Environmental correlates for tropical tree diversity and distribution patterns in Borneo

Aim Identify environmental correlates for tropical tree diversity and composition. Location Borneo, Southeast Asia. Methods A GIS‐environmental database with 5 arc minute (c. 10 × 10 km) resolution was combined with tree inventory data. Tree diversity, phylogenetic diversity (PD) and the two main compositional gradients were determined for 46 tree inventories. Akaike's information criterion and a data jackknifing procedure were used to select 50 explanatory models for diversity and composition gradients. The average of these models was used as our final diversity and compositional model. We applied Moran's I to detect spatial autocorrelation of residuals. Results Tree diversity, PD and the two main compositional gradients in Borneo were all significantly correlated with the environment. Tree diversity correlated negatively with elevation, soil depth, soil coarseness (texture) and organic carbon content, whereas it correlated positively with soil C:N ratio, soil pH, moisture storage capacity and annual rainfall. Tree PD was correlated positively with elevation and temperature seasonality and was largely determined by gymnosperms. However, angiosperm PD also correlated positive with elevation. Compositional patterns were strongly correlated with elevation but soil texture, cation‐exchange‐capacity, C:N ratio, C and N content and drainage were also important next to rainfall seasonality and El Niño Southern Oscillation drought impact. Main conclusions Although elevation is the most important correlate for diversity and compositional gradients in Borneo, significant additional variability is explained by soil characteristics (texture, carbon content, pH, depth, drainage and nutrient status) and climate (annual rainfall, rainfall seasonality and droughts). The identified environmental correlates for diversity and composition gradients correspond to those found in other tropical regions of the world. Differences between the regions are mainly formed by differences in the relative importance of the environmental variables in explaining diversity and compositional gradients.     

The effect of climatic gradients, topographic variation and species traits on the beta diversity of rain forest trees

Aim  We assessed the rates of turnover of tree species with distance (beta diversity) in wet forests of the Western Ghats (WG) complex of India to see whether climate, topographic variation or species traits influence beta diversity.
Location  The Western Ghats is a chain of mountains about 1600 km in length, running parallel to the western coast of the Indian Peninsula from above 8° N to almost 21° N latitude.
Methods  We used data from 60 small plot inventories concentrated in three regions: the southernmost part of the Western Ghats (SWG) (8°24' to 9°37' N), the Nilgiri Hills (11°12' to 11°14' N), and the central Western Ghats (CWG) (12°32' to 14°51' N). We used Sorensen's index (SI) to estimate the similarity in species composition between two plots and regressed SI against the logarithm of the distance between plots to assess beta diversity. A bootstrapping procedure provided confidence intervals for regression coefficients. To test for the effects of climate, we regressed seasonality differences between plots against SI for low-elevation (< 800 m) plots along the north–south axis, and all plots in the SWG. We assessed the impact of the rainfall gradient in the Kogar region.
Results  Among all three regions, beta diversity was highest along the latitudinal axis, and along the rainfall gradient in the Kogar region. Differences in seasonality between sites were strongly related to beta diversity along the north–south seasonality gradient and within the SWG. Within the three regions, beta diversity was highest in the region with the strongest rainfall gradient and lowest for the topographically heterogeneous SWG. Beta diversity did not differ between forest strata and dispersal modes.
Main conclusions  We conclude that climate, particularly seasonality, is probably the primary driver of beta diversity among rain forest trees of the Western Ghats complex.     

From Forest to Farmland: Species Richness Patterns of Trees and Understorey Plants along a Gradient of Forest Conversion in Southwestern Cameroon

Vegetation surveys were carried out at 24 sampling stations distributed over four land use types, namely near-primary forest, secondary forest, agroforestry systems and annual crop lands in the northeastern part of the Korup region, Cameroon, to assess the impact of forest conversion on trees and understorey plants. Tree species richness decreased significantly with increasing level of habitat modification, being highest and almost equal in secondary and near-primary forests. Understorey plant species richness was significantly higher in annual crop lands than in other land use types. The four land use types differed in tree and understorey plant species composition, the difference being smaller among natural forests. Tree and understorey plant density differed significantly between habitat types. Density was strongly correlated with species richness, both for trees and understorey plants. Five tree and 15 understorey plant species showed significant responses to habitat. A 90% average drop in tree basal area from forest to farmland was registered. Our findings support the view that agroforestry systems with natural shade trees can serve to protect many forest species, but that especially annual crop lands could be redesigned to improve biodiversity conservation in agricultural landscapes of tropical rainforest regions.     

