FATE‐HD: a spatially and temporally explicit integrated model for predicting vegetation structure and diversity at regional scale

During the last decade, despite strenuous efforts to develop new models and compare different approaches, few conclusions have been drawn on their ability to provide robust biodiversity projections in an environmental change context. The recurring suggestions are that models should explicitly (i) include spatiotemporal dynamics; (ii) consider multiple species in interactions and (iii) account for the processes shaping biodiversity distribution. This article presents a biodiversity model (FATE‐HD) that meets this challenge at regional scale by combining phenomenological and process‐based approaches and using well‐defined plant functional groups. FATE‐HD has been tested and validated in a French National Park, demonstrating its ability to simulate vegetation dynamics, structure and diversity in response to disturbances and climate change. The analysis demonstrated the importance of considering biotic interactions, spatio‐temporal dynamics and disturbances in addition to abiotic drivers to simulate vegetation dynamics. The distribution of pioneer trees was particularly improved, as were all undergrowth functional groups.     

The role of climate and plant functional trade‐offs in shaping global biome and biodiversity patterns

Aim Two of the oldest observations in plant geography are the increase in plant diversity from the poles towards the tropics and the global geographic distribution of vegetation physiognomy (biomes). The objective of this paper is to use a process‐based vegetation model to evaluate the relationship between modelled and observed global patterns of plant diversity and the geographic distribution of biomes. Location The global terrestrial biosphere. Methods We implemented and tested a novel vegetation model aimed at identifying strategies that enable plants to grow and reproduce within particular climatic conditions across the globe. Our model simulates plant survival according to the fundamental ecophysiological processes of water uptake, photosynthesis, reproduction and phenology. We evaluated the survival of an ensemble of 10,000 plant growth strategies across the range of global climatic conditions. For the simulated regional plant assemblages we quantified functional richness, functional diversity and functional identity. Results A strong relationship was found (correlation coefficient of 0.75) between the modelled and the observed plant diversity. Our approach demonstrates that plant functional dissimilarity increases and then saturates with increasing plant diversity. Six of the major Earth biomes were reproduced by clustering grid cells according to their functional identity (mean functional traits of a regional plant assemblage). These biome clusters were in fair agreement with two other global vegetation schemes: a satellite image classification and a biogeography model (kappa statistics around 0.4). Main conclusions Our model reproduces the observed global patterns of plant diversity and vegetation physiognomy from the number and identity of simulated plant growth strategies. These plant growth strategies emerge from the first principles of climatic constraints and plant functional trade‐offs. Our study makes important contributions to furthering the understanding of how climate affects patterns of plant diversity and vegetation physiognomy from a process‐based rather than a phenomenological perspective.     

Exploring the generality of associations between plant functional traits: evidence within ecological groups along an altitudinal gradient in Hyrcanian forest

We hypothesized that associations among plant functional traits may differ within different ecological assemblages and plant communities. Association among plant traits including plant maximum height, seed weight, fruit type, pollination mode, mean leaf area, and leaf type were explored within life forms, plant strategy groups along with lowland and montane forest vegetation. In total, 83 sampling plots of 400 m² were placed along a 2400 m altitudinal gradient in Hyrcanian forest. Importance‐values of species within vegetation types were used for weighting data and trait associations were explored using categorical principal component analysis. A G‐test and Fisher's exact test of independence were used to retest significance of the correlations. Different paired trait associations (association lines) including height–leaf, height–seed, height–pollination, leaf–seed, seed–fruit and fruit–pollination were observed and their ecological or physiological basis was discussed. Life forms, strategy types and vegetation types differed based on association lines. Some of the well‐known trade‐offs appear by increasing scale from ecological groups to vegetation types in Hyrcanian forest. The observed patterns of trait associations in Hyrcanian forest and several other ecosystems of the world call the generality of previously accepted trait correlations into question.     

