Effects of topography on tropical lower montane forests under different geological conditions on Mount Kinabalu,Borneo

Species composition and forest structure change with topography.However, mechanisms for topographical vegetation changes are still not wellunderstood, because a topographical gradient is a complex environmentalgradientinclusive of many factors. The foot of Mt. Kinabalu is covered with three typesof geological substrates, i.e. Quaternary and Tertiary sedimentary rocks andultrabasic (serpentine) rock. Quaternary and Tertiary sedimentaryrocks are different in site age, but controlled in primary minerals. Tertiarysedimentary and ultrabasic rocks are contrasting in primary minerals, but arecomparable in age. This setting provides an opportunity to examine thevegetation differentiation along topographical gradients that are contrastinginmajor-nutrient supply due to the difference in site age and parent rock.We established a total of nine study plots by choosing three topographicalunits(ridge, middle- and lower-slope) on each substrate inthe tropical lower montane forest. Pool size and supply of soil N and Pdecreased upslope on each substrate, and the magnitude of the reduction fromslope to ridge decreased in the order of Quaternary sedimentary > Tertiarysedimentary > ultrabasic rock. Between-substrate difference in soilnutrient condition was greater on the lower-slopes than the ridges.Maximum tree size decreased and stem density increased upslope on eachsubstrate. Detrended correspondence analysis demonstrated that speciescomposition also changed along topographical gradients on all substrates.However, the magnitude of topographical changes in forest structure and speciescomposition varied with substrate and decreased from Quaternary sedimentary>Tertiary sedimentary > ultrabasic rock. The greatest between-substratedifference in vegetation occurred on the lower-slopes. Accordingly, ourresults suggest that the magnitude of vegetation changes due to topographybecomes smaller with decreasing pool size and supply of nutrients.     

The activities of soil and root acid phosphatase in the nine tropical rain forests that differ in phosphorus availability on Mount Kinabalu, Borneo

Background and aims

Tropical rain forests on deeply weathered soils are increasingly thought to be limited by phosphorus (P), where plants and associated organisms would demonstrate adaptations to efficiently recycle P using acid phosphatase from organic matter. The activities of soil and root acid phosphatase were investigated in nine tropical rain forests that demonstrated a 20-fold difference in the soil organic P pool on Mt. Kinabalu, Borneo.

Methods

Acid phosphatase activity was measured at pH6.0 using p-nitrophenyl phosphate as substrate.

Results

The specific phosphatase activity of tree roots on a soil-surface-area basis was significantly positively related with P-use efficiency of above-ground productivity, suggesting a physiological linkage between above and below-ground systems in the adaptation to P deficiency. The phosphatase activities of soils and roots were significantly negatively correlated with the pool size of soil organic P fractions, suggesting that demand for P determines phosphatase activities.

Conclusions

It is suggested that tree roots and soil microbes develop more active phosphatases in response to the chronic shortage of soil P, which forms the basis for an important functional role for the efficient acquisition of P from soil organic matter.     

Ecological and biogeographical studies on the tropical rain forest of south Yunnan,SW China with a special reference to its relation with rain forests of tropical Asia

Abstract. Ecological and biogeographic analyses of the tropical rain forest in south Yunnan were made using data from seventeen sample plots and floristic inventories of about 1000 species of seed plants. The rain forest is shown to be a type of true tropical rain forest because it has almost the same profile, physiognomic characteristics, species richness per unit area, numbers of individuals in each tree species and diameter classes of trees as classic lowland tropical rain forests. As the area is at the northern margin of monsoonal tropics, the rain forest differs from equatorial lowland rain forests in having some deciduous trees in the canopy layer, fewer megaphanaerophytes and epiphytes but more species of lianas as well as more species of microphylls. In its floristic composition, about 80% of total families. 94% of total genera and more than 90% of total species are tropical, of which about 38% of genera and 74% of species are tropical Asian. Furthermore, the rain forest has not only almost the same families and genera, but also the same families rank in the top ten both in species richness and in dominance of stems, as lowland forests in southeast Asia. It is indisputable that the flora of the rain forest is part of the tropical Asian flora. However, most of the tropical families and genera have their northern limits in south Yunnan and most have their centre of species diversity in Malesia. More strictly tropical families and genera have relatively lower species richness and importance compared with lowland rain forests in tropical southeast Asia. Thus, the flora also shows characteristics of being at the margin of the tropics. Based mainly on physiognomy and floristic composition the tropical rain forest of Yunnan is classified into two types, i.e. seasonal rain forest and wet seasonal rain forest, the latter is further divided into two subtypes, i.e. mixed rain forest and dipterocarp rain forest. From analysis of geographic elements it is also shown that the tropical rain forest of Yunnan occurs at a geographical nexus with its flora coming mainly from four sources, i.e. Malesia, south Himalayas, Indochina and China.     

