Floristic variation within kerangas (heath) forest: re-evaluation of data from Sarawak and Brunei

The variation in species composition of trees 7.6 cm gbh in thirty-eight plots (mostly c. 0.2 ha in extent) from physiognomically-defined kerangas forest were re-analyzed by principal components analysis ordination (species centering and standardization by sample norm). Analyses were performed separately on basal area abundances, on the densities of trees in three size classes (7.6, 30.5 and 61.0 cm gbh) and on the density of small and large trees (7.6-<30.5 and 30.5-<61.0 cm gbh). A total of 636 taxa were reduced to 381 for analysis, removing those of very low density and plot frequency.Three groups of plots were identified: forest at low elevation, and generally coastal, on deep humus podzols; forest at intermediate elevation on mostly red-yellow podzols with affinities to dipterocarp forest; and forest at high elevation on mostly peaty podzols. The first group was divisible into five subgroups along a drainage gradient, while the more poorly drained plots showing affinities to peatswamp forest. Forty to eighty of the taxa, depending on the criteria for selection, were sufficient to define a stable, reduced spatial structure of the data matrix. Two subgroups, both coastal on deep podzols, represent the extreme form of kerangas forest per se. A comparison of Agathis borneensis- and Shorea albida-dominated plots revealed few other associated and differentiating taxa.Patterns were clearest from analyses of basal area data and of densities of all and small trees. Ordinations and grouping of plots for small, but not large, tree densities were similar to those for basal area. Different species were differentiated on the basis of the abundance measure, leading to group (tabular) definition of associations in a dual manner. A new system of summarization is presented which combines basal area, density and frequency in a graded hierarchical approach.The association between vegetation and soil type was difficult to unravel because of the limited environmental space sampled. Soil type was confounded with elevation, rainfall and geographical location. A major factor is clay content probably affecting nutrient status and water holding properties. Modal analysis of small tree densities showed clearest patterns in this respect. There were no patterns at the family or genus level, nor in leaf size spectra within kerangas.Problems in the treatment, analysis and summarization of tropical forest data sets are discussed. These problems centre on the scale and intensity of field sampling and the advantages of measuring small trees leading to a dual basal area and density approach. All published studies, including this one, within kerangas forest have used inadequate sampling for the purposes of revealing species changes with respect to soil type and composition.     

The diversity of 15 ha of lowland mixed dipterocarp forest, Central Kalimantan

Fifteen hectares of primary forest in Central Kalimantan were enumerated and all trees 10 cm dbh tagged and identified to species as far as possible. Tree density, relative abundance of families and species diversity were calculated, as was a mean additive species/area curve for 1 ha and a species/area curve for all 15 ha. The Jaccard index of similarity was also calculated for the 15 plots. The 15 ha enumerated contain 8771 trees with a dbh of 10 cm, belonging to 1298 morphospecies in 56 families. On average a single plot had 583 trees and 205 species. These figures are within the upper range of species diversity for the Malesian region as a whole. Dipterocarpaceae make up 14% of all trees in the plots, closely followed by Euphorbiaceae with 13%. A surprisingly low coefficient of similarity is found between the plots, the lowest being 3% and the highest 30%. Species/area curve construction showed that the asymptote is not reached in any of the 1 ha plots or when the 15 plots are added together.     

Diversity,structure and dynamics of a gallery forest in central Brazil

The structure and dynamics of approximately 64 ha of undisturbed gallery forest were studied over six years. Trees from 31 cm gbh (c. 10 cm dbh) were measured every three years from 1985. They were in 151 (10×20 m) permanent plots in the Gama forest in the Federal District of Brazil. Natural regeneration (individuals under 31 cm gbh) was measured in subplots (of 2×2 m, 5×5 m and 10×10 m) within the 200 m² plots. The total tree flora (gbh31 cm) consisted of 93 species, 81 genera and 44 families in 1985. The Leguminosae, Myrtaceae and Rubiaceae were the families richest in number of species. Most individuals and species were under 45 cm diameter and 20 m high while the maximum diameter per species ranged from 30 to 95 cm. The density structure of trees and natural regeneration was similar, in which the densities of c. 80% of the species represented less than 1% of the total density. The periodic mean annual diameter increment for trees from 10 cm dbh, was c. 0.25 cm/year. Variability was high with coefficients of variation c. 100% or more. The Gama community may maintain tree diversity and structure in undisturbed conditions. Regeneration of c. 80% of the species was found in the establishing phase (poles); the diameter structure was typical of native forests with the number of individuals decreasing with increasing size classes and showing little change over the six years; recruitment compensated for the mortality of most of the abundant species. The soils in Gama gallery forest were dystrophic with high aluminium content. Multivariate analysis suggested the stream, natural gaps and edges as the main causes of floristic differentiation at the community level.     

