Plant trait responses to grazing – a global synthesis

Herbivory by domestic and wild ungulates is a major driver of global vegetation dynamics. However, grazing is not considered in dynamic global vegetation models, or more generally in studies of the effects of environmental change on ecosystems at regional to global scale. An obstacle to this is a lack of empirical tests of several hypotheses linking plant traits with grazing. We, therefore, set out to test whether some widely recognized trait responses to grazing are consistent at the global level. We conducted a meta‐analysis of plant trait responses to grazing, based on 197 studies from all major regions of the world, and using six major conceptual models of trait response to grazing as a framework. Data were available for seven plant traits: life history, canopy height, habit, architecture, growth form (forb, graminoid, herbaceous legume, woody), palatability, and geographic origin. Covariates were precipitation and evolutionary history of herbivory. Overall, grazing favoured annual over perennial plants, short plants over tall plants, prostrate over erect plants, and stoloniferous and rosette architecture over tussock architecture. There was no consistent effect of grazing on growth form. Some response patterns were modified by particular combinations of precipitation and history of herbivory. Climatic and historical contexts are therefore essential for understanding plant trait responses to grazing. Our study identifies some key traits to be incorporated into plant functional classifications for the explicit consideration of grazing into global vegetation models used in global change research. Importantly, our results suggest that plant functional type classifications and response rules need to be specific to regions with different climate and herbivory history.     

植物功能性状与环境和生态系统功能

植物性状反映了植物对生长环境的响应和适应,将环境、植物个体和生态系统结构、过程与功能联系起来(所谓的“植物功能性状”)。该文介绍了植物功能性状的分类体系,综述了国内外植物功能性状与气候(包括气温、降水、光照)、地理空间变异(包括地形地貌、生态梯度、海拔)、营养、干扰(包括火灾、放牧、生物入侵、土地利用)等环境因素,以及与生态系统功能之间关系的研究进展,探讨了全球变化(气候变化和CO₂浓度升高)对个体和群落植物功能性状的影响。植物功能性状的研究已经取得很多成果,并应用于全球变化、古植被恢复和古气候定量重建、环境监测与评价、生态保护和恢复等研究中,但大尺度、多生境因子下的植物功能性状研究仍有待于加强,同时需要改进性状的测量手段;我国的植物功能性状研究还需要更加明朗化和系统化。     

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.     

退化高寒草地植物多样性和功能群组成与地上生产力的关系

近半个世纪以来, 气候变化和过度放牧等因素导致全球约50%的天然草地发生不同程度退化。草地退化显著改变了植物多样性和群落组成, 进而影响生态系统生产力。然而, 以往关于退化草地多样性与功能群组成对生产力的影响研究主要来自单点尺度, 仍缺乏大尺度的观测证据。为此, 该研究以青藏高原三江源退化高寒草地为研究对象, 基于不同类型草地(高寒草原、高寒草甸和高寒沼泽化草甸) 15个样点45条退化序列的标准化调查和采样, 解析了高寒草地退化过程中植物多样性与功能群组成的变化及其与地上生产力的关联。结果显示, 随退化程度的加剧, 3种类型草地的物种丰富度、Shannon-Weiner多样性指数、Simpson多样性指数和Pielou均匀度指数整体上均呈“先上升后下降”的变化趋势; 整体而言, 群落中莎草科和禾本科植物盖度沿退化梯度显著降低, 豆科植物盖度没有显著变化, 而杂类草盖度均显著增加。进一步分析发现, 退化过程中所有草地类型地上生产力的下降均主要与原生优势种盖度的降低有关, 而植物多样性对生产力的影响较弱。以上结果表明, 从植被的角度而言, 原生群落中优势种盖度的减少而不是多样性的丧失是退化高寒草地植被生产力下降的主要原因, 这一发现也为退化高寒草地恢复提供重要启示: 植被恢复过程中优先恢复原生优势物种是提升退化草地生产力的有效途径。     