Soil nitrogen mineralization rates of rainforests in a matrix of elevations and geological substrates on Mount Kinabalu, Borneo

Mt Kinabalu, Borneo, is characterized by a deep elevational gradient and mosaics of geological substrates. We chose a pair of two geological substrates (sedimentary vs ultrabasic) at five altitudes (800, 1400, 2100, 2700 and 3100 m a.s.l.). We investigated soil nitrogen (N) mineralization and nitrification rates using an incubation technique to assay the pattern and control of soil N status in this environmental matrix. In situ net mineralization rates decreased with elevation on both substrates. The decreasing pattern was linear across altitudes on ultrabasic rock, whereas on sedimentary rock it was depressed in the middle slope wet cloud zone. Sedimentary sites in this zone had low soil redox potential values and this anoxic soil condition might be related to slow N mineralization. The in situ rates were significantly greater (P < 0.05, anova) on sedimentary than on ultrabasic rock at the same altitudes except in the cloud zone. Net mineralization rates of the soils that were collected from different elevations and incubated in the same conditions were statistically invariable (P > 0.05) among the original elevations for sedimentary rock, but were variable (P < 0.05) for ultrabasic rock. Those of the soils that were collected from the same elevation and incubated at different elevations decreased significantly across altitudes (P < 0.05) for sedimentary rock, while they were invariable (P > 0.05) for ultrabasic rock. Thus, temperature had stronger effects on net N mineralization on sedimentary rock, whereas inherent soil quality had stronger effects on ultrabasic rock. Controls of soil N mineralization might be different between the two substrates, leading to diverse biogeochemical site conditions on Kinabalu.     

Symbiotic nitrogen fixation in a tropical rainforest: 15N natural abundance measurements supported by experimental isotopic enrichment

* Leguminous trees are very common in the tropical rainforests of Guyana. Here, species-specific differences in N(2) fixation capability among nodulating legumes growing on different soils and a possible limitation of N(2) fixation by a relatively high nitrogen (N) and low phosphorus (P) availability in the forest were investigated. * Leaves of 17 nodulating species and 17 non-nodulating reference trees were sampled and their delta(15)N values measured. Estimates of N(2) fixation rates were calculated using the (15)N natural abundance method. Pot experiments were conducted on the effect of N and P availability on N(2) fixation using the (15)N-enriched isotope dilution method. * Nine species showed estimates of > 33% leaf N derived from N(2) fixation, while the others had low or undetectable N(2) fixation rates. High N and low P availability reduced N(2) fixation substantially. * The results suggest that a high N and low P availability in the forest limit N(2) fixation. At the forest ecosystem level, N(2) fixation was estimated at c. 6% of total N uptake by the tree community. We conclude that symbiotic N(2) fixation plays an important role in maintaining high amounts of soil available N in undisturbed forest.     

Importance of natural licks for the mammals in Bornean inland tropical rain forests

An intensive camera-trapping study and a nutrient analysis were carried out to understand how natural licks are important for mammals in inland tropical rain forests where soil cations are usually depleted. Using camera traps, we investigated the fauna, food habits, and the frequency of visitation by species at five natural licks in the Deramakot forest reserve, Sabah, Malaysia. All food-habit types of mammals (carnivore, herbivore-frugivore, insectivore, and omnivore), which included 29 (78.4%) of 37 species known in Deramakot, were recorded at the natural licks. The sambar deer, followed by the bearded pig, the lesser mouse-deer, the Malay badger, and the orangutan were the most commonly recorded species and represented 77.5% in terms of the frequency of appearance in all photographs taken throughout the year. These results indicated that, although the proportion of species recorded at the natural licks relative to the whole mammalian fauna of the forest was high, the frequency of visitation greatly varied among the species, and only a few species dominated. The frequency of visitation seemed to reflect both the density of species and the demand for the minerals, because some endangered, low-density species were more frequently recorded by cameras than expected—for example, the orangutan which was one of the top five species among natural-lick users. The natural licks with greater concentrations of minerals in seepage soil water were significantly preferred by the sambar deer and the bearded pig than those with lower concentrations of minerals. This result suggests that the chemical properties of soil water in natural licks determine the frequency of visitation of these herbivorous species that have strong demand for minerals.     

