Elemental concentrations in a tropical montane rain forest in Sri Lanka

As a preliminary to studies on biogeochemistry of a tropical montane rain forest at about 1800 m altitude in Sri Lanka, the concentrations of 17 chemical elements in different forest compartments, i.e. in leaves of six floristically dominant tree species, undergrowth, litter, and soil at 0–10 cm and 11–60 cm depth were determined. Foliar chemical differences between species were large. Mean leaf water content, mean leaf area per leaf, and specific leaf weight of each species were also determined. Calcium seems to be used in relatively large quantities by these plants. Phosphorus concentration in all compartments was consistently low. A few species investigated appeared to concentrate aluminium and silicon at high amounts. Concentration differences between compartments were significant only for certain elements. Location effect on the concentrations in soil was considerable for most of the elements studied. When studying plant-soil chemical interactions in these forests, species-level concentrations must be taken into account as the species-specific chemical differences are obscured when treated as canopy leaves.     

海南岛热带山地雨林幼苗幼树光合与叶氮、叶磷及比叶面积的关系

以海南岛吊罗山热带山地雨林101个物种的幼苗幼树为试验材料,测定其光合、叶片氮、磷含量及比叶面积;检验其相关关系,并按乔木,乔灌木(小乔木至大灌木)和灌木3个生活型组进行分组检验。研究结果表明,单位叶面积(Aarea)和单位叶重量的光合速率(Amass)均表现出灌木>乔木>乔灌木,方差分析表明,灌木和乔灌木之间Aarea差异显著;灌木和乔木以及灌木和乔灌木之间Amass差异显著(p<0.05)。Aarea与叶氮含量之间的相关性在不同生态型组和所有物种之间均达到极显著水平(p<0.0001);与叶磷之间的相关关系在灌木(p=0.0038),乔灌木(p=0.0002)以及所有物种(p<0.0001)之间达到极显著水平,但是在乔木中未达到显著水平(p>0.05);与SLA之间在灌木(p=0.0006)、乔木(p<0.0001)和所有物种(p<0.0001)之间达到极显著水平,但是在乔灌木中未达到显著水平(p>0.05)。Amass与叶片氮含量、SLA的相关关系在不同生活型组和所有物种中都达到极显著水平(p<0.0001);与叶磷含量之间的相关性在灌木(p=0.0004),乔灌木(p=0.0018)及所有物种(p<0.0001)中极显著,在乔木生活型组中也达显著水平(p=0.0377)。逐步回归表明,与Aarea相比,Amass估计结果更接近于实际测值。由此可见,海南岛热带山地雨林林下幼苗幼树的光合和叶氮、磷含量及SLA之间相关关系与基于成树的研究非常相似,并且A比A更能稳定体现这种相关性。     

The effect of shade on leaf structure and physiology of tree seedlings from a mixed dipterocarp forest

This study builds upon past work investigating seedling leaf physiology and structure among tropical trees. We seek to explain how related and unrelated species and genera co‐occur in relation to varying amounts of shade. Seedlings of eight Sri Lankan rain forest tree species in three genera (Dipterocarpus, Mesua, Shorea section Doona) were grown for 2 years in four treatments that simulated a variety of shade environments across the understorey of a rain forest. All three genera comprise major canopy tree species of mixed dipterocarp forest, a widespread and important Asian tropical forest type. Compared with the other genera, Dipterocarpus spp. had the largest leaves, the thinnest leaf blades and relatively high rates of stomatal conductivity across all shade treatments, making them water‐loving species sensitive to droughty soils. Mesua spp. had intermediate sized leaves, with the thickest leaf blades and palisade mesophyll layers, the highest stomatal densities, the smallest aperture sizes and the lowest rates of stomatal conductance, making them the most water conservative. Shorea spp. were generally intermediate in blade and palisade mesophyll dimensions between Dipterocarpus spp. and Mesua spp., but they had the smallest leaves. Greater differences among genera than among species within genera were apparent, but species differences within genera were also apparent. Differences among genera and species conform to their known successional status and topographical affinities and provide a more comprehensive understanding of species site adaptation. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 332–343.     

