We collected two specimens of an undescribed species of Lygosoma from pitfall traps in an urban rainforest in Kuching and from the base of a forested hill in western Sarawak, East Malaysia. The new species is diagnosable from all south-east Asian congeners by morphological characters, and most closely resembles Lygosoma herberti from the Thai-Malay Peninsula. The new species shows substantial molecular divergence from its closest relatives in two protein-coding genes, one mitochondrial (ND1) and one nuclear (R35) that we sequenced for several south-east Asian congeners. We describe the new species on the basis of this distinct morphology and genetic divergence. It is the third species of Lygosoma known from Borneo, and highlights the continuing rise in lizard species diversity on the island. In addition, the discovery of this species from a small urban rainforest underscores the importance of preserving intact rainforest areas of any size in maintaining species diversity.
Depreissia is a little known genus comprising two hymenopteran-mimicking species, one found in Central Africa and one in the north of Borneo. The male of Depreissia decipiens is redescribed, the female is described for the first time. The carapace is elongated, dorsally flattened and rhombus-shaped, the rear of the thorax laterally depressed and transformed, with a pair of deep pits; the pedicel is almost as long as the abdomen. The male palp is unusual, characterized by the transverse deeply split membranous tegulum separating a ventral part which bears a sclerotized tegular apophysis and a large dagger-like retrodirected median apophysis. The female epigyne consists of one pair of large adjacent spermathecae and very long copulatory ducts arising posteriorly and rising laterally alongside the spermathecae continuing in several vertical and horizontal coils over the anterior surface. Relationships within the Salticidae are discussed and an affinity with the Cocalodinae is suggested. Arguments are provided for a hypothesis that Depreissia decipiens is not ant-mimicking as was previously believed, but is a mimic of polistinine wasps. The species was found in the canopy in the Kinabalu area only, in primary and old secondary rainforest at 200-700 m.a.s.l. Overlap of canopy-dwelling spider species with those in the understorey are discussed and examples of species richness and endemism in the canopy are highlighted. Canopy fogging is a very efficient method of collecting for most arthropods. The canopy fauna adds an extra dimension to the known biodiversity of the tropical rainforest. In southeast Asia, canopy research has been neglected, inhibiting evaluation of comparative results of this canopy project with that from other regions. More use of fogging as a collecting method would greatly improve insight into the actual species richness and species distribution in general.
Tourism is associated with leisure and travelling, and is considered one of the fastest growing sectors that contribute significantly to the country’s economy, as well as bringing benefits directly to local communities. The Rural Tourism Master Plan was established in 2001, which defines rural tourism as an attraction that offers visitors the opportunity to experience Malaysia’s traditional culture and heritage away from urban areas. This study focuses on such tourism in the Gunung Mulu National Park, a famous natural park comprising underground caves, virgin rainforests and highlands. This research uses a quantitative approach with a random sampling technique because the procedure to select a unit for inclusion in a sample is much easier, quicker and cheaper. A questionnaire is distributed via Google Form to 500 local tourists who visited the park. The results of this research may be useful in marketing the Gunung Mulu National Park worldwide, besides fine-tuning programmes to attract tourists to the national park.
Rattans (subfamily Calamoideae) are an important component of the forests of the Old World. However, few studies have been conducted on the distribution of these abundant palms within different habitats, specifically in Peninsular Malaysia. This study was aimed at assessing rattan diversity, abundance and biomass change across two different habitat types, namely, dipterocarp forests and fresh-water swamps within the Segari Melintang Forest Reserve, Perak, within two years. All rattan stems within five 100 m × 100 m sized study plots (A-E) of the two habitat types were counted in 2011 and 2013, and Shannon-Wiener diversity indices (H') and Bray-Curtis dissimilarity indices were calculated. A total of 11 species from 5 genera (161 stems ha(-1)) were sampled. Rattan abundance was higher in all swamp plots; however, rattan diversity (H') was highest in the dipterocarp plot (D: H' (2011)1.79; H' (2013)1.84). Bray-Curtis indices of rattan abundance (highest similarity in swamp: plot BC(2011) 0.484, BC(2013) 0.262) and biomass were highest for study plots with the same vegetation types in both years. For rattan biomass, the most similar plot pairs changed during the years: dipterocarp plots A and D were most similar in 2011 (0.509), and swamp plots B and C were most similar in 2013 (0.282). This study helped contribute information regarding the distribution and dynamics of rattans in a primary rainforest of Peninsular Malaysia.
