Harumanis is one of the main signatures of Perlis with regards to its delightful taste, pleasant aroma and expensive price. Harumanis authenticity and productivity had become the remarks among the farmers, entrepreneurs, consumers and plant breeders due to the existence of morphological characteristics variation among the fruits and high production cost. Assessment of Harumanis morphological characteristics of natural population and different tree ages may represent a possible source of important characteristics for development and breeding purposes of Harumanis. The aim of this study is to evaluate the morphological variation of Harumanis collected from different location in Perlis and tree age. A total of 150 Harumanis fruits from 50 trees with three different stages of development (young, middle-aged and old) were characterised using 11 traits; 10 quantitative and one qualitative morphological trait. The ANOVA analyses in combination with Dunn's pairwise and Kruskal-Wallis multiple comparison test able to point out the existence of environmental factor and age influence towards the significant different of identified morphological traits except for Total Soluble Solid (TSS) and pulp percentage. Five clusters of 50 Harumanis accessions reflect a grouping pattern which not according to neither geographical region nor age. The result of Principal Component Analysis (PCA) using the first two principal components (PCs) provided a good approximation of the data explaining 84.09% of the total variance which majorly contributed by parameters of weight, fruit dimensional characteristics, peel percentage and hue angle, h. Preliminary screening of important morphological characteristics which contribute to the phenotypic diversity of Harumanis is successfully achieved. The findings can be employed by the plant breeders and farmers for the establishment of standard grading of Harumanis and advancement of breeding crop of Harumanis in future.
This paper discusses the colonization of the stratiomyid species Ptecticus melanurus (Walker) (Diptera: Stratiomyidae) in monkey carrion and its potential for the determination of the minimum time since death (PMI). A study was conducted in a tropical forest at Bangi, Malaysia from 13 November 2009 to 8 June 2011. Twelve monkey carcasses (Macaca fascicularis Raffles) were used and divided in equal number into three different field trials. Adults of P. melanurus were first observed on monkey carrions on the second day the carcasses were placed in the field while their penultimate instar larvae were found in the wet soil under and beside carcass from day 8 to 31 days postmortem.
This study was designed to mimic homicide or suicide cases using gasoline. Six adult long-tailed macaque (Macaca fascicularis), weighing between 2.5 to 4.0 kg, were equally divided into control and test groups. The control group was sacrificed by a lethal dose of phenobarbital intracardiac while test group was force fed with two doses of gasoline LD50 (37.7 ml/kg) after sedation with phenobarbital. All carcasses were then placed in a decomposition site to observe the decomposition and invasion process of cadaveric fauna on the carcasses. A total of five decomposition stages were recognized during this study. This study was performed during July 2007. Fresh stage of control and test carcasses occurred between 0 to 15 and 0 to 39 hours of exposure, respectively. The subsequent decomposition stages also exhibited the similar pattern whereby the decomposition process of control carcasses were faster than tested one. The first larvae were found on control carcasses after 9 hours of death while the test group carcasses had only their first blowfly eggs after 15 hours of exposure. Blow flies, Achoetandrus rufifacies and Chrysomya megacephala were the most dominant invader of both carcasses throughout the decaying process. Diptera collected from control carcasses comprised of scuttle fly, Megaselia scalaris and flesh fly, sarcophagid. We concluded that the presence of gasoline and its odor on the carcass had delayed the arrival of insect to the carcasses, thereby slowing down the decomposition process in the carcass by 6 hours.
A total of 204 rodents comprising 14 host species from four different habitats were examined. Nine rodent species were trapped from the forest and another five species were trapped from the coastal, rice field and urban habitats. Rattus rattus diardii (67%) was the predominant rodent species examined. Fifty six (47.3%) rodents and shrews were found to be infested with at least one of the 20 species of ectoparasite recovered. Mites belonging to the family Trombiculidae were the predominant ectoparasite species recovered. Ticks belonging to the family Ixodidae were recovered mainly from the forest dwelling rodents. Polyplax spinulosa and Hoplopleura pacifica were the common lice species found infesting the urban rodents. Xenopsylla cheopis was the only flea species recovered. The following ecto-parasites have been incriminated as important vectors or as mechanical carriers for the transmission of zoonotic diseases: Ixodes granulatus, Dermacentor sp. Haemaphysalis sp., Amblyomma sp. Ascoschoengastia indica, Leptotrombidium deliense, Ornithonyssus bacoti, Laelaps nuttalli, H. pacifica, P. spinulosa and Xenopsylla cheopis. Urban and forest rodents were significantly higher in ecto-parasitic infestation, compared to rats from the other two habitats. However, there was no significant statistical association between male and female rodents infested with ectoparasites.
Trapping of small mammals was conducted at 5 study sites in Kuala Selangor Nature Park (KSNP) from 20-24 June 2005. A total of 11 animals comprising 2 species of rodents, Maxomys whiteheadi and Rattus exulans were caught from 3 sites, i.e from an area of mixed secondary forest and mangrove swamp; an area of mangrove swamp, and from an area of lalang fringing mangrove swamp. From these animals, the following 7 species of ectoparasites were found: Laelaps echidninus, Laelaps nuttalli, Ascoschoengastia indica, Leptotrombidium deliense, Hoplopleura pectinata, Hoplopleura pacifica and Polyplax spinulosa. One of the ectoparasites found, L. deliense is a known vector of scrub typhus and thus may pose potential health risks to visitors to KSNP.