Leaf nutrients in relation to stature and life form in tropical rain forest

Abstract. To document the relationship between a plant's position in the canopy and its leaf nutrient content, leaf nitrogen and phosphorus were determined for 30 species growing in mature evergreen lowland rain forest at La Selva Biological Station, Costa Rica. Species that grow either in the understory, midstory, or the canopy were selected. Species were further separated into three life forms: self-supporting monocots, self-supporting dicots, and climbers. Mass-based nutrient concentrations were expected to decrease with stature, as has been reported in studies of other forests. In fact, mass-based nitrogen and phosphorus did not vary significantly among the three adult-stature classes, although area-based values differed greatly: canopy plants averaged 60 % more nitrogen and 90 % more phosphorus per unit leaf area than understory plants. Differences in leaf characteristics were evident among the three life forms. Most notably, area-based phosphorus and leaf specific mass were lowest in climbers, intermediate in self-supporting dicots, and highest in self-supporting monocots. These results support the characterization of climbers as investing in inexpensive structures, perhaps in order to gain competitive advantage in light capture by allocating resources to maximize elongation rates.     

Response of recruitment to light availability across a tropical lowland rain forest community

1.  Many hypotheses about species coexistence involve differential resource use and trade-offs in species' life-history traits. Quantifying resource use across most species in diverse communities, although, has seldom been attempted.
2.  We use a hierarchical Bayesian approach to quantify the light dependence of recruitment in 263 woody species in a 50-ha long-term forest census plot in Panama. Data on sapling recruitment were obtained using the 1985–1990 and 1990–1995 census intervals. Available light was estimated for each recruit from yearly censuses of canopy density.
3.  We use a power function (linear log–log relationship) to model the light effect on recruitment. Different responses of recruitment to light are expressed by the light effect parameter b . The distribution of b had a central mode at 0.8, suggesting that recruitment of many species responds nearly linearly to increasing light.
4.  Nearly every species showed increases in recruitment with increasing light. Just nine species (3%) had recruitment declining with light, while 198 species (75%) showed increasing recruitment in both census intervals. Most of the increases in recruitment were decelerating, i.e. the increase was less at higher light ( b  < 1). In the remaining species, the response to light varied between census intervals (24 species) or species did not have recruits in both intervals (41 species).
5.   Synthesis. Nearly all species regenerate better in higher light, and recruitment responses to light are spread along a continuum ranging from modest increase with light to a rather strict requirement for high light. These results support the hypothesis that spatio-temporal variation in light availability may contribute to the diversity of tropical tree species by providing opportunities for niche differentiation with respect to light requirements for regeneration.     

Leaf dynamics of seedlings of rain forest species in relation to canopy gaps

Summary Leaf dynamics of eight tropical rain forest species seedlings was studied in three environments: the shaded forest understorey, a small gap of ±50 m², and a large gap of ±500 m². Leaf production rate and leaf loss rate were enhanced in gaps, and a large gap resulted in larger increases than a small gap. For most species net leaf gain rate was larger in gaps, although this rate was not always largest in the large gap. Leaf loss decreased, and leaf survival percentages increased with increasing shade tolerance of species, indicating a slower leaf turnover for more shade tolerant species. Leaf area growth rate was only partly determined by net leaf gain rate. Ontogenetic effects on leaf size were also important, especially in the large gap. Species which possessed leaves with high specific leaf weight (SLW) showed lower leaf loss rates and higher leaf survival percentages than species with low SLW leaves. Leaf life span seemed to be related to leafcost per unit area. The relation of specific patterns in leaf production and leaf loss to the regeneration mode of the species is briefly discussed.     

The responses to shade of seedlings of very small-seeded tree and shrub species from tropical rain forest in Singapore

1. Newly germinated seedlings of six tree and shrub species with very small seeds (31–460 μg dry mass), one light-demanding and five shade-tolerant at the stage of establishment in the wild, were grown for 5 months in neutral shade houses with 0·5, 1, 3·5 and 7·5% daylight.
2. The ratio of yield in 7·5% to that in 1% was 8:1 for the light-demanding Melastoma malabathricum but only 2:1 for the confamilial shade-tolerator Pternandra echinata. The Pternandra, Urophyllum hirsutum, Ficus chartacea, Ficus grossularioides and Pellacalyx saccardianus showed a graded series of responses to irradiance, generally consistent with their apparent demands for light in the wild. In contrast, survival of very deep shade was not clearly related to light demand in nature.
3. The results support the conclusion drawn from observational studies that large seed size is not primarily adaptive in resisting shade but in resisting the associated risks of burial by litter, desiccation during dry spells, uprooting by birds and other kinds of damage by animals or falling debris.     