陆地生态系统植物功能群研究进展

从植物功能群角度探讨陆地生态系统功能与稳定性维持机理、植物对环境变化的适应与响应以及水分、养分利用效率等成为当前生态系统生态学研究的主要技术路线.植物功能群的提出和研究,为研究复杂的生态系统提供了一个良好的方法和途径.综述了陆地生态系统功能群方面的最新研究进展,介绍了植物功能群定义的发展历程,详细比较了植物功能群划分的依据及方法,对于植物功能群与群落稳定性之间的关系、植物功能群对群落生产力的影响以及植物功能群与环境因子的动态关系等进行了深入讨论.这些研究资料表明,植物功能群整合了功能及对环境响应相似的一类植物,但植物功能特征不是绝对的、单一的,所以对植物功能群就会有不同的理解,会有不同的定义及划分方法.许多研究者从不同的角度、尺度来对植物功能群进行研究,这些研究结果有不同的针对方向和目的,使人们可以从不同的角度更全面的理解复杂的陆地生态系统.学者们在研究生态系统时,或多或少地总要与植物功能群相联系,这大大拓宽了植物功能群的应用范围.所有前人的研究使植物功能群的概念、划分、方向、应用等诸方面越来越清晰.这要求应有一个规范、统一、明确的植物功能群研究方案,这样能使对植物功能群的研究更加深入,能整合全球所有植物功能群的相关研究.     

天坑森林植物群落叶功能性状、物种多样性和功能多样性特征

以天坑内部-边缘-外部森林植物群落为研究对象,通过调查植物的群落结构、叶功能性状,探究天坑内外森林植物群落叶功能性状、物种多样性和功能多样性变化特征及其内在关联,为深入了解负地形森林生态系统的功能和恢复退化喀斯特地区的植被提供一定参考。研究结果如下：（1）比叶面积（SLA: 198.75 cm2/g)）、叶面积（LA: 42.70 cm2）、叶磷含量（LPC: 1.70 g/kg）和叶钾含量（LKC: 10.27 g/kg）在天坑内部最高,叶组织密度（LTD: 0.32 g/cm3）和叶干物质含量（LDMC: 0.41 g/g）在天坑外部最高,天坑内外森林均易受到磷限制,表明随天坑内部-边缘-外部生境变化,植物对环境的适应机制和生存策略发生了部分调整,物种的防御策略增强,生长投入策略减弱。（2）Shannon-Wiener指数（2.82）、Simpson指数（0.92）和Pielou’s均匀度指数（0.87）均以天坑外部最高,功能丰富度（1.05）、功能离散度（1.88）和Rao’s二次熵（4.52）以天坑内部最高,表明随天坑内部-边缘-外部生境的变化,植物功能性状的差异减少,物种分布及其功能性状分布总体上更为均匀、物种数量增多。（3）物种多样性指数之间、功能多样性指数之间存在较强的相关性,表明物种多样性指数之间、功能多样性指数之间存在不同的制约关系。（4）叶功能性状与物种多样性、功能多样性的相关性强,物种多样性和功能多样性之间相关性较弱,表明叶性状对生态学过程的变化较为敏感,叶功能性状与物种多样性之间存在较强的耦合关系。     

荒漠植物不同功能群性状特征及其与土壤环境的关系

研究不同功能群植物性状差异及其与土壤环境关系对于充分掌握植物的环境适应策略至关重要。以艾比湖流域为研究区,利用荒漠植物的植物高度、叶片碳、氮、磷、硫、钾、钙、钠、镁含量等9个性状,将高、低土壤水盐环境下的植物划分为5个功能群,分析不同功能群的植物组成、性状差异及其与土壤环境的关系。结果表明：(1)不同土壤水盐环境下,其植物功能群组成不同;其中白刺、胡杨和罗布麻在两个土壤水盐环境下的功能群中均存在。(2)植物的功能性状在不同土壤水盐环境下也发生了适应性的变化。高土壤水盐环境下3个功能群的植物高度、叶片碳、氮、磷和钙含量显著高于低土壤水盐环境功能群(P<0.05);低土壤水盐环境下2个功能群的植物叶片硫、钠和镁含量高于高土壤水盐环境功能群。(3)土壤含水量(SVWC)、电导率(EC)、pH以及土壤磷含量对荒漠植物功能性状影响较大。在高土壤水盐环境下,EC、pH与植物高度,叶片钾、钙含量正相关,与叶片硫含量负相关;在低土壤水盐环境下,SVWC、EC与植物高度呈显著正相关(P<0.05)。研究有助于理解荒漠植物对极端环境的适应对策,为保护荒漠地区生物多样性提供理论依据。     