Woody life-form composition and association on rainfall and soil fertility gradients in Ghana

The hypothesis that treelets/shrubs are more sensitive than trees to drought and low soil fertility was tested by analysing species densities (species per unit area) in 155 tropical forest plots (0.0625 ha) in Ghana, West Africa. Species densities of both life forms decreased with increasing dryness and with increasing soil fertility, which were correlated. Goodness of fit tests detected species preferences for four combinations of rainfall (dry: <1500 mm yr^-1, wet: 1500 mm yr^-1) and soil fertility (two classes based on a composite variable with high weightings for total exchangeable bases and pH), and a meta-analysis of these results compared life forms. The results contradicted the hypothesised response of life forms in two ways: (i) species densities of treelets/shrubs were comparably or relatively less reduced by dry conditions than were tree species densities, and (ii) treelet/shrub species did not show a more frequent preference for wet or fertile sites than did tree species. Light availability and rooting depth are discussed as explanations for the sensitivity of understorey plants to water and nutrient stress, and for the data's failure to support the hypothesis.     

Tree and shrub diversity and abundance in fragmented littoral forest of southeastern Madagascar

In the coastal littoral forest of extreme southeastern Madagascar, westudied tree diameter at breast height (DBH) 10 cm in 20, 50× 50 m plots in each of four forest fragments, andunderstorywoody vegetation (DBH < 10 cm, 1 m tall) in60,10 × 10 m plots in three of the fragments. Oneforestfragment was located in the highly degraded Lokaro region, and three in the nearbySainte-Luce forest. Atotal of 3476 trees, representing 169 species in 55 families, were recorded inthe50 × 50 m plots, and 10282 understory stems, representing195 species in 54 families, were found in the 10 × 10m plots. For each tree, DBH was recorded. Mean tree diameter andpatterns of tree size class distribution did not differ among the four forestfragments. However, the fragments differed significantly in both tree andunderstory stem densities, species richness and diversity values, and familyrichness values, with the Lokaro fragment having the lowest values for allmeasures. Furthermore, floristic patterns, family importance values, and communitysimilarity measures revealed that the species composition at theLokaro fragment was very different from the Sainte-Luce fragments. Anthropogenicdisturbance appears most pronounced in the isolated Lokaro forest, where bioticresources are limited to this single fragment.     

Tree species distribution in canopy gaps and mature forest in an area of cloud forest of the Ibitipoca Range,south-eastern Brazil

The tree community of both canopy gaps and mature forest was surveyed in a 5 ha plot of cloud forest in the Ibitipoca Range, south-eastern Brazil, aiming at: (a) comparing the tree community structure of canopy gaps with that of three strata of the mature forest, and (b) relating the tree community structure of canopy gaps with environmental and biotic variables. All saplings of canopy trees with 1–5 m of height established in 31 canopy gaps found within the plot were identified and measured. Mature forest trees with dbh 3 cm were sampled in four 40×40 quadrats laid on the four soil sites recognised in the local soil catena. All surveyed trees were identified, measured and distributed into three forest strata: understorey (<5 m of height), sub-canopy (5.1–15 m) and canopy (15.1–30 m). The following variables were obtained for each gap: mode of formation, age, soil site, slope grade, size, canopy openness and abundance of bamboos and lianas. A detrended correspondence analysis indicated that the tree community structure of gaps in all soil sites was more similar to that of the mature forest understorey, suggesting that the bank of immatures plays an important role in rebuilding the forest canopy and that gap phases may be important for understorey shade-tolerant species. There was evidence of gap-dependence for establishment for only one canopy tree species. Both canonical correspondence analysis and correlation analysis demonstrated for a number of tree species that the distribution of their saplings in canopy gaps was significantly correlated with two variables: soil site and canopy openness. The future forest structure at each gap is probably highly influenced by both the present structure of the adjacent mature forest and the gap creation event.     

