Pathways, mechanisms and predictability of vegetation change during tropical dry forest succession

The development of forest succession theory has been based on studies in temperate and tropical wet forests. As rates and pathways of succession vary with the environment, advances in successional theory and study approaches are challenged by controversies derived from such variation and by the scarcity of studies in other ecosystems. During five years, we studied development pathways and dynamics in a chronosequence spanning from very early to late successional stages (ca. 1–60 years) in a tropical dry forest of Mexico. We (1) contrasted dynamic pathways of change in structure, diversity, and species composition with static, chronosequence-based trends, (2) examined how structure and successional dynamics of guilds of trees shape community change, and (3) assessed the predictability of succession in this system. Forest diversity and structure increased with time but tree density stabilized early in succession. Dynamic pathways matched chronosequence trends. Succession consisted of two tree-dominated phases characterized by the development and dynamics of a pioneer and a mature forest species guild, respectively. Pioneer species dominated early recruitment (until ca. 10 years after abandonment), and declined before slower growing mature-forest species became dominant or reached maximum development rates (after 40–45 years). Pioneers promoted their replacement early in succession, while mature-forest species recruited and grew constantly throughout the process, with their lowest mortality coinciding with the peak of pioneer abundance. In contrast to prevailing stochastic views, we observed an orderly, community driven series of changes in this dry forest secondary succession. Chronosequences thus represent a valuable approach for revealing system-specific successional pathways, formulating hypotheses on causes and mechanisms and, in combination with repeated sampling, evaluating the effects of vegetation dynamics in pathway variation.     

Relationships between functional diversity and ecosystem functioning: A review

Functional diversity, which is the value, variation and distribution of traits in a community assembly, is an important component of biodiversity. Functional diversity is generally viewed as a key to understand ecosystem and community functioning. There are three components of functional diversity, i.e. functional richness, evenness and divergence. Functional diversity and species diversity can be either positively or negatively correlated, or uncorrelated, depending on the environmental conditions and disturbance intensity. Ecosystem functioning includes ecosystem processes, ecosystem properties and ecosystem stability. The diversity hypothesis and the mass ratio hypothesis are the two major hypotheses of explaining the effect of functional diversity on ecosystem functioning, diversity hypothesis reflects that organisms and their functional traits in a assemblage effect on ecosystem functioning by the complementarity of using resources, and mass ratio hypothesis emphasises the identify of the dominant species in a assemblage. These two hypotheses do not contradict each other and instead they reflect the two different sides of functional diversity and functional composition. The effect of functional diversity on ecosystem functioning also depends on abiotic factors, perturbation, management actions, etc. Function diversity potentially influences ecosystem service and management by effecting on ecosystem functioning. Ecosystem management groups should include functional diversity in their scheme and not just species richness.     

海南岛热带低地雨林群落水平植物功能性状与环境因子相关性随演替阶段的变化

以功能性状为基础的方法可以用来探讨植物群落中功能性状如何响应环境并揭示植物生态策略的潜在驱动力,但有关功能性状与环境因子之间的关系随植物群落演替变化的研究仍然匮乏.作者以海南岛热带低地雨林刀耕火种弃耕后处于不同演替阶段的次生林(包括弃耕后恢复15年、30年及60年的次生林)和老龄林为对象,通过群落学调查和对木本植物的功能性状及样地环境因子的测定,分析了群落水平植物功能性状与环境因子关系随演替阶段的变化规律.结果表明,随着演替的进行,林冠开阔度、土壤养分、比叶面积、叶片氮含量、叶片磷含量和叶片总有机碳含量逐渐降低,叶片干物质含量、木材密度、潜在最大高度逐渐升高,而土壤水分和叶片钾含量变化不大.多元逐步回归分析表明,影响群落水平植物功能性状的主要环境因子随演替阶段而发生显著的变化,在15年、30年和60年的次生林及老龄林中,对群落水平植物功能性状影响最大的环境因子依次为土壤有机质和pH值、林冠开阔度和土壤总磷含量、土壤总钾和有效磷含量,以及土壤有机质含量和磷含量.同一功能性状在不同演替阶段受到不同环境因子的控制,同时各功能性状又能够对不同演替阶段所处的特殊环境产生一定的适应性.     