Leaf anatomy plasticity of Alchornea triplinervia (Euphorbiaceae) under distinct light regimes in a Brazilian montane Atlantic rain forest

 Alchornea triplinervia trees occur in a montane Atlantic rainforest at the Macaé de Cima State Ecological Reserve (Nova Friburgo, RJ, Brazil). They are found in two adjacent secondary forests at distinct successional stages: a late-secondary (shaded and humid) and an early-secondary forest (drier, higher light intensities reaching the understory). Leaf samples collected in these environments were compared in regard to various anatomic parameters. Histochemical tests, anatomic measurements and counting indicated no significant variation for the basic leaf anatomy. Nevertheless, as compared to the late-secondary forest, the leaves of the early-secondary forest individuals showed an increased bulk of sclerenchyma at the main nervation and petiole, gelatinous fibers with hygroscopic walls all along the central vascular system, thicker cuticle, a higher abaxial stomatal density, abundance of tannin in the mesophyll, a significantly thicker palisade and spongy parenchyma, and compactation of the spongy parenchyma. This anatomical variation indicates an increased xeromorphism of leaves under the drier conditions of the early-secondary forest. Leaf anatomy plasticity may contribute to the high ecological plasticity of this widespread neotropical species. Received: 11 November 1996 / Accepted: 5 February 1997     

The Hemiptera (Insecta) communities of tropical rain forest in Sulawesi

The structure and composition of Hemiptera communities of tropical rain forest in Dumoga-Bone National Park, Sulawesi, Indonesia were investigated over a 1-year period. The aim was to investigate the extent to which insect samples, obtained using six different techniques, were representative of the whole community, and to explore temporal and spatial variation in Hemiptera community composition. Sampling techniques employed were Rothamsted light trapping, canopy fogging, malaise trapping at both ground and canopy level, flight interception trapping and yellow water pan trapping. Overlap between faunas collected using different techniques was surprisingly low emphasizing the limitations of using any single method to sample and thereby describe the Hemiptera community. Differences were observed at the species, family and suborder levels. Similarly, there were major differences in faunal composition between sites and diversity appeared to peak at elevations between 600–1000 m. Seasonal changes were also significant but generally lower than between methods or sites. Results are discussed in relation to factors such as food availability and host specialization and the efficacy of each sampling method is reviewed in relation to the biology of the different Hemiptera groups and the composition of the samples obtained.     

Fruit choice by red howler monkeys (Alouatta seniculus) in a tropical rain forest

This paper presents and discusses aspects of fruit selectivity by red howler monkeys (Alouatta seniculus) in relation with morphological characteristics of fruits. These data are used to provide an answer to the following questions: which are the fruit characteristics that lead fruit choice of howler monkeys and to what extent fruit characteristics play a role in seed dispersal by monkeys? The frugivorous diet of a troop of red howler monkeys was determined during a 2-year field study in French Guiana. The selection of fruit by howler monkeys was analyzed in relation to the fruit availability. Results showed that, although consumption followed availability, fruit species could be classified in three categories according to their selection ratio (percentage of consumption/percentage of abundance) as “high ranking,” “middle ranking,” and “low ranking” species. Also, the 97 species of fruit eaten by the monkeys were grouped according to the morphological characteristics thought to influence the monkeys' choice. This showed that howler monkeys consumed essentially fruits with juicy pulp, bright color, and a small number of well-protected seeds. Most of high ranking species had medium-sized fruits with yellow color, and low ranking species often had small fruits. However, howler monkeys are associated with the dispersal of seeds from fruit with a hard and indehiscent pericarp and/or large seeds, like those of the Sapotaceae family. Consequently, they can be considered as “specialized” frugivores for this fruit syndrome. © l996 Wiley-Liss, Inc.     

The role of leaf lobation in elongation responses to shade in the rosette-forming forb Serratula tinctoria (Asteraceae)

BACKGROUND AND AIMS: Lobed leaves are considered selectively advantageous in conditions of high irradiance. However, most studies have involved woody species, with only a few considering the role of leaf lobation in herbaceous rosette species. In this study, it is hypothesized that, in addition to its adaptive value in high light, leaf lobation may add to the function of petioles as vertical spacers in herbaceous species in conditions of strong competition for light. METHODS: To test this hypothesis, leaf development was examined under seasonally changing natural light conditions and a field experiment was conducted in which light climate was manipulated in a wooded meadow population of Serratula tinctoria. KEY RESULTS: No changes in leaf lobation were observed in response to experimental shading or different natural light conditions. However, in tall herbaceous vegetation, plants with highly lobed leaves achieved significantly greater vertical elongation than plants with less-lobed leaves. In contrast to herbaceous shade, tree shade had no effect on leaf elongation, suggesting differential responsiveness to competition from neighbouring herbs versus overhead shade. In shading treatments, imposed shade could only be responded to by the elongation of leaves that were produced late in development. CONCLUSIONS: The results show that extensive leaf lobation can enable greater leaf elongation in response to shade from surrounding herbaceous vegetation. The different morphological responses displayed by Serratula tinctoria to different types of shade demonstrate the importance of critically assessing experimental designs when investigating phenotypic plasticity in response to shade.     

