The role of dispersal ability, climate and spatial separation in shaping biogeographical patterns of phylogenetically distant plant groups in seasonally dry Andean forests of Bolivia

Aim To assess biogeographical patterns of Acanthaceae, Bromeliaceae, Cactaceae and Pteridophyta in Bolivian Andean seasonally dry forest islands and to explain current floristic differences between these islands by means of extrinsic (precipitation, elevation) and intrinsic (dispersal ability) factors. Location Ten isolated and disjunct seasonally dry forest areas in the Bolivian Andes and the adjacent seasonal forest areas of the Chiquitanía and Chaco regions. Methods We collated species data from recently updated and revised taxonomic treatments and herbarium collections for Acanthaceae, Bromeliaceae, Cactaceae and Pteridophyta, constructed floristic distance matrices to estimate beta diversity at the study sites and subjected them to Mantel correlation analyses. Multiple regressions on distance matrices allowed us to test the influence of geographical distance and environmental (elevation and precipitation) differences on floristic differentiation. Results Acanthaceae and Bromeliaceae, and to a lesser extent Cactaceae, showed coincident biogeographical patterns and suggested the presence of two seasonally dry forest groups in Andean Bolivia: one including all small isolated northern dry valleys and another including all southern valleys with connections to the lowland seasonal forests of the Chiquitanía and the Chaco. Most of the variation in the floristic distance matrices in these plant groups, with seed dispersal typically restricted to short distances, was explained by spatial separation between habitat islands. In contrast, pteridophytes showed a different biogeographical pattern. Their floristic differences between sites were determined by the environmental variables. The anemochorous and spore‐based dispersal system of this plant group seems to be a highly effective mechanism allowing pteridophytes to easily reach even the isolated dry valleys in inter‐Andean Bolivia. Main conclusions Current biogeographical patterns in dry Andean habitat islands can provide insights into the factors that control the processes of community assembly. We show that differences in community composition of phylogenetically distant plant groups in the understorey of seasonally dry forest islands can be explained by a combination of the habitat characteristics where the group is present (either precipitation, elevation or both) and, more interestingly, by group specific dispersal limitation (as inferred by geographical distance between island habitats).     

Gradients within gradients: The mesoscale distribution patterns of palms in a central Amazonian forest

Questions: What are the relative contributions of environmental factors and geographic distance to palm community structure at the mesoscale, and how do they depend on the length of the environmental gradient covered? How do soil and topography affect variation of the canopy and understory palm community structure at the mesoscale? How does fine‐scale variation within the broad edaphic/topographic classes affect palm community composition? Location: Reserva Ducke, terra‐firme forest, Manaus, Brazil. Methods: Palms were sampled in 72 plots 250 m × 4 m, systematically distributed over an area of 100 km². Soil, topography and distance to watercourses were measured for all plots. The relationship between community structure axes, summarized by NMDS ordinations, and environmental predictors, was analysed with multivariate regressions. Matrix regressions were used to determine the proportions of variance explained by environmental and geographic predictors. Results: Floristic variation at the mesoscale was mostly related to environmental variation, and the proportion of variance explained depended on the amplitude of the environmental gradient. Soil was the main predictor of floristic change, but its effects differed between life forms, with the understory palm community structured within one of the edaphic/topographic classes, in association with distance to watercourses. Conclusions: Dispersal limitation does not explain palm composition at the mesoscale, and the amplitude of environmental gradients covered by the analysis can be as important as the scale of analysis, in determining the relative contributions of environmental and geographical components to community structure. Soil and topography can predict a large proportion of palm composition, but gradients differ in scale, with some environmental gradients being nested within others. Therefore, although all environmental gradients are nested within distance, they do not necessarily coincide.     

