Roots of pioneer trees in the Amazonian rain forest

. On low fluvial terraces of the Caquetá River, Araracuara region, Colombia, tree root systems were examined with regard to their structure, underground stratification and biomass distribution. Excavations of skeleton roots and microscopic observation of terminal fine roots included ten species belonging to the Cecropia, Vismia, Miconia, Goupia, Clathrotropis and Brosimum genera. Roots of particular species varied in the external features of their periderm and structure of freshly cut slash. Coarse skeleton roots differed in the shape and position of the plagiotropic branches and positively geotropic taproots or sinkers. None of the observed species could be identified as a shallow-rooted tree, in spite of the general fine root concentration in the upper soil horizon. In the two seasons of study, the terminal roots displayed prevailingly secondary anatomical structure and did not maintain much primary anatomical tissue in their apices. Neither ectomycorrhizas nor endomycorrhizas were detected in the samples. In a set of regeneration stages the amount of tree roots linearly increased with the age of growth. However, fine roots below 2 mm in diameter shared 80-90% of all roots in the upper 20 cm layer of all sample plots. In an old-growth forest, the total tree root biomass amounted to 39 ton/ha, thus being comparable to the underground biomass observed in similar tropical forests.     

Variation in leaf toughness and phenolic content among five species of Australian rain forest trees

Several leaf characteristics, including toughness and total phenols and condensed tannins, were measured in Australian rain forest leaves of different ages and related to observed herbivory rates. In most cases, toughness and chemical toxicity increased as leaves aged, and corresponding insect grazing decreased. Herbivory losses ranged from 4.8% to 32.5% leaf area losses, and were more positively correlated with toughness than with phenolics. It is suggested that a suite of factors, including physical and chemical characteristics of leaves as well as spatial and temporal factors, interact to create variation in grazing intensities.     

Crown light environments of saplings of two species of rain forest emergent trees

Summary The crown light environments of saplings of two Costa Rican rain forest tree species were simultaneously compared. The species, Dipteryx panamensis (Pitt.) Record & Mell., a relatively shade-intolerant species, and Lecythis ampla Miers, a shade-tolerant species, have contrasting growth and branching patterns. Quantum sensors were placed throughout the crowns of saplings up to 2.5 m tall and quantum fluxes were recorded with microloggers for seven-day periods. The shade-intolerant species had total quantum flux densities 35% larger than those of the shade-tolerant species, but totals for both species were less than 2% of full sun. More than 90% of the quantum flux densities measured within the crowns of both species were less than 25 mol m^-2s^-1. Lateral light was an important component of daily quantum flux totals; for saplings of both species, the half-hour with the maximum average irradiance for the day frequently occurred in mid-morning or midafternoon. Despite dissimilar crown and leaf display, there was no difference in the overall variability of irradiance within the crowns of the two species. However, quantum fluxes received within the crowns differed substantially in both species. Within-crown locations differed significantly from day to day because of variation in weather conditions. Daily total quantum flux densities and totals expressed as a percent of full sun were significantly correlated with height growth over the previous 12 months.     

Reproductive allometry in Malaysian rain forest trees: Biomechanics versus optimal allocation

Summary Quantitative predictions of reproductive allometry in iteroparous plants may be derived from two bodies of theory: biomechanics and optimal allocation theory. Biomechanical theory predicts allometric scaling exponents between reproductive (R) and vegetative (V) biomass in the range of 0.44–1.33, while very general models of life history evolution predictR–V exponents > 1 in all cases. These predictions are examined in light of allometric patterns of flower and fruit production in 32 species of Malaysian rain forest trees. Among these species the mean estimatedR–V exponents are in the range 1.8–2.0 for staminate flower, pistillate flower and fruit production. This range of exponent values provides unambiguous support for some of the general predictions of optimal allocation models, but not for biomechanical theory. Optimal allocation models also predict a positive relationship between species size andR–V slope and a positive relationship between species size andR–V intercept parameters. The latter, but not the former prediction is supported by the data.R–V allometries in sexes of dioecious species were also found to differ in intercept, though not slope, reflecting smaller sizes at reproductive onset in staminate trees. Further critical examinations of reproductive allometry are encouraged as a relatively unexplored avenue for increasing the contact of theory and data in studies of life history evolution in long-lived organisms such as tropical trees.     