El Niño droughts and their effects on tree species composition and diversity in tropical rain forests

In this study I investigated the effects of the extreme, 1997/98 El Niño related drought on tree mortality and understorey light conditions of logged and unlogged tropical rain forest in the Indonesian province of East Kalimantan (Borneo). My objectives were to test (1) whether drought had a significant effect on tree mortality and understorey light conditions, (2) whether this effect was greater in logged than in undisturbed forest, (3) if the expected change in tree mortality and light conditions had an effect on Macaranga pioneer seedling and sapling densities, and (4) which (a)biotic factors influenced tree mortality during the drought. The 1997/1998 drought led to an additional tree mortality of 11.2, 18.1, and 22.7% in undisturbed, old logged and recently logged forest, respectively. Mortality was highest in logged forests, due to extremely high mortality of pioneer Macaranga trees (65.4%). Canopy openness was significantly higher during the drought than during the non-drought year (6.0, 8.6 and 10.4 vs 3.7, 3.8 and 3.7 in undisturbed, old logged and recently logged forest, respectively) and was positively correlated with the number of dead standing trees. The increase in light in the understorey was accompanied by a 30 to 300-fold increase in pioneer Macaranga seedling densities. Factors affecting tree mortality during drought were (1) tree species successional status, (2) tree size, and (3) tree location with respect to soil moisture. Tree density and basal area per surface unit had no influence on tree mortality during drought. The results of this study show that extreme droughts, such as those associated with El Niño events, can affect the tree species composition and diversity of tropical forests in two ways: (1) by disproportionate mortality of certain tree species groups and tree size classes, and (2) by changing the light environment in the forest understorey, thereby affecting the recruitment and growth conditions of small and immature trees.     

Testing indicators of sustainable forest management on understorey composition and diversity in southern Italy through variation partitioning

Tree species composition and stand structural complexity are valuable indicators of sustainable forest management. This article aims to investigate the relative influence of forest overstorey composition and structural attributes on understorey composition and diversity, taking into account also site characteristics and broad-scale environmental variables. We sampled vascular plant species composition and forest structure in 132 plots in the Cilento and Vallo di Diano National Park (southern Italy). Spearman’s non-parametric correlation coefficients were calculated between overstorey and understorey diversity indices, beech percentage, and altitude and environmental indices. A complete partitioning of the variation in understorey composition was then performed through canonical correspondence analysis considering four sets of variables: (1) overstorey composition, (2) structural attributes, (3) topography, and (4) landscape abiotic variables. Finally, we constructed a regression tree analysis of understorey species richness using the same explanatory variables. Understorey diversity indices were positively correlated with overstorey diversity indices and with environmental indices (i.e., light and soil heterogeneity). Overstorey and understorey diversity indices were negatively correlated with both altitude and the dominance of beech in the overstorey. Compositional variation was due primarily to overstorey composition and secondarily to structural attributes. Regression tree analysis revealed that altitude, overstorey species richness, and structural attributes play an important role in determining understorey species richness. According to our results, understorey composition and diversity are strongly related to overstorey composition and structural attributes. Indeed, the latter proved to be effective indicators of understorey characteristics in the study area.     

Environmental,structural, and taxonomic diversity factors drive aboveground carbon stocks in semi-deciduous tropical rainforest strata in Cameroon

Forest stratification plays a crucial role in the interception of light and plants' photosynthetic activities. However, there is still a lack of information on the contribution of tropical forest stratification to its functioning, despite the increasing number of studies. Here, we analysed from a perspective of the whole tree community (WTC) and forest strata (i.e., large trees, understory trees, and small stems), the relationship between abiotic, biotic factors and aboveground Carbon (AGC). The abiotic factors-AGC relationships were positive for all strata and WTC. However, soil factors-AGC relationship was stronger for small stems and understorey, while topography factor-AGC relationship was stronger for large trees and WTC. Tree size inequality-AGC relationship was positive and much stronger for WTC, large trees and small stems. In addition, a species diversity-AGC relationship was found positive only for large trees and WTC. These results highlight the niche complementarity effect for driving positive relationships of species diversity and individual tree size variation with aboveground biomass at large tree strata and WTC. The lack of positive effect of species diversity on AGC for understorey and small stems strata might be attributable to the selection effect or resource complementarity among species.     