Managing tropical rain forests is difficult because few long-term field data on forest growth and the impact of harvesting disturbance are available. Growth models may provide a valuable tool for managers of tropical forests, particularly if applied to the extended forest areas of up to 100,000 ha that typically constitute the so-called forest management units (FMUs). We used a stand growth model in a geographic information system (GIS) environment to simulate tropical rain forest growth at the FMU level. We applied the process-based rain forest growth model Formix 3-Q to the 55,000 ha Deramakot Forest Reserve (DFR) in Sabah, Malaysia. The FMU was considered to be composed of single and independent small-scale stands differing in site conditions and forest structure. Field data, which were analyzed with a GIS, comprised a terrestrial forest inventory, site and soil analyses (water, nutrients, slope), the interpretation of aerial photographs of the present vegetation and topographic maps. Different stand types were determined based on a classification of site quality (three classes), slopes (four classes), and present forest structure (four strata). The effects of site quality on tree allometry (height-diameter curve, biomass allometry, leaf area) and growth (increment size) are incorporated into Formix 3-Q. We derived allometric relations and growth factors for different site conditions from the field data. Climax forest structure at the stand level was shown to depend strongly on site conditions. Simulated successional pattern and climax structure were compared with field observations. Based on the current management plan for the DFR, harvesting scenarios were simulated for stands on different sites. The effects of harvesting guidelines on forest structure and the implications for sustainable forest management at Deramakot were analyzed. Based on the stand types and GIS analysis, we also simulated undisturbed regeneration of the logged-over forest in the DFR at the FMU level. The simulations predict slow recovery rates, and regeneration times far exceeding 100 years.
Dryobalanops aromatica Gaertn. f. is a major tropical canopy species in lowland tropical rain forests in Peninsular Malaysia. Diurnal changes in net photosynthetic rate (A) and stomatal conductance to water vapor (g(s)) were measured in fully expanded young and old leaves in the uppermost canopy (35 m above ground). Maximum A was 12 and 10 micro mol m(-2) s(-1) in young and old leaves, respectively; however, because of large variation in A among leaves, mean maximum A in young and old leaves was only 6.6 and 5.5 micro mol m(-2) s(-1), respectively. Both g(s) and A declined in young leaves when T(leaf) exceeded 34 degrees C and leaf-to-air vapor pressure deficit (DeltaW) exceeded 0.025, whereas in old leaves, g(s) and A did not start to decline until T(leaf) and DeltaW exceeded 36 degrees C and 0.035, respectively. Under saturating light conditions, A was linearly related to g(s). The coefficient of variation (CV) for the difference between the CO(2) concentrations of ambient air and the leaf intercellular air space (C(a) - C(i)) was smaller than the CV for A or g(s), suggesting that maximum g(s) was mainly controlled by mesophyll assimilation (A/C(i)). Minimum C(i)/C(a) ratios were relatively high (0.72-0.73), indicating a small drought-induced stomatal limitation to A and non-conservative water use in the uppermost canopy leaves.
Climate change exposes vegetation to unusual levels of drought, risking a decline in productivity and an increase in mortality. It still remains unclear how trees and forests respond to such unusual drought, particularly Southeast Asian tropical rain forests. To understand leaf ecophysiological responses of tropical rain forest trees to soil drying, a rainfall exclusion experiment was conducted on mature canopy trees of Dryobalanops aromatica Gaertn.f. (Dipterocarpaceae) for 4 months in an aseasonal tropical rain forest in Sarawak, Malaysia. The rainfall was intercepted by using a soft vinyl chloride sheet. We compared the three control and three treatment trees with respect to leaf water use at the top of the crown, including stomatal conductance (gsmax), photosynthesis (Amax), leaf water potential (predawn: Ψpre; midday: Ψmid), leaf water potential at turgor loss point (πtlp), osmotic potential at full turgor (π100) and a bulk modulus of elasticity (ε). Measurements were taken using tree-tower and canopy-crane systems. During the experiment, the treatment trees suffered drought stress without evidence of canopy dieback in comparison with the control trees; e.g., Ψpre and Ψmid decreased with soil drying. Minimum values of Ψmid in the treatment trees decreased during the experiment, and were lower than πtlp in the control trees. However, the treatment trees also decreased their πtlp by osmotic adjustment, and the values were lower than the minimum values of their Ψmid. In addition, the treatment trees maintained gs and Amax especially in the morning, though at midday, values decreased to half those of the control trees. Decreasing leaf water potential by osmotic adjustment to maintain gs and Amax under soil drying in treatment trees was considered to represent anisohydric behavior. These results suggest that D. aromatica may have high leaf adaptability to drought by regulating leaf water consumption and maintaining turgor pressure to improve its leaf water relations.
