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.     

Sapling communities in Amazonian white-water forests

Aim Structure and floristic composition of forest regeneration (trees between 1 and 10 cm diameter at breast height (d.b.h), ≥ 1 m growth height) was described in three forest types of Amazonian white‐water forests (várzea), in order to analyse whether floristic composition of saplings is related to the successional stage of the forests, whether it differs in comparision with the mature flora, and if there exists a zonation of sapling species along the gradients of flooding and irradiation. Location Mamirauá Sustainable Development Reserve, Western Brazilian Amazon. Methods The investigated forests were of the low and the high várzea type, on an annual average flooded around 4 and 1 months, respectively. The two low‐várzea forests belonged to the secondary and to the late‐successional stage, the high‐várzea forest also to the late‐successional stage. A total of 24 circular sample plots covering 1885 m² were installed. They were nested within three rectangular 1 ha permanent sample plots where individuals ≥ 10 cm d.b.h. were formerly inventoried. Average inundation and radiation [relative photosynthetically active radiation (rPAR)] at the forest floor was recorded in all inventoried plots. The Mean spatial Distribution Center (MDC; Ebdon, 1998 ) for all sapling species was calculated. Sapling species were grouped into associations with respect to their light‐demand and their location along the flood‐level gradient. Results Average flood height and average rPAR at the forest floor was highest in the secondary stage of the low várzea and lowest in the high‐várzea forest. Overall average density and basal area of the saplings averaged 2250 individuals and 2.3 m² ha^?1 in the secondary stage, 2330 individuals and 2.6 m² ha^?1 in the late‐successional stage of the low várzea and 5000 individuals and 4.8 m² ha^?1 in the high várzea. In all forest types, species richness of saplings was lower than species richness of trees ≥ 10 cm d.b.h., and amounted to 25 species in the secondary stage, to 35 species in the late‐successional stage of the low várzea and to 88 species in the high‐várzea forest. The amount of ‘immigrants’ ( Bazzaz, 1991 ) at the sapling level increased with proceeding forest succession and amounted to 24 and 29% in the low várzea and to 31% in the high várzea. Sapling species distribution, species richness and individual density was linked to both, the gradient of flooding and the gradient of irradiation. Species richness and individual density was highest in the high várzea and decreased with increasing influence of flooding. The high várzea was predominated by pronounced shade‐tolerant sapling species, whereas in the low várzea occured both, light demanding and shade‐tolerant species groups. Main conclusions Influence of flooding seems to be the main factor triggering species composition and structure of sapling communities in várzea forests. Proceeding forest succession reduces the impact of flooding because of the biogenical induced silting up of the forested sites. However, forest succession also alters forest architecture of the overstory and such the light conditions at the sapling level. Therefore, radiation is an important factor influencing species composition of sapling communities in várzea forests.     

Habitat specifity,endemism and the neotropical distribution of Amazonian white‐water floodplain trees

.The Amazon basin is covered by the most species‐rich forests in the world and is considered to house many endemic tree species. Yet, most Amazonian ecosystems lack reliable estimates of their degree of endemism, and causes of tree diversity and endemism are intense matters of debate. We reviewed the spatial distribution of 658 of the most important flood‐tolerant Amazonian white‐water (várzea) tree species across the entire Neotropics by using data from herbaria, floras, inventories and checklists. Our results show that 90% of the várzea tree species are partially or widely distributed across neotropical macro‐regions and biomes. Chi‐square analyses indicated that várzea species richness in non‐várzea macro‐regions was dependent on the flooding gradient and the longitudinal position. Cluster analysis combined with association tests indicated four significant patterns of várzea species distributions depending on species flood‐tolerance (low vs high) and spatial distribution (restricted vs widespread). We predict that the predominance of Andean substrates is the most important factor that determines the distribution of várzea tree species within and beyond the Amazon basin and explains the high floristic similarity to the Orinoco floodplains. Distribution patterns in other extra‐Amazonian macro‐regions are more likely linked to climatic factors, with rainforest climates housing more várzea species than savanna climates. 130 tree species were restricted to South‐American freshwater floodplains, and 68 (> 10%) were endemic to Amazonian várzea. We detected two centers of endemism, one in the western Amazon characterized by low and brief floods, and one in the central Amazon, characterized by high and prolonged floods. Differences in taxonomic composition of endemic centers in the western and central Amazon are the result of different abiotic factors (i.e. flood regimes), as well as the regional species pools from where the species are recruited from. We hypothesize that numerous morphological, physiological and biochemical adaptations permit survival of trees in flooded environments. Furthermore, these adaptations are independently derived across many taxa and result in a highly specialized flora. We attribute higher than expected levels of endemism to the great spatial extent and age of floodplain ecosystems in the Amazon basin, and highlight the role of Amazonian várzea as an potential driver in speciation and diversification processes.     

