The influence of tree species on canopy soil nutrient status in a tropical lowland wet forest in Costa Rica

The canopy is host to a large percentage of the flora and fauna in tropical wet forests and is distinct from the forest floor in plant richness, soil type and microclimate. In this study, we examined the influence of tree species and season on soil nutrient cycling processes in canopy soils of four tree species common to Costa Rican wet forests. We also compared the canopy soils to the associated forest floor mineral soils. Both tree species and season had strong effects on canopy soil nutrients and processes. Canopy soils from trees with high litter lignin concentrations had higher net N-mineralization rates and higher dissolved inorganic N concentrations than those with low lignin concentrations. During the dry season, net N-immobilization occurred and dissolved organic and inorganic N and available P concentrations were significantly higher than during the wet season. Overall, canopy soils had higher N levels and higher fungi + bacteria richness than forest floor mineral soils. The differences in canopy soil properties observed among tree species indicates that these species have distinct N cycles that reflect differences in both soil origin and biological controls.     

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

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

元江干热河谷植物叶片解剖和养分含量特征

研究了元江干热河谷旱田植物(旱季可浇灌,水分较好)和山坡半萨王纳植被中(自然状况,水分较差)共20种的叶片形态解剖特征,以及7种山坡植物叶片养分含量特征.结果表明,山坡植物叶片比叶重大,气孔密度大,气孔长度小,海绵组织/栅栏组织的值小等.元江干热河谷山坡植物叶片养分含量低,1.3%＞Ca＞N＞K＞1%＞Mg＞P＞S.除氮元素外,其它元素种间差别1～3倍.与热带植物群落叶片养元素含量相比,热带雨林＞元江山坡植物＞东南亚沙地旱生林和巴西矮卡廷加群落,表明元江干热河谷植物叶片具有明显的旱生性形态解剖特征,且叶片养分含量也较低.     

Effects of tree species composition on within‐forest distribution of understorey species

Question: Do tree species, with different litter qualities, affect the within‐forest distribution of forest understorey species on intermediate to base‐rich soils? Since habitat loss and fragmentation have caused ancient forest species to decline, those species are the main focus of this study. Location: Three ancient forests, along a soil gradient from acidification‐sensitive to base‐rich, were studied: Limbrichterbosch and Savelsbos in The Netherlands and Holtkrat in Denmark. Methods: Canopy and soil surveys along transects generated data for Redundancy Analysis on tree – humus relationships. We analysed the distribution of forest plant species with Canonical Correspondence Analysis. The explanatory factors were soil characteristics (pH, organic matter, loam content and thickness of the humus layers), external crown projection, ground water and canopy data. We further analysed the relationship between forest species and humus characteristics with Spearman correlations. Results: Tree species have a significant impact on humus characteristics through the nature of their litter. Humus characteristics significantly explain the distribution of forest understorey species. The pH of the first 25 cm mineral soil and the thickness of the F‐ (fermentation) layer are the primary factors affecting the distribution of ancient forest species. Conclusion: This study indicates that the species composition of the forest canopy affects the distribution of forest understorey species. Ancient forest species are more abundant and frequent underneath trees with base‐rich litter. On acidification‐sensitive soils these relationships were stronger than on more base‐rich, loamy soils.     

Understorey plant community structure in lower montane and subalpine forests, Grand Canyon National Park, USA

