Body size determines soil community assembly in a tropical forest

Tropical forests shelter an unparalleled biological diversity. The relative influence of environmental selection (i.e., abiotic conditions, biotic interactions) and stochastic–distance‐dependent neutral processes (i.e., demography, dispersal) in shaping communities has been extensively studied for various organisms, but has rarely been explored across a large range of body sizes, in particular in soil environments. We built a detailed census of the whole soil biota in a 12‐ha tropical forest plot using soil DNA metabarcoding. We show that the distribution of 19 taxonomic groups (ranging from microbes to mesofauna) is primarily stochastic, suggesting that neutral processes are prominent drivers of the assembly of these communities at this scale. We also identify aluminium, topography and plant species identity as weak, yet significant drivers of soil richness and community composition of bacteria, protists and to a lesser extent fungi. Finally, we show that body size, which determines the scale at which an organism perceives its environment, predicted the community assembly across taxonomic groups, with soil mesofauna assemblages being more stochastic than microbial ones. These results suggest that the relative contribution of neutral processes and environmental selection to community assembly directly depends on body size. Body size is hence an important determinant of community assembly rules at the scale of the ecological community in tropical soils and should be accounted for in spatial models of tropical soil food webs.     

Carbon and nutrient limitation of soil microorganisms and microbial grazers in a tropical montane rain forest

We investigated the role of carbon, nitrogen and phosphorus as limiting factors of microorganisms and microbial grazers (testate amoebae) in a montane tropical rain forest in southern Ecuador. Carbon (as glucose), nitrogen (as NH₄NO₃) and phosphorus (as NaH₂PO₄) were added separately and in combination bimonthly to experimental plots for 20 months. By adding glucose and nutrients we expected to increase the growth of microorganisms as the major food resource of testate amoebae. The response of microorganisms to experimental treatments was determined by analysing microbial biomass (SIR), fungal biomass and microbial community composition as measured by phospholipid fatty acids (PLFAs). We hypothesized that the response of testate amoebae is closely linked to that of microorganisms. Carbon addition strongly increased ergosterol concentration and, less pronounced, the amount of linoleic acid as fungal biomarker, suggesting that saprotrophic fungi are limited by carbon. Microbial biomass and ergosterol concentrations reached a maximum in the combined treatment with C, N and P indicating that both N and P also were in short supply. In contrast to saprotrophic fungi and microorganisms in total, testate amoebae suffered from the addition of C and reached maximum density by the addition of N. The results indicate that saprotrophic fungi in tropical montane rain forests are mainly limited by carbon whereas gram positive and negative bacteria benefit from increased availability of P. Testate amoebae suffered from increased dominance of saprotrophic fungi in glucose treatments but benefited from increased supply of N. The results show that testate amoebae of tropical montane rain forests are controlled by bottom–up forces relying on specific food resources rather than the amount of bacterial biomass with saprotrophic fungi functioning as major antagonists. Compared to temperate systems microbial food webs in tropical forests therefore may be much more complex than previously assumed with trophic links being rather specific and antagonistic interactions overriding trophic interactions.     

Density and community structure of soil- and bark-dwelling microarthropods along an altitudinal gradient in a tropical montane rainforest

Microarthropod communities in the soil and on the bark of trees were investigated along an elevation gradient (1,850, 2,000, 2,150, 2,300 m) in a tropical montane rain forest in southern Ecuador. We hypothesised that the density of microarthropods declines with depth in soil and increases with increasing altitude mainly due to the availability of resources, i.e. organic matter. In addition, we expected bark and soil communities to differ strongly, since the bark of trees is more exposed to harsher factors. In contrast to our hypothesis, the density of major microarthropod groups (Collembola, Oribatida, Gamasina, Uropodina) was generally low and decreased with altitude. However, as we predicted the density of each of the groups decreased with soil depth. Density of microarthropods on tree bark was lower than in soil. Overall, 43 species of oribatid mites were found, with the most abundant higher taxa being Poronota, pycnonotic Apheredermata, Mixonomata and Eupheredermata. The oribatid mite community on bark did not differ significantly from that in soil. The number of oribatid mite species declined with altitude (24, 23, 17 and 13 species at 1,850, 2,000, 2,150 and 2,300 m, respectively). Rarefaction curves indicate that overall about 50 oribatid mite species are to be expected along the studied altitudinal gradient. Results of this study indicate (1) that microarthropods may be limited by the quality of resources at high altitudes and by the amount of resources at deeper soil layers, and (2) that the bark of trees and the soil are habitats of similar quality for oribatid mites.     