Landscape patterns of understory composition and richness across a moisture and nitrogen mineralization gradient in Ohio (U.S.A.) Quercus forests

This study quantified relationships of understory vascular plant species composition and richness along environmental gradients over a broad spatial scale in second-growth oak forests in eastern North America. Species frequencies were recorded in 108 25 × 25 m plots in four study sites extending over 70 km in southern Ohio, U.S.A.. The plots were stratified into three long-term soil moisture classes with a GIS-derived integrated moisture index (IMI). In addition to the IMI, the environmental data matrix included eight soil and three overstory variables. Canonical correspondence analysis (CCA) indicated that variations in understory species composition were most strongly related to topographic variations in predicted moisture (IMI), N mineralization rate, nitrification rate, and soil pH. In addition, floristic variation at the regional scale was correlated with variations in soil texture, nitrification, pH, and PO₄ ^–, resulting from differences in the soil parent material complexes among sites. Species richness averaged 65 species/plot, and increased with moisture and fertility. Stepwise regression indicated that richness was positively correlated with N mineralization rate and nitrification rate, and inversely correlated with tree basal area. Greater richness on fertile plots was the largely the result of increasing forb richness. Forb richness per quadrat (2 m²) was most strongly and positively related to N mineralization rate. Conversely, richness of understory individuals of tree species was greatest on xeric, less-fertile plots. Our results describe general, broad-scale species-environment relationships that occurred at both the topographic scale (long-term moisture status and fertility) and the regional scale (geomorphological differences among the sites). Strong species richness-N mineralization correlations indicate an important link between below-ground processes and above-ground biodiversity. Because N availability was a strong correlate to vegetation patterns at a broad-scale, our results suggest that the increasing rates of atmospheric N deposition in the region could have a major impact on understory vegetation dynamics.     

Unexpected High Diversity of Galling Insects in the Amazonian Upper Canopy: The Savanna Out There

A relatively large number of studies reassert the strong relationship between galling insect diversity and extreme hydric and thermal status in some habitats, and an overall pattern of a greater number of galling species in the understory of scleromorphic vegetation. We compared galling insect diversity in the forest canopy and its relationship with tree richness among upland terra firme, várzea, and igapó floodplains in Amazonia, Brazil. The soils of these forest types have highly different hydric and nutritional status. Overall, we examined the upper layer of 1,091 tree crowns. Galling species richness and abundance were higher in terra firme forests compared to várzea and igapó forests. GLM-ANCOVA models revealed that the number of tree species sampled in each forest type was determinant in the gall-forming insect diversity. The ratio between galling insect richness and number of tree species sampled (GIR/TSS ratio) was higher in the terra firme forest and in seasonally flooded igapó, while the várzea presented the lowest GIR/TSS ratio. In this study, we recorded unprecedented values of galling species diversity and abundance per sampling point. The GIR/TSS ratio from várzea was approximately 2.5 times higher than the highest value of this ratio ever reported in the literature. Based on this fact, we ascertained that várzea and igapó floodplain forests (with lower GIA and GIR), together with the speciose terra firme galling community emerge as the gall diversity apex landscape among all biogeographic regions already investigated. Contrary to expectation, our results also support the “harsh environment hypothesis”, and unveil the Amazonian upper canopy as similar to Mediterranean vegetation habitats, hygrothermically stressed environments with leaf temperature at lethal limits and high levels of leaf sclerophylly.     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.     