Effects of land use on 15N natural abundance of soils in Ethiopian highlands

We used the natural abundance of ¹⁵N in soils in forests, pastures and cultivated lands in the Menagesha and Wendo-Genet areas of Ethiopia to make inferences about the N cycles in these ecosystems. Since we have described the history of these sites based on variations in ¹³C natural abundance, patterns of δ¹⁵N and δ¹³C values were compared to determine if shifts of ¹⁵N correlate with shifts of vegetation. At Menagesha, a > 500-yr-old planted forest, we found δ¹⁵N values from −8.8 to +3.5‰ in litter, from −3.5 to +4.5‰ in 0–10 cm soil layer, and from −1.5 to +6.8‰ at >20 cm soil depth. The low δ¹⁵N in litter and surface mineral soils suggests that a closed N cycle has operated for a long time. At this site, the low δ¹³C of the surface horizon and the high δ¹³C of the lower soil horizons is clear evidence of a long phase of C₄ grass dominance or cultivation of C₄ crops before the establishment of the forest >500 years ago. In contrast, at Wendo-Genet, high δ¹³C of soils reveals that most of the land has been uncovered by forests until recently. Soil δ¹⁵N was high throughout (3.4–9.8‰), and there were no major differences between forested, cultivated and pasture soils in δ¹⁵N values of surface mineral soils. The high δ¹⁵N values suggest that open N cycles operate in the Wendo-Genet area. From the points of view of soil fertility management, it is interesting that tall forest ecosystems with relatively closed N cycling could be established on the fairly steep slopes at Menagesha after a long period of grass vegetation cover or cultivation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Structure,composition and species diversity in an altitude-substrate matrix of rain forest tree communities on Mount Kinabalu,Borneo

We studied forest structure, composition and tree species diversity of eight plots in an environmental matrix of four altitudes (700, 1700, 2700 and 3100 m) and two types of geological substrates (ultrabasic and non-ultrabasic rocks) on Mount Kinabalu, Borneo. On both substrate series, forest stature, mean leaf area and tree species diversity (both 4.8 cm and 10 cm diameter at breast height [dbh]) decreased with altitude. The two forests on the different substrate series were similar at 700 m in structure, generic and familial composition and tree species diversity, but became dissimilar with increasing altitude. The decline in stature with altitude was steeper on the ultrabasic substrates than on the non-ultrabasic substrates, and tree species diversity was generally lower on ultrabasic substrates than on non-ultrabasic substrates at 1700 m. The forests on non-ultrabasic substrates at higher altitudes and those on ultrabasic substrates at the lower altitudes were similar in dbh versus tree height allometry, mean leaf area, and generic and familial composition at 1700 m. These contrasting patterns in forest structure and composition between the two substrate series suggested that altitudinal change was compressed on the ultrabasic substrates compared to the non-ultrabasic substrates. Tree species diversity was correlated with maximum tree height and estimated aboveground biomass, but was not with basal area, among the eight study sites. We suggest that forests with higher tree species diversity are characterized by greater biomass allocation to height growth relative to trunk diameter growth under more productive environment than forests with lower tree species diversity.     