Aim Our objectives were to compare understorey plant community structure among forest types, and to test hypotheses relating understorey community structure within lower montane and subalpine forests to fire history, forest structure, fuel loads and topography. Location Forests on the North Rim of Grand Canyon National Park, Arizona, USA. Methods We measured understorey (< 1.4 m) plant community structure in 0.1‐ha plots. We examined differences in univariate response variables among forest types, used permutational manova to assess compositional differences between forest types, and used indicator species analysis to identify species driving the differences between forest types. We then compiled sets of proposed models for predicting plant community structure, and used Akaike's information criterion (AIC_C) to determine the support for each model. Model averaging was used to make multi‐model inferences if no single model was supported. Results Within the lower montane zone, pine–oak forests had greater understorey plant cover, richness and diversity than pure stands of ponderosa pine (Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.). Plant cover was negatively related to time since fire and to ponderosa pine basal area, and was highest on northern slopes and where Gambel oak (Quercus gambelii Nutt.) was present. Species richness was negatively related to time since fire and to ponderosa pine basal area, and was highest on southern slopes and where Gambel oak was present. Annual forb species richness was negatively related to time since fire. Community composition was related to time since fire, pine and oak basal area, and topography. Within subalpine forests, plant cover was negatively related to subalpine fir basal area and amounts of coarse woody debris (CWD), and positively related to Engelmann spruce basal area. Species richness was negatively related to subalpine fir basal area and amounts of CWD, was positively related to Engelmann spruce basal area, and was highest on southern slopes. Community composition was related to spruce, fir and aspen basal areas, amounts of CWD, and topography. Main conclusions In montane forests, low‐intensity surface fire is an important ecological process that maintains understorey communities within the range of natural variability and appears to promote landscape heterogeneity. The presence of Gambel oak was positively associated with high floristic diversity. Therefore management that encourages lightning‐initiated wildfires and Gambel oak production may promote floristic diversity. In subalpine forests, warm southern slopes and areas with low amounts of subalpine fir and CWD were positively associated with high floristic diversity. Therefore the reduction of CWD and forest densities through managed wildfire may promote floristic diversity, although fire use in subalpine forests is inherently more difficult due to intense fire behaviour in dense spruce–fir forests.     

Biomass and nutrient distribution in eucalypt plantations growing on rehabilitated bauxite mines

The quantities of above ground biomass and nutrients accumulated by two eucalypt forests growing on rehabilitated bauxite mines in south-western Western Australia were determined. The plantation at one site was 7.5 years old and had no understorey. The plantation at the second site was 3.5 years old and had been sown with seed of leguminous understorey species, resulting in a dense understorey. Similar amounts of biomass had accumulated at the two sites. However, the second site, in which the legume understorey accounted for 67% of the total above ground biomass, had about three times as much nitrogen in its biomass as the site without an understorey. There was also more sulphur in the biomass at this site. There were similar amounts of other nutrients at both sites. The plantation with an understorey is considered more likely to develop into a forest similar to the eucalypt forest prior to mining than the plantation without an understorey.     

宁镇山区不同森林土壤生物学特性的研究

对宁镇山区不同森林土壤微生物、土壤酶活性,营养元素的变化及凋落物的分解等生物学特性进行了探讨,结果表明,土壤微生物数量和6种土壤酶的生化活性在不同的季节的不同的林型土壤中均呈现出一定的变化规律,土壤中营养元素的含量随森林的生长周期而发生规律性的变化,并与土壤中微生物数量和土壤酶活性有显著的相关性,不同林地凋落物分解速率与其营养元素归还速率存在一定的时空差异,对次生栎林、毛竹林和杉木林土壤各生物学特性的比较表明,栎林土壤营养元素的含量最丰富,并具有较强的自肥调控能力,因此种植针阔混交林有利于防止针叶纯林的地力衰退现象。     

Inhibition of an invasive plant (<Emphasis Type="Italic">Mikania micrantha</Emphasis> H.B.K.) by soils of three different forests in lower subtropical China