Overstory-Understory Relationships along Forest Type and Environmental Gradients in the Spring Mountains of Southern Nevada, USA

Isolated forested mountains in deserts have numerous ecological and societal values, but land-management practices (e.g., fire-regime alteration) and climate change can affect forest composition. We analyzed tree overstory-understory relationships on 123 sites in the Spring Mountains within the Mojave Desert near Las Vegas, Nevada, USA to assess three hypotheses. We hypothesized that: the tree species comprising understories are less tolerant of fire than species in overstories, reflecting land-management practices of fire exclusion; mid-elevation forests have the lowest overstory:understory similarity because this zone could have maximum species mixing; and overstory:understory similarity is correlated with environmental gradients (consisting of 14 topographic and soil variables). We found that Pinus monophylla comprised greater relative canopy cover in understories of juniper (32% relative cover) and pinyon-juniper (78%) forests than it did in overstories of these forests (0% and 53%). Similarly, fire-intolerant Abies concolor had 6-fold greater understory than overstory cover in forests with overstories dominated by the fire-tolerant Pinus ponderosa. Overstory:understory S?rensen similarity averaged 43%?77% among six forest types, and there was little support for the supposition that similarity was lowest in mid-elevation forests. Distributions of individual overstory and understory species more closely corresponded with environmental gradients than did overstory:understory similarity. Results suggest that there is high potential for change in at least two of the six dominant forest types of the Spring Mountains. The direction of change (species of moist, higher elevation sites establishing in understories of drier forests) is the opposite of what would be expected for forest adaptation to the warmer, drier, more fire-prone conditions projected for the next century in the southwestern USA.     

Ecological and biogeographical studies on the tropical rain forest of south Yunnan,SW China with a special reference to its relation with rain forests of tropical Asia

Abstract. Ecological and biogeographic analyses of the tropical rain forest in south Yunnan were made using data from seventeen sample plots and floristic inventories of about 1000 species of seed plants. The rain forest is shown to be a type of true tropical rain forest because it has almost the same profile, physiognomic characteristics, species richness per unit area, numbers of individuals in each tree species and diameter classes of trees as classic lowland tropical rain forests. As the area is at the northern margin of monsoonal tropics, the rain forest differs from equatorial lowland rain forests in having some deciduous trees in the canopy layer, fewer megaphanaerophytes and epiphytes but more species of lianas as well as more species of microphylls. In its floristic composition, about 80% of total families. 94% of total genera and more than 90% of total species are tropical, of which about 38% of genera and 74% of species are tropical Asian. Furthermore, the rain forest has not only almost the same families and genera, but also the same families rank in the top ten both in species richness and in dominance of stems, as lowland forests in southeast Asia. It is indisputable that the flora of the rain forest is part of the tropical Asian flora. However, most of the tropical families and genera have their northern limits in south Yunnan and most have their centre of species diversity in Malesia. More strictly tropical families and genera have relatively lower species richness and importance compared with lowland rain forests in tropical southeast Asia. Thus, the flora also shows characteristics of being at the margin of the tropics. Based mainly on physiognomy and floristic composition the tropical rain forest of Yunnan is classified into two types, i.e. seasonal rain forest and wet seasonal rain forest, the latter is further divided into two subtypes, i.e. mixed rain forest and dipterocarp rain forest. From analysis of geographic elements it is also shown that the tropical rain forest of Yunnan occurs at a geographical nexus with its flora coming mainly from four sources, i.e. Malesia, south Himalayas, Indochina and China.     

Primary and secondary vegetation patches as contributors to floristic diversity in a tropical deciduous forest landscape

The floristic diversity of Mexican tropical deciduous forests (TDF) is of critical importance given the high species richness (alpha diversity), species turnover (beta diversity), and the intense deforestation rates. Currently, most TDF landscapes are mosaics of agricultural land, secondary vegetation, and patches of relatively undisturbed primary vegetation. Here we illustrate how both primary forest remnants and secondary vegetation patches contribute to the floristic diversity of TDF in a landscape of volcanic origin in central Veracruz, Mexico. Our objectives were to assess sampling efficiency and inventory completeness, to compare mean and cumulative species richness between primary forest and secondary vegetation sites, and to analyze beta diversity between vegetation types. In an area of 12,300 m2 we recorded 105 families, 390 genera, and 682 species. Species inventories for both vegetation types were about 80% complete. Secondary vegetation is more alpha diverse than primary forest, both in terms of cumulative and mean species richness. We found a remarkably high beta diversity between vegetation types (75% of complementarity, 91.60% of mean dissimilarity). We also identified the species that contribute the most to similarity within vegetation types and to dissimilarity between vegetation types. Our results support the idea that assessing biodiversity on the landscape scale is an appropriate way to ascertain the impact of human activities. For this land mosaic, conservation of the flora would not be possible by focusing solely on primary forest remnants. We propose the implementation of a network of small conservation areas with a flexible structure, following the “archipelago reserve” model.     