Microhabitat variations and seed bank-vegetation relationships in a desert wadi ecosystem

Due to extreme variability in patterns of rainfall, plant seed banks are an important component of desert habitats. Here I report on effects of standing vegetation and three different microhabitats (channel, bank and terrace) on the soil seed bank of a desert wadi ecosystem in the Eastern Desert of Egypt. A total of 450 soil samples at 45 stands were collected to represent the different wadi microhabitats. The germinable seed bank was estimated by controlled counts of seedling emergence. The floristic composition, functional properties and diversity of the soil seed bank, as well as its similarity with the standing vegetation varied among wadi microhabitats. Such variation could be attributed to differences in disturbance intensity among microhabitats (terrace < bank < channel) and variation of soil factors along the microtopographic gradient. Channel showed the highest species richness and size of soil seed bank, followed by bank and then terrace. Moreover the Shannon index of diversity of the seed bank and its similarity with standing vegetation were significantly greater in both channel and bank microhabitats than in terrace. At the level of plant functional groups, number of seeds of annuals was higher in both channel and bank than in terrace. Shrubs were more abundant in seed banks of channel compared to terrace. The size and species richness of seed bank were increased with the total plant cover, annual/perennial ratio and species richness of the standing vegetation.     

A new graphical model for untangling complex relationships among environment, biodiversity, and ecosystem functioning

Recent advances in the research field of ‘biodiversity-ecosystem functioning’ have successfully begun to reconcile the apparent controversy on relationships between productivity and species richness. By unifying new advances into a single framework, I propose a 3D graphical model connecting the relationships among resource availability, species richness, and ‘community productivity.’ An emergent pattern from this model predicts that the effect of species richness on community productivity is maximized at intermediate levels of resource availability. This model will contribute to better understanding the relationships among environment, biodiversity, and ecosystem functioning.     

Vegetation patterns,regeneration rates and divergence in an old-field succession of the high tropical Andes

Vegetation restoration during old-field succession was studied in an alpine Andean ecosystem (paramo). 123 plots with different fallow times (1 to 12 years) and 8 plots under natural vegetation were sampled. The results indicate that secondary succession in the paramo, like in other extreme environments, can be interpreted as an autosuccession: there are mainly changes in species relative abundance and little floristic relay (i.e. species turnover). Only a few herbaceous species, mostly introduced (e.g. Rumex acetosella), act as strict pioneers and strongly dominate the early stages. Then, they undergo a progressive decline, while native forbs (e.g. Lupinus meridanus) and grasses (e.g. Vulpia myuros) have their peak abundance in intermediate stages. The characteristic paramo life forms, sclerophilous shrubs (e.g. Baccharis prunifolia, Hypericum laricifolium) and giant rosettes (e.g. Espeletia schultzii), appear very early and gradually increase in abundance during succession, becoming dominant in the late stages and showing a dual behaviour, both as ruderal and stress tolerant species. The 1st axis of a Detrended Correspondence Analysis arranges the sites according to their fallow time. The 2nd and 3rd axes, associated with diverging pathways of regeneration, are correlated with topographic factors and physio-chemical soil characteristics. Hence, structural divergence between plots increases along succession as community composition starts to reflect the conditions of each site. We found evidence of a constant rate of succession during the first 12 years, contradicting the generally accepted hypothesis in the succession literature of a continous slow down up to the climax. Regeneration of vegetation physiognomy is relatively fast, questioning the prevailing idea of slow restoration in alpine ecosystems. However, 12 years of fallow are insufficient to attain the species richness of the natural paramo. Under the current trend of fallow length reduction observed in traditional potato cultivation in the Andes, our results raise doubts about the conservationist value of this management strategy.     