Nitrogen‐fixing tree abundance in higher‐latitude North America is not constrained by diversity

The rarity of nitrogen (N)‐fixing trees in frequently N‐limited higher‐latitude (here, > 35°) forests is a central biogeochemical paradox. One hypothesis for their rarity is that evolutionary constraints limit N‐fixing tree diversity, preventing N‐fixing species from filling available niches in higher‐latitude forests. Here, we test this hypothesis using data from the USA and Mexico. N‐fixing trees comprise only a slightly smaller fraction of taxa at higher vs. lower latitudes (8% vs. 11% of genera), despite 11‐fold lower abundance (1.2% vs. 12.7% of basal area). Furthermore, N‐fixing trees are abundant but belong to few species on tropical islands, suggesting that low absolute diversity does not limit their abundance. Rhizobial taxa dominate N‐fixing tree richness at lower latitudes, whereas actinorhizal species do at higher latitudes. Our results suggest that low diversity does not explain N‐fixing trees' rarity in higher‐latitude forests. Therefore, N limitation in higher‐latitude forests likely results from ecological constraints on N fixation.     

Diurnal foraging ant–tree co-occurrence networks are similar between canopy and understorey in a Neotropical rain forest

Discussion of the vertical stratification of organisms in tropical forests has traditionally focused on species distribution. Most studies have shown that, due to differences in abiotic conditions and resource distribution, species can be distributed along the vertical gradient according to their ecophysiological needs. However, the network structure between distinct vertical strata remains little-explored. To fill this gap in knowledge, we used baits to sample ants in the canopy and understorey trees of a Mexican tropical rain forest to record the ant–tree co-occurrences. We examined the ant–tree co-occurrences in the canopy and understorey using complementary network metrics (i.e., specialization, interaction diversity, modularity, and nestedness). In addition, we evaluated co-occurrence patterns between ant species on trees, using C-score analysis. In general, we found no differences in the network structure, although the interaction diversity was greater in the understorey than in the canopy networks. We also observed that co-occurrence networks of each vertical stratum featured four ant species in the central core of highly co-occurring species, with three species unique to each stratum. Moreover, we found a similar trend toward ant species segregation in the both strata. These findings reveal a similar pattern of ant–ant co-occurrences in both vertical strata, probably due to the presence of arboreal-nesting ants in the understorey. Overall, we showed that despite the marked differences in species composition and environmental conditions between understorey and canopy strata, ant–tree co-occurrences in these habitats could be governed by similar mechanisms, related to dominance and resource monopolization by ants.     

Density and species diversity of trees in four tropical forests of the Albertine rift, western Uganda

We assessed tree species density and diversity in 12 1‐ha plots in four forests of the Albertine rift, western Uganda. There were 5747 trees of diameter ≥ 10 cm in 53 families, 159 genera, and 212 species. Density ranged between 344 and 557 trees ha^?1 (average 479 trees ha^?1). Tree species diversity was highest in the Bwindi and Budongo forests. The Euphorbiaceae family was the most species rich (25 species) followed by Rubiaceae and Meliaceae with 16 species each. Canonical Correspondence Analysis (CCA) showed that major gradients in environmental variables influenced tree species distribution. Sample scores on ordination axes 1 and 2 were strongly correlated with pH and altitude, respectively. Correlated with rainfall and other soil factors, pH and altitude are presumed to be among the most important in influencing the distribution of tree species in the Albertine rift forests. Strategies that take account of variations in pH and elevation are required to conserve tree species in forests of the Albertine rift.     

