Local adaptation of mycorrhizae communities changes plant community composition and increases aboveground productivity

Soil microbial communities can have an important role in the adaptation of plants to their local abiotic soil conditions and in mediating plant responses to environmental stress. This has been clearly demonstrated for individual plant species, but it is unknown how locally adapted microbes may affect plant communities. It is possible that the adaptation of microbial communities to local conditions can shape plant community composition. Additionally, it is possible that the effects of locally adapted microorganisms on individual plant species could be altered by co-occurring plant species. We tested these possibilities in plant community mesocosms with soils and mycorrhizal fungi (AMF) from three locations. We found that plant community biomass responded positively to local adaptation of AMF to soil conditions. Plant community composition also changed in response to local adaptation of AMF. Unexpectedly, the strongest benefits of locally adapted AMF went to early successional plant species that have the highest relative growth rates and the lowest responsiveness to the presence of AMF. Late successional plants that responded positively overall to the presence of AMF were often suppressed in communities with local AMF, perhaps because of strong competition from fast growing plant species. These results show that local adaptation of soil microbial communities can shape plant community composition, and the benefits that plants derive from locally adapted microorganisms can be reshaped by the competitive context in which these associations occur.     

陕西子午岭生态因素对植物群落的影响

为探讨生境对植被格局分布的影响,对黄土高原马栏林区60块样地进行植被学调查的基础上,采用17个环境指标刻画植物群落的空间位置、地形和土壤特征;利用双向指示种分析(TWINSPAN)划分了该区不同演替阶段的植物群落类型;利用前向选择法(forward selection)及Monte Carlo检验对不同演替阶段植物群落物种组成影响显著(p＜0.05)的环境因子进行筛选;利用去势典范对应分析(DCCA)排序方法分析不同演替阶段植物群落分布格局与环境的关系;利用偏典范对应分析(partial CCA)定量分离环境、空间及其交互作用对植被格局总体变异的影响.结果表明:(1)马栏林区的植物群落可划分为13个类型,分别属于4个不同的演替阶段;(2)对演替初期阶段群落影响显著的环境因子是土壤含盐量和碱解氮,对演替过渡阶段群落影响显著的环境因子是海拔和腐殖质厚度,对演替亚顶级阶段群落影响显著的因子是海拔、坡向、枯落物厚度、腐殖质厚度和pH值,而对演替顶级阶段群落分异影响显著的因子是海拔、坡向、pH值和速效磷;(3)不同演替阶段群落的生态学特性和分布规律与环境空间的生态梯度格局吻合较好;(4)随着演替的进行,环境因子单独对植物群落的影响越来越大,而样地位置单独作用和样地位置与环境因子的交互作用之和越来越小.     

Temporal variation in the influence of forest succession on caterpillar communities: A long‐term study in a tropical dry forest

Forest succession can influence herbivore communities through changes in host availability, plant quality, microclimate, canopy structure complexity and predator abundance. It is not well known, however, if such influence is constant across years. Caterpillars have been reported to be particularly susceptible to changes in plant community composition across forest succession, as most species are specialists and rely on the presence of their hosts. Nevertheless, in the case of tropical dry forests, plant species have less defined successional boundaries than tropical wet forests, and hence herbivore communities should be able to persist across different successional stages. To test this prediction, caterpillar communities were surveyed during eight consecutive years in a tropical dry forest in four replicated successional stages in Chamela, Jalisco and Mexico. Lepidopteran species richness and diversity were equivalent in mature forests and early successional stages, but a distinctive caterpillar community was found for the recently abandoned pastures. Species composition tended to converge among all four successional stages during the span of eight years. Overall, our results highlight the importance of both primary and secondary forest for the conservation of caterpillar biodiversity at a landscape level. We also highlight the relevance of long‐term studies when assessing the influence of forest succession to account for across year variation in species interactions and climatic factors. Abstract in French is available with online material.     

Primary succession of Bistorta vivipara (L.) Delabre (Polygonaceae) root‐associated fungi mirrors plant succession in two glacial chronosequences

Glacier chronosequences are important sites for primary succession studies and have yielded well‐defined primary succession models for plants that identify environmental resistance as an important determinant of the successional trajectory. Whether plant‐associated fungal communities follow those same successional trajectories and also respond to environmental resistance is an open question. In this study, 454 amplicon pyrosequencing was used to compare the root‐associated fungal communities of the ectomycorrhizal (ECM) herb Bistorta vivipara along two primary succession gradients with different environmental resistance (alpine versus arctic) and different successional trajectories in the vascular plant communities (directional replacement versus directional non‐replacement). At both sites, the root‐associated fungal communities were dominated by ECM basidiomycetes and community composition shifted with increasing time since deglaciation. However, the fungal community's successional trajectory mirrored the pattern observed in the surrounding plant community at both sites: the alpine site displayed a directional‐replacement successional trajectory, and the arctic site displayed a directional‐non‐replacement successional trajectory. This suggests that, like in plant communities, environmental resistance is key in determining succession patterns in root‐associated fungi. The need for further replicated study, including in other host species, is emphasized.     