Tree competition and species coexistence in a warm-temperate old-growth evergreen broad-leaved forest in Japan

The growth dynamics and mode of competition between adult trees 5.0cm in diameter at breast height (DBH) of nine abundant treespeciesoccupying ca. 85% of the total basal area were investigated in a 4ha study plot (200 m × 200 m) of awarm-temperate old-growth evergreen broad-leaved forest in the Tatera ForestReserve of Tsushima Island, southwestern Japan. In the plot, adult trees 5.0 cm DBH co-occurred with 35 woody plant species (except forwoody vine species). The most dominant and largest species,Castanopsis cuspidata var. sieboldiiexhibited a bimodal DBH distribution; it was found in both the upper and lowervertical layers. Other tree species had unimodal DBH distributionscorrespondingmostly to the lower vertical layer. We developed a model for individual growthincorporating both intra- and interspecific competition and degree ofcompetitive asymmetry. One-sided interspecific competition was detected in 17cases out of the 66 possible combinations on the scale of the 4 hastudy plot. The direction of interspecific competition was generally one-sidedfrom layer-I species to layer-II and III ones. The effects of two-sidedcompetition were detected only in layer-II and III species. OnlyDistylium racemosum exhibited one-sided intraspecificcompetition. We also found 11 cases of positive interspecific relationships.Generally, competitive relationships prevailed over positive relationshipsbetween adult trees in this warm-temperate evergreen broad-leaved forest.Competition between adult trees 5.0 cm in DBH did not occurinthe same vertical layer, but occurred only between trees in different verticallayers. This suggests that competition between adult trees 5.0cm in DBH plays a key role in the variation in species coexistencebetween different vertical layers on the 4 ha scale of thewarm-temperate evergreen broad-leaved forests. Moreover, it was found bycomparing with three different forest types that interspecific competition ismore intense in warm-temperate forests than in cool-temperate or sub-borealforests. We conclude that, compared to cool-temperate or sub-boreal forests(which have little interspecific competition), warm-temperate forests supportmore complex interspecific relationships and species-specific habitatpreferences that result in higher species diversity.     

Diversity of forest tree species in Yanbaru,the northern part of Okinawa Island

The natural forests of Yanbaru, in the northern part of Okinawa Island, harbor many endemic and endangered birds and mammals, and are dominated by an evergreen oak, Castanopsis sieboldii. The Simpson diversity (D) and equitability index (J) were calculated using survey data on number of stems ( 4.5 cm DBH) of each species found in sample plots.Near-climax old forests (age 50 yr, without pine trees) showed high species diversity of trees, 0.92 ± 0.01 in D and 0.83 ± 0.05 in J for trees of which DBH 4.5 cm, and 0.81 ± 0.04 in D and 0.75 ± 0.05 in J for trees of DBH 10 cm.These high values are comparable to those of tropical rain forests. Although even young forests showed high species diversity, diversity indices tended to increase with forest age.The U.S. Marine Corps leases the eastern half of Yanbaru which contains most of these near-climax forests. Conservation of natural forests in this area is recommended.     

High tree alpha-diversity in Amazonian Ecuador

In a 1 ha square plot of terra firme forest at 260 m elevation in Amazonian Ecuador, all trees with diameter at breast height (dbh) 5 cm were studied. There were 1561 individuals, 473 species, 187 genera and 54 families. Of these, 693 individuals, 307 species, 138 genera and 46 families had a dbh 10 cm. This is the highest number of tree species ever recorded for a tropical rain forest sample of this size. In both dbh classes, the most species-rich families were: Fabaceae sensu lato (including Mimosaceae and Caesalpiniaceae), Lauraceae and Sapotaceae; the most species-rich genera, were Pouteria, Inga and Protium. The vertical space was partitioned among species: 166 species were found only in the 5–10 dbh cm class and were mostly sub-canopy treelets, and 307 species with dbh 10 cm were mostly large canopy trees.     

