Commonness,rarity, and intraspecific variation in traits and performance in tropical tree seedlings

One of the few rules in ecology is that communities are composed of many rare and few common species. Trait‐based investigations of abundance distributions have generally focused on species‐mean trait values with mixed success. Here, using large tropical tree seedling datasets in China and Puerto Rico, we take an alternative approach that considers the magnitude of intraspecific variation in traits and growth as it relates to species abundance. We find that common species are less variable in their traits and growth. Common species also occupy core positions within community trait space indicating that they are finely tuned for the available conditions. Rare species are functionally peripheral and are likely transients struggling for success in the given environment. The work highlights the importance of considering intraspecific variation in trait‐based ecology and demonstrates asymmetry in the magnitude of intraspecific variation among species is critical for understanding of how traits are related to abundance.     

Species abundance is jointly determined by functional traits and negative density dependence in a subtropical forest in southern China

植物物种多度受功能性状和负密度依赖共同影响——以中国南亚热带黑石顶森林样地为例 影响植物群落中植物物种多度的因素较多,确定关键影响因素及阐明其具体机制一直是群落生态学的研究重点之一。目前确定性的影响因素主要有两大类：植物功能性状和负密度依赖。功能性状通过影响植物的竞争能力、资源的获取效率、对环境的适应能力等方面进而影响植物的多度;负密度依赖表现在同种或异种植物个体在空间上聚集,特异性的寄主(病原菌、昆虫等)传播容易导致植物个体(主要为幼苗和小树)死亡,种群发生自疏,进而影响植物的多度。到目前为止,单一探究植物功能性状或负密度依赖对植物多度影响机制的研究已有大量报道,而将两者结合来探讨对植物多度的影响的工作相对较少。为了更深入地探索影响植物多度的机制,本研究中我们综合分析了功能性状与负密度依赖对植物多度的影响。研究地点为广东省黑石顶亚热带森林50 ha大样地;功能性状方面选取了叶面积(LA)、比叶面 积(SLA)、叶片干物质含量(LDMC)、叶片氮含量(LNC)和最大电子传输速率(ETR_max);植物物种所受负密度依赖强度通过构建小树在样地两次普查期间的存活状况受邻体密度影响的层次贝叶斯模型计算;通过结构方程模型构建功能性状-负密度依赖-物种多度的内在联系框架。结构方程模型结果显示,物种多度受功能性状和负密度依赖共同作用,其中功能性状对物种多度的影响包括直接作用和通过负密度依赖的间接作用。具体来说,SLA对多度的影响包括直接和间接两种;LDMC和LNC仅间接影响物种多度;LA和ETR_max对多度只有直接影响;负密度依赖与物种多度之间存在直接的负相关关系,说明多度较高的物种受到的负密度依赖效应更强。结构方程模型对物种多度差异的解释度达到20%。以上结果表明,黑石顶植物多度分布是功能性状与负密度依赖共同作用的结果;比叶面积在所研究的因素中贡献最大。该项工作有助于提高我们对亚热带森林植物常见种和稀有种分布格局的理解。     

高山草地植物群落数量特征沿海拔梯度变化及其影响因素

为了探索青藏高原湿润地区草地的演变规律和未来对气候变化的响应,以雅家埂高山带不同海拔草地为对象,研究了草地群落特征沿海拔梯度的变化,并分析了气候因子和土壤因子对群落特征的影响。结果表明:(1)沿海拔梯度草地群落特征具有明显分异,地上生物量随海拔升高呈现显著降低,而地下生物量和群落的根冠比呈现显著增加。群落多样性指数在较低海拔间呈现显著变化,而在较高海拔间变化不明显。总体而言,随海拔升高群落多样性增大,均匀度增大。(2)不同海拔群落特征差异主要由气候因子控制,且温度起主导作用,土壤因子的解释度可忽略。(3)整个海拔梯度上群落多样性指数与生物量呈负相关关系,支持根竞争和光竞争假说。     