Conservation value of timber quality versus associated non-timber quality stands for understory diversity in Nothofagus forests

Conservation strategies of forested landscapes must consider biodiversity of the included site types, i.e. timber-quality forests and associated non-timber-quality stands. The objectives were to characterize forest overstory structure in timber-quality versus associated non-timber-quality stands; and to compare their understory communities. Six forest types were sampled in Nothofagus forests of Tierra del Fuego (Argentina): two timber-quality N. pumilio forests, and four associated non-timber-quality stands (edge, N. antarctica, wetlands and streamside forests). Overstory structure and understory vegetation (species richness, frequencies, cover and biomass) were characterized during spring and summer seasons. Analysis of variance and multivariates were carried out. Overstory structure differed across the site types, with higher tree size, canopy closure and tree volume in timber-quality stands. Fifty-one understory plant species were observed, but understory variables varied with site types, especially wetlands (highest native and exotic richness, cover and biomass, and 25% of exclusive species). Site types were grouped in three: N. antarctica stands, streamside stands and the other N. pumilio forests according to multivariate analysis. Forty three percent of plants were distributed in all site types, and all timber-quality forest understory species were present in some associated non-timber-quality stands. Timber-quality N. pumilio forests have a marginal value for understory conservation compared to associated non-timber-quality stands, because these last include all the plants observed in timber-quality forests and also possess many exclusive species. Therefore, protection of associated non-timber-quality stands during forest management planning could increase understory conservation at landscape level, and these could be better reserves of understory diversity than retentions of timber-quality stands.     

Successional Models as Guides for Restoration of Riparian Forest Understory

We compare two successional models as guides for restoring native riparian understory species along a 160‐km stretch of the Sacramento River in California. In 2001 and 2007, we surveyed cover, frequency, and richness of native and exotic understory species in 15 sites planted (1989–1996) with overstory species to determine whether native understory species colonized naturally (passive relay floristics model). In 2007, we surveyed 20 additional sites (planted 1997–2003) in 14 of which understory species were planted (initial floristics model) to evaluate whether planting accelerated community recovery. We surveyed 10 remnant forests as references for successional trajectories. Mean cover and frequency of natives changed little over time in sites where they were not planted initially; increases in native cover in a few sites were primarily due to a single common species (Galium aparine). Species composition shifted from light‐demanding to shade‐adapted species, both exotic and native, in response to a doubling of overstory cover. Sites with high intensity understory plantings had greater cover and frequency of native understory species than unplanted sites, but were still low relative to reference forests. Light‐demanding natives (e.g., Artemisia douglasiana, Rubus ursinus, and grasses) established in sites where they were planted; however, a shade‐adapted species (Carex barbarae) did not survive well. Our research indicates that the passive relay floristics and the initial floristic composition approaches serve to restore a few common native understory species, but that planting species as site conditions become appropriate (active relay floristics model) will be needed to restore entire native understory communities.     

Diversity and biogeography of vascular epiphytes in Western Amazonia, Yasuní, Ecuador