New light on the floristic composition and diversity of indigenous territory and national park Isiboro-Sécure, Bolivia

A floristic inventory of woody plants was conducted in the tropical forest vegetation of Indigenous Territory and National Park Isiboro-Sécure, Bolivia. All stems with a diameter at breast height ≥2.5 cm were sampled in two terra firme and two floodplain 0.1 ha forest transects, resulting in a total of 447 species and 1,564 individuals. Results correspond fairly well with other Bolivian and neotropical studies in terms of species distribution patterns, physiognomy, family distribution, growth forms, as well as species and family richness. Our finding that only 6% of all species occurred in all four transects, while 69% was found in only one transect, supports the hypothesis that Amazonian plant communities are dominated by a limited set of species, whereas the large majority of species follows a patchy distribution. However, Fisher’s Alpha index values were much higher than was expected from literature data. It is hypothesized that this might be related to high local precipitation levels and/or natural and (historical) anthropogenic disturbance regimes. Although much more research is necessary, our results suggest that TIPNIS represents a strongly underestimated local center of plant diversity.     

Influence of clay licks on the diversity and structure of an Amazonian forest

The spatial heterogeneity of resource availability is a major driver of biodiversity patterns. Some environmental conditions and resources are characterized by large‐scale patterns of variation within the landscape. Clumped local discontinuities or discrete elements also increase spatial heterogeneity, promoting local ‘biodiversity hot spots’ by modifying habitat characteristics and promoting plant–animal interactions. Clay licks are faunal attractors owing to their role in the nutritional ecology of the user species; nevertheless, the effect of their presence on the surrounding vegetation has been poorly quantified. Here, we use data from 100 × 10 m transects and evaluate the effects of the presence of clay licks on forest diversity and structure at local and landscape scales. In clay lick areas, there was a higher abundance of certain species, which helps to homogenize species composition between localities counteracting the natural distance‐decay of compositional similarity between transects without clay lick influence (controls). Compared to control sites, clay lick′s forests had higher palm densities, shorter but more variable individuals in the canopy and understory, a thinner canopy layer, and denser herbaceous and ground level covers. These differences were found along the whole length of transects in both sampled areas types. These results reveal that the presence of discrete elements (i.e., clay licks) may help to explain the compositional and structural heterogeneity of Amazonian forests influencing ecological processes such as seed dispersal and trampling. These considerations may be relevant for other biomes where clay licks are present and give weight to their inclusion in conservation initiatives in tropical forests.     

Contrasting effects of mosaic structure on alpha and beta diversity of bird assemblages in a human‐modified landscape

Habitat loss and fragmentation are key processes causing biodiversity loss in human‐modified landscapes. Knowledge of these processes has largely been derived from measuring biodiversity at the scale of ‘within‐habitat’ fragments with the surrounding landscape considered as matrix. Yet, the loss of variation in species assemblages ‘among’ habitat fragments (landscape‐scale) may be as important a driver of biodiversity loss as the loss of diversity ‘within’ habitat fragments (local‐scale). We tested the hypothesis that heterogeneity in vegetation cover is important for maintaining alpha and beta diversity in human‐modified landscapes. We surveyed bird assemblages in eighty 300‐m‐long transects nested within twenty 1‐km² vegetation ‘mosaics’, with mosaics assigned to four categories defined by the cover extent and configuration of native eucalypt forest and exotic pine plantation. We examined bird assemblages at two spatial scales: 1) within and among transects, and 2) within and among mosaics. Alpha diversity was the mean species diversity within‐transects or within‐mosaics and beta diversity quantified the effective number of compositionally distinct transects or mosaics. We found that within‐transect alpha diversity was highest in vegetation mosaics defined by continuous eucalypt forest, lowest in mosaics of continuous pine plantation, and at intermediate levels in mosaics containing eucalypt patches in a pine matrix. We found that eucalypt mosaics had lower beta diversity than other mosaic types when ignoring relative abundances, but had similar or higher beta diversity when weighting with species abundances. Mosaics containing both pine and eucalypt forest differed in their bird compositional variation among transects, despite sharing a similar suite of species. This configuration effect at the mosaic scale reflected differences in vegetation composition among transects. Maintaining heterogeneity in vegetation cover could help to maintain variation among bird assemblages across landscapes, thus partially offsetting local‐scale diversity losses due to fragmentation. Critical to this is the retention of remnant native vegetation.     