松嫩草地4种植物功能群土壤微生物碳源利用的差异

为了探讨草地不同植物功能群土壤微生物碳源利用差异,利用Biolog-ECO微平板检测法,研究了松嫩放牧草地禾草(Grass)、羊草(Leymus chinensis)、杂类草(Forb)和豆科牧草(Legume)4种植物功能群土壤微生物碳源代谢的多样性变化特征。结果表明,在培养的240h内,4种不同植物功能群的土壤微生物对碳源的利用程度均随着时间的延长而升高,表明微生物代谢活性随着时间而增强;不同植物功能群土壤微生物总体活性(AWCD)(P0.001)、Shannon-Wiener指数(H)(P0.001)、物种丰富度(R)(P=0.005)、Pielou均匀度指数(E)(P0.001)差异显著;其中禾草的各项指数明显高于其他3种(P0.01),杂类草的各指标均最低。禾草、羊草对糖类、氨基酸类、羧酸类、多聚物类有较好的利用,豆科牧草除羧酸类外对其他碳源都有更好的利用,杂类草只对酚类碳源利用率最高,而对其他碳源利用率极低。总体得出4种功能群土壤微生物的碳源利用率顺序为:禾草羊草豆科牧草杂类草。     

基于能量属性的植物功能群划分方法探索——以内蒙古锡林河流域草原植物群落为例

植物功能群(plant functional groups, PFGs) 是具有确定的植物功能特征的一系列植物的组合,是生态学家为研究植被对气候变化和干扰的响应而引入的生态学概念.目前功能群研究中最核心的问题仍在于决定植物功能群划分的植物特征的选择上.以内蒙古锡林河流域草原植物群落为例,选取3个草原类型(羊草草原、大针茅草原和羊草草甸草原)及其退化梯度系列(未退化、轻度退化、中度退化、重度退化),在对植物热值进行分析测定的基础上,依据植物的能量属性-单位重量干物质在完全燃烧后所释放出来的热量值,采用人为分段的方法对草原植物进行了能量功能群的划分(高能值植物功能群、中能值植物功能群和低能值植物功能群).并探讨了这种能量功能群划分方法在草原植被动态研究中的客观性与可行性.     

Environmental filtering predicts plant‐community trait distribution and diversity: Kettle holes as models of meta‐community systems

• Meta‐communities of habitat islands may be essential to maintain biodiversity in anthropogenic landscapes allowing rescue effects in local habitat patches. To understand the species‐assembly mechanisms and dynamics of such ecosystems, it is important to test how local plant‐community diversity and composition is affected by spatial isolation and hence by dispersal limitation and local environmental conditions acting as filters for local species sorting.We used a system of 46 small wetlands (kettle holes)—natural small‐scale freshwater habitats rarely considered in nature conservation policies—embedded in an intensively managed agricultural matrix in northern Germany. We compared two types of kettle holes with distinct topographies (flat‐sloped, ephemeral, frequently plowed kettle holes vs. steep‐sloped, more permanent ones) and determined 254 vascular plant species within these ecosystems, as well as plant functional traits and nearest neighbor distances to other kettle holes.Differences in alpha and beta diversity between steep permanent compared with ephemeral flat kettle holes were mainly explained by species sorting and niche processes and mass effect processes in ephemeral flat kettle holes. The plant‐community composition as well as the community trait distribution in terms of life span, breeding system, dispersal ability, and longevity of seed banks significantly differed between the two habitat types. Flat ephemeral kettle holes held a higher percentage of non‐perennial plants with a more persistent seed bank, less obligate outbreeders and more species with seed dispersal abilities via animal vectors compared with steep‐sloped, more permanent kettle holes that had a higher percentage of wind‐dispersed species. In the flat kettle holes, plant‐species richness was negatively correlated with the degree of isolation, whereas no such pattern was found for the permanent kettle holes.Synthesis: Environment acts as filter shaping plant diversity (alpha and beta) and plant‐community trait distribution between steep permanent compared with ephemeral flat kettle holes supporting species sorting and niche mechanisms as expected, but we identified a mass effect in ephemeral kettle holes only. Flat ephemeral kettle holes can be regarded as meta‐ecosystems that strongly depend on seed dispersal and recruitment from a seed bank, whereas neighboring permanent kettle holes have a more stable local species diversity.

     