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。     

长期氮添加对典型草原植物多样性与初级生产力的影响及途径

氮(N)沉降对陆地生态系统的结构和功能已产生了重要的影响, N也是中国北方草原植物生长和初级生产力的主要限制性元素。物种多样性和功能多样性是揭示生物多样性对生态系统功能维持机制的关键指标, 然而, 关于长期N添加下草原物种多样性与功能多样性的关系, 及其对初级生产力的影响途径及机制, 尚不十分清楚。为此, 该研究依托在内蒙古典型草原建立的长期N添加实验平台, 实验处理包括1个完全对照(不添加任何肥料)和6个N添加水平(0、1.75、5.25、10.50、17.50和28.00 g·m^-2·a^-1), 研究了长期N添加对典型草原物种多样性、功能多样性和初级生产力的影响大小及途径。结果表明: 1) N添加显著降低了典型草原的物种丰富度和Shannon-Wiener指数, 但对功能多样性(包括功能性状多样性指数和群落加权性状值)无显著的影响。2)结构方程模型分析表明, 功能多样性主要受物种丰富度的影响, 但是物种多样性减少并没有导致功能多样性降低, 其原因主要是功能群组成发生了改变, 即群落内多年生根茎禾草所占比例显著增加, 以致群落加权性状值变化不大。3) N通过影响物种丰富度和功能群组成, 间接影响群落加权性状值, 进而影响群落净初级生产力。其中, 群落加权性状值是最重要的影响因子, 可解释48%的初级生产力变化, 表明初级生产力主要是由群落内优势物种的生物量及功能性状所决定, 因此该研究的结果很好地支持了质量比假说。     

Functional diversity of crustacean zooplankton communities: towards a trait-based classification

1. While studies of phytoplankton and terrestrial plant communities have increasingly emphasised the use of functional traits in ecological research, few have yet to apply this approach to zooplankton communities.
2. This study reviews laboratory and observational studies on zooplankton feeding and life history and provides a series of functional trait tables for the North American freshwater zooplankton. Qualitative and quantitative trait tables highlight areas where data were more scarce and point to which types of studies could fill in gaps in our knowledge of zooplankton niches.
3. Data were most complete for the Cladocera across most traits, while feeding information for cyclopoids was most sparse. Qualitative data that distinguished congeneric species were lacking for most groups.
4. A regional community dendrogram for common north-eastern North American zooplankton species was generated and shows that taxonomic differences between species do not capture fully functional differences based on the traits of body length, habitat, trophic group and feeding type.
5. The data collected here, combined with readily measurable species attributes, can be used to generate a multivariate measure of the functional niche of each species found in a community. Armed with this information, functional relationships that are useful for ecological studies of lake ecosystems can be more easily conducted.     

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。     

功能多样性及其研究方法

功能多样性是联系生物多样性和生态系统功能的关键性因素。植物群落内功能多样性的量化描述对于研究生物多样性响应环境变化及其对生物多样性-生态系统功能关系的影响至关重要。由于功能多样性的概念含混不清,使得对其范围的确定、定量描述及其对生态系统功能的作用的评价存在较大的难度。在查阅大量文献的基础上,以植物群落为例,介绍了功能多样性的定义、计算方法及研究中应注意的问题。在植物群落功能多样性指数的选择和测定过程中,对获取相关物种必要的功能特征信息、确定其权重、确定不同物种在群落中的相对丰富度、利用所得的功能多样性指数解释和预测生态系统过程的变化等问题,进行了研究。     

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.     

刈割和施肥对高寒草甸物种多样性和功能多样性时间动态及其关系的影响

强烈扰动和环境胁迫对植物群落的物种多样性(SD)和功能多样性(FD)有重要影响, 但SD和FD随时间的变化模式及其关系至今少有研究。该研究通过对高寒矮生嵩草(Kobresia humilis)草甸为期7年(2007-2013年)的刈割(3个水平: 不刈割、留茬3 cm和留茬1 cm)和施肥(2个水平: 施肥和不施肥)控制实验探讨了SD和FD随时间的变化模式及其关系。研究结果显示: (1)刈割显著增加了SD和FD, 施肥处理则降低了SD, 对FD增加仅有微弱影响; (2)各处理群落的SD随着时间而下降, FD则随时间增加; (3)随着刈割强度的增加, SD(x)-FD(y)关系由正线性相关变为无相关, 斜率大小为slope_l ≥ slope_m > slope_h (下标l、m和h分别表示轻度、中度和重度扰动强度), 施肥并不会改变此关系形式和斜率; (4)刈割与施肥对SD和FD的互作效应都不显著, 且不施肥群落SD(x)-FD(y)关系的斜率也为slope_l ≥ slope_m > slope_h。上述结果说明, 刈割并不一定导致植物功能性状的趋同构建, 也能引发趋异构建, 而施肥引起的强烈种间竞争也并未显著增强趋异构建过程, 这与植物群落构建理论的预测不完全一致。与施肥相比, 刈割扰动是决定群落中SD-FD时间关系形式的主要因素, 并决定着SD-FD关系斜率的变化。