Tree species diversity of tropical forest vegetation in Xishuangbanna, SW China

Tree species diversity of four tropical forest vegetation types was investigated in Xishuangbanna, southwestern China. These are: tropical seasonal rain forest, tropical montane rain forest, evergreen broad-leaved forest and monsoon forest over limestone. A total of 17 samples were taken and four species diversity indices were calculated: Shannon-Wiener's H, the complement of Simpson's index, d, Fisher's and evenness index E. The results reveal the long-tailed rank/abundance diagrams of these forests. However, this feature is greatly reduced in the samples of monsoon forest over limestone. Tropical seasonal rain forest shows the highest tree species diversity of all four vegetation types. Owing to the variation of microenvironment, diversity values within the same vegetation type vary between the samples from different patches. The tree species diversity of single-dominant rain forest is not significantly lower than that of mixed rain forest, because the dominant species of some single-dominant rain forests are principally in the emergent layer. This is composed of sparse and huge trees of one species and, consequently, creates a unique canopy architecture and more heterogeneous microenvironments for the more diversified species composition under the emergent layer. The occurrence of tree species with small population sizes, particularly of species represented by only one individual, is highly correlated with the tree species diversity of the local forest vegetation. They are crucial elements in the richness of local biodiversity.     

Tree species richness and turnover throughout New Zealand forests

Abstract. Patterns of mortality, recruitment, and forest turnover were investigated using permanent plot data from temperate forests in 14 localities throughout New Zealand. Tree mortality and recruitment rates were calculated from tagged trees ≥ 10 cm diameter at 1.4 m on individual 400 m² plots, and turnover rates were calculated as the mean of mortality and recruitment rates. Turnover rates (1.4% per year) were very similar to those recorded for tropical forests (i.e. 1.5% per year). As was shown in tropical forests, we also found significant relationships between forest turnover and species richness. In New Zealand forests there was also a decrease in species richness and turnover rates with increasing latitude. Although species richness is well known to decline with latitude, our study provides support for a possible link between seasonality and disturbance with tree turnover and species diversity. While tree mortality and recruitment rates were approximately in balance at some localities, in others there were imbalances between mortality and recruitment rates.     

The Relationship between Local Abundance and Distribution of Rain Forest Trees across Environmental Gradients in India

We tested whether local abundance of rain forest trees in the medium elevation wet forests of the southern Western Ghats (WG) was related to environmental tolerance, life form, and geographical range. We selected trees in medium elevation wet forests (750–1700 m asl) of the southern WG, using two data bases: a small plot (30 × 30 m) data base of 288 species of trees (≥ 3 cm dbh) in 33 plots totaling 2.97 ha, and a data base of 135 species of tree (≥ 10 cm dbh) in larger plots of 1 ha each, totaling 4.84 ha. The species density per hectare and number of records in the plot network was used in a factor analysis to give a measure of the local abundance of each species. The altitude and seasonality ranges of these species in the WG was assessed from independent data bases and used to generate an environmental tolerance score. Results indicated that as a species became locally more abundant, it occurred across a wider range of environmental gradients, but regional distribution was not related to geographical distribution. Understory species tended to be rarer with smaller range sizes and lower environmental tolerances than overstory species. Climate change is predicted to have drastic effects on restricted range species with limited environmental tolerances.     

Contrasting patterns of vertical stratification in two moth families in a Costa Rican lowland rain forest

The vertical stratification of two species-rich moth families (Lepidoptera: Arctiidae, Geometridae) was analysed in a lowland rain forest in Costa Rica. Moths were trapped with automatic 8 W ultraviolet light traps at three sites in the understorey and canopy of a primary forest at the La Selva Biological Station (10.4° N, 84.0° W). A total of 846 arctiid moths (148 species) and 946 geometrid moths (140 species) were analysed. Species richness and diversity of arctiid moths was significantly lower in the understorey than in the canopy (Fisher's alpha: 24 vs. 49) whereas geometrid moths showed an inverse pattern (Fisher's alpha: 44 vs. 27). This resulted in an overall increase in the proportion of conspicuously coloured species towards the canopy. Moth ensembles were clearly separated in multidimensional scaling ordinations, and differed significantly in their faunal composition and dominance between the strata. The available host plant data suggest that the flight height of moths was determined by larval resource availability. Examples include understorey flyers such as the geometrid genus Eois feeding on Piper, and canopy flyers such as the arctiid moth genera Aclytia, Macrocneme and Poliopastea which feed on lianas.     

