生态学的多样性指数:功能与系统发育

生物多样性是生态学的核心问题。传统的多样性指数仅包含物种数和相对多度的信息,这类基于分类学的多样性指数并不能很好地帮助理解群落构建和生态系统功能。不同物种对群落构建和生态系统功能所起到的作用类型和贡献也不完全相同,且物种在生态过程中的作用和贡献往往与性状密切相关,因此功能多样性已经成为反映物种群落构建、干扰以及环境因素对群落影响的重要指标。同时,由于亲缘关系相近的物种往往具有相似的性状,系统发育多样性也可以作为功能多样性的一个替代。功能多样性和系统发育多样性各自具有优缺点,但二者均比分类多样性更能揭示群落和生态系统的构建、维持与功能。     

毛乌素沙地植被不同恢复阶段植物群落物种多样性、功能多样性和系统发育多样性

进入21世纪以来, 中国荒漠化恢复取得显著成效, 荒漠化、沙化土地面积持续减少, 植被覆盖度大幅提升, 但关于植被恢复过程中生物多样性如何变化的研究不足, 这制约着对荒漠化恢复成效的全面评估。本文基于群落调查和叶功能性状(叶片厚度、叶片干物质含量、比叶面积和叶片密度)的测定, 分析了毛乌素沙地不同恢复阶段(半固定沙地、固定沙地、结皮覆盖沙地和草本植物覆盖沙地)的植物群落物种多样性、功能多样性和系统发育多样性特征。结果表明: (1)多数叶功能性状的系统发育信号不显著, 表明环境因子对研究区植物功能性状的塑造作用很强。(2)对于α多样性, 结皮覆盖沙地的物种多样性(Shannon-Wiener多样性, H)、物种丰富度(S)、功能丰富度(FRic)及系统发育多样性(PD)指数均显著低于其他恢复阶段, 而其他3个阶段间无显著差异; 这些指数间均显著正相关, 表明物种多样性、功能多样性和系统发育多样性在植被恢复过程中协同变化。(3) β多样性指数随恢复阶段间隔增加而逐渐增大, 表明物种组成、功能属性及系统发育关系随植被恢复持续变化, 且半固定沙地到固定沙地的群落物种组成、功能属性及系统发育关系更替最快, 导致群落间差异最大。(4)固定沙地、结皮覆盖沙地和草本植物覆盖沙地群落系统发育结构均趋向于发散, 表明竞争排斥是群落构建的主要驱动力; 而半固定沙地群落系统发育结构无一致规律, 表明群落构建可能受到生境过滤和竞争排斥的综合作用。研究结果可为植被建设与管理提供参考, 为毛乌素沙地生态保育和生物多样性的保护提供科学依据。     

Tree diversity drives associational resistance to herbivory at both forest edge and interior

Tree diversity is increasingly acknowledged as an important driver of insect herbivory. However, there is still a debate about the direction of associational effects that can range from associational resistance (i.e., less damage in mixed stands than in monocultures) to the opposite, associational susceptibility. Discrepancies among published studies may be due to the overlooked effect of spatially dependent processes such as tree location within forests. We addressed this issue by measuring crown defoliation and leaf damage made by different guilds of insect herbivores on oaks growing among conspecific versus heterospecific neighbors at forest edges versus interior, in two closed sites in SW France forests. Overall, oaks were significantly less defoliated among heterospecific neighbors (i.e., associational resistance), at both forest edge and interior. At the leaf level, guild diversity and leaf miner herbivory significantly increased with tree diversity regardless of oak location within stands. Other guilds showed no clear response to tree diversity or oak location. We showed that herbivore response to tree diversity varied among insect feeding guilds but not between forest edges and interior, with inconsistent patterns between sites. Importantly, we show that oaks were more defoliated in pure oak plots than in mixed plots at both edge and forest interior and that, on average, defoliation decreased with increasing tree diversity from one to seven species. We conclude that edge conditions could be interacting with tree diversity to regulate insect defoliation, but future investigations are needed to integrate them into the management of temperate forests, notably by better understanding the role of the landscape context.     

Beyond pairwise distances: neighbor-joining with phylogenetic diversity estimates

The "neighbor-joining algorithm" is a recursive procedure for reconstructing trees that is based on a transformation of pairwise distances between leaves. We present a generalization of the neighbor-joining transformation, which uses estimates of phylogenetic diversity rather than pairwise distances in the tree. This leads to an improved neighbor-joining algorithm whose total running time is still polynomial in the number of taxa. On simulated data, the method outperforms other distance-based methods. We have implemented neighbor-joining for subtree weights in a program called MJOIN which is freely available under the Gnu Public License at http://bio.math.berkeley.edu/mjoin/.     