Weak relationships between leaf phenology and isohydric and anisohydric behavior in lowland wet tropical forest trees

Wet tropical forest trees display a wide range of leaf phenology dynamics. However, the interrelation between deciduousness, water status, and leaf and stem characteristics have been poorly investigated compared with dry forests. We studied wet forest trees to answer the following questions: (1) do water regulation modes (iso/anisohydric behavior) of evergreen species differ from those found in deciduous species? (2) Does leaf water potential (Ψ_L) influences leaffall and emergence dynamics? (3) Are leaf and stem characteristics consistent across evergreen and deciduous trees? We evaluated vegetative phenology, Ψ_L, and leaf and stem characteristics of six evergreen and three deciduous species monthly for 2 yr. Species exhibited different leaffall and emergence dynamics, as well as different water regulation modes, independent of their deciduousness. Thus, the relationship between leaf phenology and water regulation behaviors appears to be a species‐specific property rather than a functional group attribute. Ψ_L had no direct influence on the dynamics of leaffall and/or emergence, indicating that this process is not modulated by water availability alone. Individual groups of evergreen and deciduous species could not be identified using principal component analysis (PCA), but a decoupling was observed in the leaf and stem economics spectra. The lack of an interrelation between deciduousness and iso/anisohydry, as well as the independence of leaf and stem trade‐offs, emphasizes that more systematic measurements of vegetative phenology and ecophysiological characteristics are necessary to advance our knowledge of leaf habit and water regulation behaviors based on the functional traits of wet forest plants.     

植物遗传多样性与生态系统功能关系的研究进展

全球变化和人类活动导致物种生境的萎缩, 造成很多植物种群数量缩减, 遗传多样性快速丧失。对于物种多样性低的生态系统, 优势种的遗传多样性可能比物种多样性对生态系统功能产生更大的影响。因此, 了解遗传多样性和生态系统功能的关系(GD-EF)及其机制对生物多样性保护、应对环境变化和生态修复具有指导意义。该文综述了植物遗传多样性对生态系统结构(高营养级生物群落结构)和生态系统功能(初级生产力、养分循环和稳定性)的影响及机制、功能多样性对GD-EF的影响、遗传多样性效应和物种多样性效应的比较, 以及GD-EF在生态修复等实际应用的研究进展。最后指出当前研究的不足之处, 以期为后续研究提供参考: 1)还需深入研究GD-EF机制; 2)未评估遗传多样性对生态系统多功能性的影响; 3)不同遗传多样性测度对生态系统功能的影响不明确; 4)缺少长期的和多空间尺度结合的GD-EF实验; 5)遗传多样性效应相对于其他因子的作用不清楚。     

The relationship between tree biodiversity and biomass dynamics changes with tropical forest succession

Theory predicts shifts in the magnitude and direction of biodiversity effects on ecosystem function (BEF) over succession, but this theory remains largely untested. We studied the relationship between aboveground tree biomass dynamics (Δbiomass) and multiple dimensions of biodiversity over 8–16 years in eight successional rainforests. We tested whether successional changes in diversity–Δbiomass correlations reflect predictions of niche theories. Diversity–Δbiomass correlations were positive early but weak later in succession, suggesting saturation of niche space with increasing diversity. Early in succession, phylogenetic diversity and functional diversity in two leaf traits exhibited the strongest positive correlations with Δbiomass, indicating complementarity or positive selection effects. In mid‐successional stands, high biodiversity was associated with greater mortality‐driven biomass loss, i.e. negative selection effects, suggesting successional niche trade‐offs and loss of fast‐growing pioneer species. Our results demonstrate that BEF relationships are dynamic across succession, thus successional context is essential to understanding BEF in a given system.     

植物群落多样性与可入侵性关系研究进展

综述了植物群落的多样性和可入侵性关系的研究进展.对模型研究、观察性研究和实验性研究得出植物群落的多样性和可入侵性关系不同结论的原因进行了分析,并提出了该方面进一步研究的建议.模型研究的群落多样性形成机制单一、群落状态达到平衡、物种特征差异不明显等简化前提假设,削弱了其结论的可信性.观察性研究的尺度不同导致不同结论,在小尺度上的主要影响因素可能是群落内部的生物因素,但在较大尺度上很多外部因素共同影响多样性和可入侵性,会导致两者出现正相关.实验性研究通常是在小尺度下进行,排除了外部因素的干扰,但是在人工组建群落多样性梯度时,会出现取样效应、相同物种多样性的群落丰富度和密度不同等问题.未来研究应重点考虑的几个方面,即观察性研究与实验相结合,分析不同尺度上入侵的机理和过程;平衡实验设计,避免抽样效应等误来源;模型与实验设计相耦合.     