践踏干扰对碧塔海高寒草甸植被茎叶性状、物种多样性和功能多样性的影响

高寒草甸具有重要的生态服务功能,然而固有脆弱性使其极易遭受气候变化和人为干扰等多重因素的影响。作为滇西北旅游资源中重要的组分之一,高寒草甸吸引了大批游客前往开展徒步旅行活动,但伴随着的践踏干扰作用会不可避免地对高寒草甸生态系统带来负面影响。然而,目前关注践踏干扰对滇西北高寒草甸植被的影响,特别是植被功能性状和功能多样性如何发生变化方面的研究还十分欠缺。以云南省香格里拉市碧塔海自然保护区内的典型高寒草甸生态系统为研究对象,采用实验践踏的方式(一共5种不同强度的践踏处理)来模拟旅游活动对草甸植被的干扰作用,并以草甸植被的茎叶性状特征为切入点,重点探讨践踏干扰对茎叶性状的平均大小和变异程度的影响,以及物种丰富度(以物种形态分类为基础)和功能丰富度(以功能性状为基础)之间的关系。研究结果显示,随着践踏强度的增加,植株高度和叶片大小的平均值,而不是茎叶性状的变异程度,出现明显下降趋势。此外,物种丰富度和功能丰富度均随践踏强度的增强而减小,且两者之间呈现显著正相关关系。然而,较之轻度践踏实验组,重度践踏实验组中的功能均匀度和功能分离度水平均有所增加,表明践踏干扰可能会在短期内打破优势种对资源的绝对占有格局和减少物种间的生态位重叠程度。尽管高寒草甸对人类践踏活动有一定的承受能力,但气候变化和人为干扰等多重因素势必会改变和影响高寒草甸群落的结构和功能可持续性,这也对高寒草甸的保护与管理工作提出了更加紧迫的要求。     

海南尖峰岭热带山地雨林林冠层树种功能多样性特征

以海南尖峰岭热带山地雨林3块1 hm²样地为研究对象,利用11个林冠功能性状结合样地地形及林冠乔木树种样地清查数据,分别基于单维性状和多维性状比较物种多度加权对群落功能离散度指数——平均成对距离(MPD)和平均最近类群距离(MNTD)的影响;同时分析林冠层功能丰富度(FRic)与物种丰富度之间的关系,最后利用零模型探讨不同生境类型下标准化效应MPD和MNTD(经过物种多度加权且剔除群落物种丰富度差异影响)的变化,进而评价林冠层群落水平功能多样性格局及其对局域生境异质性的响应.结果表明: 功能性状维度和物种多度对MPD的影响强烈,不同维度功能性状多度加权前后MPD相关性较弱(R=0.359～0.628);但对MNTD的影响相对较弱,不同维度功能性状多度加权前后MNTD相关性较强(R=0.746～0.820);未经物种多度加权的MPD和MNTD均普遍高估了林冠层的功能离散度.林冠层功能丰富度与物种丰富度基本呈指数相关关系(F=128.20;R²=0.632;AIC=97.72;P＜0.001),且功能丰富度很有可能存在一定的物种丰富度阈值.基于不同维度功能性状的林冠层功能多样性格局及其生境响应存在一定的差异性.在生物竞争激烈的低沟生境中,林冠层功能多样性倾向于比预期零模型随机产生的功能多样性高,林冠树种功能性状表现出离散分布;而在其他生境类型中,林冠层功能多样性倾向于接近或低于随机产生的功能多样性,林冠树种功能性状随机或聚集分布.     

The biogeography and filtering of woody plant functional diversity in North and South America

Aim In recent years evidence has accumulated that plant species are differentially sorted from regional assemblages into local assemblages along local‐scale environmental gradients on the basis of their function and abiotic filtering. The favourability hypothesis in biogeography proposes that in climatically difficult regions abiotic filtering should produce a regional assemblage that is less functionally diverse than that expected given the species richness and the global pool of traits. Thus it seems likely that differential filtering of plant traits along local‐scale gradients may scale up to explain the distribution, diversity and filtering of plant traits in regional‐scale assemblages across continents. The present work aims to address this prediction. Location North and South America. Methods We combine a dataset comprising over 5.5 million georeferenced plant occurrence records with several large plant functional trait databases in order to: (1) quantify how several critical traits associated with plant performance and ecology vary across environmental gradients; and (2) provide the first test of whether the woody plants found within 1° and 5° map grid cells are more or less functionally diverse than expected, given their species richness, across broad gradients. Results The results show that, for many of the traits studied, the overall distribution of functional traits in tropical regions often exceeds the expectations of random sampling given the species richness. Conversely, temperate regions often had narrower functional trait distributions than their smaller species pools would suggest. Main conclusion The results show that the overall distribution of function does increase towards the equator, but the functional diversity within regional‐scale tropical assemblages is higher than that expected given their species richness. These results are consistent with the hypothesis that abiotic filtering constrains the overall distribution of function in temperate assemblages, but tropical assemblages are not as tightly constrained.     