The phylogenetic and functional diversity of regional breeding bird assemblages is reduced and constricted through urbanization

Aim

Urbanization broadly affects the phylogenetic and functional diversity of natural communities through a variety of processes including habitat loss and the introduction of non‐native species. Due to the challenge of acquiring direct measurements, these effects have been studied primarily using “space‐for‐time” substitution where spatial urbanization gradients are used to infer the consequences of urbanization occurring across time. The ability of alternative sampling designs to replicate the findings derived using space‐for‐time substitution has not been tested.

Location

Global.

Methods

We contrasted the phylogenetic and functional diversity of breeding bird assemblages in 58 cities worldwide with the corresponding regional breeding bird assemblages estimated using geographic range maps.

Results

Compared to regional assemblages, urban assemblages contained lower phylogenetic diversity, lower phylogenetic beta diversity, a reduction in the least evolutionary distinct species and the loss of the most evolutionarily distinct species. We found no evidence that these effects were related to the presence of non‐native species. Urban assemblages contained fewer aquatic species and fewer aquatic foraging species. The distribution of body size and range size narrowed for urban assemblages with the loss of species at both tails of the distribution, especially large bodied and broadly distributed species. Urban assemblages contained a greater proportion of species classified as passerines, doves or pigeons; species identified as granivores; species that forage within vegetation or in the air; and species with more generalized associations with foraging strata.

Main conclusions

Urbanization is associated with the overall reduction and constriction of phylogenetic and functional diversity, results that largely replicate those generated using space‐for‐time substitution, increasing our confidence in the quality of the combined inferences. When direct measurements are unavailable, our findings emphasize the value of developing independent sampling methods that broaden and reinforce our understanding of the ecological implications of urbanization.

     

Restoration goals: Why are fauna still overlooked in the process of recovering functioning ecosystems and what can be done about it?

Despite the evidence that fauna play complex and critical roles in ecosystems (e.g. pollination and nutrient cycling) and the knowledge that they need to be considered in restoration, fauna often remain poorly represented in restoration goal setting, monitoring and assessments of restoration success. Fauna clearly are integral to the aspirations of achieving full ecosystem recovery. However, over‐reaching assumptions about the unassisted return of fauna to restored sites, low investment in fauna monitoring, and minimal consideration of the requirements for fauna monitoring in regulatory guidance and standards appear to have led to the historically vegetation‐centric approaches to rehabilitation and ecological restoration. We argue that ecological complexities render assumptions of unassisted fauna return inappropriate in many situations and may represent a missed opportunity to enhance ecological outcomes and improve restoration trajectories. We advocate for greater consideration of fauna as facilitators of ecological restoration and, particularly for well‐funded projects, for monitoring to place greater emphasis on examining the behaviour and resilience of restored fauna communities. There is a clear need for both industry and regulators to recognise that fauna can be crucial facilitators of restoration and appreciate that the return and monitoring of functional faunal communities can be costly, challenging and may require detailed study across a wide range of taxonomic groups. Failure to advance from business as usual models may risk leaving a legacy of ostensibly functional, but biodiversity‐depauperate, restored ecosystems.     

Spatial patterns of phylogenetic diversity

Ecologists and conservation biologists have historically used species-area and distance-decay relationships as tools to predict the spatial distribution of biodiversity and the impact of habitat loss on biodiversity. These tools treat each species as evolutionarily equivalent, yet the importance of species' evolutionary history in their ecology and conservation is becoming increasingly evident. Here, we provide theoretical predictions for phylogenetic analogues of the species-area and distance-decay relationships. We use a random model of community assembly and a spatially explicit flora dataset collected in four Mediterranean-type regions to provide theoretical predictions for the increase in phylogenetic diversity - the total phylogenetic branch-length separating a set of species - with increasing area and the decay in phylogenetic similarity with geographic separation. These developments may ultimately provide insights into the evolution and assembly of biological communities, and guide the selection of protected areas.     