Cursorial spiders and succession: age or habitat structure?

Summary The structure of cursorial spider assemblages was examined along a gradient of four temperature successional communities. Species diversity (H), richness (S), and evenness (J) exhibited a dichotomy between herbaceous and woody communities rather than a progressive change with community age: all three parameters were higher in the two younger fields than in the two older woodlands, which is contrary to conventional successional theory. Species importance curves were steeper in the two woody communities. The breadth of the distribution of adult body lengths was greater in the two herbaceous communities. Indices of community similarity revealed neither a successional trend nor the vegetative dichotomy. We suggest the hypothesis that habitat structure is a more important determinant of cursorial spider diversity than successional age per se, and that the switch in dominance from herbaceous to woody vegetation is the critical change. We further suggest that competition for prey is more important to cursorial spiders in early successional (herbaceous) communities, because of a switch in the limiting resource from prey in these communities to the amount of accumulating litter (a spatial resource) in older woody stands. This may explain the greater variation in adult body size of these generalist predators in the two younger communities.     

Contrasting Responses of Soil Microbial and Nematode Communities to Warming and Plant Functional Group Removal Across a Post-fire Boreal Forest Successional Gradient

Global warming is causing increases in surface temperatures and has the potential to influence the structure of soil microbial and faunal communities. However, little is known about how warming interacts with other ecosystem drivers, such as plant functional groups or changes associated with succession, to affect the soil community and thereby alter ecosystem functioning. We investigated how experimental warming and the removal of plant functional groups along a post-fire boreal forest successional gradient impacted soil microbial and nematode communities. Our results showed that warming altered soil microbial communities and favored bacterial-based microbial communities, but these effects were mediated by mosses and shrubs, and often varied with successional stage. Meanwhile, the nematode community was generally unaffected by warming and was positively affected by the presence of mosses and shrubs, with these effects mostly independent of successional stage. These results highlight that different groups of soil organisms may respond dissimilarly to interactions between warming and changes to plant functional groups, with likely consequences for ecosystem functioning that may vary with successional stage. Due to the ubiquitous presence of shrubs and mosses in boreal forests, the effects observed in this study are likely to be significant over a large proportion of the terrestrial land surface. Our results demonstrate that it is crucial to consider interactive effects between warming, plant functional groups, and successional stage when predicting soil community responses to global climate change in forested ecosystems.     

Ecological response hides behind the species abundance distribution: Community response to low‐intensity disturbance in managed grasslands

Land‐use and management are disturbance factors that have diverse effects on community composition and structure. In traditional rural grasslands, such as meadows and pastures, low‐intensity management is maintained to enhance biodiversity. Maintenance of road verges, in turn, creates habitat, which may complement traditional rural grasslands. To evaluate the effect of low‐intensity disturbance on insect communities, we characterized species abundance distributions (SAD) for Carabidae, Formicidae, and Heteroptera in three grassland types, which differed in management: meadows, pastures, and road verges. The shape of SAD was estimated with three parameters: abundance decay rate, dominance, and rarity. We compared the SAD shape among the grassland types and tested the effect of environmental heterogeneity (plant species richness) and disturbance intensity (trampling in pastures) on SADs. The shape of SADs did not differ among the grassland types but among the taxonomic groups instead. Abundance decay rate and dominance were larger for Formicidae, and rarity smaller, than for Carabidae and Heteroptera. For Carabidae and window‐trapped Heteroptera, rarity increased with increasing plant species richness. For Formicidae, dominance increased with trampling intensity in pastures. Although the SAD shape remained largely unchanged, the identity of the dominant species tended to vary within and among grassland types. Our study shows that for a given taxonomic group, the SAD shape is similar across habitat types with low‐intensity disturbances resulting from different management. This suggests that SADs respond primarily to the intensity of disturbance and thus could be best used in monitoring communities across strong disturbance and environmental gradients. Because taxonomic groups can inherently have different SADs, taxon‐specific SADs for undisturbed communities must be empirically documented before the SAD shape can be used as an indicator of environmental change. Because the identity of the dominant species changes from management type to another, the SAD shape alone is not an adequate monitoring tool.     