Structure and dynamics of a tropical dry forest in Ghana

Forest under low rainfall (averaging 745 mm yr^-1) on the Shai Hills in S.E. Ghana has redeveloped following cessation of farming in the 1890s. Forest stature is low, with a canopy at about 11 m, principally of three species, Diospyros abyssinica, D. mespiliformis and Millettia thonningii. Drypetes parvifolia and Vepris heterophylla are common understorey trees. Twelve species of woody liane were recorded. Species of thicket vegetation in the area were also present at low density. Most species are evergreen.Tree mortality averaged 2.3% yr^-1 and exceeded recruitment (1.5% yr^-1). Differences between species in mortality and recruitment were pronounced: canopy species showed a small decline in density; understorey species increased markedly and the thicket species declined. Seed production was very variable, but seedling establishment was very poor for all species. Seedling mortality was high (11% yr^-1) especially for small seedlings. These population trends probably represent the latter stages of succession of forest regrowth after farming about 100 years ago.Compared with tropical rain forest, Shai Hills forest has similar relative tree diameter growth (1–3.5% yr^-1), mortality and recruitment rates, and small-litter fall (5.52 t ha^-1 yr^-1).Shai Hills forest differs from rain forest by its short stature, relatively few (evergreen) tree species, poor regeneration from seed, high soil nutrient status and low rainfall. Similar forests have been reported in east Africa and in parts of New Guinea.Abbreviations dbh diameter at breast height (1.3 m) - gbh girth at breast height died May 1984     

Influence of pasture understoreys and tree management on soil moisture under a young New Zealand stand of Pinus radiata

Soil moisture content (SMC) and throughfall were monitored along transects between rows of radiata pine (Pinus radiata D. Don) planted at 7 m row spacing and grown with ryegrass/clover or lucerne pastures or with no understorey. SMC was recorded to 1 or 2 m depths for years 2–6 or 4–6, respectively. The predominant rainfall from the south–south–west (SSW) direction created rain shadows (60% of open pasture) on the northern sunny side of the trees. The rain shadows increased as the trees grew but their position were modified by stand pruning and thinning in years 4 and 5. SMC in the top meter showed strong positional effects with the lowest SMC under the tree crowns and on the northern side. There were strong seasonal patterns driven by high spring and summer evapotranspiration, modified by rain shadows and stand age. For the no-understorey treatment, SMC remained high midway between tree rows until the trees were 4 years old and started utilizing this moisture. For the two pasture understoreys, the difference in SMC between the north and south sides decreased with time after trees were thinned and pruned. In the no-understorey there was little difference between the two aspects. Understorey evapotranspiration and fine rooting patterns also helped explain soil moisture use patterns. No difference was detected between two contrasting radiata pine tree types. The effective rooting depth of the ryegrass/clover understorey was <0.7 m while that of lucerne and trees were deeper. Alternative tree planting practices may reduce some of the negative effects of tree-understory moisture competition.     

Ecological impacts of desert plantation forests on biodiversity

This investigation involves the comparison of the diversity of understorey vegetation of four desert planted forests with the adjacent desert areas. Spatial and temporal variations in species composition and structure were compared, and alpha and beta diversities were compared for the field data collected from sampled sites. The diversity of native desert species decreased from 66% in desert areas to 44% of the total recorded plants inside the forests. Meanwhile, the percentage of agricultural weed species increased in forests to >twofold more than that recorded in desert areas. Plant communities in forest stands shared <50% of their species with adjacent vegetation in desert stands. The percentage of variation in species composition was >340% in some forests compared with the desert areas. Alpha diversity and β‐diversity were significantly higher in forest stands than in the desert. Spatial and temporal variations in species diversity were correlated with type of tree canopy and irrigation system. Planted forests had significant negative effects on the diversity of native desert shrubs and trees. Using flood irrigation and more spacing between trees might help in conserving the floristic diversity of desert shrubs and trees at the forest floor.     

Vertical distribution of epiphytic bryophytes in an Indonesian rainforest

We studied species richness, composition and vertical distribution of epiphytic bryophytes in submontane rainforest of Central Sulawesi. Bryophytes were sampled on eight canopy trees and on eight trees in the forest understorey. Microclimate was measured at trunk bases and at crown bases. The total recorded number of 146 epiphytic bryophyte species is among the highest ever reported for tropical forests and underlines the importance of the Malesian region as a global biodiversity hotspot. Species composition differed significantly between understorey trees and canopy tree trunks on the one hand, and the forest canopy on the other. Fourty-five percent of the bryophyte species were restricted to canopy tree crowns, 12% to the understorey. Dendroid and fan-like species mainly occurred in the forest understorey whereas tufts were most species rich in the tree crowns. The findings reflect the different microclimatic regimes and substrates found in the understorey and in the forest canopy. The results indicate that assessments of the bryophyte diversity of tropical forests are inadequate when understorey trees and tree crowns are excluded.     