Forest canopies are dynamic interfaces between organisms and atmosphere, providing buffered microclimates and complex microhabitats. Canopies form vertically stratified ecosystems interconnected with other strata. Some forest biodiversity patterns and food webs have been documented and measurements of ecophysiology and biogeochemical cycling have allowed analyses of large-scale transfer of CO2, water, and trace gases between forests and the atmosphere. However, many knowledge gaps remain. With global research networks and databases, and new technologies and infrastructure, we envisage rapid advances in our understanding of the mechanisms that drive the spatial and temporal dynamics of forests and their canopies. Such understanding is vital for the successful management and conservation of global forests and the ecosystem services they provide to the world.
Soil hardness plays a vital role in evaluating the physical properties of soil structure. With regards to the impact of compaction on practical forest management issues, most report and review forms were available. Thus, the aim of this study was to evaluate the soil condition in riparian forest restoration planted with indigenous species along Kayan Ulu River with special reference to soil hardness. Soil hardness was measured by using Hasegawa-type cone penetrometer from the surface soils to 100 cm depth, with a total of 48 random points for both study sites surveyed; restoration sites planted with Shorea macrophylla in year 1996 and 1998 (SPD96 and SPD98, respectively) for both on and between planting lines. Our findings indicated that, soil hardness in SPD98 was harder as compared to SPD96 at shallow depth presented in one drop penetrability. Likewise, soil penetration resistance on planting line in SPD98 was significantly higher than SPD96 at surface soils (0-20 cm) and subsurface soils (20-40 cm). A high number of strikes and soil penetration resistance indicate that the soils were highly compacted. However, there was no significant difference in term of soil penetration resistance between planting line. In order to avoid effects on tree productivity, it is recommended that in future, the evaluation of soil hardness should be determined during the early establishment for future restoration of riparian ecosystem.
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.
Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.
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.
Variations in leaf photosynthetic, morphological and biochemical properties with increasing plant height from seedlings to emergent trees were investigated in five dipterocarp species in a Malaysian tropical rain forest. Canopy openness increased significantly with tree height. Photosynthetic properties, such as photosynthetic capacity at light saturation, light compensation point, maximum rate of carboxylation and maximum rate of photosynthetic electron transport, all increased significantly with tree height. Leaf morphological and biochemical traits, such as leaf mass per area, palisade layer thickness, nitrogen concentration per unit area, chlorophyll concentration per unit dry mass and chlorophyll to nitrogen ratio, also changed significantly with tree height. Leaf properties had simple and significant relationships with tree height, with few intra- and interspecies differences. Our results therefore suggest that the photosynthetic capacity of dipterocarp trees depends on tree height, and that the trees adapt to the light environment by adjusting their leaf morphological and biochemical properties. These results should aid in developing models that can accurately estimate carbon dioxide flux and biomass production in tropical rain forests.
Tree autotrophic respiratory processes, especially stem respiration or stem CO2 efflux (Estem), are important components of the forest carbon budget. Despite efforts to investigate the controlling processes of Estem in recent years, a considerable lack in our knowledge remains on the abiotic and biotic drivers affecting Estem dynamics. It has been strongly advocated that long-term measurements would shed light onto those processes. The expensive scientific instruments needed to measure gas exchange have prevented Estem measurements from being applied on a larger temporal and spatial scale. Here, we present an automated closed dynamic chamber system based on inexpensive and industrially broadly applied CO2 sensors, reducing the costs for the sensing system to a minimum. The CO2 sensor was cross-calibrated with a commonly used gas exchange system in the laboratory and in the field, and we found very good accordance of these sensors. We tested the system under harsh tropical climatic conditions, characterized by heavy tropical rainfall events, extreme humidity and temperatures, in a moist lowland forest in Malaysia. We recorded Estem of three Dyera costulata (Miq.) trees with our prototype over various days. The variation of Estem was large among the three tree individuals and varied by 7.5-fold. However, clear diurnal changes in Estem were present in all three tree individuals. One tree showed high diurnal variation in Estem, and the relationship between Estem and temperature was characterized by a strong hysteresis. The large variations found within one single tree species highlight the importance of continuous measurement to quantify ecosystem carbon fluxes.
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.