Diversity and structure of tropical rain forest of Tutuila,American Samoa: effects of site age and substrate

We report tree community diversity, guild composition, and forest structure from three 1.2 ha (100 m × 120 m) permanent forest research plots on Tutuila, American Samoa, an isolated volcanic island in the South Pacific Ocean. Plots were established in three habitat types of lowland hill forest: two in mature tracts that differed in substrate type (talus vs non-talus), and a third in a 30–40 year-old abandoned plantation on non-talus soil. We encountered a total of 57 tree species 10 cm dbh. Richness was similar across sites, but composition differed substantially. We were able to classify with confidence 24 tree species into four distinct guilds based on forest-type preference: early successional or persistent successional (disturbed forest), generalist (no preference), or mature-phase. Sample size limitations or interactions between site age and substrate precluded categorization of all species. Thirty-eight percent (9/24) of the tree species were successional, a result which contrasts sharply with data from (formerly) continental forest in Panama. Spatial distributions of 33 species revealed 17 species exhibiting clumping or hyperdispersion (i.e., regular spacing) in at least one site. Possible non-anthropogenic mechanisms promoting clumping in the plots were (a) topography (edaphic), (b) gap affiliation, (c) inefficient or altered patterns of propagule dispersal, and (d) lack of natural seedling predators. Forest structure differed across site type, with stem densities highest in regenerating forest; conversely, regenerating forest had the lowest basal areas. Steep talus forest sequestered the most carbon (344.3 Mg ha^–1), and secondary forest sustained only 42% of levels found in talus forest (145.5 Mg ha^–1). Mature forest on non-talus soil sequestered the majority of carbon in mid-sized trees (30–50 cm dbh). Future assessments of land-use cover and biomass will provide for a complete estimate of the carbon budget of Tutuila. Finally, the results of this study suggest that conservation of the native fauna is essential in retaining the potential for regeneration of native forest after large-scale disturbance.     

Plant biodiversity and conservation of tropical semi-evergreen forest in the Shervarayan hills of Eastern Ghats, India

Species diversity, density, population structure and dispersion patterns of all trees and lianas (30cm gbh) were inventoried in a tropical semi-evergreen forest in the Shervarayan hills of Eastern Ghats, south India. Such data are necessary for ecosystem conservation of the under-studied Eastern Ghats, as extensive forests here have already been converted to coffee and orange plantations and the landscape changed due to aluminium ore mining and quarrying. Four 1-ha plots were established in Sanyasimalai (SM) reserve forest of the Shervarayan hills, one plot (SM1) located close to mining and quarrying area, two other contiguous plots (SM2 and SM3) located in selective felling area and the fourth (SM4) in a relatively undisturbed forest. These are 1 to 4km apart in the same semi-evergreen forest tract. In the four study plots a total of 3260 stems (mean density 815ha–1) covering 80 species in 71 genera and 44 plant families were recorded. Species richness was greatest in the undisturbed plot SM4 (50), while lowest (33) in the selectively felled site SM2. The forest stand (SM4) was also denser (986 stemsha–1) and more voluminous (basal area 44.3m2ha–1 as compared with the site mean of 35m2ha–1) than the other plots. Four trees, Chionanthus paniculata, Syzygium cumini, Canthium dicoccum and Ligustrum perrottetii dominated the stand, collectively contributing to >50% of the total density. Species richness and stand density decreased with increasing tree girths. The forest stand contained a growing population, but there was considerable variation in basal area distribution between the plots. Trends in species population structure varied, particularly for selective-felled species. Most species exhibited clumped dispersion of individuals both at 0.25ha and 1-ha scales. Variation in plant diversity and abundance are related to site attributes and human impacts.     

Primary forest dynamics in lowland dipterocarp forest at Danum Valley, Sabah, Malaysia, and the role of the understorey