浙江天童常绿阔叶林演替阶段共有种的树木构型

树木构型是木本植物为响应光照变化在其空间建造结构上的配置模式和形态体现。研究演替不同阶段共有种构型的变化可以剔除植物谱系的影响, 反映植物构型特征与光资源供给性的关系。该研究在浙江宁波天童、南山和北仑3个次生演替序列上选择了5个演替共有种, 分4个群落高度层级, 对照分析了树高、冠幅深度和面积、枝条伸展方向、基径、叶片盖度和聚集度构型性状随演替的变化, 并分析了与冠幅曝光指数的线性关系。结果表明: 1)随着演替进行, 冠幅厚度和面积、叶片盖度、叶片聚集度和基径逐步增加, 但在个别相邻演替阶段增加不显著; 2)随着演替进行, 植物的垂直枝比例降低, 水平枝比例增加; 3)演替过程中植物冠幅曝光指数在各层级内都呈现出减小趋势; 4)构型性状和植物冠幅曝光指数间存在显著的线性回归关系(p < 0.001)。总之, 随着常绿阔叶林演替进行, 演替共有种构型的变化反映了物种功能类群由阳性先锋植物向耐阴植物的转化, 其中, 植物对光资源的适应是导致构型变化的主要原因。     

Iceberg Disturbance and Successional Spatial Patterns: The Case of the Shelf Antarctic Benthic Communities

High-latitude, shelf Antarctic benthic communities are highly diversified and structured, dominated by benthic suspension feeders, and are subject to major natural disturbances. This study focuses on spatial patterns of the Antarctic benthos emphasizing the succession process after iceberg disturbance. For this purpose, underwater photographs (1 m² each) from the southeastern Weddell Sea shelf (<300 m depth) were analyzed using techniques from the field of landscape ecology. Here, we examine measurements of spatial patterns (landscape indices) to describe changes in structural patterns along successional stages on these Antarctic benthic communities. We show a gradual separation from the early to older stages of succession based on sessile benthic cover area, size, shape, diversity, and interspersion and juxtaposition indices. Conceptually, the results describe a gradient from samples belonging to first stages of recovery with low cover area, low complexity of patch shape, small patch size, low diversity and patches poorly interspersed to samples from later stages with higher values of these indices. Cover area was the best predictor of recovery. We conclude that a variety of factors affect the observed successional sequences of Antarctic shelf benthic communities after iceberg disturbance, including the existence and dispersal abilities of propagules, growth rates, and competition between species. Overall, changes in the magnitude, frequency, and duration of disturbance regimes and alterations of ecosystem resilience pose major challenges for conservation of Antarctic benthos. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of earthworms and arbuscular mycorrhizal fungi depend on the successional stage of a grassland plant community

Earthworms and arbuscular mycorrhizal fungi (AMF) have profound impacts on plant performance. However, it is largely unknown if and how earthworms and AMF may affect plant succession. We planted mesocosms with an early-mid successional and a mid-late successional grassland plant community and added endogeic earthworms and commercial AMF in a full-factorial way to natural background soil. Earthworms had a positive effect on the total root and shoot biomass of both plant communities, with the effect on the shoots being slightly enhanced by co-inoculation with AMF. Surprisingly, the earthworm effect on the mid successional plant species depended on the successional stage of the plant community. Earthworms had a positive effect on the mid successional plant species when they were growing in the mid-late successional plant community, but no effect when the same plant species were growing in the early-mid successional plant community. Addition of AMF alone tended to reduce the shoot biomass of the early successional plant species, while the addition of earthworms in the presence or absence of AMF increased their shoot biomass. We conclude that the impacts of earthworms on plant species may depend on the successional stage of the plant community, while the effect of AMF addition depends on the successional stage of the plant community and may be changed by the presence of earthworms. Earthworms and AMF addition affect plants and plant communities of different successional stages differently with potential effects on plant succession.     

Microbial community composition but not diversity changes along succession in arctic sand dunes

The generality of increasing diversity of fungi and bacteria across arctic sand dune succession was tested. Microbial communities were examined by high‐throughput sequencing of 16S rRNA genes (bacteria) and internal transcribed spacer (ITS) regions (fungi). We studied four microbial compartments (inside leaf, inside root, rhizosphere and bulk soil) and characterized microbes associated with a single plant species (Deschampsia flexuosa) across two sand dune successional stages (early and late). Bacterial richness increased across succession in bulk soil and leaf endosphere. In contrast, soil fungal richness remained constant while root endosphere fungal richness increased across succession. There was, however, no significant difference in Shannon diversity indices between early and late successional stage in any compartment. There was a significant difference in the composition of microbial communities between early and late successional stage in all compartments, although the major microbial OTUs were shared between early and late successional stage. Co‐occurrence network analysis revealed successional stage‐specific microbial groups. There were more co‐occurring modules in early successional stage than in late stage. Altogether, these results emphasize that succession strongly affects distribution of microbial species, but not microbial diversity in arctic sand dune ecosystem and that fungi and bacteria may not follow the same successional trajectories.     