Species composition of an ectomycorrhizal fungal community along a local nutrient gradient in a boreal forest

Soil abiotic factors are considered to be important in determining the distribution of ectomycorrhizal (ECM) fungal species; however, there are few field data to support this. Here, we relate ECM species distributions to changes in soil chemistry along a short (90-m), natural nutrient gradient. The ECM community was characterized, using morphological and molecular techniques, in soil samples collected at 10-m intervals. There were pronounced changes in ECM fungal community structure along the transect, with many taxa showing discrete distributions. Although there was a change of host from Pinus to Picea along the gradient, host-specific fungi did not account for the observed change in community structure. Ordination analyses showed that community structure was strongly correlated with soil characteristics, in particular extractable ammonium and base saturation. However, autocorrelation among soil parameters makes it difficult to isolate the effects of individual parameters. The distinctive changes in soil and vegetation along the transect used in this study provided an exceptional opportunity to examine the local-scale impact of natural spatial heterogeneity on an ECM fungal community.     

Local and regional dominance of woody plants along an elevational gradient in a tropical montane forest of northwestern Bolivia

The aim of this study was to investigate the dominance patterns of woody plants in a tropical montane forest of northwestern Bolivia and to understand underlying processes at the local and regional scales. We inventoried three elevation ranges: lower (1,200–1,500 m), intermediate (2,000–2,300 m), and upper montane forests (2,800–3,100 m). At each elevation, we inventoried two sites that were ~100 km apart. Specifically, we asked the following questions: (1) are dominant taxa distributed locally, or are they also dominant at larger scales? And (2) is the local importance of shared taxa congruent among sites at the same elevation range? We inventoried 18,876 woody plant individuals with a diameter at breast height ≥2.5 cm belonging to 877 species, 286 genera, and 100 families in 54 0.1-ha plots. A strong floristic congruence was found at the family and genus levels within and across elevations, but not at the species level. The pattern of species dominance for the whole study regions was similar to that reported for similar scales in the Amazonia (10–15 % of species accounted for 50–75 % of individuals), although dominant species were not oligarchic across the whole elevational gradient due to the high environmental heterogeneity. Dominant taxa were shared to a larger degree between sites at the same elevational range than non-dominant taxa, indicating that oligarchy does not mean uniformity. Finally, the shared taxa exhibited similar importance between sites at the lower elevation range but dissimilar importance at the higher elevation range, reflecting an increase in the relative importance of local processes versus regional processes with increasing elevation.     

Effects on nutrient cycling of conifer restoration in a degraded tropical montane forest

Background and aims

Exotic coniferous species have been used widely in restoration efforts in tropical montane forests due to their tolerance to adverse conditions and rapid growth, with little consideration given to the potential ecological benefits provided by native tree species. The aim of this study was to elucidate differences in litterfall and nutrient flow between a montane oak forest (Quercus humboldtii Bonpl.) and exotic coniferous plantations of pine (Pinus patula Schltdl. & Cham.) and cypress (Cupressus lusitanica Mill.) in the Colombian Andes.

Methods

Litter production, litter decomposition rate, and element composition of leaf litter were monitored during 3 years.

Results

Litter production in the oak forest and pine plantation was similar, but considerably lower in the cypress plantation . Similar patterns were observed for nutrient concentrations in litterfall, with the exception of Ca which was three times higher in the cypress plantation. The annual decay rate of litter was faster in the montane oak forest than in either of the exotic coniferous plantations. The potential and net return of nutrients to the forest floor were significantly higher in oak forest than in the exotic coniferous plantations.