Biological invasion represents one of the most serious threats to biodiversity, and invasion ecology research has become one of the central issues of contemporary environmental science. However, the relative role of soil development as correlated with succession in influencing variation in invasion resistance has seldom been examined. We hypothesized that the invasion potential of exotic plants depends on soil conditions. In this study, we explored variation among soils of three forest types in their resistance to invasion by Mikania micrantha H.B.K. in lower subtropical China; this species is one of the top ten most invasive weeds in the world. In model systems of soils from different forests, M. micrantha performed worst on undisturbed late-successional stage forest (monsoon evergreen broadleaf forest, MEBF) soils and did not appear to take advantage of abundant soil nutrients. These results could reflect that a potential inhibitory effect was driven by MEBF soil. Plant litter is an important component of soils, litter bioassay and addition experiments demonstrated native plant species from different forests differentially confer invasion resistance and affect the performance of exotic plants. Naturally, other factors such as the relative abundances of nutrients, microbial feedbacks, could have contributed to these results as well. Our results challenge the notion that forests are not immune to invasion and suggest that maintaining late-successional plant communities in lower subtropical China, which could be related to increased productivity and stability, should be a primary objective of land managers.     

Rainforest expansion reduces understorey plant diversity and density in open forest of eastern Australia

The expansion of rainforest pioneer trees into long‐unburnt open forests has become increasingly widespread across high rainfall regions of Australia. Increasing tree cover can limit resource availability for understorey plant communities and reduce understorey diversity. However, it remains unclear if sclerophyll and rainforest trees differ in their competitive exclusion of understory plant communities, which contain most of the floristic diversity of open forests. Here, we examine dry open forest across contrasting fire histories (burnt and unburnt) and levels of rainforest invasion (sclerophyll or rainforest midstorey) to hindcast changes in understorey plant density, richness and composition. The influence of these treatments and other site variables (midstorey structure, midstorey composition and soil parameters) on understorey plant communities were all examined. This study is the first to demonstrate significantly greater losses of understorey species richness, particularly of dry open‐forest specialists, under an invading rainforest midstorey compared to a typical sclerophyll midstorey. Rainforest pioneers displaced over half of the understorey plant species, and reduced ground cover and density of dry forest specialists by ~90%. Significant understorey declines also occurred with increased sclerophyll midstorey cover following fire exclusion, although losses were typically less than half that of rainforest‐invaded sites over the same period. Understorey declines were closely related to leaf area index and basal area of rainforest and wattle trees, suggesting competitive exclusion through shading and potentially belowground competition for water. Around 20% of displaced species lacked any capacity for population recovery, while transient seed banks or distance‐limited dispersal may hinder recovery for a further 68%. We conclude that rainforest invasion leads to significant declines in understorey plant diversity and cover in open forests. To avoid elimination of local native plant populations in open forests, fires should occur with sufficient frequency to prevent overstorey cover from reaching a level where shade‐intolerant species fail to thrive.     

Restoration of the understorey layer of recent forest bordering ancient forest

Abstract. Recently established forests are commonly characterized by an impoverished understorey. Restoration is mostly based on spontaneous secondary succession, but little is known about the time period needed to achieve a community species pool with species composition equal to that of ancient forests. Vegetation in transects of 197 plots in 13 recent forest stands contiguous to the Meerdaalwoud ancient forest complex was surveyed. The recent forest stands ranged in age from 36 to 132 yr. The community species pool was described with an ecological, functional and phytosociological approach and based on groups derived from a CCA. Differences in community species pool between age classes of recent forest stands were analysed. During establishment of a new forest competitive species, forest edge species and species with high Ellenberg values for light and nitrogen and a more persistent seed bank will dominate the understorey. After 90 yr of succession the cover by these species decreases and reaches equal values to ancient forest after ca. 105 yr. A large number of forest species will be able to colonize the forest in less than 90 yr. Some typical forest species, however, have very low colonization rates and still have low cover in recent forest more than 105 yr old, so that complete restoration of the understorey requires a time period of over a century. Anthropogenic introduction of forest plant species may reduce the time required for ancient forest vegetation equality.     