Aim Although vascular epiphytes are important components of species richness and complexity of Neotropical forests, vascular epiphytes are under‐represented in large scale biogeographical analyses. We studied the diversity, biogeography and floristic relationships of the epiphytic flora of the Yasuní region (Western Amazonia) in a Neotropical context, with special emphasis on the influence of the Andean flora on floristic composition and diversity of surrounding lowland forests. Location Western Amazonian lowland rainforest, Tiputini Biodiversity Station (0°38′ S 76°09′ W, 230 m a.s.l., 650 ha), Yasuní National Park, Ecuador. Methods We compared the vascular epiphyte flora of Yasuní with 16 published Neotropical epiphyte inventories. Secondly, based on a floristic database with records of more than 70,000 specimens of vascular epiphytes from the Neotropics the elevational composition of eight selected inventories was analysed in detail. Results The vascular epiphyte flora of Yasuní is characterized by a very high species richness (313 spp.). A moderate portion of species is endemic to the Upper Napo region (c. 10%). However, this figure is much higher than previous analyses primarily based on woody species suggested. Geographical ranges of these species match with a proposed Pleistocene forest refuge. Compared with Northern and Central Amazonian sites, Western Amazonian epiphyte communities are characterized by a higher portion of montane and submontane species. Species richness of vascular epiphytes at the sites was correlated with the amount of rainfall, which is negatively correlated with the number of dry months. Main conclusion Recent and historic patterns of rainfall are the driving forces behind diversity and floristic composition of vascular epiphytes in Western Amazonia: high annual rainfall in combination with low seasonality provides suitable conditions to harbour high species richness. The proximity to the Andes, the most important centre of speciation for most Neotropical epiphytic taxa, in combination with the climatic setting has allowed a continuous supply of species richness to the region. At least for epiphytes, the borderline between the Andean and Amazonian flora is much hazier than previously thought. Moreover, the comparatively moist climate in Western Amazonia during the Pleistocene has probably led to fewer extinctions and/or more speciation than in more affected surrounding lowlands.     

Vegetation diversity, composition and structure in a cattle agro-landscape of Matiguás, Nicaragua

The diversity, composition and structure of vegetation in a cattle landscape in Matiguás, Nicaragua was characterized, and the floristic and structural differences of six types of habitats (secondary forests, riparian forests, charrales, live fences and pastures with high and low tree cover) were compared. A total of 3 949 trees of 180 species and 52 families were recorded. Forty six percent of the total trees reported for the landscape were represented by Guazuma ulmifolia (18.5%), Bursera simaruha (13.2%), Tabebuia rosea (6.3%), Enterolobium cyclocarpum (4.2%) and Albizia saman (3.4%). Many of the dominant species in the landscape were typical of open and disturbed areas. There were significant differences between the different habitats in the patterns of tree species richness, abundance, diversity, structure and floristic composition. The riparian forests had greater tree richness (p=0.0001) and diversity (p=0.0009) than other habitats. The floristic composition varied across habitats. with pairs of habitats sharing between 18.4 and 51.6% of the same tree species, and with clear differences in composition between the forested (riparian and secondary forests) and agricultural habitats. Of the habitats studied, the riparian forests and secondary forests seem to have greatest value for the conservation of the flora in the agropaisaje because they have the greatest species richness, and maintain small populations of endangered species. On the basis of the study, we recommend including agricultural landscapes in strategies to conserve tree diversity and suggest measures to ensure the maintenance of tree diversity in the Matiguas landscape.     

Species richness and soil properties in Pinus ponderosa forests: A structural equation modeling analysis

Question: How are the effects of mineral soil properties on understory plant species richness propagated through a network of processes involving the forest overstory, soil organic matter, soil nitrogen, and understory plant abundance? Location: North‐central Arizona, USA. Methods: We sampled 75 0.05‐ha plots across a broad soil gradient in a Pinus ponderosa (ponderosa pine) forest ecosystem. We evaluated multivariate models of plant species richness using structural equation modeling. Results: Richness was highest at intermediate levels of understory plant cover, suggesting that both colonization success and competitive exclusion can limit richness in this system. We did not detect a reciprocal positive effect of richness on plant cover. Richness was strongly related to soil nitrogen in the model, with evidence for both a direct negative effect and an indirect non‐linear relationship mediated through understory plant cover. Soil organic matter appeared to have a positive influence on understory richness that was independent of soil nitrogen. Richness was lowest where the forest overstory was densest, which can be explained through indirect effects on soil organic matter, soil nitrogen and understory cover. Finally, model results suggest a variety of direct and indirect processes whereby mineral soil properties can influence richness. Conclusions: Understory plant species richness and plant cover in P. ponderosa forests appear to be significantly influenced by soil organic matter and nitrogen, which are, in turn, related to overstory density and composition and mineral soil properties. Thus, soil properties can impose direct and indirect constraints on local species diversity in ponderosa pine forests.     

