Distribution patterns of tree species in a Malaysian tropical rain forest

Abstract. Spatial patterns of tree species were studied in a 50-ha tropical rain forest plot in the Pasoh forest, Malaysia. This forest is characterized by a high diversity and very high number of rare species. Out of the 745 species occurring with > five individuals, 80.4 % had an aggregated distribution, 19.5% were randomly distributed and one species had a regular distribution. The spatial patterns of rare vs. common species, juvenile vs. adult trees, and coarse vs. fine scales were compared. Rare species are generally less aggregated than common ones and most of the randomly distributed species are rare. Spatial patterns shift from high clumping to looser intensity or random distribution when moving from juveniles to adults for the same species. No adult tree species display a regular pattern, however. Regular distributions were rarely found; this is probably due to intraspecific competition at a local scale. There is a negative correlation between per capita death rate and population density. This study suggests that the Pasoh forest and its high diversity are subjected to multiple controlling factors, e.g., topography, spacing effect, density-dependent processes and species rarity. The importance of any factor changes across spatial and temporal scales.     

Plurality of tree species responses to drought perturbation in Bornean tropical rain forest

Drought perturbation driven by the El Niño Southern Oscillation (ENSO) is a principal stochastic variable determining the dynamics of lowland rain forest in S.E. Asia. Mortality, recruitment and stem growth rates at Danum in Sabah (Malaysian Borneo) were recorded in two 4-ha plots (trees ≥ 10 cm gbh) for two periods, 1986–1996 and 1996–2001. Mortality and growth were also recorded in a sample of subplots for small trees (10 to <50 cm gbh) in two sub-periods, 1996–1999 and 1999–2001. Dynamics variables were employed to build indices of drought response for each of the 34 most abundant plot-level species (22 at the subplot level), these being interval-weighted percentage changes between periods and sub-periods. A significant yet complex effect of the strong 1997/1998 drought at the forest community level was shown by randomization procedures followed by multiple hypothesis testing. Despite a general resistance of the forest to drought, large and significant differences in short-term responses were apparent for several species. Using a diagrammatic form of stability analysis, different species showed immediate or lagged effects, high or low degrees of resilience or even oscillatory dynamics. In the context of the local topographic gradient, species’ responses define the newly termed perturbation response niche. The largest responses, particularly for recruitment and growth, were among the small trees, many of which are members of understorey taxa. The results bring with them a novel approach to understanding community dynamics: the kaleidoscopic complexity of idiosyncratic responses to stochastic perturbations suggests that plurality, rather than neutrality, of responses may be essential to understanding these tropical forests. The basis to the various responses lies with the mechanisms of tree-soil water relations which are physiologically predictable: the timing and intensity of the next drought, however, is not. To date, environmental stochasticity has been insufficiently incorporated into models of tropical forest dynamics, a step that might considerably improve the reality of theories about these globally important ecosystems.     

Seasonality of weather and tree phenology in a tropical evergreen mountain rain forest

Flowering and fruiting as phenological events of 12 tree species in an evergreen tropical mountain rain forest in southern Ecuador were examined over a period of 3–4 years. Leaf shedding of two species was observed for 12 months. Parallel to the phenological recordings, meteorological parameters were monitored in detail and related to the flowering and fruiting activity of the trees. In spite of the perhumid climate of that area, a high degree of intra- and inter-specific synchronisation of phenological traits was apparent. With the exception of one species that flowered more or less continuously, two groups of trees could be observed, one of which flowered during the less humid months (September to October) while the second group started to initiate flowers towards the end of that phase and flowered during the heavy rains (April to July). As reflected by correlation coefficients, the all-time series of meteorological parameters showed a distinct seasonality of 8–12 months, apparently following the quasi-periodic oscillation of precipitation and related cloudiness. As revealed by power spectrum analysis and Markov persistence, rainfall and minimum temperature appear to be the only parameters with a periodicity free of long-term variations. The phenological events of most of the plant species showed a similar periodicity of 8–12 months, which followed the annual oscillation of relatively less and more humid periods and thus was in phase or in counter-phase with the oscillations of the meteorological parameters. Periods of unusual cold or dryness, presumably resulting from underlying longer-term trends or oscillations (such as ENSO), affected the homogeneity of quasi-12-month flowering events, fruit maturation and also the production of germinable seeds. Some species show underlying quasi-2-year-oscillations, for example that synchronise with the development of air temperature; others reveal an underlying decrease or increase in flowering activity over the observation period, influenced for instance by solar irradiance. As Ecuador suffers the highest rate of deforestation in South America, there is an urgent need for indigenous plant material for reforestation. A detailed knowledge of the biology of reproduction in relation to governing external factors (mainly climate) is thus required.     