Recovery of mammal diversity in tropical forests: a functional approach to measuring restoration

Ecological restoration is increasingly applied in tropical forests to mitigate biodiversity loss and recover ecosystem functions. In restoration ecology, functional richness, rather than species richness, often determines community assembly, and measures of functional diversity provide a mechanistic link between diversity and ecological functioning of restored habitat. Vertebrate animals are important for ecosystem functioning. Here, we examine the functional diversity of small‐to‐medium sized mammals to evaluate the diversity and functional recovery of tropical rainforest. We assess how mammal species diversity and composition and functional diversity and composition, vary along a restoration chronosequence from degraded pasture to “old‐growth” tropical rainforest in the Wet Tropics of Australia. Species richness, diversity, evenness, and abundance did not vary, but total mammal biomass and mean species body mass increased with restoration age. Species composition in restoration forests converged on the composition of old‐growth rainforest and diverged from pasture with increasing restoration age. Functional metrics provided a clearer pattern of recovery than traditional species metrics, with most functional metrics significantly increasing with restoration age when taxonomic‐based metrics did not. Functional evenness and dispersion increased significantly with restoration age, suggesting that niche complementarity enhances species' abundances in restored sites. The change in community composition represented a functional shift from invasive, herbivorous, terrestrial habitat generalists and open environment specialists in pasture and young restoration sites, to predominantly endemic, folivorous, arboreal, and fossorial forest species in older restoration sites. This shift has positive implications for conservation and demonstrates the potential of tropical forest restoration to recover rainforest‐like, diverse faunal communities.     

戴云山物种多样性与系统发育多样性海拔梯度分布格局及驱动因子

生物多样性的海拔分布格局是生态学研究的热点。海拔作为综合性因子驱动着植物群落的物种、系统发育与功能多样性的空间分布。以戴云山南坡900-1600 m森林植物群落为研究对象,探讨物种多样性、系统发育指数与环境驱动因子的相互关系以及环境因子在群落构建与多样性维持中的重要意义。结果表明：（1）森林植物群落的系统发育多样性与物种多样性沿海拔均呈现中间高度膨胀格局。（2）物种多样性Margalef指数、Shannon-Wiener指数与系统发育多样性指数呈显著正相关,表明物种多样性越高,系统发育多样性也越高。Shannon-Wiener指数与物种多样性指数（Margalef、Pielou、Simpson指数）、系统发育多样性及系统发育结构都存在显著相关性,一定程度上Shannon-Wiener指数可以代替其他指数。Pielou指数、Simpson指数、Shannon-Wiener指数与系统发育结构NRI （Net relatedness index）指数、NTI （Net nearest taxa index）指数存在显著正相关,表明群落优势度、均匀度与系统发育结构相关性较强。（3）土壤全磷含量是影响系统发育多样性和物种多样性的主要驱动因子,土壤含水量是影响Shannon-Wiener、Pielou、Simpson指数的最显著因子,海拔是影响群落系统发育结构的主要因素。海拔是影响系统发育结构变化的主要环境因子,而土壤因子是影响物种多样性与系统发育多样性的主要因素,进一步验证了物种多样性与系统发育多样性的高度相关,结果旨在揭示物种群落空间分布规律。     

Geographic patterns and environmental correlates of taxonomic and phylogenetic diversity of aquatic plants in China

Studies on large-scale geographic patterns of aquatic plant diversity can promote research on the generality of macroecological patterns in different ecosystems. Here, we compiled a checklist of 889 aquatic angiosperms in China, including 738 helophytes (emergent and marshy plants) and 151 hydrophytes (submerged, free-floating, and floating-leaved plants). We explore the geographic patterns and environmental correlates of aquatic plant diversity based on six metrics including species richness (SR), weighted endemism (WE), phylogenetic diversity (PD), phylogenetic endemism (PE), the standardized effect size of phylogenetic diversity (PDses), and the standardized effect size of mean phylogenetic distance (MPDses). Our results show that the diversity of aquatic plants in China is extremely uneven, with high diversity in southeastern China and low diversity in northwestern China, and the geographic patterns of taxonomic and PD are generally consistent. The pattern of helophytes differs from that of hydrophytes. Notably, the wavy-shaped pattern of aquatic plant diversity (especially SR and PD for hydrophytes) across the latitude observed in this study is not consistent with those previously observed for aquatic plants in other continents. Climatic variables and water environmental variables are the main drivers of aquatic plant diversity in China; however, the effects of individual variables differ between helophytes and hydrophytes. Water environmental variables have a greater impact on PDses and MPDses of hydrophytes than those of helophytes. Overall, our work provides insight into understanding the large-scale patterns of aquatic plant diversity and is a critical addition to previous studies on the macroecological pattern of terrestrial organisms.     