Litter fall and nutrient cycling in karri (Eucalyptus diversicolor F. Muell.) forest in relation to stand age

Annual litter fall, nutrient concentrations in litter components and annual weight of nutrients in litter fall have been estimated for karri forest stands aged 2, 6, 9 and 40 years and in mature forest. The weight of litter falling annually increases with stand age, ranging from 1.13 t/ha in 2-year-otd regeneration to 9.45 t/ha in mature forest. This increase is due mainly to greater amounts of twigs, bark and fruit falling in older stands. Leaf fait is relatively independent of stand age once the canopy of regenerating stands closes and the understorey has developed. Concentrations of N, P, K, S and Mn in karri leaf litter differ significantly between sites and the differences appear to be related to stand age. Highest levels of these elements are found in karri leaf litter from the youngest stand and the concentrations decrease with increasing stand age. The amounts of annual litter fall and of nutrients cycling in litter are among the largest reported for Australian forests. In particular cycling of Ca, K and Mg in mature karri forest is greater than has been reported for any other eucalypt forest. Karri forest understorey plays a key rote in nutrient cycling in these ecosystems, contributing 30–70% of the weight of many of the nutrients in the leaf component of titter. Understorey leaf material is particularly important in the cycling of N, S and the micro-nutrients Cu and Zn.     

Do nutrient-poor soils inhibit development of forests? A nutrient stock analysis

Nutrient-poor soils often support low-stature grasslands, savannas and shrublands where the climate is warm enough and wet enough for closed forests. Though this pattern has long been recognised, the causes are debated and poorly explored. I tested the hypothesis that forest fails to develop where the total nutrient pool is too small to construct both the foliage and the wood. I estimated potential woody biomass from the difference between soil nutrient stocks and forest foliage stocks. Nutrient stocks required for foliage were estimated from leaf tissue concentrations and foliage biomass typical of Amazon forests. Potential wood biomass was estimated from wood nutrient concentrations typical of Amazon forests. Data on soil nutrient stocks were assembled from studies from South American and African forests and savannas and from south-western Australian and south-west African heathlands. According to these calculations, estimated nutrient stocks (kg ha^?1) to build a forest would need to be > = P: 20–30, K 200–350, Ca 300–600 and Mg 55–65. Many surface soil horizons from both savanna and heathland sites were below these thresholds. However when deeper soil layers were included, most soils had adequate nutrient stocks. The nutrients in shortest supply were Ca and K and not P. This study suggests that nutrient stocks are usually adequate for constructing the wood needed to build a forest, except where soils are highly leached and very shallow. The implication is that, at steady state, low nutrient stocks seldom constrain forest development. The apparent failure of low nutrient stocks to explain the missing forests on nutrient-poor soils emphasises the need for new ideas on how nutrients, alone or in combination with other factors such as fire, influence vegetation structure.     

Herbaceous Understorey: An Overlooked Player in Forest Landscape Dynamics?

Dense herbaceous understorey layers can impact tree regeneration and thereby affect forest succession. However, the implications of this interaction on large spatial and temporal scales are not well understood. To analyse the role of overstorey–understorey interactions for forest dynamics, we implemented an understorey layer (composed of the plant functional types grasses, forbs, ferns, herbs and shrubs) in the forest landscape model LandClim, focusing on competition for light as the main mode of interaction. The model was used to simulate post-disturbance dynamics over an elevational gradient of 560–2800 m a.s.l. in Central Europe. Simulation results showed strong impacts of the herbaceous understorey on tree regeneration within the first decades, but generally little effect on late-successional forests, i.e. not providing any evidence for ‘arrested’ succession. The results also demonstrated varying overstorey–understorey interactions across the landscape: strongest effects were found at low to mid elevations of the study landscapes, where tree establishment was substantially delayed. At high elevations, tree growth and establishment were more limited by low temperatures, and the effect of light competition from the understorey was negligible. Although the inclusion of large windthrow disturbances increased the biomass of herbaceous understorey across the landscape, this had only a small impact on the overstorey due to the presence of advance regeneration of trees. Overall, our results demonstrate that the herbaceous understorey can have a significant impact for forest landscape dynamics through light competition, and that non-woody plants should not be neglected in forest modelling.     