Conclusions

Future restoration programs should consider new species that can emulate the nutrient flow of native broadleaf species instead of exotic species that tend to impoverish soil nutrient stocks in tropical montane forests.     

Biogeochemical processes along a nutrient gradient in a tropical ombrotrophic peatland

Biogeochemical properties, including nutrient concentrations, carbon gas fluxes, microbial biomass, and hydrolytic enzyme activities, were determined along a strong nutrient gradient in an ombrotrophic peatland in the Republic of Panama. Total phosphorus in surface peat decreased markedly along a 2.7 km transect from the marginal Raphia taedigera swamp to the interior sawgrass swamp, with similar trends in total nitrogen and potassium. There were parallel changes in the forest structure: basal area decreased dramatically from the margins to the interior, while tree diversity was greatest at sites with extremely low concentrations of readily-exchangeable phosphate. Soil microbial biomass concentrations declined in parallel with nutrient concentrations, although microbes consistently contained a large proportion (up to 47%) of the total soil phosphorus. Microbial C:P and N:P ratios and hydrolytic enzyme activities, including those involved in the cycles of carbon, nitrogen, and phosphorus, increased towards the nutrient-poor wetland interior, indicating strong below-ground nutrient limitation. Soil CO₂ fluxes and CH₄ fluxes did not vary systematically along the nutrient gradient, although potential soil respiration determined on drained soils was lower from nutrient-poor sites. Soil respiration responded strongly to drainage and increased temperature. Taken together, our results demonstrate that nutrient status exerts a strong control on above and below-ground processes in tropical peatlands with implications for carbon dynamics and hence long term development of such ecosystems.     

Soil‐based habitat partitioning in understorey palms in lower montane tropical forests

Aim Dispersal assembly and niche assembly are two competing theories proposed to explain the maintenance of species diversity in tropical forests. Dispersal theory emphasizes the role of chance colonization events and distance‐limited seed dispersal in explaining species abundance and distribution, whereas niche theory emphasizes differences among species in requirements for potentially limiting resources. Species distribution patterns in tropical forests often correlate with geology and topography, but tests of the relative importance of dispersal and niche partitioning have been hampered by an inadequate characterization of resource availability. The aim of this study was to explore how soil chemical and physical properties, climate, and geographic distance affect understorey palm communities in lower montane forests. Location Fortuna Forest Reserve, Chiriqui Province, and Palo Seco Forest Reserve, Bocas del Toro Province, in western Panama. Methods Understorey palms and soil nutrient concentrations were surveyed within 10 sites on different soil types across a 13‐km transect. Variation in palm community composition was examined in relation to spatial and environmental variables. Results The 25 understorey palm species recorded in the study were non‐randomly distributed among forests differing in soil nutrient availability. In support of dispersal theory, floristic similarity decreased predictably with increasing geographic distance. However, environmental and soil variables were also correlated with geographic distance. Floristic similarity was also highly associated with a subset of environmental variables. Variation in palm community similarity was most strongly correlated with inorganic nitrogen availability and cation concentration. A subset of soil variables had a stronger relationship with floristic similarity when geographic distance was controlled for than did geographic distance when differences in soils were controlled for. Main conclusions Both dispersal and niche processes affect palm species distribution patterns. Although spatially limited dispersal may influence species distribution patterns, soil‐based habitat associations, particularly with respect to soil nitrogen, cation availability and aluminium concentrations, remain important factors influencing palm community composition at the mesoscale level in this tropical montane forest.     