Considerations on structure and function of the “Central-Amazonian inundation forest ecosystem” with particular emphasis on selected soil animals

Summary Studies were carried out in Central-Amazonian inundation forests having seasonal inundations during the emersion phase (e.p.) and the submersion phase (s.p.). Two várzea forests (white water areas), one in the Amazon valley on Ilha de Curarí and one at Lago Janauarí, and an igapó (black water area) situated in the Rio Negro valley at Rio Tarumã Mirím were investigated. A community analysis consisting of the species diversity and species similarity of the aquatic and terrestrial soil fauna was executed in these forests. Each forest is differentiated into an inner, middle, and outer part. The annual development of the soil invertebrates is dictated by the periodic changes in water level. Data on species diversity indicate an alternate occurrence of the soil- and trunk-dwelling carabids and staphylinids (Coleoptera). Low species diversity is obtained for the aquatic soil fauna during the high water period. This is correlated with both oxygen concentration and relative abundance of a few specialized species.A function model approach was tried. Investigations showed that both the phytophages and the saprophages consumed about 9.4% and 13% respectively of the yearly litter fall. Litter decomposition is retarded during the inundation period (170 g·m^-2·s.p.^-1 is decomposed) but is accelerated during the emersion phase (420 g·m^-2·e.p.^-1 is decomposed). Nearly 14% of the yearly litter fall is presumed to be exported by the current from the inundation forest during the inundation phase, probably into the surrounding rivers and lakes. A rough estimation of a nutrient budget containing some selected elements was given. The estimation allowed us to suppose, that in the igapó the loss of some elements cannot be compensated for by the input from the river water and the precipitation. Unlike the igapó, the várzea forest may compensate for this loss of litter through input of nutrients from the inflowing white water. The igapó may thus be considered as a source of nutrients while the várzea forest is a nutrient sink.     

Changes in riparian forest composition along a sedimentation rate gradient

Riparian forests are highly valued for maintaining water quality through the retention of sediments and nutrients. They also provide some of the most diverse and species-rich habitats in the world. What is largely unknown, however, is how sediment deposition affects plant community composition in these forests. The objective of this study was to examine changes in plant community composition across a gradient of increasing rates of sedimentation in riparian forests in the southeastern Coastal Plain, USA. Seventeen plots were established within riparian forests receiving between 0 and 5.5 cm year⁻¹ of sediment deposits. Species density and biomass estimates were collected annually from 2002 to 2006 for overstory and mid-story plant species within each plot. Percent cover and nested frequency of understory plant species were determined annually during 2004–2006. Measures of community composition in the understory, mid-story, and overstory layers of forests were compared to changes in environmental factors associated with increased sedimentation. In the understory, annual, exotic, and upland species had higher importance values in plots receiving high sediment deposition. The densities of shade-intolerant and N-fixing species in the mid-story also increased with increasing sedimentation rates. Increased overstory mortality was associated with high sedimentation rates, though increases in understory light levels in these gaps were not the main driver of understory species changes. Edaphic factors, such as soil texture, moisture, and temperature, were significantly correlated to species composition in all three forest layers, suggesting that changes in soil physical structure due to sedimentation may drive community-level changes in these forests.     

Natural Regeneration in Plantations of Native Trees in Lowland Brazilian Atlantic Forest: Community Structure,Diversity, and Dispersal Syndromes

Plantations of native‐tree species are often recommended for ecological restoration, but the understanding of how these techniques catalyze natural ecological processes is limited. We investigated natural regeneration in five plantations of native trees in the Poço das Antas Biological Reserve (PABR) in the state of Rio de Janeiro, Brazil. The plantations were 9–11 years old, and contained 8–14 native‐tree species with different compositions and relative density of species. We analyzed floristic composition, structure (density and basal area) of overstory and understory strata, as well as other ecological attributes (dispersal syndromes, fruit or seed size, and the availability of fruit for frugivores). Zoochorous species comprised 77% of the community, with a prevalence of the two smallest size classes of propagules (< 0.6 and 0.6–1.6 cm) in natural regeneration. The density of zoochorous plants in the understory was positively correlated with their density in the overstory, indicating their influence on natural regeneration (r² = 0.36; p < 0.0002). Fruit availability for frugivores (density and richness of plants fruiting during the year) was also positively correlated with the density of stems in the understory. Therefore, attributes such as dispersal syndrome and fruiting season should be considered in selecting species to be planted. The differences in natural regeneration observed in each of the native‐tree plantations indicated that the performance of plantations as a restoration strategy may differ, depending on initial species composition, planting density, and site conditions.     