Tree species composition and diversity gradients in white-water forests across the Amazon Basin

Aim Attention has increasingly been focused on the floristic variation within forests of the Amazon Basin. Variations in species composition and diversity are poorly understood, especially in Amazonian floodplain forests. We investigated tree species composition, richness and α diversity in the Amazonian white‐water (várzea) forest, looking particularly at: (1) the flood‐level gradient, (2) the successional stage (stand age), and (3) the geographical location of the forests. Location Eastern Amazonia, central Amazonia, equatorial western Amazonia and the southern part of western Amazonia. Methods The data originate from 16 permanent várzea forest plots in the central and western Brazilian Amazon and in the northern Bolivian Amazon. In addition, revised species lists of 28 várzea forest inventories from across the Amazon Basin were used. Most important families and species were determined using importance values. Floristic similarity between plots was calculated to detect similarity variations between forest types and over geographical distances. To check for spatial diversity gradients, α diversity (Fisher) of the plots was correlated with stand age, longitudinal and latitudinal plot location, and flood‐level gradient. Results More than 900 flood‐tolerant tree species were recorded, which indicates that Amazonian várzea forests are the most species‐rich floodplain forests worldwide. The most important plant families recorded also dominate most Neotropical upland forests, and c. 31% of the tree species listed also occur in the uplands. Species distribution and diversity varied: (1) on the flood‐level gradient, with a distinct separation between low‐várzea forests and high‐várzea forests, (2) in relation to natural forest succession, with species‐poor forests in early stages of succession and species‐rich forests in later stages, and (3) as a function of geographical distance between sites, indicating an increasing α diversity from eastern to western Amazonia, and simultaneously from the southern part of western Amazonia to equatorial western Amazonia. Main conclusions The east‐to‐west gradient of increasing species diversity in várzea forests reflects the diversity patterns also described for Amazonian terra firme. Despite the fine‐scale geomorphological heterogeneity of the floodplains, and despite high disturbance of the different forest types by sedimentation and erosion, várzea forests are dominated by a high proportion of generalistic, widely distributed tree species. In contrast to high‐várzea forests, where floristic dissimilarity increases significantly with increasing distance between the sites, low‐várzea forests can exhibit high floristic similarity over large geographical distances. The high várzea may be an important transitional zone for lateral immigration of terra firme species to the floodplains, thus contributing to comparatively high species richness. However, long‐distance dispersal of many low‐várzea trees contributes to comparatively low species richness in highly flooded low várzea.     

Diversity partitioning of moorland plant communities across hierarchical spatial scales

Understanding of the scaling of diversity is critical to enhance conservation strategies for subalpine moorland ecosystems vulnerable to future environmental changes. However, a paucity of quantitative data strongly limits such attempts. In this study, we used an additive diversity partitioning framework and quantified diversity patterns of moorland plant communities across hierarchical spatial scales, within- and between-sample transects, and between sites (corresponding to α and two levels of β diversity). Moorland sites markedly differed in size (range 1,000–160,000 m²) and were isolated from each other to varying extents within an inhospitable matrix (i.e., forests). We found that β diversity components were consistently higher, whereas the local α diversity component was consistently lower than expected by chance. We observed substantial contribution at the between-site scale to total species richness. By focusing on diversity patterns of moorland plant communities across multiple hierarchical spatial scales, we could thus identify the scale at which regional diversity is maximized. Our results suggest that protection of as many moorland sites as possible, to ensure beta diversity between sites, will effectively conserve total diversity. The use of additive diversity partitioning is a major step forward in providing strategies for the biological conservation of subalpine moorland ecosystems vulnerable to future environmental changes.     