Spatial covariance between ecosystem services and biodiversity pattern at a national scale (France)

We present at a national-scale estimates of spatial covariance between areas important for ecosystem services (ES) and biodiversity, at a fixed spatial scale in France. We calculated different diversity and community metrics for common bird communities: taxonomic diversity (TD), functional diversity (FD), community specialization index (CSI), trophic index (TI), phylogenetic diversity (PD) and community evolutionary distinctiveness (CED). The ES multifunctionality (ES_MF) was measured using a combination of ES evenness and overall ES estimator. Spatially explicit tests were used to compare the spatial patterns of ES and diversity metrics. Mixed models were used for comparisons.We found low spatial congruence between ES and both diversity and community metrics in France. However, we detected even opposite associations between ES and each biodiversity component. Crop production was negatively associated with CED, it was positively correlated with CSI. No positive associations were found between ES_MF and any diversity metric, independently of farming systems. The only significant association was negative: lower values of CED were associated with hotspots of ES_MF. We found also a negative effect of crop production on bird CED. The conservation implication is remarkable, because conservation policies focusing solely on the economic value of ES will fail to protect overall biodiversity.     

Vegetation differentiation and secondary succession on a limestone hill in southern Poland

Abstract. The vegetation of the SkoHzanka Reserve, situated on a limestone hill near Krakow, was examined over a 30 year period, using the Braun-Blanquet method. With the help of a numerical classification eight types of grassland communities were distinguished, belonging to the classes Sedo-Scleranthetea, Nardo-Callunetea and Festuco-Brometea, and 11 types of woodland communities, both remnants of ancient woodlands from the classes Querco-Fagetea and Vaccinio-Piceetea and more recent woods. In the case of grasslands the results of detrended correspondence analysis (DCA) indicated that the largest variation was associated with the soil type. Woods on sandy soils are very different from those on calcareous soils, and the younger recent woods are very different from the ancient woodlands. An analysis of historical cartographic material and aerial photographs shows that in the last few decades the area of recent woods has increased considerably at the expense of grasslands. Changes in species diversity in the woods during secondary succession and their successional convergence were examined and discussed. It was found that young and mature woods on sandy soils are more similar to each other than young and mature woods on calcareous soils. These differences are mainly related to the fact that many woodland herb species growing in rich, broadleaved ancient woods are unable to colonize recent woods since they have a limited colonization capacity.     

Functional diversity and traits assembly patterns of lichens as indicators of successional stages in a tropical rainforest

Understanding community assembly patterns with regard to functional traits, which may be common to different species, allows us to compare ecological communities in a wider range of environmental and phytogeographic conditions. Moreover, a functional approach may facilitate the comprehension of the relation between biotic changes and ecosystem functions in complex systems such as tropical forests. Considering the lack of information in relation to the influence of environmental conditions on lichen functional traits, on lichen functional patterns in tropical forests and the potential usage of this approach in bioindication studies, this paper aims at (i) determining the functional structure of the lichen communities along the forest succession gradient, (ii) assessing the relation of lichen functional traits to this gradient and (iii) verifying the potential of using lichen functional traits as indicators of successional stages in tropical rainforests. Lichens were sampled in 24 sampling units of three successional stages (6–10; 12–20; and 40–60 years of recovering). The results corroborated our main hypothesis that the functional structure of the lichen communities responds to structural changes along a forest regeneration gradient. Growth forms (foliose, fruticose and crustose) as well as some reproductive trait states (lirella, isidia and perithecia) were the most suitable lichen traits to be used as indicators of forest succession in the southern edge of the Atlantic Rainforest. Lirella, fruticose and foliose were more related to earlier successional stages; isidia was more related to intermediate stages, while perithecia and crustose were more related to older stages. These results reinforce the ability of lichens to be used as bioindicators of forest conditions.     