喀斯特森林不同演替阶段植物群落物种多样性、功能性状、化学计量及其关联

探讨森林植物群落物种多样性、功能性状与化学计量特征及其内在关联,有助于构建结构-性状-功能的研究脉络,深入理解生态系统的过程与功能,为群落结构优化配置和调控提供科学依据。以贵州喀斯特高原峡谷区森林草灌、灌木、灌乔和乔木4个演替阶段的植物群落为对象,通过群落学调查和植物功能性状、生态化学计量测度,研究其物种多样性、功能性状和化学计量特征及其互作关系。结果表明:(1)乔木阶段的Shannon-Wiener指数与Pielou均匀度指数均为最高,依次达8.62、2.41,表明伴随着群落演替,物种分布的均匀程度增加。(2)植物群落物种多样性指数之间仅Margalef丰富度指数与Simpson优势度指数未呈显著相关(-0.644),其余均为显著或极显著相关关系,表明物种多样性指数之间存在较强的促进或抑制作用。(3)叶片厚度与其余指标,以及群落物种多样性指数与功能性状、化学计量之间均未呈显著相关,比叶面积(SLA)仅与土壤N∶P呈显著正相关(0.742),表明群落物种多样性、功能性状和生态化学计量特征之间的相关性较弱。(4)SLA为灌木阶段的272.13 cm2/g为最大,δ13C值在演替后期略高,草灌、灌木阶段倾向于受N、P共同限制,说明随演替进展,植物的适应策略和资源利用对策等均发生了一定的调整,表明植物和环境之间存在较强的协变关系。(5)该区森林培育时,应构建完整的乔木、灌木和草本层片,提高生态系统对养分矿化、转化和循环等的自调控能力。     

Seed germination traits of two plant functional groups in the saline deltaic ecosystems

Aims Salt stress resulting from soil salinization is one of the driving forces of the land degradation throughout the world. The modern Yellow River delta is one of the most saline areas in China. Phytoremediation can be an effective way to restore the salinized ecosystems, which requires selecting appropriate plant species. This study explored the germination responses of common plant species from contrasting habitats in the Yellow River delta to varying salinity, offering experimental information for ecosystem restoration in the Yellow River delta.Methods In this study, 15 common plant species from the Yellow River delta were divided into two groups (high-salinity and low-salinity groups) by their natural habitats using Canonical Correlation Analysis. Seeds of each species were treated with five salinity levels (0, 5, 10, 20 and 30 ppt), using a randomized complete block design, and germinated seeds were counted and removed daily for 28 days to calculate the final germination proportion and mean time to germination. The germination responses of seeds to salinity treatments were compared between the two groups.Important findings In relation to salinity, seed germination behavior of the test species was closely related to the salinity level of the habitats over which they were distributed. Species from the habitats with higher salinity had generally higher final germination proportion but shorter mean time to germination than those from the habitats with lower salinity in all of five salinity treatments used. The final germination proportion and mean time to germination of low-salinity group species were more sensitive to salinity than those of high-salinity group species. Selecting the species with high final germination proportion and short mean time to germination is important for restoration of salinized land.     

Plant chemical traits define functional and phylogenetic axes of plant biodiversity

To determine which types of plant traits might better explain ecosystem functioning and plant evolutionary histories, we compiled 42 traits for each of 15 perennial species in a biodiversity experiment. We used every possible combination of three traits to cluster species. Across these 11,480 combinations, clusters generated using tissue %Ca, %N and %K best mapped onto phylogeny. Moreover, for the 15 best combinations of three traits, 82% of traits were chemical, 16% morphological and 2% metabolic. The diversity-dependence of ecosystem productivity was better explained by the %Ca, %N and %K clusters: compared to adding a new species at random, adding a species from an absent cluster/clade better-explained gains in productivity. Species number impacted productivity only when all clusters were present. Our results suggest that tissue elemental chemistry might be more phylogenetically conserved and more strongly related to ecosystem functioning than commonly measured morphological and physiological traits, a possibility that merits exploration.     

嘉峪关草湖湿地植物功能群组成及其性状对不同生境的响应

植物功能群与生境的关系是生态学研究的重要内容,对于揭示植物的环境适应机制、群落构建机制、预测植被变化等具有重要意义。为研究局域尺度上干旱区湿地、沙漠、戈壁等生态系统功能群的环境适应机制,在嘉峪关草湖国家湿地公园内,基于植物功能性状数据,研究了荒漠区不同生境条件下的功能群组成和功能性状特性。结果表明:随着生境条件的改变,土壤含水量、含盐量、容重、pH值等土壤性质发生显著变化;植物的功能群组成和功能性状会发生适应性的变化,沼泽湿地生境内植被的功能性状以克隆、C₄光合途径、根茎型等特征为主,盐沼湿地以叶片被蜡、叶茎肉质、CS策略等特征为主,沙丘生境以叶片具毛、风播、具增厚或增长的根、一年生、CR策略等特征为主,荒漠戈壁以叶片少、具增厚或增长的根、虫媒、地上芽等特征为主。局域尺度上,生境异质性尤其是土壤条件的异质性是影响植物群落组成和植物功能性状变化的重要环境因子。     