15N natural abundance of foliage and soil across boreal forests of Finland

This study presents the latitudinal variation (from 60° 30′ N to 68° 2′ N latitude) of natural abundances of ¹⁵N in the foliage, humus and soils of boreal forests of Finland. Our results clearly showed that N concentration of the foliage did not change significantly with latitudes but their ¹⁵N values were significantly higher in higher latitude sites relative to that of the mid and lower latitude sites, indicating the different forms of N uptake at the higher latitudes compared to the lower latitudes. We assume that the higher foliage δ¹⁵N values of the higher latitudes trees might be due to either more openness of N cycle (greater proportional N losses) in these latitudes compared to the sites of southern latitudes (lower N losses) or the differences in their mycorrhizal associations. Regression analysis showed that the temperature was the main factor influencing the ¹⁵N natural abundance of humus and soil of all forest ecosystems, both before and after clear-cut, whereas rainfall was the main controlling factor to the foliage ¹⁵N. Possible reasons behind the increasing δ¹⁵N natural abundances of plants and soils with increasing latitudes are discussed in this paper. The clear-cut did not show any specific trend on the ¹⁵N fractionation in humus and soil, i.e. both ¹⁵N-enrichment and -depletion occurred after clear-cut.     

Foraging in tropical rain forests: The case of the penan of Sarawak,East Malaysia (Borneo)

Bailey et al. (1989) and Headland (1987) have recently proposed hypotheses stating that human foragers are unable to live in undisturbed tropical rain forests without some reliance on cultivated foods. The present discussion considers these hypotheses, as well as some of the evidence by which they have been tested. Four conceptual problems in the way these hypotheses have been formulated are identified: (1) assumptions about the relationship between key features of tropical forest ecosystems and human subsistence potential, (2) in-consistencies in the definition of pure foraging, (3) adherence to a dichotomy between foraging and agriculture, the result being that conscious and unconscious effects of exploitation on the demographic parameters of key resources is ignored, and (4) problems in defining the significance of ecotones. I consider the case of Penan hunter-gatherers of Borneo, a population which, by virtue of their reliance on the sago palm Eugeissona utilis, contradicts the conclusions of Bailey et al. and Headland. I consider salient aspects of Penan reliance on Eugeissona, and describe how Penan exploitation of this resource may positively effect its availability. This case is seen to provide a challenge to the hypotheses of Bailey et al. and Headland, not only in the extent to which it contradicts their conclusions but, more significantly, in what it reveals about the assumptions upon which their hypotheses are based. This points to the need for greater precision in the definition of future hypotheses about foraging in tropical forests.     

15N natural abundance in forest and pasture soils of the Brazilian Amazon Basin

The natural abundance of ¹⁵N was examined in soil profiles from forests and pastures of the Brazilian Amazon Basin to compare tropical forests on a variety of soil types and to investigate changes in the sources of nitrogen to soils following deforestation for cattle ranching. Six sites in the state of Rondônia, two sites in Pará and one in Amazonas were studied. All sites except one were chronosequences and contained native forest and one or more pastures ranging from 2 to 27 years old. Forest soil ¹⁵N values to a depth of 1 m ranged from 8 to 23 and were higher than values typically found in temperate forests. A general pattern of increasing ¹⁵N values with depth near the soil surface was broadly similar to patterns in other forests but a decrease in ¹⁵N values in many forest profiles between 20 and 40 cm suggests that illuviation of ¹⁵N-depleted nitrate may influence total soil ¹⁵N values in deeper soil where total N concentrations are low. In four chronosequences in Rondônia, the ¹⁵N values of surface soil from pastures were lower than in the original forest and ¹⁵N values were increasingly depleted in older pastures. Inputs of atmospheric N by dinitrogen fixation could be an important N source in these pastures. Other pastures in Amazonas and Pará and Rondônia showed no consistent change from forest values. The extent of fractionation that leads to ¹⁵N enrichment in soils was broadly similar over a wide range of soil textures and indicated that similar processes control N fractionation and loss under tropical forest over a broad geographic region. Forest ¹⁵N profiles were consistent with conceptual models that explain enrichment of soil ¹⁵N values by selective loss of ¹⁴N during nitrification and denitrification.     