A new method for measuring tree height in tropical rain forest

Abstract. A new method is described for measuring tree height in tropical forest with a dense undergrowth, the so-called angles-measure method, which is related to the height-matching method formerly used by British foresters.     

Species diversity,susceptibility to disturbance and tree population dynamics in tropical rain forest

Abstract. In 1964 a census of all trees > 9.7 cm diameter at breast height (DBH) was conducted on 22 plots totalling 13.2 ha in lowland tropical evergreen rain forest on Kolombangara, Solomon Islands. Over the following 30 yr (1964–1994), populations of all individuals > 4.85 cm DBH of the 12 most common tree species and amounts of disturbance have been monitored on a declining number of these plots (in 1994, nine plots totalling 5.4 ha were still being recorded). Between November 1967 and April 1970, Kolombangara was struck by four cyclones, although only two of these caused substantial amounts of damage to the canopy structure. Multivariate analysis has identified six forest types on Kolombangara (Greig-Smith et al. 1967). The species richness and diversity of trees in the 1964 census, turnover rates of the populations monitored over 1964–1975, and the amount of disturbance sustained during a cyclone in 1970, were all positively correlated across five of the forest types. The sixth forest type was a consistent outlier in these analyses and is believed to have been seriously disturbed by humans about a century ago. The floristics, turnover and disturbance data support Connell's intermediate disturbance hypothesis. The most species-rich forest types contained a higher proportion of fast-growing individuals and species that are early-successional and which have low density timber. Properties of these species rendered them more susceptible to damage when struck by the 1970 cyclone. They showed higher turnover rates because disturbance-dependent species are also characterised by higher mortality and recruitment rates. Thus, periodic cyclones appear to favour the maintenance of differences in species diversity and composition between forest types.     

Nitrogen conservation in a tropical rain forest

Summary A series of plots in an Amazonian Rain Forest were trenched and treated with calcium to determine the effects of perturbation on numbers and activity of nitrifying bacteria. Although treatment resulted in between 2 and 26% of the nitrogen being lost from the humus layer, virtually all of it was in the NH ₄ ⁺ form. Numbers of nitrifying bacteria in the plots were relatively low. The low numbers and low activity of nitrifying bacteria is attributed to the low pH and high concentration of tannins in the root mat. It is hypothesized that the suppression of nitrifying bacteria results in nitrogen conservation in the rain forest.     

Seed rain and advanced regeneration in a tropical rain forest

By comparing seed rain, seedling and sapling community structures we assessed the possible role played by vertebrate seed dispersal as a structuring factor in advanced regeneration of closed-canopied sites in the tropical rain forest of Los Tuxtlas, Mexico. Seed weight, initial morphology of seedlings and species abundance were also analyzed to determine if these traits influenced the probability of establishment in the shade.About half of the seed species falling in five closed forest sites (25×25 m) during one year came from fruiting trees growing within the sites (local seeds) and half from fruiting trees found outside the sites (immigrant seeds). Seeds of liana and upper-canopy species were over-represented among immigrant seeds compared with seeds of understory tree species. This probably reflects the activity of frugivorous arboreal mammals, bats, and birds. Species with immigrant seeds had both a lower abundance and a narrower spatial distribution than locally produced seeds. Therefore, immigrant seeds showed higher diversity values than locally produced seeds.Average seed size and the proportion of epigeous seedlings were similar in local and immigrant species. Under closed-canopied sites, factors affecting community organization seem to operate selectively, favoring the establishment of large-seeded, local abundant species in the advanced regeneration. However, the fact that some saplings of immigrant species were found in the plots suggest that a slow species infiltration may be occurring leading to a slow shift in the advanced regeneration species composition. We propose that the influence of seed dispersal on advanced regeneration structure depends on the disturbance history of the patches where seeds land.     