The root systems of forest trees are composed of different diameters and heterogeneous physiological traits. However, the pattern of root respiration rates from finer and coarser roots across various tropical species remains unknown. To clarify how respiration is related to the morphological traits of roots, we evaluated specific root respiration and its relationships to mean root diameter (D) of various diameter and root tissue density (RTD; root mass per unit root volume; gcm(-3)) and specific root length (SRL; root length per unit root mass; mg(-1)) of the fine roots among and within 14 trees of 13 species from a primary tropical rainforest in the Pasoh Forest Reserve in Peninsular Malaysia. Coarse root (2-269mm) respiration rates increased with decreasing D, resulting in significant relationships between root respiration and diameter across species. A model based on a radial gradient of respiration rates of coarse roots simulated the exponential decrease in respiration with diameter. The respiration rate of fine roots (<2mm) was much higher and more variable than those of larger diameter roots. For fine roots, the mean respiration rates for each species increased with decreasing D. The respiration rates of fine roots declined markedly with increasing RTD and increased with increasing SRL, which explained a significant portion of the variation in the respiration among the 14 trees from 13 species examined. Our results indicate that coarse root respiration in tree species follows a basic relationship with D across species and that most of the variation in fine root respiration among species is explained by D, RTD and SRL. We found that the relationship between root respiration and morphological traits provides a quantitative basis for separating fine roots from coarse roots and that the pattern holds across different species.
We investigated effects of heterogeneous stomatal behavior on diurnal patterns of leaf gas exchange in 10 tree species. Observations were made in middle and upper canopy layers of potted tropical rainforest trees in a nursery at the Forest Research Institute Malaysia. Measurements were taken from 29 January to 3 February 2010. We measured in situ diurnal changes in net photosynthetic rate and stomatal conductance in three leaves of each species under natural light. In both top-canopy and sub-canopy species, midday depression of net assimilation rate occurred in late morning. Numerical analysis showed that patchy bimodal stomatal behavior occurred only during midday depression, suggesting that the distribution pattern of stomatal apertures (either uniform or non-uniform stomatal behavior) varies flexibly within single days. Direct observation of stomatal aperture using Suzuki's Universal Micro-Printing (SUMP) method demonstrated midday patchy stomatal closure that fits a bimodal pattern in Shorea leprosula Miq., Shorea macrantha Brandis. and Dipterocarpus tempehes V.Sl. Inhibition of net assimilation rate and stomatal conductance appears to be a response to changes in vapor pressure deficit (VPD). Variable stomatal closure with increasing VPD is a mechanism used by a range of species to prevent excess water loss from leaves through evapotranspiration (viz., inhibition of midday leaf gas exchange). Bimodal stomatal closure may occur among adjacent stomata within a single patch, rather than among patches on a single leaf. Our results suggest the occurrence of patches at several scales within single leaves. Further analysis should consider variable spatial scales in heterogeneous stomatal behavior between and within patches and within single leaves.
We observed diurnal and seasonal patterns of leaf-scale gas exchange within the crown of a Dipterocarpus sublamellatus Foxw. tree growing in a lowland dipterocarp forest at Pasoh, Peninsular Malaysia. Observations were carried out nine times over 6 years, from September 2002 to December 2007. Observation periods included both wet and mild-dry periods, and natural and saturated photosynthetic photon flux density (PPFD) light conditions. In situ measurements of the diurnal change in net photosynthetic rate and in stomatal conductance were carried out on canopy leaves of a 40-m-tall D. sublamellatus tree, which was accessed from a canopy corridor. A diurnal change in electron transport rate was observed under saturated PPFD conditions. The maximum net assimilation rate was approximately 10 micromol m(-2) s(-1). There was a clear inhibition of the net assimilation rate coupled with stomatal closure after late morning and this inhibition occurred year-round. Although the electron transport rate decreased alongside this inhibition, it sometimes followed on. Numerical analysis showed that the main factor in the inhibition of the net assimilation rate was patchy bimodal stomatal closure, which occurred in both mild-dry and wet periods. The midday depression occurred year-round, though there are fluctuations in soil moisture during the mild-dry and wet periods. The magnitude of the inhibition was not related to soil water content but was related to vapor pressure deficit (VPD): that is, whether the days were sunny and hot or cloudy and cool. On cloudy, cool days in the wet period, the net photosynthesis was only moderately inhibited, but it still decreased in the afternoon and was coupled with patchy stomatal closure, even in quite moderate VPD, leaf temperature and PPFD conditions. Our results suggest that patchy stomatal closure signaled by the increase in VPD, in transpiration and by circadian rhythms, was the key factor in constraining midday leaf gas exchange of the D. sublamellatus canopy leaves.
Forest growth is an important factor both economically and ecologically, and it follows a predictable trend with age. Generally, growth accelerates as canopies develop in young forests and declines substantially soon after maximum leaf area is attained. The causes of this decline are multiple and may be linked to age- or size-related processes, or both. Our objective was to determine the relative effects of tree age and tree size on the physiological attributes of two broadleaf species. As age and size are normally coupled during growth, an approach based on grafting techniques to separate the effects of size from those of age was adopted. Genetically identical grafted seedlings were produced from scions taken from trees of four age classes, ranging from 4 to 162 years. We found that leaf-level net photosynthetic rate per unit of leaf mass and some other leaf structural and biochemical characteristics had decreased substantially with increasing size of the donor trees in the field, whereas other gas exchange parameters expressed on a leaf area basis did not. In contrast, these parameters remained almost constant in grafted seedlings, i.e., scions taken from donor trees with different meristematic ages show no age-related trend after they were grafted onto young rootstocks. In general, the results suggested that size-related limitations triggered the declines in photosynthate production and tree growth, whereas less evidence was found to support a role of meristematic age.
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.