Landscape-Scale Environmental and Floristic Variation in Costa Rican Old-Growth Rain Forest Remnants

Studies of tropical rain forest beta-diversity debate environmental determinism versus dispersal limitation as principal mechanisms underlying floristic variation. We examined the relationship between soil characteristics, terrain, climate variation, and rain forest composition across a 3000 km² area in northeastern Costa Rica. Canopy tree and arboreal palm species abundance and soils were measured from 127 0.25-ha plots across Caribbean lowlands and foothills. Plot elevation, slope, temperature, and precipitation variation were taken from digital grids. Ordination of forest data yielded three floristic groups with strong affinities to foothills and differing lowland environments. Variation in floristics, soil texture, and climate conditions showed parallel patterns of significantly positive spatial autocorrelation up to 13 km and significantly negative correlation beyond 40 km. Partial Mantel tests resulted in a significant correlation between floristic distance and terrain, climate and soil textural variables controlling the effect of geographical distance. Separate comparisons for palm species showed significant correlation with Mg and Ca concentrations among other soil factors. Arboreal palm species demonstrated a stronger relationship with soil factors than did canopy trees. Correlation between floristic data and geographical distance, related to seed dispersal or unmeasured variables, was not significant after controlling for soil characteristics and elevation. Canopy trees and palms showed differing relationships to soil and other environmental factors, but lend greater support for a niche-assembly hypothesis than to a major role for dispersal limitation in determining species turnover for this landscape.     

Geographical patterns in the beta diversity of China's woody plants: the influence of space,environment and range size

Beta diversity (i.e. species turnover rate across space) is fundamental for understanding mechanisms controlling large‐scale species richness patterns. However, the influences on beta diversity are still a matter of debate. In particular, the relative role of environmental and spatial processes (e.g. environmental niche versus dispersal limitation of species) remains elusive, and the influence of species range size has been poorly tested. Here, using distribution maps of 11 405 woody species in China (ca 9.6 × 10⁶ km²), we investigated 1) the geographical and directional patterns of beta diversity for all woody species and species with different range sizes, and 2) compared the effects of environmental and spatial processes on these patterns. Beta diversity was calculated as the decay of similarity in species composition with increasing distance. Variables representing environmental energy, water availability, climatic seasonality, habitat heterogeneity and human activities were used to evaluate the effects of environmental processes, while spatial distance was used to assess the influence of spatial processes. The results indicated significant directional patterns of beta diversity: the similarity decay along the latitudinal gradient was 1.6–2.3 times faster than that along the longitudinal gradient. Beta diversity also increased with the decrease of species range size. As compared with spatial processes, environmental processes had stronger effects on longitudinal beta diversity and on the beta diversity of widely‐ranged species. This was opposite to the larger influence of spatial processes on latitudinal beta diversity and the beta diversity of narrowly‐ranged species. These results suggest that the distributions of narrowly‐ranged woody species in China may have not reached equilibrium with their environmental niches due to dispersal limitation induced by China's topography and/or their low dispersal ability. The projected rapid climatic changes will likely endanger such species. Species dispersal processes should be taken into account in future conservation strategies in China.     

Tree species composition and environmental relationships in a Neotropical swamp forest in Southeastern Brazil