Variation in seed and seedling traits in Pithecellobium pedicellare,a tropical rain forest tree

Summary Pithecellobium pedicellare, a mimosoid legume, is a large canopy tree in the tropical rain forests of Costa Rica. We examined the pattern of variation in seed weight, germination date, hypocotyl length (stem), and rachis length (the first leaf) of the seedlings in this species. Seeds collected from widely dispersed individual trees at the La Selva Biological Station, Costa Rica, were randomly planted in blocks, and grown under controlled, indoor conditions for about 2 weeks. There-fore, we were able to quantify the effects of maternal family on mean seed weight and the effects of maternal family and microenvironment on the remaining traits examined. A significant effect of maternal family was detected for all traits. In particular, the maternal effects on germination date and seedling size traits which were consistently significant even after controlling the initial seed weight may indicate that the maternal effects reflect, at least to some extent, maternal genetic control over these traits. Despite overall strong maternal effects, the performance of maternal siblings, such as the rachis length, differed among blocks. The sensitivity of maternal siblings to the local environments may contribute to the maintenance of genetic variability in this highly outcrossing tropical species.     

Wood mechanics, allometry, and life-history variation in a tropical rain forest tree community

Wood density plays a central role in the life-history variation of trees, and has important consequences for mechanical properties of wood, stem and branches, and tree architecture. Wood density, modulus of rupture, modulus of elasticity, and safety factors for buckling and bending were determined for saplings of 30 Bolivian rain forest tree species, and related to two important life-history axes: juvenile light demand and maximum adult stature. Wood density was strongly positively related to wood strength and stiffness. Species safety factor for buckling was positively related to wood density and stiffness, but tree architecture (height : diameter ratio) was the strongest determinant of mechanical safety. Shade-tolerant species had dense and tough wood to enhance survival in the understorey, whereas pioneer species had low-density wood and low safety margins to enhance growth in gaps. Pioneer and shade-tolerant species showed opposite relationships between species traits and adult stature. Light demand and adult stature affect wood properties, tree architecture and plant performance in different ways, contributing to the coexistence of rain forest species.     

Molecular systematic analysis reveals cryptic tertiary diversification of a widespread tropical rain forest tree

The broad geographic range of many Neotropical rain forest tree species implies excellent dispersal abilities or range establishment that preceded the formation of current dispersal barriers. In order to initiate historical analyses of such widespread Neotropical trees, we sequenced the nuclear ribosomal spacer (ITS) region of Symphonia globulifera L. f. (Clusiaceae) from populations spanning the Neotropics and western Africa. This rain forest tree has left unmistakable Miocene fossils in Mesoamerica (15.5-18.2 Ma) and in South America ( approximately 15 Ma). Although marine dispersal of S. globulifera is considered improbable, our study establishes three marine dispersal events leading to the colonization of Mesoamerica, the Amazon basin, and the West Indies, thus supporting the paleontological data. Our phylogeographic analysis revealed the spatial extent of the three Neotropical S. globulifera clades, which represent trans-Andes (Mesoamerica+west Ecuador), cis-Andes (Amazonia+Guiana), and the West Indies. Strong phylogeographic structure found among trans-Andean populations of S. globulifera stands in contrast to an absence of ITS nucleotide variation across the Amazon basin and indicates profound regional differences in the demographic history of this rain forest tree. Drawing from these results, we provide a historical biogeographic hypothesis to account for differences in the patterns of beta diversity within Mesoamerican and Amazonian forests.     