Biodiversity and carbon storage are correlated along a land use intensity gradient in a tropical montane forest watershed,Mexico

Tropical montane cloud forest landscapes are changing, and forest conversion to other land uses is a major driver of biodiversity loss. Land use intensification can lead to significant losses in biodiversity and carbon storage (C); however, the impacts may vary greatly depending on land use type, management practices, and environmental context. We investigated how biodiversity and C are related along a gradient of land use intensification characterized by four dominant land uses in the upper part of Antigua River watershed, Mexico. The land uses were montane cloud forest, secondary forest, and traditional and intensive shade coffee plantations. We determined tree species composition, diversity, ecosystem structure, wood density and C content in dominant tree species to assess aboveground biomass (AGB) and C storage within eight study sites across the land use intensity gradient. A total of 83 tree species was recorded. A canonical correspondence analysis indicated that land uses are separated by particular tree species assemblages. Forests had higher basal area, density, and biomass than coffee plantations, however, the traditional shade coffee plantation had values similar to secondary forest. Calculating C using the standard estimate of 50% of AGB resulted in an overestimation of stored C by 5.8 to 4.1% compared to calculations based on actual measurements. Carbon storage in AGB and biodiversity were strongly and positively related across the land use intensity gradient, although the distinction between the two different intensities of coffee plantation management was not consistently as clear as we had expected. Carbon was highest in forest, but secondary forests and traditional shade coffee plantation had similar C, while intensive coffee had the lowest C content. These results highlight the importance of considering the potential of low intensity land uses such as traditional coffee plantations to mitigate biodiversity loss and preserve ecosystem functions as part of conservation efforts.     

Productive phenology of tropical montane forests: Fertilization experiments along a moisture gradient

Phenological responses of leaves and roots were studied in the tropical montane forests of Mount Kinabalu, Borneo. Soil nutrient supply, in addition to the supply of light and water, is a potential abiotic factor influencing plant phenology in the tropics. The main objective of this study was to evaluate the contribution of soil nutrient supply to plant productive phenology. Fertilization experiments, including controls, nitrogen fertilized and nitrogen and phosphorus fertilized treatments, were conducted on three vegetation types in different moisture environments. Responses of leaves and roots were compared among treatments and among vegetation types. Leaf flushing was induced by nitrogen fertilization in the upper montane forest, where extremely wet moisture conditions are associated with cloud cover. This induction of leaf flushing by fertilization was not observed in the other forests. Root growth was suppressed by fertilization when leaf flushing was not induced by fertilization. These results indicate that soil pulsed nutrient release could be a cue for leaf flushing in a tropical wet environment, and that leaf phenology could be regulated by external abiotic factors and root phenology could be regulated by internal plant demands.     

Tree community structure and stem mortality along a water availability gradient in a Mexican tropical dry forest

We document spatial changes in species diversity, composition, community structure, and mortality of trees across a gradient of water availability in a tropical dry forest in western Mexico. This gradient occurs along the main stream of a small watershed of less than 1 km in length. Four 30 × 80 m plots were established systematically to include the driest (ridge top of the watershed) to the wettest sites (watershed bottom) within this watershed. All stems larger than 5 cm were identified, and measured for diameter and height. Dead stems larger than 5 cm were measured and classified as: a) found on live or dead trees, and b) standing (“snags”) or lying (“downlogs”) on the ground. The number of recorded species per plot declined from 73 to 44 species as water availability decreased. A decline in estimated total richness, and in Shannon-Wiener and Simpson diversity indices was also observed in the drier plots. Species composition strongly changed along the gradient, with the two ends of the gradient sharing only 11% of the species. Stem density and percentage of dead stems and trees increased in abundance and basal area from the wetter to the drier sites. Tree and stem size (basal area, height and stem diameter) showed the opposite trend. Nonetheless, total basal area of live trees was largest at the two end gradient locations and oscillated between 12.22 m² ha⁻¹ and 7.93 m² ha⁻¹. Proportion of snags increased towards the driest site (from 46 to 72%), while that of down logs decreased. Overall, our results suggest that small-scale gradients of water availability play a paramount role in the spatial organization of tree communities in seasonal tropical environments. This revised version was published online in June 2006 with corrections to the Cover Date.     