Changes in species composition in two 4-ha plots of lowland dipterocarp rainforest at Danum, Sabah, were measured over ten years (1986-1996) for trees > or = 10 cm girth at breast height (gbh). Each included a lower-slope to ridge gradient. The period lay between two drought events of moderate intensity but the forest showed no large lasting responses, suggesting that its species were well adapted to this regime. Mortality and recruitment rates were not unusual in global or regional comparisons. The forest continued to aggrade from its relatively (for Sabah) low basal area in 1986 and, together with the very open upper canopy structure and an abundance of lianas, this suggests a forest in a late stage of recovery from a major disturbance, yet one continually affected by smaller recent setbacks. Mortality and recruitment rates were not related to population size in 1986, but across subplots recruitment was positively correlated with the density and basal area of small trees (10-< 50cm gbh) forming the dense understorey. Neither rate was related to topography. While species with larger mean gbh had greater relative growth rates (rgr) than smaller ones, subplot mean recruitment rates were correlated with rgr among small trees. Separating understorey species (typically the Euphorbiaceae) from the overstorey (Dipterocarpaceae) showed marked differences in change in mortality with increasing gbh: in the former it increased, in the latter it decreased. Forest processes are centred on this understorey quasi-stratum. The two replicate plots showed a high correspondence in the mortality, recruitment, population changes and growth rates of small trees for the 49 most abundant species in common to both. Overstorey species had higher rgrs than understorey ones, but both showed considerable ranges in mortality and recruitment rates. The supposed trade-off in traits, viz slower rgr, shade tolerance and lower population turnover in the understorey group versus faster potential growth rate, high light responsiveness and high turnover in the overstorey group, was only partly met, as some understorey species were also very dynamic. The forest at Danum, under such a disturbance-recovery regime, can be viewed as having a dynamic equilibrium in functional and structural terms. A second trade-off in shade-tolerance versus drought-tolerance is suggested for among the understorey species. A two-storey (or vertical component) model is proposed where the understorcy-overstorey species' ratio of small stems (currently 2:1) is maintained by a major feedback process. The understorey appears to be an important part of this forest, giving resilience against drought and protecting the overstorey saplings in the long term. This view could be valuable for understanding forest responses to climate change where drought frequency in Borneo is predicted to intensify in the coming decades.     

Effects of fire and selective logging on the tree species composition of lowland dipterocarp forest in East Kalimantan, Indonesia

Tree species composition (diameter at breast height (dbh) 10 cm) was studied in primary, selectively logged and heavily burnt forests in East Kalimantan, Indonesia. The number of trees, tree species, and the Fishers's- diversity index were determined for the first 15 years (burnt forest) and 25 years (selectively logged forest) after disturbance. Additionally the population structure of six common and typical Macaranga pioneer tree species was compared through time between selectively logged, burnt and primary forest. Both selectively logged and burnt forest showed a significant reduction in number of trees and tree species per surface area directly after disturbance. Fire especially affected dominant tree species, while for selective logging the opposite was observed. In selectively logged forest the number of trees, tree species and the Fishers's- index reached pre-disturbance levels within c. 15 years. For burnt forest, only the number of trees recovered to pre-disturbance levels. The number of tree species stayed constant after disturbance, while the Fishers's- index decreased. The six studied Macaranga pioneer tree species seedlings were present in all forest types. Their density seems to be unrelated to light levels in the forest understorey but strongly related to the number of mature parent trees. Their sapling densities were strongly related to light levels in the forest understorey. The studied Macaranga species formed an important part of both under- and over-storey in burnt forest 15 years after disturbance, while they were almost absent in the understorey and only moderately common in the overstorey of selectively logged forest.     

Liana diversity and host relationships in a tropical evergreen forest in the Indian Eastern Ghats

In tropical evergreen forest in the Kolli Hills of the Indian Eastern Ghats, four 2 ha (100 m × 200 m) replicate plots (two plots each in undisturbed and human-impacted sites), were inventoried for species diversity of lianas 5 cm girth at breast height (g.b.h.) and their relationships with 30 cm g.b.h. host trees. Liana diversity included 26 species from 18 families and 24 genera. The population density and basal area of lianas in the study plots were 48 individuals ha^–1 and 0.23 m² ha^–1, respectively, while those of the trees were 478 stems ha^–1 and 43.6 m² ha^–1, respectively. As the lianas and their hosts had often been cut in the disturbed sites, their diversity was less there than in the undisturbed sites. Five (19%) liana species were common to all four sites. Three lianas, Hiptage benghalensis (Malpighiaceae), Elaeagnus indica (Elaeagnaceae) and Gnetum ula (Gnetaceae) were dominant. The twining mechanism (54% of liana species and 71% of individuals) and zoochorous diaspores (73% of species and 77% of individuals) predominated. A total of 336 trees from 39 species, 34 genera and 22 families hosted 345 lianas. The ratio of liana : host for species was 1 : 1.5 and for individuals was 1 : 1. Liana preferences for certain host trees, host girth classes and trellis heights were evident.     