Morpho‐anatomical and physiological differences between sun and shade leaves in Abies alba Mill. (Pinaceae,Coniferales): a combined approach

Morphology, anatomy and physiology of sun and shade leaves of Abies alba were investigated and major differences were identified, such as sun leaves being larger, containing a hypodermis and palisade parenchyma as well as possessing more stomata, while shade leaves exhibit a distinct leaf dimorphism. The large size of sun leaves and their arrangement crowded on the upper side of a plagiotropic shoot leads to self‐shading which is explainable as protection from high solar radiation and to reduce the transpiration via the lamina. Sun leaves furthermore contain a higher xanthophyll cycle pigment amount and Non‐Photochemical Quenching (NPQ) capacity, a lower amount of chlorophyll b and a total lower chlorophyll amount per leaf, as well as an increased electron transport rate and an increased photosynthesis light saturation intensity. However, sun leaves switch on their NPQ capacity at rather low light intensities, as exemplified by several parameters newly measured for conifers. Our holistic approach extends previous findings about sun and shade leaves in conifers and demonstrates that both leaf types of A. alba show structural and physiological remarkable similarities to their respective counterparts in angiosperms, but also possess unique characteristics allowing them to cope efficiently with their environmental constraints.     

Fine partitioning of epiphyte habitat within Johansson zones in tropical Australian rain forest trees

For over three decades, the Johansson zones have been widely used in epiphyte studies as a way of stratifying the host tree into habitat zones. The usefulness of this system, however, has been questioned. We test the effectiveness of the Johansson zones by grouping epiphyte species by the substrate and microclimatic attributes of their individual occurrences and assessing the fidelity of these groups to the Johansson zones. Habitat characteristics were recorded for every individual epiphyte on 30 trees in the lower montane rain forests of northeastern Australia. Twenty‐four epiphyte species were agglomerated into four groups using Ward's method. Group 4 was highly distinct and included shade‐loving species and nomadic vines from the lower zones of the host trees. Group 3 contained species from the most exposed habitats. Group 1 had higher light levels and lower substrate thickness than Group 2, yet both groups had close to identical distributions over the Johansson zones. This suggests that groups of epiphyte species may utilize different micro‐sites within the same zone. While the Johansson zones are a useful tool in epiphyte studies, finer partitioning of habitat within the host tree may be missed.     

西双版纳热带季节雨林的树种组成和群落结构动态

研究了西双版纳热带季节雨林1 hm2(hectare)动态监测样地1993年与2007年之间树种组成和群落结构的变化。对样地中胸径≥5 cm的乔木进行了每木调查。目前其树种组成的热带分布科、属所占比例分别为91%和94%,具有较高比例的热带植物区系性质。在1993年与2007年两次调查之间,树种数量由145种增至179种,仅有1到2个个体的稀有树种所占比例从54%降为51.1%。从森林的垂直结构来看,A、B、C三层的个体死亡率分别为12.8%、12.9%和19.0%,各层树木的增长率分别为-8.5%、-1.4%和44.8%。与此相对应,C层小径级的树木所占比例有较大提高。虽然小径级的树木在种类和数量上比例增大,但个体数量和种类组成相对稳定的A、B层优势树种变化不大,维持了群落结构的稳定性。14 a间,群落中新增加的具有先锋性质的树种不超过5个。1993年时,A、B两层尚有先锋树种存在,2007年已经从A、B两层中退出。因此,从14 a间树种组成和群落结构的变化来看,虽然具有树木的死亡和增补,但其物种成分和群落结构的总体格局没有明显的变化,处于动态平衡过程中。     

Growth of tree seedlings in a tropical dry forest in relation to soil moisture and leaf traits