Phenology of a tropical rain forest in Malaya

During 1960-69, observations were made of flowering, fruiting and foliar activity of 61 canopy trees (representing 45 species) in a largely undisturbed tract of Hill Dipterocarp Forest. Rainfall was measured by an accumulating gauge and, at a nearby station, the number of rainy days recorded. Annually recurrent floral cycles were displayed by only ten species. In these and in 11 additional species, foliar cycles were also circannual. Gregarious flowering, apparently in response to drought, occurred in many species in 1963 and 1968. In other years, interspecific differences in periodicity maintained a minimal incidence of flowering at 44%, and fruiting at 27% of species in the sample. Despite specific variations in phenology, the community as a whole exhibited regular seasonality, with single annual peaks of flowering and fruiting and a double peak of leaf production.     

Effect of tree host species on fungal community composition in a tropical rain forest in Panama

Abstract.  To determine whether host species influence the composition of fungal communities, the ascomycetes and basidiomycetes present on three tree species ( Prioria copaifera (Fabaceae), Quararibea asterolepis (Bombacaceae), and Trichilia tuberculata (Meliaceae)) were sampled on the 50-ha Forest Dynamics Project plot in lowland moist tropical forest on Barro Colorado Island, Panama. The most abundant fungal morphotypes of both ascomycetes and basidiomycetes were generalists found on all three hosts, but detrended correspondence analysis revealed distinct differences in fungal community composition among host trees. These differences among hosts were constant across census years. Randomization tests revealed that there were significantly fewer host-generalist fungi than expected for ascomycetes but not for basidiomycetes. These results indicate that host composition plays a role in structuring both ascomycete and basidiomycete fungal communities, but that the most successful fungal morphotypes are capable of colonizing multiple host species.     

Shade tolerance and regeneration of some tree species of a tropical rain forest in Western Kenya

Shadetolerance and regeneration of some heavily exploited timber species(Uvariopsis congensis, Antiaris toxicaria, Funtumia africana,Prunus africana and Aningeria altissima)were studied in two adjacent sites in Kakamega tropical rain forest. The twosites have been subjected to different logging intensities. Within one of thesites, relatively intact areas were compared with adjoining more disturbedones.Seedling presence or absence in mature forest phase, sapling proportions in gapand the mature phase and diameter distributions were used to classify thespecies into shade tolerance groups. Results indicate that except forUvariopsis, the species are non-pioneer lightdemanders and could fit into three shade tolerance groups: A groupresembling pioneers but with seedling in shade, a mid-tolerant group andone showing a high shade tolerance. Amount of regeneration and disturbance wererelated: regeneration being higher in the more disturbed site except forUvariopsis. The intersite differences were mirrored whendisturbed and adjoining more intact areas within one of the sites werecompared.Species more abundant in the disturbed site also generally scored low in shadetolerance. However, the mid-tolerant Funtumia andnot Prunus or Aningeria that hadlowershade tolerance scores, dominated regeneration in the disturbed forest parts.The shade tolerance data suggest that Uvariopsis, Funtumiaand Antiaris could be amenable to natural forestmanagement. For Prunus and Aningeria,artificial regeneration in buffer plantations around the natural forest may beabetter option.     

A tropical rain forest feast

An Introduction to Tropical Rain Forests (2nd edn) by T.C. Whitmore Oxford Science Publications, 1998. ￡45 hbk, ￡19.50 pbk (224 pages) ISBN 0 19 850148 X/0 19 850147 1 Tropical Rain Forest: A Wider Perspective edited by F.B. Goldsmith Chapman and Hall (Conservation Biology Series) 1998. ￡69.00 hbk (xix+416 pages) ISBN 0 412 81510 9 Vanishing Treasures of the Philippine Rain Forest by L.R. Heaney and J.C. Rigalado Jr University of Chicago Press, 1998. ￡19.25/$24.00 pbk (96 pages) ISBN 0 91486819 5.     