We examined the relationships between topography, soil properties and tree species composition in a Neotropical swamp forest in southeastern Brazil. Plots were sampled in the forest, encompassing three different soil ground water regimes along the topographical declivity. All non-climbing plant individuals with trunk height >1.3 m were sampled. A canonical correspondence analysis—CCA—of the species–environmental relationships grouped tree species according to drainage and chemical soil conditions. A total of 86 species were found, being 77 species in the inferior, 40 species in the intermediate and 35 species in the superior topographic section. Some species were among the 10 most abundant ones, both in the overall sampled area and in each topographical section, with alternation events occurring only with their abundance position. However, substantial differences in floristic composition between sections were detected in a fine spatial scale, due to higher number of species, diversity index (H′) and species unique (exclusives) in the inferior topographic section. These higher values can be attributed to its higher spatial heterogeneity that included better drained and seasonally waterlogged soils, higher soil fertility and lower acidity. The increase of the soil water saturation and the uniform conditions derived from the superficial water layer has led to a lower number of species and an increase on the palm trees abundance in the intermediate and superior sections. Our results showed that at a small spatial scale niche differentiation must be an important factor related to the increase of the local diversity. The wide distribution of the most abundant species in the studied area and the increase of local diversity corroborate the pattern of distribution of species in larger scales of swamp forests, in which the most abundant species repeat themselves in high densities in different remnants. However, the floristic composition of each remnant is strongly variable, contributing to the increase of regional diversity.     

Relationships between tree component structure,topography and soils of a riverside forest,Rio Botucaraí, Southern Brazil

The relationships among floristic, structural and physiognomic variables of the tree component, flooding regime variations and soil fertility were investigated in a riparian forest fragment (ca. 43 ha) in the Rio Botucaraí watershed near the confluence with the Rio Jacuí, southern Brazil. All the trees with a dbh ≥ 15 cm were surveyed in 100 contiguous 10 × 10 m plots and soil chemical and textural variables were obtained from the analyses of 15 topsoil samples (0–20 cm depth) collected at different locations. The sample sites encompass all the topographic variation of the area. We used canonical correspondence analysis (CCA) to seek correlations between environmental variables and tree component distribution. The 1,547 surveyed individuals belonged to 30 species and 16 botanical families. The Shannon diversity index (H′) and the Pielou equability (J′) were 1.995 and 0.586 nats ind.⁻¹, respectively. Sebastiania commersoniana, Casearia sylvestris, Eugenia uniflora and Eugenia hyemalis presented the highest importance values. Species’ densities chiefly correlated with site elevation, flooding regime and soil chemical fertility. The analyses of ecological categories (EC) revealed that most species are heliophylous and typical of early successional stages. Richness and diversity (P < 0.001) variations from different topographic positions reflect that at the local scale, flooding was a limiting factor for the establishment of certain species suggesting that local processes control species richness and diversity.     

The Role of Urban Structures in the Distribution of Wasteland Flora in the Greater Paris Area,France

Wastelands are likely to host a significant part of urban floristic diversity but have received limited attention because they are not considered interesting green zones. Here, we explore the potential role of wastelands in maintaining urban biodiversity to help define effective urban management plans. We quantified floristic diversity in 98 wasteland sites of Hauts-de-Seine, one of the most densely populated areas in France, and characterized the environmental parameters and spatial distribution of sites to identify some of the factors that influence plant species composition and to explore the impact of urban environment on the floristic interest of wastelands. Their floristic richness represented 58% of the total richness observed in the whole study area. Site richness depended on site area (the largest sites were the richest) and site age, with a maximum in sites of intermediate age (4–13 years). In the largest sites only (>2,500 m²), the floristic distance among sites was positively correlated with geographic distance, which suggests that migration of species among large sites partly controls local floristic composition. In contrast, the environmental distance among sites was not correlated with floristic distance. Finally, we showed that the presence of collective and individual dwellings within 200 m of a wasteland decreased its floristic rarity, whereas the presence of rivers or ponds increased it. We derive several recommendations to optimize the management of wastelands with respect to conservation of urban biodiversity.     