Succession of Ephemeral Secondary Forests and Their Limited Role for the Conservation of Floristic Diversity in a Human-Modified Tropical Landscape

Both local- and landscape-scale processes drive succession of secondary forests in human-modified tropical landscapes. Nonetheless, until recently successional changes in composition and diversity have been predominantly studied at the patch level. Here, we used a unique dataset with 45 randomly selected sites across a mixed-use tropical landscape in central Panama to study forest succession simultaneously on local and landscape scales and across both life stages (seedling, sapling, juvenile and adult trees) and life forms (shrubs, trees, lianas, and palms). To understand the potential of these secondary forests to conserve tree species diversity, we also evaluated the diversity of species that can persist as viable metapopulations in a dynamic patchwork of short-lived successional forests, using different assumptions about the average relative size at reproductive maturity. We found a deterministic shift in the diversity and composition of the local plant communities as well as the metacommunity, driven by variation in the rate at which species recruited into and disappeared from the secondary forests across the landscape. Our results indicate that dispersal limitation and the successional niche operate simultaneously and shape successional dynamics of the metacommunity of these early secondary forests. A high diversity of plant species across the metacommunity of early secondary forests shows a potential for restoration of diverse forests through natural succession, when trees and fragments of older forests are maintained in the agricultural matrix and land is abandoned or set aside for a long period of time. On the other hand, during the first 32 years the number of species with mature-sized individuals was a relatively small and strongly biased sub-sample of the total species pool. This implies that ephemeral secondary forests have a limited role in the long-term conservation of tree species diversity in human-modified tropical landscapes.     

针茅草原放牧衰退演替阶段的模糊聚类分析

本文利用模糊聚类分析的系统聚类法和ISODATA分类法,对短花针茅草原因过度放牧而形成的衰退演替系列的各阶段进行了分类;比较成功地在群落分类上联合使用了PCA排序与ISODATA分类。结果表明:这两种方法都将衰退演替系列分成4个阶段,它们是短花针茅群落、扁穗冰草群落、冷蓠群落和亚氏旋花群落。这与前人用其他方法分类的结果相似。本文为群落分类提供了两种合理、可行的数学分类方法。     

SPECIES-AREA RELATIONSHIP DURING PRIMARY SUCCESSION IN GRANITE OUTCROP PLANT COMMUNITIES

The species-area relationship and its underlying explanatory mechanisms were investigated in a primary successional sere on a southeastern (United States) granite outcrop. There, plant communities occupy soil-filled depressions separated from one another by areas of bare rock. They have been termed “island communities.” Soil depth and area increase as succession proceeds from Sedum smallii, to lichen-annual, to annual-perennial, and to herb-shrub-tree stages. Although plant species richness is significantly and positively correlated with island area in the system studied (all successional stages considered), the relationship between species richness and island maximum soil depth is stronger. However, island maximum soil depth and area are significantly and positively correlated. The exponential function describes the speciesarea relationship better than the transformed power function. Within successional stage, species richness shows no significant relationship with area or depth, except for late-successional island communities. Processes related to community successional development may explain the species-area relationship that exists when islands of all stages are considered. However, mechanisms related to equilibrium between extinction and immigration may be responsible for the speciesarea relationship for late-successional island communities.     

Early succession of butterfly and plant communities on set-aside fields

 Hypotheses on secondary succession of butterfly and plant communities were tested using naturally developed 1- to 4-year-old set-aside fields (n = 16), sown fields (n = 8) and old meadows (n = 4) in 1992 in South Germany. Pioneer successional fields (1st and 2nd year of succession, dominated by annuals) and early successional fields (3rd and 4th year of succession where perennials, especially grasses became dominant) had fewer plant species than mid-successional fields (old meadows). In contrast to established hypotheses, mean number of plant species decreased from 1- to 4-year-old set-aside fields. Species richness of butterfly communities did not change during the first four years of succession, but species composition changed greatly. Pioneer successional fields were characterized by (1) specialized butterflies depending on annual pioneer foodplants (e.g. Issoria lathonia), and (2) species preferring the pioneer successions despite their host plants being more abundant on early and mid-successional fields (e.g. Papilio machaon). The variability in butterfly species richness was best explained by flower abundance which was closely correlated with plant species richness. Species whose abundance was correlated with habitat connectivity were significantly smaller than species which correlated with flower abundance. Numbers of caterpillar species were correlated with numbers of adult butterfly species. Life-history features of butterflies changed significantly from pioneer to early and mid-successional fields. We found decreasing body size and migrational ability, decreasing numbers of species hibernating as imago, decreasing numbers of generations and increasing larval stage duration with age of succession, but, contrary to expectation, host plant specialization, numbers of egg-cluster laying species and egg diameter did not change with successional age. Received 18 September 1995 / Accepted: 17 July 1996     