城市建成区自生草本植物群落的物种多样性与功能多样性——以深圳市为例

城市自生草本植物的物种多样性和功能多样性直接影响城市生态系统的功能和稳定性,但目前相关研究依然很缺乏。以深圳市为例,基于建成区600个1hm²样地的现场调查数据,分析自生草本植物的群落结构、物种多样性、群落功能特征及功能多样性。结果显示:①调查共记录自生草本植物61科178属273种,其中多年生草本占54.6%,乡土植物占65.6%。②占明显优势的功能特征有中小型叶(91.85%)、草质叶(42.59%)、纸质叶(32.96%)、叶片无毛或近无毛(56.67%)、中等密度绒毛(40.37%)、叶面较粗糙(52.59%)、花期4-6个月(50.78%)、果期4-6个月(49.22%)、干果(86.03%)。③群落类型间功能丰富度FDp、功能离散度Rao二次熵差异极显著(P ＜ 0.01)、功能均匀度FEve差异显著(P ＜ 0.05)。④公园绿地的物种多样性指数明显高于其他三类绿地;不同绿地类型间的功能丰富度和功能离散度在滞尘、降噪、降温增湿及生物多样性保护方面都有极显著差异(P ＜ 0.01),功能均匀度只在滞尘及生物多样性保护方面差异极显著(P ＜ 0.01)。⑤各功能多样性指数均与Margalef指数、Simpson指数及Shannon-Wiener指数呈极显著正相关关系(P ＜ 0.01)。研究结论和方法为维护城市生态系统稳定性、建设生态宜居城市、促进城市可持续发展提供理论依据。     

Spatiotemporal patterns and dynamics of species richness and abundance of woody plant functional groups in a tropical forest landscape of Hainan Island, South China

Tropical forests are among the most species-diverse ecosystems on Earth. Their structures and ecological functions are complex to understand. Functional group is defined as a group of species that play similar roles in an ecosystem. The functional group approach has been regarded as an effective way of linking the compositions of complex ecosystems with their ecological functions. To understand the variation of functional groups in species-rich ecosystems after disturbance, the present study investigated the spatial pattern and temporal dynamics of woody plants in a typically fragmented natural forest landscape of Hainan Island in South China. The study area was classified into eight landscape types based on vegetation type, disturbance manner and the time of recovery. The woody plant species were aggregated into seven functional groups based on the growth form, successional status and plant size. The results gained from the present study showed that all functional groups, except for the emergent and canopy tree species, were present in all eight landscape types. Each landscape type had different numbers of dominant functional groups. There are similar species richness and stem abundance structure among functional groups between mid-successional clear cut lowland rainforest and old growth tropical coniferous forest. This similarity exists in selective logged lowland rainforest and old-growth lowland rainforest, as well as among landscape types of montane rainforest. The functional groups with the same successional status had similar patterns of species richness and stem abundance ratios among different landscape types. The variation patterns of functional groups along the successional stages in terms of species richness and stem abundance among the tropical lowland rainforest landscape types were more similar to each other than those in the tropical montane reinforest landscape types. This study provides further support for the competition-colonization tradeoff and successional niche theory as opposed to models of neutrality and ecological equivalence.     

Environment differentially affects the functional and phylogenetic structures of plant communities in a dry evergreen Afromontane tropical forest

Testing how local environmental conditions influence plant community assembly is important to understand the underlying mechanisms that promote and/or maintain biodiversity. Functional traits are used to find the broad spectrum of resource use strategies that plants use to respond to environmental variation. The patterns and drivers of plant community assembly through the lens of traits and phylogeny; however, remain to be studied in a uniquely biodiversity rich but poorly known fragmented dry Afromontane forest of Ethiopia. Here, we combined trait and community phylogenetic data from thirty sampling plots of 20 × 20 m size to determine the functional and phylogenetic structures and their drivers in a fragmented, human-dominated dry evergreen Afromontane forest. We found phylogenetic and functional clustering of plants in which the effect of environment was found to be trait specific. A weak phylogenetic signal for traits was detected suggesting that species resource use strategies may not be inferred using species phylogenetic distance. Additionally, we found functional traits to be weak in predicting species abundance distribution. Overall, while this study shows a non-random community assembly pattern, it also highlights the importance of deterministic processes being trait specific.     