Floristic composition across a climatic gradient in a neotropical lowland forest

Abstract. This study deals with the floristic composition of lowland tropical forest in the watershed of the Panama Canal. The floristic composition of large trees in 54 forest plots was analysed with respect to environmental factors, including precipitation, geologic parent material, stand age, topography, and soils. The plots contain 824 species of trees with a diameter at breast height ≥10 cm and represent a regional flora with exceptional β‐diversity. Plot data indicate that the Panamanian forest is strongly spatially structured at the landscape scale with floristic similarity decreasing rapidly as a function of inter‐plot geographic distance, especially for distances <5 km. The ordinations and patterns of endemism across the study area indicate broad floristic associations well correlated with Holdridge life zones. The results indicate the positive aspects of life zone classification at regional scales, while simultaneously highlighting its inadequacy for finer scales of analysis and resource management. Multivariate gradient analysis techniques (Non‐metric Multidimensional Distance Scaling and Detrended Correspondence Analysis) show clear patterns of floristic variability correlated with regional precipitation trends, surficial geology, and local soil attributes. Geologic and edaphic conditions, such as acidic soils or excessively drained limestone substrates, appear to override the effects of precipitation and modify forest composition. We conclude that the Panamanian forest shows clear patterns of spatial organization along environmental gradients, predominantly precipitation. The rapid decline in floristic similarity with distance between stands also suggests a role for dispersal limitation and stochastic events.     

A floristic analysis of the lowland dipterocarp forests of Borneo

Aim To (1) identify floristic regions in the lowland (below 500 m a.s.l.) tropical dipterocarp rain forest of Borneo based on tree genera, (2) determine the characteristic taxa of these regions, (3) study tree diversity patterns within Borneo, and (4) relate the floristic and diversity patterns to abiotic factors such as mean annual rainfall and geographical distance between plots. Location Lowland tropical dipterocarp rain forest of Borneo. Methods We used tree (diameter at breast height ≥ 9.8 cm) inventory data from 28 lowland dipterocarp rain forest locations throughout Borneo. From each location six samples of 640 individuals were drawn randomly. With these data we calculated a Sørensen and Steinhaus similarity matrix for the locations. These matrices were then used in an UPGMA clustering algorithm to determine the floristic relations between the locations (dendrogram). Principal coordinate analysis was used to ordinate the locations. Characteristic taxa for the identified floristic clusters were determined with the use of the INDVAL method of Dufrene & Legendre (1997) . Finally, Mantel analysis was applied to determine the influence of mean annual rainfall and geographical distance between plots on floristic composition. Results A total of 77 families and 363 genera were included in the analysis. On average a random sample of 640 trees from a lowland dipterocarp forest in Borneo contains 41.6 ± 3.8 families and 103.0 ± 12.7 genera. Diversity varied strongly on local scales. On a regional scale, diversity was found to be highest in south‐east Borneo and central Sarawak. The most common families were Dipterocarpaceae (21.9% of trees) and Euphorbiaceae (12.2% of trees). The most common genera were Shorea (12.3% of trees) and Syzygium (5.0% of trees). The 28 locations were clustered in geographically distinct floristic regions. This was related to the fact that floristic similarity depended strongly on the geographical distance between plots and similarity in mean annual rainfall. Conclusions We identified five main floristic regions within the lowland dipterocarp rain forests of Borneo, each of which had its own set of characteristic genera. Mean annual rainfall is an important factor in explaining differences in floristic composition between locations. The influence of geographical distance on floristic similarity between locations is probably related to the fact that abiotic factors change with distance between plots. Borneo's central mountain range generally forms an effective dispersal barrier for the lowland tree flora. Diversity patterns in Borneo are influenced by the mid‐domain effect, habitat size and the influence of past climatic changes (ice ages during the Pleistocene).     