Influence of overstory composition on understory colonization by native species in plantations on a degraded tropical site

Abstract. Patterns of understory colonization by native and naturalized trees and shrubs were evaluated in 4.5-year-old plantations of three exotic tree species, Casuarina equisetifolia, Eucalyptus robusta, and Leucaena leucocephala, on a degraded coastal grassland site with reference to overstory composition and understory environmental conditions. 19 secondary forest species were established in the plantation understories (with a total area of 0.52 ha), while no natural regeneration occurred in unplanted, though protected, control areas. The majority of these species (90 %) and the total seedling population (97 %) were zoochorous, indicating the importance of frugivorous bats and particularly birds as facilitators of secondary forest species colonization. Understory species richness and seedling densities were affected significantly by overstory composition, the most abundant regeneration occurring beneath Leucaena and least under Casuarina. Understory colonization rates within mixed-species stands were intermediate between those of single-species stands of the trees comprising their overstories. Significant negative correlations were found between understory species richness and seedling density, and forest floor depth and dry mass, especially for small-seeded ornithochorous species. Higher colonization rates near the peripheries of plantation plots relative to plot interiors were due in part to roosting site preferences by frugivores, particularly bats. The study results indicate that overstory species selection can exert a significant influence on subsequent patterns of colonization by secondary forest species and is an important consideration in the design of plantations for ‘catalyzing’ succession on deforested, degraded sites.     

Influence of light and soil moisture on Sierran mixed-conifer understory communities

Sierra Nevada forests have high understory species richness yet we do not know which site factors influence herb and shrub distribution or abundance. We examined the understory of an old-growth mixed-conifer Sierran forest and its distribution in relation to microsite conditions. The forest has high species richness (98 species sampled), most of which are herbs with sparse cover and relatively equal abundance. Shrub cover is highly concentrated in discrete patches. Using overstory tree cover and microsite environmental conditions, four habitats were identified; tree cluster, partial canopy, gap, and rock/shallow soil. Herb and shrub species were strongly linked with habitats. Soil moisture, litter depth and diffuse light were the most significant environmental gradients influencing understory plant distribution. Herb cover was most strongly influenced by soil moisture. Shrub cover is associated with more diffuse light, less direct light, and sites with lower soil moisture. Herb richness is most affected by conditions which influence soil moisture. Richness is positively correlated with litter depth, and negatively correlated with direct light and shrub cover. Disturbance or management practices which change forest floor conditions, shallow soil moisture and direct light are likely to have the strongest effect on Sierran understory abundance and richness.     

Understory vegetation response to mountain pine beetle disturbance in northern Colorado lodgepole pine forests

Understory plants are an important element of forests, having a considerable influence on ecosystem functioning and canopy-tree development following disturbance. Recent bark beetle outbreaks across western North American forests have caused extensive canopy mortality, creating new growing conditions that provide the opportunity for changes within the intact understory. Over a five-year period following peak mountain pine beetle (MPB) activity across lodgepole pine-dominated forests in Rocky Mountain National Park, Colorado, we measured the changes in plant diversity, cover, and dominance by lifeform and quantified tree regeneration rates. Average species richness and diversity increased, but overall plant cover did not change. Graminoids appeared to benefit the most, increasing in average cover, richness, and relative dominance. The rise in graminoid dominance was largely at the expense of shrubs, which showed little ability to benefit from overstory mortality within the first years following attack. Most plant responses were positively related to the total tree basal area lost since the peak of the outbreak, suggesting that increased resource availability following tree death may benefit understory plants. However, a negative relationship between several understory variables and tree sapling density provides evidence that understory plants compete with saplings for the newly available resources. Tree seedling density nearly doubled over the duration of the study, indicating a strong regeneration pulse. Among species, lodgepole pine displayed the greatest tree seedling establishment. This is one of the first studies to use repeated measurements to describe this often-overlooked component of forest change associated with MPB disturbance.