Turnover of plant lineages shapes herbivore phylogenetic beta diversity along ecological gradients

Understanding drivers of biodiversity patterns is of prime importance in this era of severe environmental crisis. More diverse plant communities have been postulated to represent a larger functional trait‐space, more likely to sustain a diverse assembly of herbivore species. Here, we expand this hypothesis to integrate environmental, functional and phylogenetic variation of plant communities as factors explaining the diversity of lepidopteran assemblages along elevation gradients in the Swiss Western Alps. According to expectations, we found that the association between butterflies and their host plants is highly phylogenetically structured. Multiple regression analyses showed the combined effect of climate, functional traits and phylogenetic diversity in structuring butterfly communities. Furthermore, we provide the first evidence that plant phylogenetic beta diversity is the major driver explaining butterfly phylogenetic beta diversity. Along ecological gradients, the bottom up control of herbivore diversity is thus driven by phylogenetically structured turnover of plant traits as well as environmental variables.     

Restoring species diversity: assessing capacity in the U.S. native plant industry

Large quantities of diverse and appropriately adapted native plant germplasm are required to facilitate restoration globally, yet shortages can prevent restorations from attaining desired species diversity and structure. An extensive native plant industry has developed in the United States to help meet these demands, yet very little is known about its capacity to support germplasm needs. To better understand current capacity and germplasm availability, we report results of the first comprehensive and quantitative assessment of the native plant industry in the United States, which includes at least 841 vendors nationwide and the species they make available for restoration. We synthesized lists of commercially available species from native plant vendors across the United States and identified gaps in species availability to inform germplasm research, development, and production. Of the approximately 25,000 vascular plant taxa native to the United States, 26% are sold commercially, with growth form, conservation status, distribution, and taxonomy significantly predicting availability. In contrast, only 0.07% of approximately 3,000 native nonvascular taxa are sold commercially. We also investigated how demand for germplasm to support high‐quality restoration efforts is met by vendors in the Midwestern tallgrass prairie region, which has been targeted extensively by restoration efforts for decades. In this well‐developed native plant market, 74% of more than 1,000 target species are commercially available, often from vendors that advertise genetically diverse, locally sourced germplasm. We make recommendations to build on the successes of regional markets like the tallgrass prairie region, and to fill identified gaps, including investing in research to support production, ensuring more consistent and clear demand, and fostering regional collaboration.     

高寒湿地植物群落的物种多样性保护及生态恢复对策

以物种组成较为复杂的青藏高原东部地区玛曲县高寒湿地为背景．从恢复生态学角度,系统地探讨了影响湿地植物群落物种多样性与生产力变化的主要因素,从不同的时间和空间尺度研究了典型样区植物群落中物种多样性与生产力的恢复机理与过程,以及生境异质性对其的影响。结果表明：在导致湿地植被退化的人类干扰诸因素中．超载放牧是主要因素。该地区超载量高达24％;其次．气候变化对湿地物种多样性与生产力也有较大影响。典型样区的恢复试验结果表明：实行围栏轮牧、控制载畜量、鼠虫害防治和建植人工草地等人工综合措施,能使样区植被得以卓有成效的恢复与保护。其中．黄河滩地的植物群落中．物种丰富度从23种／m^2上升到29种／m^2;Shannon多样性指数从3．82上升到4．62;草场生产力(地上生物量)从729．2g／m^2上升到935．38g／m^2。阳坡和阴坡样地植被也有类同的恢复效果。在以上三类样地中．以滩地的生态恢复效果最为显著。据此,我们提出了相应的湿地物种多样性保护和生态恢复对策．即：加强自然保护区建设、协调人类活动与物种多样性保护的关系、严格控制载畜量、发展人工草地等主要措施与建设。     