Leaf and photosynthetic characteristics of pioneer and forest species in tropical montane habitats

Gas exchange and chlorophyll a fluorescence measurements of expanding and adult leaves of four plant species were compared under field conditions. The pioneer species (PS) tended to have thinner leaves with lower nitrogen content and higher stomatal density compared to forest species (FS). Expanding leaves featured lower photosynthetic pigment contents and gas exchange capacity than adult leaves consistent with an immature photosynthetic apparatus. At the time of maximum irradiance, sun-exposed leaves of both PS and FS showed alteration of initial, variable, and maximum fluorescence as well as their ratios indicating photoinhibition. However, leaves recovered to some extent at predawn, suggesting the activation of photoprotective mechanisms. Sun-exposed leaves had comparable responses to high irradiance.     

Seedling growth of five tropical dry forest tree species in relation to light and nitrogen gradients

Aims Increasing anthropogenic nitrogen (N) deposition has been claimed to induce changes in species composition and community dynamics. A greenhouse experiment was conducted to examine the effect of increased N availability on growth and functional attributes of seedlings of five tree species with different life history characteristics under varying irradiances. The following questions have been addressed: (i) how do the pioneer and non-pioneer species respond in absolute growth and relative growth rate (RGR) to the interaction of light and nitrogen? (ii) how does the interaction between irradiance and nitrogen availability modulate growth attributes (i.e. functional attributes)? (iii) is there any variation in growth responses between leguminous and non-leguminous species along the light and nitrogen gradients?Methods Seedlings of five tree species (Acacia catechu, Bridelia retusa, Dalbergia sissoo, Lagerstroemia parviflora and Terminalia arjuna) were subjected to twelve combinations of irradiance and N levels. Various growth traits, including height (HT), basal area (BA), whole plant dry biomass (M D), leaf mass per unit area (LMA), leaf area ratio (LAR), net assimilation rate (NAR), RGR, biomass fractions, root-to-shoot ratio (R:S) and leaf nitrogen content, were studied to analyse intra- and inter-specific responses to interacting light and N gradients.Important findings Significant interactions for irradiance and N availability for majority of growth attributes indicates that growth and biomass allocation of seedlings were more responsive to N availability under high irradiance. However, species responded differentially to N addition and they did not follow successional status. Slow growers (B. retusa, a shade-tolerant species and L. parviflora, a light demander) exhibited greater response to N enrichment than the fast growers (A. catechu, D. sissoo and T. arjuna). However, N-mediated increment in growth traits was greater in non-legumes (B. retusa, L. parviflora and T. arjuna) compared with that of legumes (A. catechu and D. sissoo). Allocation of biomass to root was strongly suppressed at the highest N supply across species; however, at high irradiance and high N availability, a greater suppression in R:S ratio was observed for B. retusa. NAR was a stronger determinant of RGR relative to LAR, suggesting its prominent role in increased RGR along increasing irradiances. Overall, a higher growth response of slow-growing species to elevated N levels, particularly the non-pioneers (B. retusa and L. parviflora) suggests that future N deposition may lead to perturbations in competition hierarchies and species composition, ultimately affecting community dynamics in nutrient-poor tropical dry forests.     

Tree species distributions across five habitats in a Bornean rain forest

We examined the distribution of tree species across five habitats in 69 small plots within a single watershed of the Gunung Palung National Park, West Kalimantan, Indonesia (GPNP). The spatially complex distribution and close proximity of habitats provided an opportunity to test habitat specificity of tree species across strong environmental gradients, in a situation where dispersal into ‘inappropriate’ habitat should not be a rare event. Habitat had a weak influence on community structure, although species diversity was lower in the alluvium and peat habitats. Association tests based on two randomization models (spatially independent and explicit) were used to examine habitat distribution of 55 ‘common’ and 142 ‘frequent’ taxa. The general patterns were similar in the two models but the interpretation of specific patterns depended greatly on assumptions about dispersal ability. A majority (67%) of the common species was significantly associated with a single habitat, while few were restricted to one habitat. A small proportion (16%) of the species appear to be habitat generalists. The peat habitat had the most profound effect on species distribution. Overall, a large amount of variation was found in the degree of habitat specificity, even within speciose groups. No obvious evolutionary or ecological correlates with degree of habitat specificity were found. These results suggest that a mixture of stochastic and deterministic processes determine species distribution even across strong environmental gradients.     