Effect of forest fragmentation on the woody flora of the highlands of Chiapas, Mexico

This study was conducted in the Chiapas Highlands, a tropical mountain region where traditional agricultural practices have resulted in a mosaic landscape of forest fragments embedded in a matrix of secondary vegetation and crop fields. The question addressed was how may woody species richness be affected by forest fragment attributes derived from traditional land-use patterns. Species inventories of total woody species, canopy and understorey trees, and shrubs were obtained in 22 forest fragments (5 ha). Multiple regression analyses were applied to examine the effects of size, matrix, isolation and shape of the forest fragments on richness of these species guilds. Fragment size was correlated with shape (r = 0.75) and isolation (r = –0.69), and isolation was correlated with shape (r = –0.75). Total species richness, and number of shrubs and understorey trees in fragments were related to isolation; moreover, additive effects of fragment shape were found for shrubs. The number of canopy species was not related to any fragment variable. Matrix did not help to explain species richness, possibly due to the landscape structure created by the traditional land-use patterns. In addition to size and isolation, we point out the need of considering shape and matrix as additional fragmentation attributes, along with social and economic factors, if we are ever going to be successful in our management and conservation actions.     

Aboveground biomass of tropical rain forest stands in Indonesian Borneo

Aboveground plant biomass was examined in a tall virgin tropical lowland evergreen rain forest dominated by Dipterocarpaceae in Sebulu, East Kalimantan, Indonesia, with special reference to the gap-, building- and mature phases of the forest growth cycle. From the records of dimensions of sample trees examined by the stratified clip technique and DBH inventory data of trees in a study plot, the biomass of larger trees (DBH 4.5 cm) was estimated by the allometric correlation method. The biomass of smaller plants (DBH < 4.5 cm) was estimated by harvesting the plants in small quadrat plots. Although large differences were found between aboveground-biomass-estimates in different patches of different growth stages, the aboveground biomass in a 1.0 ha plot was 509 t/ha, and the one-sided LAI was 7.3 ha/ha. These values seem to result from the tall forest architecture with huge emergent trees (over 70 m high) and a moderate packing of plant mass indicated by the basal area value of 38.8 m²/ha for trees with DBH 4.5 cm.This study was financed through a grant to H. Ogawa from the Overseas Scientific Research Funds of the Ministry of Education, Science, and Culture, Tokyo. Sponsorship from the Lembaga Ilumu Pengetahuan Indonesia (LIPI), Jakarta, the Lembaga Biologi National (LBN), Bogor, and the Herbarium Bogoriense, Bogor is gratefully acknowledged. We are also grateful to Drs Soetiyati, M. Rifai, K. Kartawinata, and the staffs of P. T. Kutai Timber Indonesia for their kind support, Dr K. Ogino for his advice and cooperation in field work, and Dr H. Kataoka for providing us with the geological map of Samarinda Province.     

Biodiversity inventory of trees in a large-scale permanent plot of tropical evergreen forest at Varagalaiar, Anamalais, Western Ghats, India

A permanent plot of 30 ha (600 × 500 m²) was established for long-term ecological research on biodiversity and forest functioning in a tropical evergreen forest at Varagalaiar, Anamalais, Western Ghats. This paper gives the results of the first census of trees in the 30 ha plot. All trees 30 cm gbh were permanently tagged with sequentially numbered aluminium tags and their girth measured. 148 tree species (in 120 genera and 49 families) were enumerated in a total sample of 13,393 individuals. Species diversity, density and dispersion patterns were determined. The mean species richness, density and basal area per hectare were 65 species, 446 stems and 36.26 m² respectively. The Sorensen similarity index for thirty 1-ha subplots varied from 0.7 to 0.9, indicating the homogeneity in species composition of the stand. The pronounced species hierarchy has resulted in the dominance of four species that occupy four different forest storeys: Drypetes longifolia (lower storey) Reinwardtiodendron anamallayanum (middle storey), Poeciloneuron indicum (upper storey) and Dipterocarpus indicus (emergent). The forest was dominated by Euphorbiaceae in terms of richness (18 species) and abundance of species (3788 stems), and Dipterocarpaceae based on the contribution to basal area (18.4%). Species richness and density decreased with increasing girth-class. Girth class frequency of the stand exhibited an expanding population of trees. Twenty-nine tree species, analysed for spatial dispersion at 1-ha scale, exhibited both clumped and uniform patterns, except Vateria indica which was randomly distributed in one hectare. Repeat census scheduled for subsequent years, will elucidate the behaviour of the individual species and the population dynamics of the tree flora and will be useful for forest conservation and management. It will also help to monitor human activity within this forest.     

