to that measured at 43 m above ground during the periods 0800-0810 and 1750-1800 h, was significantly higher in the small gap than in the understory plot. Diffuse PFD transmittance was also positively correlated with microsite total daily PFD. To examine the effects of the subtle heterogeneity of light regimes on leaf carbon gain, we simulated carbon gain by sun and shade leaves in a typical shade-tolerant species, Brosimum aticastrum Sw. (Moraceae). Despite the similarity in total daily PFD, total daily carbon gain was considerably higher in the gap than in the understory for both sun and shade leaves. This study suggests that frequency distribution of PFD is critical in describing microsite PFD regimes and determining leaf carbon gain in the tropical forest floor.     

Continuous and Fluctuating Leaf Phenology of Adults and Seedlings of a Shade‐tolerant Emergent Tree,Dipterocarpus sublamellatus,in Malaysia

Continuous leaf phenology, with sequential production of metamers, is usually found in pioneer species, but is rare in shade‐tolerant species. Because of the nature of ‘continuous’ growth, continuous phenology has seldom been related to meteorological factors. We studied the leaf demography of seedlings and tall trees of a shade‐tolerant emergent species, Dipterocarpus sublamellatus (Dipterocarpaceae), in Malaysia. Although leaf phenology of D. sublamellatus was classified as continuous, the rates of leaf production and leaf fall were not constant and greatly fluctuated during the census period. Seasonal patterns of leaf production and fall were not synchronized across tall trees in most combinations or between seedlings and tall trees. Mean number of leaves produced per month and its seasonal fluctuation were greater in tall trees than in seedlings. Moreover, relationships between leaf phenology and meteorological factors (monthly rainfall, net radiation, maximum and minimum temperature, and minimum relative humidity) differed among trees. This suggests that endogenous rhythms, i.e., differences in allocation of resources to growth and storage among trees, are important to fluctuations in leaf phenology. These patterns are in contrast to patterns found in various pioneer species with continuous phenology, in which rates of leaf production and fall are generally constant. Different patterns might be caused by differences in allocation to growth and storage between pioneer and shade‐tolerant species, and might be related to their regeneration strategies.     

Non‐zero‐sum neutrality test for the tropical rain forest community using long‐term between‐census data

For community ecologists, “neutral or not?” is a fundamental question, and thus, rejecting neutrality is an important first step before investigating the deterministic processes underlying community dynamics. Hubbell''s neutral model is an important contribution to the exploration of community dynamics, both technically and philosophically. However, the neutrality tests for this model are limited by a lack of statistical power, partly because the zero‐sum assumption of the model is unrealistic. In this study, we developed a neutrality test for local communities that implements non‐zero‐sum community dynamics and determines the number of new species (N _sp) between observations. For the non‐zero‐sum neutrality test, the model distributed the expected N _sp, as calculated by extensive simulations, which allowed us to investigate the neutrality of the observed community by comparing the observed N _sp with distributions of the expected N _sp derived from the simulations. For this comparison, we developed a new “non‐zero‐sum N _sp test,” which we validated by running multiple neutral simulations using different parameter settings. We found that the non‐zero‐sum N _sp test rejected neutrality at a near‐significance level, which justified the validity of our approach. For an empirical test, the non‐zero‐sum N _sp test was applied to real tropical tree communities in Panama and Malaysia. The non‐zero‐sum N _sp test rejected neutrality in both communities when the observation interval was long and N _sp was large. Hence, the non‐zero‐sum N _sp test is an effective way to examine neutrality and has reasonable statistical power to reject the neutral model, especially when the observed N _sp is large. This unique and simple approach is statistically powerful, even though it only employs two temporal sequences of community data. Thus, this test can be easily applied to existing datasets. In addition, application of the test will provide significant benefits for detecting changing biodiversity under climate change and anthropogenic disturbance.     

Incidence of Extrafloral Nectaries and Their Relationship with Growth and Survival of Lowland Tropical Rain Forest Trees

Mutualistic relationships between organisms have long captivated biologists, and extrafloral nectaries, or nectar‐producing glands, found on many plants are a good example. The nectar produced from these glands provides food for ants, which may defend the plant from potential herbivores in turn. However, relatively little is known about their impact on the long‐term growth and survival of plants that produce them. To better understand the ecological significance of extrafloral nectaries, we examined their incidence on lowland tropical rain forest trees in Yasuní National Park in Amazonian Ecuador, and collated data from two other tropical lowland forest sites (Barro Colorado Island, Panamá and Pasoh Forest Reserve, Malaysia). At Yasuní, extrafloral nectaries were found on 137 of 1123 species censused (12.2%), widely distributed among different angiosperm families. This rate of incidence is high but consistent with other tropical locations. Furthermore, this study adds 18 new genera and two new families (Urticaceae and Caricaceae) to the list of taxa exhibiting extrafloral nectaries. Using demographic data from long‐term forest dynamics plots at each site, we compared the growth and mortality rates of species with extrafloral nectaries to those without. After controlling for phylogeny, no general relationship between extrafloral nectary presence and demographic rates could be detected, suggesting little demographic signal from any community‐wide ecological effects.     