Natural abundance of 15N in tropical plants with emphasis on tree legumes

Natural abundance of ¹⁵N ( ¹⁵N) of leaves harvested from tropical plants in Brazil and Thailand was analyzed. The ¹⁵N values of non-N₂-fixing trees in Brazil were +4.5±1.9, which is lower than those of soil nitrogen (+8.0±2.2). In contrast, mimosa and kudzu had very low ¹⁵N values (–1.4+0.5). The ¹⁵N values of Panicum maximum and leguminous trees, except Leucaena leucocephala, were similar to those of non-N₂-fixing trees, suggesting that the contribution of fixed N in these plants is negligible. The ¹⁵N values of non-N₂-fixing trees in Thailand were +4.9±2.0. Leucaena leucocephala, Sesbania grandiflora, Casuarina spp. and Cycas spp. had low ¹⁵N values, close to the value of atmospheric N₂ (0), pointing to a major contribution of N₂ fixation in these plants. Cassia spp. and Tamarindus indica had high ¹⁵N values, which confirms that these species are non-nodulating legumes. The ¹⁵N values of Acacia spp. and Gliricidia sepium and other potentially nodulating tree legumes were, on average, slightly lower than those of non-N₂-fixing trees, indicating a small contribution of N₂ fixation in these legumes.     

Breeding birds in the tropical rain forests of Kibale National Park, Uganda

A combination of trapping and foot surveys was used to assess the breeding status of birds in unlogged and logged sites in the tropical rain forest of Kibale National Park, Uganda. Breeding of forest edge and gap species was greatly enhanced by logging, whereas crevice and hole nesting specialist breeders were adversely affected. Egg‐laying periods corresponded to higher rainfall regimes at the nearby field station. The relevance of these findings to bird conservation is discussed.     

A natural hybrid ofMonophyllaea (Gesneriaceae) in the tropical rain forests of West Sumatra

A natural hybrid (2n = 21) between the parapatric rain forest speciesMonophyllaea hirtella (2n = 20) andM. horsfieldii (2n = 22) (Gesneriaceae) has been observed at Sg. Lubuk Paraku, Padang, W. Sumatra. The hybrids showed intermediary characters between the parental species in the inflorescence structure, flower size and colour, indumentum, chromosome numbers, and habitat. The 29% pollen fertility of a single hybrid specimen suggests that the genetical isolation between the parental species may break down and reproduction over subsequent generations may occur.     

Habitat associations with topography and canopy structure of tree species in a tropical montane forest on Mount Kinabalu,Borneo

Habitat associations with topography and canopy structure of 42 abundant tree species were studied in a 2.74-ha plot of tropical montane forest on Mount Kinabalu, Borneo. Many of these species belong to the same higher taxa including eight families and four genera. Analysis of intraspecific spatial distributions for stems ≥ 10 cm diameter revealed that 28 species (including all six species of Fagaceae) showed aggregated distributions at the 100-m² and/or 400-m² scales, and that 20 species showed habitat associations with topography by torus-translation tests; 17 species showed both characteristics. Species' associations with the local canopy structure were characterized by crown position index (CPI), which was defined relative to neighbour trees. The CPI differed greatly among individual stems at 10–40 cm diameter, and 19 species showed significantly different frequencies of crowns exposed vertically versus those shaded beneath the canopy. Mean growth rates at 10–40 cm diameter and size distributions of species were not related to topographic associations, but were explained by the associations with canopy structure; species with more exposed crowns grew faster and had less positively skewed distributions. Diversity in habitat associations was manifest between two genera (Syzygium and Tristaniopsis) in the family Myrtaceae and among species in these genera, but was less evident in other families and two genera (Garcinia and Lithocarpus). This revised version was published online in June 2006 with corrections to the Cover Date.     

Simulating the effects of N availability, straw particle size and location in soil on C and N mineralization

Predicting the C and N mineralization of straw added to soil is important for forecasting subsequent soil N availability during and between crop growth cycles. The decomposition module of the STICS model, parameterized under optimal conditions, was used to predict straw decomposition in sub-optimal conditions, i.e. when contact between soil and residue was poor (due to large size residues or surface placement) or when mineral N availability was restricted. The data used in the simulations were obtained from published studies of effects of residue size, location and N availability on C and N mineralization from straw under controlled laboratory conditions. We selected studies in which the dynamics of C and N mineralization were measured simultaneously. The dynamics of straw mineralization could be well predicted by the model under optimal conditions with standard parameter values as derived from measured C/N ratios of the residues, but not under sub-optimal conditions which required a new parameterization. A good fit could be obtained on these treatments by a marked reduction in the rate constants of residue and microbial biomass decomposition and a marked increase in the microbial biomass C/N ratio. Our results show the need to include in decomposition models routines for simulating effects of spatial heterogeneity of residue distribution, different particle sizes and limiting N availability.