Intraspecific variation in plant size,secondary plant compounds,herbivory and parasitoid assemblages during secondary succession

During secondary succession on abandoned agricultural fields the diversity and abundance of insect communities often increases, whereas the performance and nutritional quality of early successional plants often declines. As the diversity and abundance of insects on a single plant are determined by characteristics of the environment as well as of the host plant, it is difficult to predict how insects associated with a single plant species will change during succession. We examined how plant characteristics of the early successional plant species ragwort (Jacobaea vulgaris), and the herbivores and parasitoids associated with these plants change during secondary succession. In ten grasslands that differed in time since abandonment (3–26 years), we measured the size and primary and secondary chemistry of individual ragwort plants. For each plant we also recorded the presence of herbivores in flowers, leaves and stems, and reared parasitoids from these plant parts. Ragwort plants were significantly larger but had lower nitrogen concentrations in recently abandoned sites than in older sites. Pyrrolizidine alkaloid (PA) composition varied among plants within sites but also differed significantly among sites. However, there was no relationship between the age of a site and PA composition. Even though plant size decreased with time since abandonment, the abundance of stem-boring insects and parasitoids emerging from stems significantly increased with site age. The proportion of plants with flower and leaf herbivory and the number of parasitoids emerging from flowers and leaves was not related to site age. Parasitoid diversity significantly increased with site age. The results of our study show that ragwort and insect characteristics both change during secondary succession, but that insect herbivore and parasitoid abundances are not directly related to plant size or nutritional quality.     

Quantifying relationships between traits and explicitly measured gradients of stress and disturbance in early successional plant communities

Questions: How can one explicitly quantify, and separately measure, stress and disturbance gradients? How do these gradients affect functional composition in early successional plant communities and to what extent? Can we accurately predict trait composition from knowledge of these gradients? Location: Southern Quebec, Canada. Methods: Using eight environmental variables measured in 48 early successional plant communities, we estimated stress and disturbance gradients through structural equation modelling. We then measured 10 functional traits on the most abundant species of these 48 communities and calculated their community‐level mean and variance weighted by the relative abundance of each species. Finally, we related these community‐weighted means and variances to the estimated stress and disturbance gradients using general linear models or generalized additive models. Results: We obtained a well‐fitting measurement model of the stress and disturbance gradients existing in our sites. Of the 10 studied traits, only average plant reproductive height was strongly correlated with the stress (r²=0.464) and disturbance (r²=0.543) gradients. Leaf traits were not significantly related to either the stress or disturbance gradients. Conclusions: The well‐fitting measurement model of the stress and disturbance gradients, combined with the generally weak trait–environment linkages, suggests that community assembly in these early successional plant communities is driven primarily by stochastic processes linked to the history of arrival of propagules and not to trait‐based environmental filtering.     

Interactions between mycorrhizal colonization and plant life forms along the successional gradient of coastal sand dunes in the eastern Mediterranean, Turkey

The mycorrhizal status of dune plant species in relation to their plant life forms was surveyed along a successional gradient of sand dune on the southern Mediterranean coast of Turkey. Roots of 64 dune plant species belonging to 30 families were collected from sand dune communities at four different successional stages: embryonic dunes (ED), mobile dunes (MD), fixed dunes (FD), and remnant dunes (RD). Of the plant species surveyed in all successional stages, 54 (84%) had formed mycorrhizal associations. Nonmycorrhizal plants with cryptophyte life forms predominated in the earlier successional stages (ED and MD), whereas the number and percent coverage of mycorrhizal plant species belonging to hemicryptophytes, phanerophytes, and chamaephytes generally increased with the stabilization of sand dunes. Arbuscular mycorrhizal (AM) colonization was found to be the dominant mycorrhizal type in ED, MD, and RD. But phanerophytes with dual colonization, AM and ectomycorrhizal, became the dominant life form with high plant coverage in the FD stage. Total percentage of mycorrhizal root length colonization showed significant positive correlations relating to soil parameters such as organic matter and nitrogen content, while negatively correlating to high soil reaction (pH).     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.