南方红壤区3种典型森林恢复方式对植物群落多样性的影响

合理的森林恢复方式能提高植物多样性,进而提高生态系统服务功能.在我国南方红壤区研究了3种典型森林恢复方式(引进种恢复的湿地松(Pinus elliottii)人工林本地种恢复的马尾松(Pinus massoniana)人工林和自然恢复的天然次生林)的植物区系构成和植物群落多样性.结果表明:(1)湿地松人工林有155种植物,隶属66科118属,马尾松人工林有137种植物,隶属59科97属;天然次生林有226种植物,隶属86科160属,3种森林恢复方式的乔木层、灌木层和草本层的优势科属明显不同,马尾松人工林的优势物种和天然次生林更相似;(2)天然次生林的植物区系基本构成、植物区系类型种类高于马尾松人工林和湿地松人工林,并且天然次生林的温带成分比例高于湿地松人工林;(3)恢复方式对植物群落的多样性指数有显著影响,天然次生林的物种丰富度、辛普森指数明显高于马尾松人工林和湿地松人工林,两种人工林之间差异不显著;(4)3种森林恢复方式的植物群落结构存在显著差异,相比湿地松人工林,马尾松人工林的植物群落组成与天然次生林更相似.总之,自然恢复的天然次生林植物群落多样性高于人工恢复的马尾松人工林和湿地松人工林,本地种马尾松人工林在维持区域植物群落结构功能上优于引进种湿地松人工林.     

功能多样性和谱系多样性对热带云雾林群落构建的影响

近年来, 功能多样性和谱系多样性为探究群落构建机制提供了新方法。为了更准确地了解海南岛高海拔热带云雾林群落构建机制, 该研究以海南岛霸王岭热带云雾林为对象, 测定7个环境因子和13个植物功能性状。利用主成分分析(PCA)筛选环境因子, 以霸王岭、尖峰岭和黎母山热带云雾林分布物种建立区域物种库, 结合模型, 分析Rao二次熵(RaoQ)和平均成对谱系距离(MPD)变化对植物群落构建的影响。结果表明: 林冠开阔度、土壤全磷含量和坡度是影响植物群落构建的关键环境因子。多数功能性状的系统发育信号很低且不显著, 说明热带云雾林群落的系统发育关系与功能性状随历史进程变化不一致。RaoQ和MPD的实际观测值都显著低于期望值, 且其标准效应值与土壤磷含量显著相关, 说明生境过滤是驱动热带云雾林群落构建的关键因子, 土壤磷含量是群落构建的关键环境筛。     

Contrasting patterns of phylogenetic diversity across climatic zones of Western Ghats: A biodiversity hotspot in peninsular India

This study attempts to understand the biogeographic history of the Western Ghats forests by investigating decoupling between phylogenetic and taxonomic diversity. We specifically test whether the deciduous forests have been recently established, whether the southern region was a refuge, and whether the deciduous and evergreen forest species have disparate evolutionary histories. We used species composition data from 23 forest types along the Western Ghats for all woody angiosperms above 10‐cm diameter at breast height. Forests were broadly grouped as either evergreen or deciduous. Mean phylogenetic distances corrected for species richness and mean phylogenetic beta diversity corrected for shared species were assessed using z‐scores from null distributions. Null distributions were generated by randomizing the species relationships on the phylogeny. We found that all evergreen forests showed a greater phylogenetic diversity as compared with null expectations. Deciduous forests showed the inverse pattern. Within the evergreen belt, there was a decreasing phylogenetic diversity from south to north, as predicted by the southern refuge hypothesis. The phylogenetic beta diversity across evergreen–deciduous forests was lesser than the null expectation, whereas it was much higher across forests within the evergreen belt. This study provides the first phylogenetic evidence for the antiquity of evergreen forests as well as the southern refuge hypothesis in the Western Ghats. The deciduous forests species have shared evolutionary histories with the evergreen forest species, suggesting multiple shifts between evergreen and deciduous states through the lineages. Conversely, the evergreen species exhibited a disparate evolutionary history across these forests, possibly owing to sharper ecological or climatic gradients.     

Restoration Success: How Is It Being Measured?

The criteria of restoration success should be clearly established to evaluate restoration projects. Recently, the Society of Ecological Restoration International (SER) has produced a Primer that includes ecosystem attributes that should be considered when evaluating restoration success. To determine how restoration success has been evaluated in restoration projects, we reviewed articles published in Restoration Ecology (Vols. 1[1]–11[4]). Specifically, we addressed the following questions: (1) what measures of ecosystem attributes are assessed and (2) how are these measures used to determine restoration success. No study has measured all the SER Primer attributes, but most studies did include at least one measure in each of three general categories of the ecosystem attributes: diversity, vegetation structure, and ecological processes. Most of the reviewed studies are using multiple measures to evaluate restoration success, but we would encourage future projects to include: (1) at least two variables within each of the three ecosystem attributes that clearly related to ecosystem functioning and (2) at least two reference sites to capture the variation that exist in ecosystems.