Bioassays of nutrient limitation in a tropical rain forest soil

Summary Six speices of shrubs and one large herb with contrasting life history patterns were used as bioassays of nutrient availability in a Costa Rican lowland rain forest soil. Growth responses of the herb (Phytolacca rivinoides, Phytolaccaceae) confirmed soil measurements indicating high availability of N and potentially limiting levels of P, K, Mg and Ca. Growth responses of the shrub species (Miconia spp., Melastomataceae and Piper spp., Piperaceae) to a complete nutrient fertilizer were generally less than that of Phytolacca. Lack of a strong shrub response to +P fertilization is probably due to mycorrhizal associations and slower growth rates of woody species. In general, increased growth did not occur at the expense of phenolic production in the leaves. The results emphasize that assessment of specific nutrient limitations to plant growth vary depending on species selected for the bioassay, even among species from the same community.     

Seasonal changes in soil phosphorus dynamics and root mass along a flooded tropical forest gradient in the lower Orinoco river,Venezuela

The effects of natural hydrological fluctuations on the nature and bioavailability of soil phosphorus (P) in relation to iron (Fe) and aluminum (Al) chemistry and root mass were studied along a flooded tropical forest gradient in Mapire river, Venezuela. Soil samples were collected following a complete natural hydroperiod: end of the dry season (May 2004), end of the rainy season (November 2004) and end of the subsequent dry season (May 2005), and from three zones subjected to different flooding intensities: MAX, MED and MIN zones inundated for 8, 5 and 2 months per year respectively. The results showed that flood induced the increase of resin-Pi in the MAX zone, but not in the MED and MIN zones. Flood in the soil of the MAX zone also induced the increase of the NaOH-Pi fraction, which removes inorganic P sorbed onto secondary Fe and Al minerals. Changes in this redox-sensitive P form can be considered indirect evidence that P in the MAX zone can be released from the dissolution of iron oxyhydroxide. This field study also showed that along the flooded forest gradient, fine root mass declined during the flood event. However, such decline was more pronounced in the MIN zone than in the MAX zone. In this zone fine root mass was higher than in the other zones.     

中亚热带森林土壤微生物群落多样性随海拔梯度的变化

运用Biolog EcoPlate技术, 对武夷山不同海拔植被带(常绿阔叶林(EBF)、针叶林(CF)、亚高山矮林(DF)、高山草甸(AM))土壤微生物群落多样性差异进行了研究。结果表明: 不同海拔植被带土壤微生物群落功能多样性差异显著。土壤平均颜色变化率(AWCD)随培养时间延长而逐渐增加, 同一深度土层的AWCD值随海拔升高而逐渐降低, 大小顺序依次为EFB > CF > DF > AM。同一海拔植被带, 不同深度土层的AWCD值总体趋势依次为0-10 cm > 10-25 cm > 25-40 cm。土壤微生物群落Simpson指数、Shannon-Wiener指数、丰富度指数和McIntosh指数的总体趋势为EBF最高, CF和DF次之, AM最低。不同海拔植被带土壤微生物对不同碳源利用强度存在较大差异, 其中EBF利用率最高, AM利用率最低, 碳水化合物和羧酸类碳源是各海拔植被带土壤微生物的主要碳源。主成分分析结果表明, 从31个因素中提取的与碳源利用相关的主成分1、主成分2分别能解释变量方差的75.27%和16.14%, 在主成分分离中起主要贡献作用的是胺类和氨基酸类碳源。土壤微生物群落多样性随着海拔上升、土层加深而逐渐下降的原因, 可能是生物量、林分凋落物、土壤养分、微小动物、植物根系等多种因素共同作用的结果。     

Light use efficiency of California redwood forest understory plants along a moisture gradient

We investigated photosynthesis of five plant species growing in the understory at three sites (1,170-, 1,600- and 2,100-mm annual moisture inputs), along the geographical range of coastal California redwood forest, to determine whether greater inputs of rain and fog at northern sites enhance photosynthetic utilization of fluctuating light. Measurements of understory light environment and gas exchange were carried out to determine steady state and dynamic photosynthetic responses to light. Leaf area index ranged from 4.84 at the 2,100-mm site to 5.98 at the 1,170-mm site. Maximum rates of net photosynthesis and stomatal conductance (g) did not vary appreciably within species across sites. Photosynthetic induction after a change from low to high light was significantly greater in plants growing in lower light conditions regardless of site. Photosynthetic induction also increased with the rate of g in diffuse light, prior to the increase to saturating light levels. Post-illumination CO₂ assimilation was the largest factor contributing to variation in C gain during simulated lightflecks. The duration of post-illumination photosynthetic activity, total CO₂ assimilation per light received, and light use efficiency during simulated lightflecks increased significantly with moisture inputs in four out of five species. Increasing leaf N concentration with increasing moisture inputs in three out of five species, coupled with changes in leaf N isotopic composition with the onset of the summer fog season suggest that natural N deposition increases with rain and fog inputs and contributes to greater utilization of fluctuating light availability in coastal California redwood forests.     