Leaf traits determine the growth-survival trade-off across rain forest tree species

A dominant hypothesis explaining tree species coexistence in tropical forest is that trade-offs in characters allow species to adapt to different light environments, but tests for this hypothesis are scarce. This study is the first that uses a theoretical plant growth model to link leaf trade-offs to whole-plant performances and to differential performances across species in different light environments. Using data of 50 sympatric tree species from a Bolivian rain forest, we observed that specific leaf area and photosynthetic capacity codetermined interspecific height growth variation in a forest gap; that leaf survival rate determined the variation in plant survival rate under a closed canopy; that predicted height growth and plant survival rate matched field observations; and that fast-growing species had low survival rates for both field and predicted values. These results show how leaf trade-offs influence differential tree performance and tree species' coexistence in a heterogeneous light environment.     

Leaf and soil nitrogen and phosphorus availability in a neotropical rain forest of nutrient-rich soil

The nitrogen and phosphorus supply in a lowland rain forest with a nutrient-rich soil was investigated by means of the leaf N/P quotient. It was hypothesised a high N and P supply to the forest ecosystem with a N and P rich soil. Total N and extractable P were determined in the surface (10 cm) soil of three plots of the forest. Total N was analysed by the Kjeldahl method, and P was extracted with HCI and NH4F. The leaf N/P quotient was evaluated from the senesced leaves of 11 dominant tree species from the mature forest. Samples of 5 g of freshly fallen leaves were collected from three trees of each species. Nitrogen was analysed by microkjeldahl digestion with sulphuric acid and distilled with boric acid, and phosphorus was analysed by digestion with nitric acid and perchloric acid, and determined by photometry. Concentrations of total N (0.50%, n = 30) and extractable P (4.11 microg g(-1), n = 30) in the soil were high. As expected, P supply was sufficient, but contrary to expected, N supply was low (N/P = 11.8, n = 11).     

Fruit trees in a Malaysian rain forest

An inventory was made of 50 ha of primary lowland rain forest in Peninsular Malaysia, in which ca. 340,000 trees 1 cm dbh or larger were measured and identified to species. Out of a total plot tree flora of 820 species, 76 species are known to bear edible fruit. Especially diverse were the wild species of mango (Mangifera, Anacardiaceae, 12 spp.), mangosteen (Garcinia, Clusiaceae, 13 spp.), breadfruit (Artocarpus, Moraceae, 10 spp.) and rambutan (Nephelium, Sapindaceae, 5 spp.). Median population size for all species of fruit trees was 3.0 trees per ha and 0.2 adult trees per ha. Direct economic value of wild fruit trees was small; only one species has been very much collected and sold, Parkia speciosa (Fabaceae), amounting to less than US$20 per ha per year. The potential value of the species as genetic resources is very large: 24 species are cultivated, 38 edible species are congeneric with cultivated crops and at least 10 other species bear inedible fruit but are related to cultivated crops. We conclude that the Peninsular Malaysian rain forest is exceedingly rich in wild fruit trees, that these normally live at low densities, and that their principal economic value is as genetic resources.     

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.     

Fire survival of lowland tropical rain forest trees in relation to stem diameter and topographic position

The objective of this study was to relate patterns in forest structure, tree species diversity, and tree species composition to stem diameters and topography in unburned, once burned and twice burned lowland dipterocarp rain forests in East Kalimantan, Indonesia. To do this four unburned old growth forests were compared with three forests that burned once (1997/1998) and three forests that burned twice (1982/1983 and 1997/1998). Fire resulted in a strong reduction of climax tree density which was negatively related to tree diameter. However, a disproportionate reduction in small diameter understorey climax tree species occurred only after repeated fires. Climax tree species in both burned forest types were most common in swamps, river valleys and on lower slopes, while their density was much lower on places higher along hillsides. In unburned forest the opposite was observed, with climax tree density increasing steadily from swamp and river valleys to upper slopes and ridges. In contrast to climax trees, pioneer trees were abundant throughout the burned forest, with highest numbers on hill sides and ridges. Our results indicate that both diameter and topographic position of trees strongly affect their fire survival chances in tropical lowland forests.     