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。     

Genetic diversity of Couratari multiflora and Couratari guianensis (Lecythidaceae): consequences of two types of rarity in central Amazonia

We quantified the within-population genetic variation of Couratari multiflora and C. guianensis, two tree species found in terra firme forests of central Amazonia. Both species have some ecological features in common, but they differ in population abundance across their geographic ranges. While C. multiflora has been found only in low-density populations in all sites studied to date, C. guianensis is relatively common in some sites and very scarce in others. In a 400-ha plot, we found 41 and 29 adults of C. multiflora and C. guianensis, respectively. Twenty-two saplings of C. guianensis and 103 seedlings of C. multiflora were also examined. The mean expected heterozygosities (H_em) of seedlings and adults of C. multiflora were 0.431 and 0.436, and the mean fixation indices (F_m), 0.114 and 0.176, respectively. For C. guianensis, saplings and adults presented H_em equal to 0.425 and 0.429, and the F_m were 0.393 and 0.527, respectively. These low-density populations of two congeneric species did not differ in terms of genetic diversity, but rather they differed in terms of mean observed heterozygosity (H_om), and therefore F_m. The species with variable population density had lower H_om and greater F_m relative to the species that is always found in low-density.     

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。     

Biogeographic context is related to local scale tree demography,co-occurrence and functional differentiation

Identifying the drivers of community structure and dynamics is a major pursuit in ecology. Emphasis is typically placed on the importance of local scale interactions when attempting to explain these fundamental ecological patterns. However, regional scale phenomena are also important predictors. The importance of regional scale context should be more evident in assemblages where multiple species are close to their range margins. Here, we test the importance of regional scale context using data from a temperate forest plot that contains two species groups – one near its northern range limit and one near its southern range limit. We show the proximity of species to their southern or northern range margins is linked to local scale co-occurrence, similarity in gene expression responses to a key environmental driver, demographic performance and inter-specific variation in conspecific negative density dependence. In sum, many of the key local scale patterns and processes of interest to community ecologists are linked to biogeographic context that is frequently ignored.     

Common spatial patterns of trees in various tropical forests: Small trees are associated with increased diversity at small spatial scales

Tropical forests are notable for their high species diversity, even on small spatial scales, and right‐skewed species and size abundance distributions. The role of individual species as drivers of the spatial organization of diversity in these forests has been explained by several hypotheses and processes, for example, stochastic dilution, negative density dependence, or gap dynamics. These processes leave a signature in spatial distribution of small trees, particularly in the vicinity of large trees, likely having stronger effects on their neighbors. We are exploring species diversity patterns within the framework of various diversity‐generating hypotheses using individual species–area relationships. We used the data from three tropical forest plots (Wanang—Papua New Guinea, Barro Colorado Island—Panama, and Sinharaja—Sri Lanka) and included also the saplings (DBH ≥ 1 cm). Resulting cross‐size patterns of species richness and evenness reflect the dynamics of saplings affected by the distribution of large trees. When all individuals with DBH ≥1 cm are included, ~50% of all tree species from the 25‐ or 50‐ha plot can be found within 35 m radius of an individual tree. For all trees, 72%–78% of species were identified as species richness accumulators, having more species present in their surroundings than expected by null models. This pattern was driven by small trees as the analysis of DBH >10 cm trees showed much lower proportion of accumulators, 14%–65% of species identified as richness repellers and had low richness of surrounding small trees. Only 11%–26% of species had lower species evenness than was expected by null models. High proportions of species richness accumulators were probably due to gap dynamics and support Janzen–Connell hypothesis driven by competition or top‐down control by pathogens and herbivores. Observed species diversity patterns show the importance of including small tree size classes in analyses of the spatial organization of diversity.