Anthropogenic impacts on tropical forest biodiversity: a network structure and ecosystem functioning perspective

Huge areas of diverse tropical forest are lost or degraded every year with dramatic consequences for biodiversity. Deforestation and fragmentation, over-exploitation, invasive species and climate change are the main drivers of tropical forest biodiversity loss. Most studies investigating these threats have focused on changes in species richness or species diversity. However, if we are to understand the absolute and long-term effects of anthropogenic impacts on tropical forests, we should also consider the interactions between species, how those species are organized in networks, and the function that those species perform. I discuss our current knowledge of network structure and ecosystem functioning, highlighting empirical examples of their response to anthropogenic impacts. I consider the future prospects for tropical forest biodiversity, focusing on biodiversity and ecosystem functioning in secondary forest. Finally, I propose directions for future research to help us better understand the effects of anthropogenic impacts on tropical forest biodiversity.     

The relationship between biodiversity and ecosystem functioning in food webs

Recent theoretical and experimental work provides clear evidence that biodiversity loss can have profound impacts on functioning of natural and managed ecosystems and the ability of ecosystems to deliver ecological services to human societies. Work on simplified ecosystems in which the diversity of a single trophic level is manipulated shows that diversity can enhance ecosystem processes such as primary productivity and nutrient retention. Theory also strongly suggests that biodiversity can act as biological insurance against potential disruptions caused by environmental changes. However, these studies generally concern a single trophic level, primary producers for the most part. Changes in biodiversity also affect ecosystem functioning through trophic interactions. Here we review, through the analysis of a simple ecosystem model, several key aspects inherent in multitrophic systems that may strongly affect the relationship between diversity and ecosystem processes. Our analysis shows that trophic interactions have a strong impact on the relationships between diversity and ecosystem functioning, whether the ecosystem property considered is total biomass or temporal variability of biomass at the various trophic levels. In both cases, food-web structure and trade-offs that affect interaction strength have major effects on these relationships. Multitrophic interactions are expected to make biodiversity–ecosystem functioning relationships more complex and non-linear, in contrast to the monotonic changes predicted for simplified systems with a single trophic level.     

Structure and floristics of secondary and old-growth forest stands in lowland Costa Rica

We characterized stand structure and floristic composition of woody life forms in three, 16–18 yr old secondary stands that regenerated after pasture abandonment, and three nearby old-growth stands of tropical rain forest in lowland Costa Rica. Basal area and stem density for each of four plant size classes (seedlings, saplings, treelets, trees) were similar among stand types, but density of adult canopy palms (individuals 10 cm DBH), was lower in the secondary stands. We estimate that 15% of the basal area of stems 10 cm DBH correspond to remnant trees in our secondary stands. The observed rapid woody regrowth compared to other published studies in the lowland neotropics, can be attributed to moderate land use and possibly, to the influence of nutrient-rich volcanic soils in the study area. Overall, plant species richness was lower in the secondary stands, but this difference was less pronounced in the smallest size classes (seedlings, saplings). Median percent similarity of all pairwise stand comparisons showed that floristic composition of saplings (stems 1 m tall and 5 cm DBH) was more similar between secondary and old-growth stands than for trees (stems 10 cm DBH). Because the potential value of secondary forests in conserving woody plant diversity appears highest for the young size classes, we suggest that further studies on floristic composition, especially those addressing the dynamics of the understory component, are needed to refine our understanding of the role of this natural resource in the maintenance of plant biodiversity in disturbed landscapes.     

Fire survival of lowland tropical rain forest trees in relation to stem diameter and topographic position

The objective of this study was to relate patterns in forest structure, tree species diversity, and tree species composition to stem diameters and topography in unburned, once burned and twice burned lowland dipterocarp rain forests in East Kalimantan, Indonesia. To do this four unburned old growth forests were compared with three forests that burned once (1997/1998) and three forests that burned twice (1982/1983 and 1997/1998). Fire resulted in a strong reduction of climax tree density which was negatively related to tree diameter. However, a disproportionate reduction in small diameter understorey climax tree species occurred only after repeated fires. Climax tree species in both burned forest types were most common in swamps, river valleys and on lower slopes, while their density was much lower on places higher along hillsides. In unburned forest the opposite was observed, with climax tree density increasing steadily from swamp and river valleys to upper slopes and ridges. In contrast to climax trees, pioneer trees were abundant throughout the burned forest, with highest numbers on hill sides and ridges. Our results indicate that both diameter and topographic position of trees strongly affect their fire survival chances in tropical lowland forests.