Vertically stratified frugivore community composition and interaction frequency in a liana fruiting across forest strata

Vertical stratification is a key feature of tropical forests and structures plant–frugivore interactions. However, it is unclear whether vertical differences in plant-frugivore interactions are due to differences among strata in plant community composition or inherent preferences of frugivores for specific strata. To test this, we observed fruit removal of a diverse frugivore community on the liana Marcgravia longifolia in a Peruvian rain forest. Unlike most other plants, Marcgravia longifolia produces fruits across forest strata. This enabled us to study effects of vertical stratification on fruit removal without confounding effects of plant species and stratum. We found a high number of visits of a few frugivore species in the understorey and a low number of visits of many different frugivores in the canopy and midstorey. Whereas partial and opportunistic frugivores foraged across strata with differing frequencies, obligate frugivores were only found eating fruits in the higher strata. Avian frugivores foraging in the canopy were mainly large species with pointed wings, whereas under- and midstorey avian foragers were smaller with rounded wings. Our findings suggest a continuous shift in the frugivore community composition along the vertical gradient, from a few generalized frugivores in the understorey to a diverse set of specialized frugivores in the canopy. This shift in the frugivore community leads to correlated, reciprocal changes from specialized to generalized plant-frugivore interactions. Thus, we conclude that vertical niche differentiation between species in tropical forests persists even when food resources are available across strata. This highlights its role for promoting biodiversity and ecosystem functioning.     

Impacts of land use on riparian forest along an urban – rural gradient in southern Manitoba

Extensive landscape modification by humans has led to the fragmentation of riparian forests across North America. We compared the vegetation of extant riparian forest along an urban-rural disturbance gradient. In 1999, twenty-five sites along Assiniboine River in Manitoba, Canada were categorized according to land use: urban, suburban, high intensity rural, low intensity rural, and relatively high quality reference forest. Differences in herbaceous, shrub, and tree species composition and diversity were related to the proportion of surrounding land use, forest patch size, connectivity, and area:perimeter ratio. Urban riparian forests were more disturbed and isolated. They were smaller and characterized by drier, more alkaline soils. Moreover, they had significantly lower native and overall understorey species diversity, and had a higher proportion of exotics including Solanum dulcamara and Hesperis matronalis. Suburban forests were less disturbed, faced greater development pressure, and had sandier soils. Although suburban understorey diversity was similar to that of rural forests, suburban sites had a higher proportion of exotic species, especially escaped horticultural and invasive species including Caragana arborescens and Rhamnus cathartica. Reference sites were relatively large and exhibited greater connectivity, but there was little difference in species composition and diversity among high intensity rural, low intensity rural, and reference sites. These site types were less disturbed than either urban or suburban forests, and reference sites were characterized by hydrophilic species including Scirpus fluviatilis and Carex aquatilis. Our results suggest that landscape measures of disturbance, and related changes in environment, may be confidently used to assess impacts of land use on vegetation along urban-rural gradients. This revised version was published online in June 2006 with corrections to the Cover Date.     

Overstorey tree species regulate colonization by native and exotic plants: a source of positive relationships between understorey diversity and invasibility

The North American woody species, Prunus serotina Ehrh., is an aggressive invader of forest understories in Europe. To better understand the plant invasion process, we assessed understorey plants and Prunus serotina seedlings that have colonized a 35-year-old replicated common-garden experiment of 14 tree species in south-western Poland. The density and size of established (> 1 year old) P. serotina seedlings varied among overstorey species and were related to variation in light availability and attributes of the understorey layer. In a multiple regression analysis, the density of established P. serotina seedlings was positively correlated with light availability and understorey species richness and negatively correlated with understorey species cover. These results suggest that woody invader success is adversely affected by overstorey shading and understorey competition for resources. Simultaneously, however, invader success may generally be positively associated with understorey species richness because both native and invasive plant colonization respond similarly to environmental conditions, including those influenced by overstorey tree species. Identification of characteristics of forests that increase their susceptibility to invasion may allow managers to target efforts to detect invasives and to restore forests to states that may be less invasible.     