Habitat differentiation of Lauraceae species in a tropical lower montane forest in northern Thailand

Dependency on topographical habitat was examined for Lauraceae tree species in a lower montane forest using a large-scale research plot established at Doi Inthanon National Park, northern Thailand. Twenty species of 10 genera of Lauraceae were recorded in a 7.5-ha part of the plot; Lauraceae accounted for 18% of the total basal area. Lauraceae was the most species-rich and most abundant family in the plot. In a cluster analysis based on the matrix of spatial associations between species, two clusters were recognized. Members of one cluster seemed to associate with lower-elevation habitats, and members of the other associated with habitats on ridges. By subdividing the study plot into 20m×20m squares, a discriminant analysis could be applied to the presence–absence data for the 17 species that had sufficient population density. The predictor variables used were the relative elevation, slope inclination, slope direction (transformed to deviation from SSW) and slope convexity for each of the squares. The discriminant models were tested statistically by applying the random shift technique. The models were significant for 11 of the species (65% of the species examined) and were associated with the topographical condition of the habitat. Stepwise selection of the predictor variables for these 11 species revealed that relative elevation and slope convexity were the most important factors for predicting the presence or absence of the Lauraceae species. Both these variables were considered to indicate the hydrological condition of the habitat.     

Changes in diversity along a successional gradient in a Costa Rican upper montane Quercus forest

Changes in terrestrial vascular plant diversity along a successional gradient were studied in a Costa Rican upper montane Quercus forest. In 1991 and 1992 species presence and cover were recorded in 12 successional 0.1 ha forest plots. A total of 176 species in 122 genera and 75 families were found. Asteraceae was the most speciose family. With the help of TWINSPAN three successional phases were classified: (i) Early Secondary Forest (ESF, 145 spp.), (ii) Late Secondary Forest (LSF, 130 spp.) and (iii) Primary Forest (PF, 96 spp.). Detrended Correspondence Analysis (DCA) species ordination using DECORANA illustrates that different ecological species groups can be distinguished along the time sequence. Alpha diversity (Shannon-Wiener index, among others) in ESF and LSF was significantly greater than in PF. This is probably explained by downslope migration of numerous sub(alpine) species to cleared and recently abandoned montane sites. Beta diversity applying Sørensen's similarity coefficients declined during succession. Using linear regression, the minimum time required for floristic recovery following disturbance and abandonment was calculated at 65.9 years. A comparison with other studies shows that secondary forests in upper montane Costa Rica can be as diverse as in neotropical lowlands.     

Hypolithic and soil microbial community assembly along an aridity gradient in the Namib Desert

The Namib Desert is considered the oldest desert in the world and hyperarid for the last 5 million years. However, the environmental buffering provided by quartz and other translucent rocks supports extensive hypolithic microbial communities. In this study, open soil and hypolithic microbial communities have been investigated along an East–West transect characterized by an inverse fog-rainfall gradient. Multivariate analysis showed that structurally different microbial communities occur in soil and in hypolithic zones. Using variation partitioning, we found that hypolithic communities exhibited a fog-related distribution as indicated by the significant East–West clustering. Sodium content was also an important environmental factor affecting the composition of both soil and hypolithic microbial communities. Finally, although null models for patterns in microbial communities were not supported by experimental data, the amount of unexplained variation (68–97 %) suggests that stochastic processes also play a role in the assembly of such communities in the Namib Desert.