Environment versus dispersal in the assembly of western Amazonian palm communities

Aim It is a central issue in ecology and biogeography to understand what governs community assembly and the maintenance of biodiversity in tropical rain forest ecosystems. A key question is the relative importance of environmental species sorting (niche assembly) and dispersal limitation (dispersal assembly), which we investigate using a large dataset from diverse palm communities. Location Lowland rain forest, western Amazon River Basin, Peru. Methods We inventoried palm communities, registering all palm individuals and recording environmental conditions in 149 transects of 5 m × 500 m. We used ordination, Mantel tests and indicator species analysis (ISA) to assess compositional patterns, species responses to geographical location and environmental factors. Mantel tests were used to assess the relative importance of geographical distance (as a proxy for dispersal limitation) and environmental differences as possible drivers of dissimilarity in palm species composition. We repeated the Mantel tests for subsets of species that differ in traits of likely importance for habitat specialization and dispersal (height and range size). Results We found a strong relationship between compositional dissimilarity and environmental distance and a weaker but also significant relationship between compositional dissimilarity and geographical distance. Consistent with expectations, relationships with environmental and geographical distance were stronger for understorey species than for canopy species. Geographical distance had a higher correlation with compositional dissimilarity for small‐ranged species compared with large‐ranged species, whereas the opposite was true for environmental distance. The main environmental correlates were inundation and soil nutrient levels. Main conclusions The assembly of palm communities in the western Amazon appears to be driven primarily by species sorting according to hydrology and soil, but with dispersal limitation also playing an important role. The importance of environmental characteristics and geographical distance varies depending on plant height and geographical range size in agreement with functional predictions, increasing our confidence in the inferred assembly mechanisms.     

Mapping landscape beta diversity of plants across KwaZulu-Natal,South Africa,for aiding conservation planning

Collective properties of biodiversity, such as beta diversity, are suggested as complementary measures of species richness to guide the prioritisation and selection of important biodiversity areas in regional conservation planning. We assessed variation in the rate of plant species turnover along and between environmental gradients in KwaZulu-Natal, South Africa using generalised dissimilarity modelling, in order to map landscape levels of floristic beta diversity. Our dataset consisted of 434 plots (1000 m²) containing 997 grassland and savanna matrix species. Our model explained 79 % of the null deviance observed in floristic dissimilarities. Variable rates of turnover existed along the major environmental gradients of mean annual temperature, median rainfall in February, and soil cation exchange capacity, as well as along gradients of geographical distance. Beta diversity was highest in relatively warm, drier summer regions and on dystrophic soils. Areas of high beta diversity identify areas that should be included in conservation plans to maximise representation of diversity and highlight areas best suited to protected area expansion. Biome transition areas in high beta diversity areas may be susceptible to climate variability. Including beta diversity turnover rates in regional conservation plans will help to preserve evolutionary and ecological processes that create and maintain diversity.     

Biodiversity assessment in incomplete inventories: leaf litter ant communities in several types of Bornean rain forest

Biodiversity assessment of tropical taxa is hampered by their tremendous richness, which leads to large numbers of singletons and incomplete inventories in survey studies. Species estimators can be used for assessment of alpha diversity, but calculation of beta diversity is hampered by pseudo-turnover of species in undersampled plots. To assess the impact of unseen species, we investigated different methods, including an unbiased estimator of Shannon beta diversity that was compared to biased calculations. We studied alpha and beta diversity of a diverse ground ant assemblage from the Southeast Asian island of Borneo in different types of tropical forest: diperocarp forest, alluvial forest, limestone forest and heath forests. Forests varied in plant composition, geology, flooding regimes and other environmental parameters. We tested whether forest types differed in species composition and if species turnover was a function of the distance between plots at different spatial scales. As pseudo-turnover may bias beta diversity we hypothesized a large effect of unseen species reducing beta diversity. We sampled 206 ant species (25% singletons) from ten subfamilies and 55 genera. Diversity partitioning among the four forest types revealed that whereas alpha species richness and alpha Shannon diversity were significantly smaller than expected, beta-diversity for both measurements was significantly higher than expected by chance. This result was confirmed when we used the unbiased estimation of Shannon diversity: while alpha diversity was much higher, beta diversity differed only slightly from biased calculations. Beta diversity as measured with the Chao-Sørensen or Morisita-Horn Index correlated with distance between transects and between sample points, indicating a distance decay of similarity between communities. We conclude that habitat heterogeneity has a high influence on ant diversity and species turnover in tropical sites and that unseen species may have only little impact on calculation of Shannon beta diversity when sampling effort has been high.     