Clarifying the dynamics of fine roots is critical to understanding carbon and nutrient cycling in forest ecosystems. An optical scanner can potentially be used in studying fine-root dynamics in forest ecosystems. The present study examined image analysis procedures suitable for an optical scanner having a large (210 mm × 297 mm) root-viewing window. We proposed a protocol for analyzing whole soil images obtained by an optical scanner that cover depths of 0-210 mm. We tested our protocol using six observers with different experience in studying roots. The observers obtained data from the manual digitization of sequential soil images recorded for a Bornean tropical forest according to the protocol. Additionally, the study examined the potential tradeoff between the soil image size and accuracy of estimates of fine-root dynamics in a simple exercise. The six observers learned the protocol and obtained similar temporal patterns of fine-root growth and biomass with error of 10-20% regardless of their experience. However, there were large errors in decomposition owing to the low visibility of decomposed fine roots. The simple exercise revealed that a smaller root-viewing window (smaller than 60% of the original window) produces patterns of fine-root dynamics that are different from those for the original window size. The study showed the high applicability of our image analysis approach for whole soil images taken by optical scanners in estimating the fine-root dynamics of forest ecosystems.
Limited knowledge about vertical variation in wood CO2 efflux (Rwood) is still a cause of uncertainty in Rwood estimates at individual and ecosystem scales. Although previous studies found higher Rwood in the canopy, they examined several tree species of similar size. In contrast, in the present study, we measured vertical variation in Rwood for 18 trees including 13 species, using a canopy crane for a more precise determination of the vertical variation in Rwood, for various species and sizes of trees in order to examine the factors affecting vertical variation in Rwood and thus, to better understand the effect of taking into account the vertical and inter-individual variation on estimates of Rwood at the individual scale. We did not find any clear pattern of vertical variation; Rwood increased significantly with measurement height for only one tree, while it decreased for two more trees, and was not significantly related with measurement height in 15 other trees. Canopy to breast height Rwood ratio was not related to diameter at breast height or crown ratio, which supposedly are factors affecting vertical variation in Rwood. On average, Rwood estimates at individual scale, considering inter-individual variation but ignoring vertical variation, were only 6% higher than estimates considering both forms of variation. However, estimates considering vertical variation, while ignoring inter-individual variation, were 13% higher than estimates considering both forms of variation. These results suggest that individual measurements at breast height are more important for estimating Rwood at the individual scale, and that any error in Rwood estimation at this scale, due to the absence of any more measurements along tree height, is really quite negligible. This study measured various species and sizes of trees, which may be attributed to no clear vertical variation because factors causing vertical variation can differ among species and sizes.
Difficult access to 40-m-tall emergent trees in tropical rainforests has resulted in a lack of data related to vertical variations in wood CO2 efflux, even though significant variations in wood CO2 efflux are an important source of errors when estimating whole-tree total wood CO2 efflux. This study aimed to clarify vertical variations in wood CO2 efflux for emergent trees and to document the impact of the variations on the whole-tree estimates of stem and branch CO2 efflux. First, we measured wood CO2 efflux and factors related to tree morphology and environment for seven live emergent trees of two dipterocarp species at four to seven heights of up to ∼ 40 m for each tree using ladders and a crane. No systematic tendencies in vertical variations were observed for all the trees. Wood CO2 efflux was not affected by stem and air temperature, stem diameter, stem height or stem growth. The ratios of wood CO2 efflux at the treetop to that at breast height were larger in emergent trees with relatively smaller diameters at breast height. Second, we compared whole-tree stem CO2 efflux estimates using vertical measurements with those based on solely breast height measurements. We found similar whole-tree stem CO2 efflux estimates regardless of the patterns of vertical variations in CO2 efflux because the surface area in the canopy, where wood CO2 efflux often differed from that at breast height, was very small compared with that at low stem heights, resulting in little effect of the vertical variations on the estimate. Additionally, whole-tree branch CO2 efflux estimates using measured wood CO2 efflux in the canopy were considerably different from those measured using only breast height measurements. Uncertainties in wood CO2 efflux in the canopy did not cause any bias in stem CO2 efflux scaling, but affected branch CO2 efflux.