Hyper‐abundance of Native Wild Pigs (Sus scrofa) in a Lowland Dipterocarp Rain Forest of Peninsular Malaysia1

This study reports extraordinarily high density estimates for the wild pig (Sus scrofa) from an aseasonal tropical forest site within the species'native range. At Pasoh Forest Reserve, a 2500 ha area of lowland dipterocarp rain forest in Peninsular Malaysia, line transects were used to estimate pig density from May to October in 1996 and 1998. In 1996, 44 sightings of S. scrofa consisting of 166 individuals were recorded along 81 km of transects. In 1998, 39 sightings documented 129 individuals along 79.9 km of transects. Estimated population density was 47.0 pigs/km² in 1996 and 27.0 pigs/km²‐ in 1998. Sus scrofa biomass in this forest was estimated at 1837 kg/km² and 1346 kg/ km² in 1996 and 1998, respectively. Differences between years were attributed to changes in the density of young pigs, coincident with a mast‐seeding year of dipterocarp trees in 1996. Pig densities at Pasoh Forest Reserve were much higher than at other forest locations within the species' native range in Europe and Asia. Because Pasoh Forest Reserve is a forest fragment, two factors likely account for the extremely high pig densities: (1) local extinction of natural predators (mainly tigers and leopards) and (2) an abundant year‐round food supply of African oil palm fruits from extensive plantations bordering the reserve.     

Clidemia hirta Invasion at the Pasoh Forest Reserve: An Unexpected Plant Invasion in an Undisturbed Tropical Forest1

Clidemia hirta is a highly invasive shrub in tropical forests throughout the world, but has had little success invading mainland sites and undisturbed forests. In the early 1990s, this plant was found to have invaded an unexpected site, an undisturbed continental tropical forest at Pasoh, Peninsular Malaysia. In 1997, a study was conducted of the C. hirta population at the Pasoh Forest Reserve. A demographic survey of the 50–ha long–term research plot at Pasoh located 1002 C. hirta individuals, 69 of which were reproductive at the time of the study. All but 8 individuals were located in high light gaps or gap edges. Relative growth rates were significantly higher in gaps and gap edges than in the understory, and no reproductive individuals were found in the understory. Mean plant size and dry biomass density increased steadily over the course of the study, while the observed mortality rate was 0 percent over two months. The biomass density of Clidemia at Pasoh was <500 g/ha at the conclusion of this study, but because the population is confined almost exclusively to high light environments, its density in these sites is much higher. The location of C. hirta plants in gaps was significantly correlated with past disturbance by wild pigs, suggesting that soil disturbance and light availability are essential for their establishment. The implication of this study is that by competing with native species in gaps, C. hirta invasion has the potential to alter forest regeneration at Pasoh. Changing land use practices near the reserve have increased the number of wild pigs, and thus the level of disturbance, which may explain the recent success of C. hirta at Pasoh.     

Branch architecture,light interception and crown development in saplings of a plagiotropically branching tropical tree,Polyalthia jenkinsii (Annonaceae)

To investigate crown development patterns, branch architecture, branch-level light interception, and leaf and branch dynamics were studied in saplings of a plagiotropically branching tree species, Polyalthia jenkinsii Hk. f. & Thoms. (Annonaceae) in a Malaysian rain forest. Lengths of branches and parts of the branches lacking leaves ('bare' branches) were smaller in upper branches than in lower branches within crowns, whereas lengths of 'leafy' parts and the number of leaves per branch were larger in intermediate than in upper and lower branches. Maximum diffuse light absorption (DLA) of individual leaves was not related to sapling height or branch position within crowns, whereas minimum DLA was lower in tall saplings. Accordingly, branch-level light interception was higher in intermediate than in upper and lower branches. The leaf production rate was higher and leaf loss rate was smaller in upper than in intermediate and lower branches. Moreover, the branch production rate of new first-order branches was larger in the upper crowns. Thus, leaf and branch dynamics do not correspond to branch-level light interception in the different canopy zones. As a result of architectural constraints, branches at different vertical positions experience predictable light microenvironments in plagiotropic species. Accordingly, this pattern of carbon allocation among branches might be particularly important for growth and crown development in plagiotropic species.