Biomass and nutrient distribution in two eucalypt forest ecosystems

The distribution of biomass and nutrients (N, P, K, Na, Mg, and Ca) among components of a Eucalyptus regnans forest and a mixed Eucalyptus obliqua-Eucalyptus dives forest near Melbourne in southern Victoria have been determined and are discussed. Both forests were found to have relatively low root biomass. Trees and soil accounted for more than 80% of each nutrient present in both ecosystems. The results of nutrient distribution studies in Australian eucalypt forests and in temperate forests elsewhere in the world are compared. These comparisons suggest that the above-ground living biomass in eucalypt forests contains no abnormally high or low quantities of macronutrients in relation to coniferous or deciduous hardwood forests. Detailed comparison among ecosystems is rendered difficult by the variable treatment given to roots and soils.     

Vertical distribution of epiphytic bryophytes in an Indonesian rainforest

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

Functional identity of overstorey tree height and understorey conservative traits drive aboveground biomass in a subtropical forest

The niche complementarity hypothesis has received empirical support but species differ in functional strategies for their contribution to ecosystem function, as predicted by the mass ratio hypothesis. Our understanding of how functional identity of conservative and acquisitive strategies of trees predicts aboveground biomass across forest strata (i.e. overstorey and understorey) remains unclear. Aboveground biomass, community-weighted mean (CWM − functional identity) of trait values (6 leaf and 2 stem traits), and soil physicochemical properties were estimated for 125 plots in a 5-ha subtropical forest in Eastern China. We used multiple linear regressions models to relate aboveground biomass to CWM indices at overstorey and understorey strata separately, and whole-community level. We finally employed the structural equation model to test for the effects of overstorey on understorey strata, in addition to the effects of soil physicochemical properties. Forest strata optimal models showed that overstorey strata had high aboveground biomass when they are dominated by functional identity of tree height, whereas high aboveground biomass in understorey strata was driven by functional identity of dense-wooded conservative strategy. Whole-community optimal model showed that communities dominated by functional identity of leaf dry matter content and mean leaf area had high aboveground biomass. Aboveground biomass was negatively related to soil nutrients across forest strata and whole-community level. The structural equation model showed that CWM of overstorey tree height did not affect understorey functional identity and aboveground biomass, when soil physicochemical properties were accounted. Soil nutrients had positive effect on functional identity of overstorey tree height whereas negative effect on functional identity of understorey dense-wooded strategy. This study highlights the fundamental roles of forest strata where overstorey and understorey strata contribute to their corresponding aboveground biomass with contrasting functional strategies across a range of soil nutrients. High aboveground biomass was potentially driven by functional identity of tree height through making use of plentiful soil nutrients at overstorey strata, whereas by conservative strategy at understorey strata through enduring nutrient-poor soils. To better understand the roles of functional identity of conservative and acquisitive strategies in driving ecosystem functions, it is worth to analyse forest strata separately.     

广西森林土壤主要养分的空间异质性

通过对广西区7大森林片区的11个主要森林类型样地土壤养分含量进行分析,利用地统计学方法,研究了广西区森林主要土壤养分的空间变异状况及分布格局,探讨了其相关的生态学过程,以期为广西区森林分区与林间采取不同措施管理提供科学依据。结果表明:广西区森林土壤主要养分基本属于中等变异,速效养分变异大于全量养分,其中AP、AK变异程度最大;广西森林土壤主要养分的半变异函数均表现出一定的空间结构特征,TN、TP、AP表现为中等强度的空间自相关,TK、AN、AK表现为强烈的空间自相关;不同土壤养分空间结构不同,Kriging等值线图表明广西区氮素含量比较丰富、K含量中等、P含量较少,北部片区土壤养分含量普遍大于南部片区,这可能与气候、降雨、人工种植森林树种、地形、林地管理措施等有关。