In this study, we demonstrated the occurrence of stomatal patchiness and its spatial scale in leaves from various sizes of trees grown in a lowland dipterocarp forest in Peninsular Malaysia. To evaluate the patterns of stomatal behavior, we used three techniques simultaneously to analyze heterobaric or homobaric leaves from five tree species ranging from 0.6 to 31 m in height: (i) diurnal changes in chlorophyll fluorescence imaging, (ii) observation and simulation of leaf gas-exchange rates and (iii) a pressure-infiltration method. Measurements were performed in situ with 1000 or 500 μmol m(-2) s(-1) photosynthetic photon flux density. Diurnal patterns in the spatial distribution of photosynthetic electron transport rate (J) mapped from chlorophyll fluorescence images, a comparison of observed and simulated leaf gas-exchange rates, and the spatial distribution of stomatal apertures obtained from the acid-fuchsin-infiltrated area showed that patchy stomatal closure coupled with severe midday depression of photosynthesis occurred in Neobalanocarpus heimii (King) Ashton, a higher canopy tree with heterobaric leaves due to the higher leaf temperature and vapor pressure deficit. However, subcanopy or understory trees showed uniform stomatal behavior throughout the day, although they also have heterobaric leaves. These results suggest that the occurrence of stomatal patchiness is determined by tree size and/or environmental conditions. The analysis of spatial scale by chlorophyll fluorescence imaging showed that several adjacent anatomical patches (lamina areas bounded by bundle-sheath extensions within the lamina) may co-operate for the distributed patterns of J and stomatal apertures.
Understory plants in tropical forests often experience a low-light environment combined with high CO2 concentration. We hypothesized that the high CO2 concentration may compensate for leaf carbon loss caused by the low light, through increasing light-use efficiency of both steady-state and dynamic photosynthetic properties. To test the hypothesis, we examined CO2 gas exchange in response to an artificial lightfleck in Dipterocarpus sublamellatus Foxw. seedlings under contrasting CO2 conditions: 350 and 700 μmol CO2 mol(-1) air in a tropical rain forest, Pasoh, Malaysia. Total photosynthetic carbon gain from the lightfleck was about double when subjected to the high CO2 when compared with the low CO2 concentration. The increase of light-use efficiency in dynamic photosynthesis contributed 7% of the increased carbon gain, most of which was due to reduction of photosynthetic induction to light increase under the high CO2. The light compensation point of photosynthesis decreased by 58% and the apparent quantum yield increased by 26% at the high CO2 compared with those at the low CO2. The study suggests that high CO2 increases photosynthetic light-use efficiency under both steady-state and fluctuating light conditions, which should be considered in assessing the leaf carbon gain of understory plants in low-light environments.
Malaysian Borneo has a lower population density and is an area known for its lush rainforests. However, changes in pollutant profiles are expected due to increasing urbanisation and commercial-industrial activities. This study aims to determine the variation of surface O3concentration recorded at seven selected stations in Malaysian Borneo. Hourly surface O3data covering the period 2002 to 2013, obtained from the Malaysian Department of Environment (DOE), were analysed using statistical methods. The results show that the concentrations of O3recorded in Malaysian Borneo during the study period were below the maximum Malaysian Air Quality Standard of 100ppbv. The hourly average and maximum O3concentrations of 31 and 92ppbv reported at Bintulu (S3) respectively were the highest among the O3concentrations recorded at the sampling stations. Further investigation on O3precursors show that sampling sites located near to local petrochemical industrial activities, such as Bintulu (S3) and Miri (S4), have higher NO2/NO ratios (between 3.21 and 5.67) compared to other stations. The normalised O3values recorded at all stations were higher during the weekend compared to weekdays (unlike its precursors) which suggests the influence of O3titration by NO during weekdays. The results also show that there are distinct seasonal variations in O3across Borneo. High surface O3concentrations were usually observed between August and September at all stations with the exception of station S7 on the east coast. Majority of the stations (except S1 and S6) have recorded increasing averaged maximum concentrations of surface O3over the analysed years. Increasing trends of NO2and decreasing trends of NO influence the yearly averaged maximum of O3especially at S3. This study also shows that variations of meteorological factors such as wind speed and direction, humidity and temperature influence the concentration of surface O3.