A First Quantitative Census of Vascular Epiphytes in Rain forests of Colombian Amazonia

Epiphytism in Colombian Amazonia was described by counting vascular epiphytes in thirty 0.025-ha (5 × 50 m) plots, well-distributed over the main landscape units in the middle Caquetá area of Colombian Amazonia. Each plot was directly adjacent to a 0.1-ha plot at which the species composition of trees and lianas (diameter at breast height (DBH) 2.5 cm) had been recorded 3 years earlier. The purpose of the study was to explore abundance, diversity, and distribution of epiphytes between the principal landscape units. A total of 6129 individual vascular epiphytes were recorded belonging to 27 families, 73 genera, and 213 species (which included 59 morpho-species). Araceae, Orchidaceae, and Bromeliaceae were the most speciose and abundant families. A total of 2763 phorophytes were registered, 1701 (62%) of which with DBH 2.5 cm. About 40–60% of the woody plants with DBH 2.5 cm carried epiphytes, which points at low phorophyte limitation throughout all landscapes. Epiphytism was concentrated on stem bases. Just as trees, epiphyte species assemblages were well associated with the main landscapes. Contrary to trees, however, epiphyte abundance and diversity (species richness, Fishers alpha index) hardly differed between the landscapes. This calls for caution when explanations for distribution and dynamics of tree species are extrapolated to growth forms with a totally different ecology.     

Canopy dieback in lower montane forests of Alto Urimán,Venezuelan Guayana

Canopy dieback in patches was discovered in an otherwise undisturbed very humid lower montane forest on a quartzitic sandstone plateau of the middle Caroní river basin, Venezuela. The patches vary in size from about 0.1 to 2 ha (50 to 700 upper story dead trees per ha). Preliminary inventories were carried out at 3 selected sites comparing 4 dieback patches (M) with adjacent not affected (V) forests. In the M plots, 40 to 61% of all trees with dbh 10 cm were dead. They consisted mostly (81–100%) of the endemic tree species Terminalia quintalata of the upper story (dbh > 20 to 60 cm, height up to 30 m), which is much sparser in the V stands. Data on stand structure and species composition are presented. They seem to indicate a spatial variation in the density of Terminalia quintalata, but also of other dominant species, and reproductive problems of Terminalia, which is hardly present in the diameter classes below 10 cm dbh. V and M stands grow on similar soils with a perched high water table. They are practically free of clay and rich in humus. The exchange complex is dominated by H⁺. Low pH and dilution of base cations represent the main chemical stress factors. Different concepts of the dieback process are discussed, including cyclic reproductive patterns, nutrient sequestering and drought frequency.     

Structure and floristics of secondary and old-growth forest stands in lowland Costa Rica

We characterized stand structure and floristic composition of woody life forms in three, 16–18 yr old secondary stands that regenerated after pasture abandonment, and three nearby old-growth stands of tropical rain forest in lowland Costa Rica. Basal area and stem density for each of four plant size classes (seedlings, saplings, treelets, trees) were similar among stand types, but density of adult canopy palms (individuals 10 cm DBH), was lower in the secondary stands. We estimate that 15% of the basal area of stems 10 cm DBH correspond to remnant trees in our secondary stands. The observed rapid woody regrowth compared to other published studies in the lowland neotropics, can be attributed to moderate land use and possibly, to the influence of nutrient-rich volcanic soils in the study area. Overall, plant species richness was lower in the secondary stands, but this difference was less pronounced in the smallest size classes (seedlings, saplings). Median percent similarity of all pairwise stand comparisons showed that floristic composition of saplings (stems 1 m tall and 5 cm DBH) was more similar between secondary and old-growth stands than for trees (stems 10 cm DBH). Because the potential value of secondary forests in conserving woody plant diversity appears highest for the young size classes, we suggest that further studies on floristic composition, especially those addressing the dynamics of the understory component, are needed to refine our understanding of the role of this natural resource in the maintenance of plant biodiversity in disturbed landscapes.