Abundance and Production of Riparian Trees in the Lowland Floodplain of the Queets River, Washington

Riparian zones associated with alluvial rivers are spatially dynamic, forming distinct vegetative mosaics that exhibit sharp contrasts in structure and processes related to the underlying biophysical template. The productivity of riparian plants, especially trees, influences streamside community characteristics as well as the forms and fluxes of organic matter to adjacent streams – thereby strongly impacting patterns of channel morphology, water flow, sedimentation, and habitat in rivers. As part of a comprehensive investigation of riparian dynamics in coastal rain forest rivers of the Pacific Northwest (USA), we examined riparian tree abundance (density, basal area, and biomass) and rates of production (basal area growth [BAI] and bole wood biomass increase [P]) of seven common species – red alder (Alnus rubra), Sitka spruce (Picea sitchensis), bigleaf maple (Acer macrophyllum), western hemlock (Tsuga heterophylla), black cottonwood (Populus trichocarpa), vine maple (Acer circinatum) and willow (Salix spp.) – in the lowland floodplain of the Queets River (Olympic National Park), Washington. Measurements were made annually for three years (1999 – 2001) in 16 permanent plots on three biophysical templates that formed a toposequence – active floodplain, young terrace and mature terrace. Stem density was highest in the active floodplain (∼27,000 stems/ ha), decreasing in the young terrace (∼2,700 stems /ha) and the mature terrace (∼500 stems/ha). Basal area and total stem biomass were lowest in the active floodplain (∼16 m2/ha and ∼18 Mg dry weight/ha, respectively) and higher on the young terrace (∼32 m2/ha and ∼134 Mg dry weight/ha) and on the mature terrace (∼69 m2/ha and ∼540 Mg dry weight /ha). Total plot-scale BAI was not significantly different among the physical templates with mean values ranging from approximately 1.4 (low terrace) to approximately 2.8 m2/ha/y (active floodplain). In contrast, P was significantly higher on the mature terrace (10.3 Mg/ha) than the active floodplain (3.2 Mg/ha) but there was no significant difference between young terrace (6.5 Mg/ha) and mature terrace. For the entire Queets River floodplain (57 km² over 77 km of river length), the mature terrace contributed 81% of the total annual production (28,764 Mg) whereas the active floodplain and young terrace accounted only for 5 and 14%, respectively. Overall, we show that riparian trees grow quickly in this coastal Pacific Northwest system and that the older riparian forests on mature terraces are the main contributors to stem production at the plot and floodplain scales for at least 350 years after stand initiation. This suggests that, in combination with the rapid lateral migrations of many alluvial rivers, the older riparian forests on those terraces are important and sustained sources of organic matter (especially large woody debris, LWD) that, over decades to centuries, shape the character of coastal rivers in the Pacific Northwest.     

Tree changes in a mature rainforest with high diversity and endemism on the Brazilian coast

The tree changes of 1.02 ha of montane forest at the Santa Lúcia Biological Station, southeastern Brazil, were analyzed using two surveys separated by an interval of 11 years with the aim of confirming the patterns of stability of structure and diversity over time. In the original survey all trees with diameter at breast height ≥6.4 cm were sampled. In second survey (this study), dead trees, survivors and recruits in the same forest were reported. The data suggest a dynamic balance of the forest structure because mortality (−1.06% year⁻¹ for number of trees and −0.85% year⁻¹ for basal area) was very close to recruitment (0.89% year⁻¹) and ingrowth (1.05% year⁻¹). The high diversity of the original survey (H′ > 5.2) was maintained by the turnover species. The main tree populations also showed stability of number of trees and basal area. This pattern was shared by most of the 28 local endemic species, ensuring the maintenance of their populations in the plot.