Tropical rainforests control the exchange of water and energy between the land surface and the atmosphere near the equator and thus play an important role in the global climate system. Measurements of latent (LE) and sensible heat exchange (H) have not been synthesized across global tropical rainforests to date, which can help place observations from individual tropical forests in a global context. We measured LE and H for four years in a tropical peat forest ecosystem in Sarawak, Malaysian Borneo using eddy covariance, and hypothesize that the study ecosystem will exhibit less seasonal variability in turbulent fluxes than other tropical ecosystems as soil water is not expected to be limiting in a tropical forested wetland. LE and H show little variability across seasons in the study ecosystem, with LE values on the order of 11 MJ m-2 day and H on the order of 3 MJ m-2 day-1. Annual evapotranspiration (ET) did not differ among years and averaged 1579 ± 47 mm year-1. LE exceeded characteristic values from other tropical rainforest ecosystems in the FLUXNET2015 database with the exception of GF-Guy near coastal French Guyana, which averaged 8-11 MJ m-2 day-1. The Bowen ratio (Bo) in tropical rainforests in the FLUXNET2015 database either exhibited little seasonal trend, one seasonal peak, or two peaks. Volumetric water content (VWC) and VPD explained a trivial amount of the variability of LE and Bo in some of the tropical rainforests including the study ecosystem, but were strong controls in others, suggesting differences in stomatal regulation and/or the partitioning between evaporation and transpiration. Results demonstrate important differences in the seasonal patterns in water and energy exchange across different tropical rainforest ecosystems that need to be understood to quantify how ongoing changes in tropical rainforest extent will impact the global climate system.
Canarium odontophyllum Miq. fruit, popularly recognized as dabai fruit in Sarawak, is a
seasonal fruit found in the tropical rainforest of East Malaysia. A dabai fruit can be divided into
several anatomical parts, and different parts of the fruit have different valuable phytochemicals. Due
to the lack of promotion, dabai fruit is viewed as nutritionally inferior fruit by the public. On the
contrary, the fruit is rich in nutrients such as protein, fat, carbohydrates, sodium, calcium and iron.
Many phytochemicals have been detected from different parts of dabai fruit, and these molecules have
been linked to beneficial properties such as hypolipidemic, anti-atherosclerotic, anti-cholinesterase,
antimicrobial and potentially anti-diabetic. The aim of this article is to review research studies on this
fruit in order to provide adequate baseline information for commercial exploitation as well as for
future studies.
Arbuscular mycorrhizal fungi (AMF) and ectomycorrhizal fungi (EMF) produce contrasting plant-soil feedbacks, but how these feedbacks are constrained by lithology is poorly understood. We investigated the hypothesis that lithological drivers of soil fertility filter plant resource economic strategies in ways that influence the relative fitness of trees with AMF or EMF symbioses in a Bornean rain forest containing species with both mycorrhizal strategies. Using forest inventory data on 1245 tree species, we found that although AMF-hosting trees had greater relative dominance on all soil types, with declining lithological soil fertility EMF-hosting trees became more dominant. Data on 13 leaf traits and wood density for a total of 150 species showed that variation was almost always associated with soil type, whereas for six leaf traits (structural properties; carbon, nitrogen, phosphorus ratios, nitrogen isotopes), variation was also associated with mycorrhizal strategy. EMF-hosting species had slower leaf economics than AMF-hosts, demonstrating the central role of mycorrhizal symbiosis in plant resource economies. At the global scale, climate has been shown to shape forest mycorrhizal composition, but here we show that in communities it depends on soil lithology, suggesting scale-dependent abiotic factors influence feedbacks underlying the relative fitness of different mycorrhizal strategies.
Southeast Asian rainforests at upper hill elevations are increasingly vulnerable to degradation because most lowland forest areas have been converted to different land uses. As such, understanding the genetics of upper hill species is becoming more crucial for their future management and conservation. Shorea platyclados is an important, widespread upper hill dipterocarp in Malaysia. To elucidate the genetic structure of S. platyclados and ultimately provide guidelines for a conservation strategy for this species, we carried out a comprehensive study of the genetic diversity and demographic history of S. platyclados. Twenty-seven populations of S. platyclados across its range in Malaysia were genotyped at 15 polymorphic microsatellite loci and sequenced at seven noncoding chloroplast DNA (cpDNA) regions. A total of 303 alleles were derived from the microsatellite loci, and 29 haplotypes were identified based on 2892 bp of concatenated cpDNA sequences. The populations showed moderately high genetic diversity (mean HE = 0.680 for microsatellite gene diversity and HT = 0.650 for total haplotype diversity) and low genetic differentiation (FST = 0.060). Bayesian clustering divided the studied populations into two groups corresponding to western and eastern Malaysia. Bottleneck analysis did not detect any recent bottleneck events. Extended Bayesian skyline analyses showed a model of constant size for the past population history of this species. Based on our findings, priority areas for in situ and ex situ conservation and a minimum population size are recommended for the sustainable utilization of S. platyclados.