延河流域森林草原区不同植物功能型适应策略及功能型物种数量随退耕年限的变化

不同退耕年限退耕地的环境差异以及不同生物间的相互作用导致各阶段植物功能型物种数量不同。研究退耕地植被自然恢复过程中不同植物功能型适应策略及功能型物种数量随退耕年限的变化,对于理解植物对环境的响应机制及植物的适应策略具有重要意义。采用空间序列代替时间序列的方法,以延河流域森林草原区不同退耕年限、自然恢复的植物群落为研究对象,调查了不同退耕年限的植物群落33个,共44种植物,涉及16个科35个属,分别测定了每个物种的叶厚度、比叶面积、叶组织密度、叶片氮含量、比根长、根组织密度、细根氮含量等7项能够反映植物生存对策且易于测量的功能性状。依据这7项植物功能性状,采用数量分类方法将全部物种划分为3个功能型。结果表明:(1)根据C-S-R理论,功能型Ⅰ植物用于防御的投资较多,生长速率处于中间水平,偏向于"胁迫-干扰型",功能型Ⅱ植物能够通过维持体内的养分平衡的方式对抗资源贫瘠或干旱的环境,偏向于"胁迫-竞争型"对策,而功能型Ⅲ植物吸收大量的营养和资源用于生长,偏向于"竞争型";(2)功能型Ⅰ在整个植被恢复时间序列中占据优势地位(61%—80%),并呈增加趋势,功能型Ⅱ则由恢复初期的25%降低为恢复后期的15%,功能型Ⅲ从恢复初期的14%降低到恢复后期的5%。同时,在功能型Ⅰ内部,优势物种也发生着相应的相互替代。虽然土壤养分含量整体上随着植被自然恢复时间的延长而呈上升趋势,但是植物的生存环境并未改善到不存在干扰与胁迫的程度。因此,在植被恢复初期的四、五十年内,"胁迫-干扰型"策略的植物占据着绝对优势。随着植被恢复时间的延长,能够高效利用资源且抗胁迫能力强的物种代替了以快速生长和传播为适应策略的物种。     

鼎湖山森林群落多样性垂直分布格局的研究

根据鼎湖山10 m×1 200 m的植物固定样带调查,沿海拔梯度对植物群落分布格局及生物多样性特性进行了分析。结果表明,本固定样带有3个森林类型,5个群落类型,构成一演替序列,但在垂直梯度上有交错现象。生物多样性指数为:乔木层,中生性阔叶林>针阔混交林>阳生性阔叶林;草本层,针阔混交林>阔叶林。阳性草本植物为主的草本群大部分生长在阳性树种为主的阔叶林和针阔混交林下,但是林下微环境变化对草本层植物优势度的影响也较大,以耐荫植物为主的草本群在阳性树种为主的阔叶林和针阔混交林也有分布,表明相同群落类型林下小环境的变化。     

A global distribution of biodiversity inferred from climatic constraints: results from a process‐based modelling study

We investigated the connection between plant species diversity and climate by using a process‐based, generic plant model. Different ‘species' were simulated by different values for certain growth‐related model parameters. Subsequently, a wide range of values were tested in the framework of a ‘Monte Carlo' simulation for success; that is, the capability of each plant with these parameter combinations to reproduce itself during its lifetime. The range of successful parameter combinations approximated species diversity. This method was applied to a global grid, using daily atmospheric forcing from a climate model simulation. The computed distribution of plant ‘species' diversity compares very well with the observed, global‐scale distribution of species diversity, reproducing the majority of ‘hot spot' areas of biodiversity. A sensitivity analysis revealed that the predicted pattern is very robust against changes of fixed model parameters. Analysis of the climatic forcing and of two additional sensitivity simulations demonstrated that the crucial factor leading to this distribution of diversity is the early stage of a plant's life when water availability is highly coupled to the variability in precipitation because in this stage root‐zone storage of water is small. We used cluster analysis in order to extract common sets of species parameters, mean plant properties and biogeographic regions (biomes) from the model output. The successful ‘species' cannot be grouped into typical parameter combinations, which define the plant's functioning. However, the mean simulated plant properties, such as lifetime and growth, can be grouped into a few characteristic plant ‘prototypes', ranging from short‐lived, fast growing plants, similar to grasses, to long‐lived, slow growing plants, similar to trees. The classification of regions with respect to similar combinations of successful ‘species' yields a distribution of biomes similar to the observed distribution. Each biome has typical levels of climatic constraints, expressed for instance by the number of ‘rainy days' and ‘warm days'. The less the number of days favourable for growth, the greater the level of constraints and the less the ‘species' diversity. These results suggest that climate as a fundamental constraint can explain much of the global scale, observed distribution of plant species diversity.