"AimsThe growth of plant species in tropical dry forest (TDF) is expected to be largely governed by the availability of soil moisture. In this study we attempt to identify mechanisms by which seedlings of dry tropical trees cope with water stress by adjusting their leaf characteristics to water availability and micro environments, and address following questions: How are leaf traits and relative growth rate (RGR) of the dominant seedling species of TDF affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional traits with each other? Can leaf traits singly or in combination predict the growth rate of seedling species of TDF? The study was conducted in situ on four sites (viz., Hathinala, Gaighat, Harnakachar and Ranitali, listed in order of decreasing SMC) within the tropical dry deciduous forest in northern India. Methods Five leaf traits viz., specific leaf area (SLA), leaf dry matter content (LDMC), concentrations of leaf nitrogen (leaf N), phosphorus (leaf P) and chlorophyll (Chl) and two physiological processes, viz., stomatal conductance (Gs net) and photosynthetic rate (A net), and RGR, of four dominant tree seedling species of a TDF (viz., Buchanania lanzan, Diospyros melanoxylon, Shorea robusta and Terminalia tomentosa) on four sites were analysed for species, site and season effects over a 2-year period. Step-wise multiple regression was performed to predict RGR from mean values of SMC, leaf traits and physiological processes. Principal component analysis (PCA) was performed to observe the extent of intra- vs. inter-specific variability in the leaf traits and physiological rates.Important findings All the traits and physiological rates were interrelated and showed significant positive relationship with RGR except for the correlation of LDMC with RGR which was not significant. Further, relationships of SMC with all leaf traits, physiological rates and RGR were significant, except for that between SMC and SLA for B. lanzan and D. melanoxylon. The slope of seedling trait:SMC relationship, a measure of phenotypic plasticity in response to soil moisture gradient, varied among species. Among the four species, T. tomentosa was the most plastic and S. robusta the least. In conclusion, leaf traits and physiological processes were strongly related to soil water availability on the one hand and seedling growth on the other. Gs net is the most important variable which accounted for the greatest amount of variability (62%) in RGR, emphasizing the role of stomatal conductance in shaping growth patterns across spatial and temporal gradients of soil water availability. Gs net and SMC together explained 64% variability in RGR, indicating that other traits/factors, not studied by us are also important in modulating the growth of tropical tree seedlings.     

Acclimation to irradiance in seedlings of three tropical rain forest Garcinia species after simulated gap formation

We investigated the acclimation of seedlings of three tropical rain forest sub-canopy Garcinia species (G. xanthochymus, G. cowa, and G. bracteata) after transfer from 4.5 (LI) to 40 % (HI) sunlight and 12.5 (MI) sunlight to HI (LH₁ and LH₂ denoting transfer from LI to HI and MI to HI transfer, respectively). The changes of chlorophyll (Chl) fluorescence, net photosynthetic rate (P _N), dark respiration rate (R _D), Chl content per unit area (Chl_area), leaf mass per unit area (LMA), and seedling mortality were monitored over two months after transfer. These parameters together with leaf anatomy of transferred and control seedlings (kept in LI, MI, and HI) were also examined after two months. No seedlings died during the two months. F_v/F_m, P _N, and Chl_area of the transferred seedlings decreased in the first 3 to 12 d. LH₁ leaves showed larger reduction in F_v/F_m (>23 % vs. <16 %) and slower recovery of F_v/F_m than LH₂ leaves. P _N started to recover after about one week of I transfer and approached higher values in all G. cowa seedlings and G. xanthochymus LH₁ seedlings than those before the transfer. However, P _N of G. bracteata seedlings approached the values before transfer. The final P _N values in leaves of transferred G. xanthochymus and G. cowa seedlings approached that of leaves kept in HI, while the final P _N values of transferred leaves of G. bracteata were significantly lower than that of leaves grown under HI (p<0.05). R _D of G. xanthochymus LH₁ seedlings and all G. cowa seedlings increased and approached the value of the seedlings in HI. The final Chl_area of both G. xanthochymus and G. cowa approached the values before transfer, but that of G. bracteata did not recover to the level before transfer. The final Chl_area of all transferred seedlings was not significantly different from that of seedlings in HI except that G. cowa LH₁ seedlings had higher Chl_area than that in HI. LMA decreased within 2 d and then increased continuously until about 30 d and approached the value under HI. Spongy/palisade mesophyll ratio decreased after transfer because of the increase in palisade thickness. Leaf thickness did not change, so LMA increase of transferred seedlings was mainly due to the increase of leaf density. Thus the mature leaves under LI and MI of G. xanthochymus and G. cowa are able to acclimate to HI by leaf physiological and anatomical adjustment, while G. bracteata had limited ability to acclimate to HI.