Sandfly distribution and abundance in a tropical rain forest

1. The distribution patterns of sandflies (Diptera: Psychodidae) upon tree buttresses were studied in tropical rain forest at Finca la Selva in the Caribbean lowlands of Costa Rica. 2. Four species of sandfly, Lutzomyia shannoni Dyar, L. trapidoi F. & H., L. ylephiletor F. & H. and L. vespertilionis F. & H. comprising 97% of those caught, used tree buttresses as diurnal resting sites. Their distribution on the buttresses was aggregated. 3. During the dry season tree species had no significant effect upon the distribution of the sandflies. However, during the wet season the distributions of two of the species, L. trapidoi and L. ylephiletor, were significantly affected by the species of tree; it is suggested that some species of tree may provide greater protection from rainfall than others. 4. L. vespertilionis was restricted to a single buttress on each positive tree. Distribution of this species is evidently determined by the distribution of its host animal, the bat (Emballonuridae). Female flies feed upon the bat's blood and male flies may be attracted to the bat as it provides a source of female sandflies. It is suggested, therefore, that tree buttresses serve as sandfly swarming sites. 5. Within a large buttress the sandflies are not randomly distributed but are aggregated in particular areas. Within these aggregations, the sandflies are vertically zoned upon the buttress with a shift in species composition with height. Two hypothesis were suggested to account for this distribution pattern: a response to an environmental gradient or an interaction between the four species of fly.     

Pollen movement to flowering canopies of pistillate individuals of three rain forest tree species in tropical Australia

This study measured the quantities of effective pollen vectors and their pollen loads arriving at the canopies of dioecious tropical rain forest trees in north-east Queensland. Population flowering synchrony, effective pollinator populations and pollen loads transferred between staminate and pistillate trees were compared among three insect-pollinated tree species. All three were visited by a wide range of insects, 75% of which (mostly 3–6 mm long) carried conspecific pollen. Fewer than 8% of individual insects were found to be carrying single-species pollen exclusively and none could be described as specialist pollen foragers. The introduced honeybee carried greater quantities of pollen than any native species but was not necessarily a reliable pollinator. The brief flowering periods in Neolitsea dealbata (3–4 weeks) and Litsea leefeana (4–5 weeks) populations were synchronized among individuals. Flowering in the Diospyros pentamera population extended over 15 weeks and most individuals were in flower for most of this period. Staminate trees began flowering earlier, produced more flowers and attracted relatively more insects than did pistillate trees, suggesting a density-dependent response of pollinators to flowering performance. Pollen was trapped in greater quantities on insects at staminate trees than at pistillate trees. Insect numbers increased at peak flowering periods and Diptera were the most abundant flower visitors. Anthophilous Coleoptera were more numerous at staminate than at pistillate trees in all three tree species populations. Larger quantities of pollen were mobilized during peak flowering times although the greatest quantities were transferred to pistillate canopies towards the end of the population flowering periods. Diptera carried pollen more often to pistillate N. dealbata and L. leefeana trees than did other groups whereas Coleoptera carried pollen more often to pistillate D. pentamera trees. The two contrasting flowering performances in the three tree species are discussed with reference to mechanisms that facilitate pollen transfer between staminate and pistillate trees.     

云南热带季雨林及其与热带雨林植被的比较

在中国植物学文献中, 对热带季雨林的解释和运用是不一致的, 特别是易于把季雨林与热带雨林相混淆。季雨林是在具有明显干、湿季变化的热带季风气候下发育的一种热带落叶森林植被, 是介于热带雨林与热带稀树草原(savanna)之间的一个植被类型。云南的热带季雨林在分布生境、生态外貌特征、植物种类组成和地理成分构成上, 均与热带雨林有明显区别, 季雨林主要分布在海拔1 000 m以下的几大河流开阔河段两岸和河谷盆地, 其群落结构相对简单, 乔木一般仅有1至2层, 上层树种在干季落叶或上层及下层树种在干季都落叶; 在生活型组成上, 季雨林的木质藤本相对较少, 大高位芽植物及地上芽植物很少, 但地面芽植物很丰富, 地下芽植物和一年生植物也相对丰富; 在叶级和叶型特征上, 季雨林植物的小叶和复叶比例相对较高, 分别占到24%和44%; 在植物区系地理成分构成上, 季雨林的热带分布属合计也占绝对优势, 但以泛热带分布属的比例相对较高, 约占到总属数的30%, 热带亚洲至热带非洲分布属的比例也较高, 约占总属数的12%。季雨林的地理成分更为多样性, 起源与发展历史也更复杂和古老。