Phylogeography of a species complex of lowland Neotropical rain forest trees (Carapa,Meliaceae)

Aim Many tropical tree species have poorly delimited taxonomic boundaries and contain undescribed or cryptic species. We examined the genetic structure of a species complex in the tree genus Carapa in the Neotropics in order to evaluate age, geographic patterns of diversity and evolutionary relationships, and to quantify levels of introgression among currently recognized species. Location Lowland moist forests in the Guiana Shield, the Central and Western Amazon Basin, Chocó and Central America. Methods Genetic structure was analysed using seven nuclear simple sequence repeats (nuSSR), five chloroplast SSRs (cpSSR), and two chloroplast DNA (cpDNA) intergenic sequences (trnH–psbA and trnC–ycf6). Bayesian clustering analysis of the SSR data was used to infer population genetic structure and to assign 324 samples to their most likely genetic cluster. Bayesian coalescence analyses were performed on the two cpDNA markers to estimate evolutionary relationships and divergence times. Results Two genetic clusters (nu_guianensis and nu_surinamensis) were detected, which correspond to the Neotropical species C. guianensis (sensu latu) and C. surinamensis. Fourteen cpDNA haplotypes clustered into six haplogroups distributed between the two nuclear genetic clusters. Divergence between the haplogroups was initiated in the Miocene, with some haplotype structure evolving as recently as the Pleistocene. The absence of complete lineage sorting between the nuclear and chloroplast genomes and the presence of hybrid individuals suggest that interspecific reproductive barriers are incomplete. NuSSR diversity was highest in C. guianensis and, within C. guianensis, cpDNA diversity was highest in the Central and Western Amazon Basin. Regional genetic differentiation was strong but did not conform to an isolation‐by‐distance process or exhibit a phylogeographical signal. Main conclusions The biogeographical history of Neotropical Carapa appears to have been influenced by events that took place during the Neogene. Our results point to an Amazonian centre of origin and diversification of Neotropical Carapa, with subsequent migration to the Pacific coast of South America and Central America. Gene flow apparently occurs among species, and introgression events are supported by inconsistencies between chloroplast and nuclear lineage sorting. The absence of phylogeographical structure may be a result of the ineffectiveness of geographical barriers among populations and of reproductive isolation mechanisms among incipient and cryptic species in this species complex.     

Species number, species abundance and body length relationships of arboreal beetles in Bornean lowland rain forest trees

ABSTRACT.

• 1 The relationships between number of species, abundance per species, and body length are examined for 859 species of beetles in samples of arthropods collected from ten Bornean lowland forest trees by insecticide fogging. Similar relationships are examined for different feeding guilds of these beetles, and for those beetles from different species of trees.
• 2 The data are used to construct four interrelated graphs, namely species: abundance, species: body length, population abundance: body length and total number of individuals: body length distributions.
• 3 In contrast to a number of previous studies, no consistent linear relationship between population density and body length was found for the Bornean beetles and it is suggested that, as in birds, the added dispersal ability of flight reduces critical population densities necessary for persistence in small species. Previous relationships between body weight and population abundance may also be artefacts of the way in which data were gathered.
• 4 Despite large samples, we failed to locate the mode in plots of the number of species in each abundance category (species: abundance distribution).
• 5 Species: body length and total number of individuals: body length plots were similar to those found in previous studies, although using data for Coleoptera alone may have produced a steeper decline in the total number of individuals as body size increases than is apparent in samples of all arthropods.
• 6 We present the first three-dimensional graph relating numbers of species, body lengths and population abundances. The surface of this three-dimensional relationship is relatively simple.

     

Reproductive patterns of selected understorey trees in the Malaysian rain forest: the sexual species

HA, C. O., SANDS, V. E., SOEPADMO, E. & JONG, K., 1988. Reproductive patterns of selected understorey trees in the Malaysian rain forest: the sexual species. Investigations were carried out on the floral, fruit and seed biology, and the embryology of representatives of the Sapindaceae and Bombacaceae which occur in the lower canopy of the lowland rain forest of Peninsular Malaysia. Pollination studies indicated cross-compatibility and outbreeding in all species examined, associated with self-compatibility in the polygamous monecious Pometia pinnata and Allophylus cobbe and, to a limited extent, in the effectively dioecious Xerospermum intermedium . In these sapindaceous species, together with the self-incompatible androdioecious Nephelium lappaceum , the structurally hermaphrodite flowers were determined embryologically to be functionally female with no anther dehiscence. The self-incompatible Durio griffithii of the Bombacaceae, and X. intermedium, N. lappaceum and P. pinnata were shown by embryological studies to form a sexual zygote. The observations on these outcrossing species are considered in relation to their potential for gene exchange and their contribution to the maintenance of species diversity in the lowland rain forest.