Differences in wing morphology are predicted to reflect differences in bat foraging strategies. Experimental tests of this prediction typically assess the relationship between wing morphology and a measures of flight performance on an obstacle course. However, studies have lacked measures of obstacle avoidance ability true scores, which may confound interpretation of ability across the range of presented tasks. Here, we used Rasch analysis of performance in a collision-avoidance experiment to estimate the ability of bat species to fly through vegetative clutter. We refer to this latent trait as 'clutter negotiating ability' and determined the relationships between clutter negotiating ability and wing morphology in 15 forest insectivorous bat species that forage in the densely cluttered rainforests of Malaysia. The clutter negotiating ability scores were quantified based on individual responses of each species to 11 different obstacle arrangements (four banks of vertical strings 10-60 cm apart). The tasks employed for the collision-avoidance experiment were reliable and valid, although Rasch analysis suggested that the experiment was too easy to discriminate completely among the 15 species. We found significant negative correlations between clutter negotiating ability and body mass, wingspan, wing loading and wing area but a positive significant correlation with wingtip area ratio. However, in stepwise multiple regression analyses, only body mass and wing loading were significant predictors of clutter negotiating ability. Species fell into clusters of different clutter negotiating ability, suggesting a potential mechanism for resource partitioning within the forest interior insectivorous ensemble.
Ants are diverse and abundant, especially in tropical ecosystems. They are often cited as the agents of key ecological processes, but their precise contributions compared with other organisms have rarely been quantified. Through the removal of food resources from the forest floor and subsequent transport to nests, ants play an important role in the redistribution of nutrients in rainforests. This is an essential ecosystem process and a key energetic link between higher trophic levels, decomposers and primary producers. We used the removal of carbohydrate, protein and seed baits as a proxy to quantify the contribution that ants, other invertebrates and vertebrates make to the redistribution of nutrients around the forest floor, and determined to what extent there is functional redundancy across ants, other invertebrate and vertebrate groups. Using a large-scale, field-based manipulation experiment, we suppressed ants from c. 1 ha plots in a lowland tropical rainforest in Sabah, Malaysia. Using a combination of treatment and control plots, and cages to exclude vertebrates, we made food resources available to: (i) the whole foraging community, (ii) only invertebrates and (iii) only non-ant invertebrates. This allowed us to partition bait removal into that taken by vertebrates, non-ant invertebrates and ants. Additionally, we examined how the non-ant invertebrate community responded to ant exclusion. When the whole foraging community had access to food resources, we found that ants were responsible for 52% of total bait removal whilst vertebrates and non-ant invertebrates removed the remaining 48%. Where vertebrates were excluded, ants carried out 61% of invertebrate-mediated bait removal, with all other invertebrates removing the remaining 39%. Vertebrates were responsible for just 24% of bait removal and invertebrates (including ants) collectively removed the remaining 76%. There was no compensation in bait removal rate when ants and vertebrates were excluded, indicating low functional redundancy between these groups. This study is the first to quantify the contribution of ants to the removal of food resources from rainforest floors and thus nutrient redistribution. We demonstrate that ants are functionally unique in this role because no other organisms compensated to maintain bait removal rate in their absence. As such, we strengthen a growing body of evidence establishing ants as ecosystem engineers, and provide new insights into the role of ants in maintaining key ecosystem processes. In this way, we further our basic understanding of the functioning of tropical rainforest ecosystems.
Tree species in tropical rain forests exhibit a rich panoply of spatial patterns that beg ecological explanation. The analysis of tropical census data typically relies on spatial statistics, which quantify the average aggregation tendency of a species. In this article we develop a cluster-based approach that complements traditional spatial statistics in the exploration and analysis of ecological hypotheses for spatial pattern. We apply this technique to six study species within a fully mapped 50-ha forest census in peninsular Malaysia. For each species we identify the scale(s) of spatial aggregation and the corresponding tree clusters. We study the correlation between cluster locations and abiotic variables such as topography. We find that the distribution of cluster sizes exhibits equilibrium and nonequilibrium behavior depending on species life history. The distribution of tree diameters within clusters also varies according to species life history. At different spatial scales, we find evidence for both niche-based and dispersal-limited processes producing spatial pattern. Our methodology for identifying scales of aggregation and clusters is general; we discuss the method's applicability to spatial problems outside of tropical plant ecology.