Characteristics of soil fauna community in the Dongjiao coconut plantation ecosystem in Hainan, China

From July 2002 to July 2004, we investigated the soil fauna in the Dongjiao coconut plantation of Hainan Island. A total of 5,378 specimens were obtained. These species represented 4 phyla, 12 classes and 27 genera. The soil animal clusters are as follows: All major animal groups presented in a tropical rain forest are present in Dongjiao coconut plantation's soil community. The Dongjiao coconut plantation demonstrated the typical characteristics of a tropical soil animal community. The number of species and the diversity index (H) increase from the high latitude areas toward the equator; The dominance index (C) decreased from the high latitude areas toward the equator; the ratio of Acarina/Collembola and the percentage occurrence of increase along the declination of latitude; the occurrence of termites, a typical member of tropical community, varies from absent to present and a dramatic increase along the declination of latitude. Compared to primary tropical rainforests, the Dongjian coconut plantation community is relatively low in species diversity, and has a high dominance index and low diversity index. This may partially due to some characters of the plantation: singular tree species, monsoon climate, seashore location, high pH and salinity of the soil, soil moisture and other environmental factors. A seasonal change occurs in the community structure but is not obvious. Soil around human residences has a higher organic material content, and has higher counts of specimens and numbers of species, compared to other three sampling sites. Coconut production at locations around human residences is higher than at any other microhabitats. The high production is positively correlated with the richness of animal community in the soil.     

Characteristics of soil fauna community in the Dongjiao coconut plantation ecosystem in Hainan, China

From July 2002 to July 2004, we investigated the soil fauna in the Dongjiao coconut plantation of Hainan Island. A total of 5,378 specimens were obtained. These species represented 4 phyla, 12 classes and 27 genera. The soil animal clusters are as follows: All major animal groups presented in a tropical rain forest are present in Dongjiao coconut plantation's soil community. The Dongjiao coconut plantation demonstrated the typical characteristics of a tropical soil animal community. The number of species and the diversity index (H) increase from the high latitude areas toward the equator; The dominance index (C) decreased from the high latitude areas toward the equator; the ratio of Acarina/Collembola and the percentage occurrence of increase along the declination of latitude; the occurrence of termites, a typical member of tropical community, varies from absent to present and a dramatic increase along the declination of latitude. Compared to primary tropical rainforests, the Dongjian coconut plantation community is relatively low in species diversity, and has a high dominance index and low diversity index. This may partially due to some characters of the plantation: singular tree species, monsoon climate, seashore location, high pH and salinity of the soil, soil moisture and other environmental factors. A seasonal change occurs in the community structure but is not obvious. Soil around human residences has a higher organic material content, and has higher counts of specimens and numbers of species, compared to other three sampling sites. Coconut production at locations around human residences is higher than at any other microhabitats. The high production is positively correlated with the richness of animal community in the soil.     

Effects of rain, nitrogen, fire and grazing on tree recruitment and early survival in bush-encroached savanna, South Africa

Moisture, nutrients, fire and herbivory are the principal factors governing tree–grass cover ratios of savannas. We investigated tree (Acacia mellifera) recruitment after fire and under conditions of maximum-recorded rainfall, nitrogen addition and grazing in a completely-crossed field experiment. We employed a similar garden experiment with the exception of the fire treatment. Tree germination in the field was extremely low, probably due to below-average natural rainfall in plots that only received natural rain, and insufficient watering frequency in irrigated plots. Due to low germination in the field experiment, no treatment significantly affected tree recruitment. In the garden experiment, frequent watering, nutrient control (i.e. no nitrogen addition) and grazing enhanced tree recruitment with significant interactions between rain, nitrogen and grazing. We infer that above-average rainfall years with frequent rainfall events are required for mass tree recruitment. Grass defoliation makes space and resources available for tree seedlings. Nitrogen enrichment increases the competitive ability of fast-growing grasses more than that of the N₂-fixing tree component. In contrast to conventional wisdom that grazing alone causes encroachment, we suggest that there are complex interactions between the above-mentioned factors and ‘triggering’ events such as unusually high rainfall.