群落中克隆植物的重要性

综述了群落水平上克隆植物的重要性,克隆植物的生态习性,克隆植物的竞争关系等方面的研究进展,并试图在克隆植物竞争关系的背景下,结合其生境状况,探讨植物克隆性与植物物种多样性的关系。克隆植物的等级系统（基株—分株系统—分株）使其具有克隆性所赋予的多样的生活史、资源利用以及空间占据方式。列举了克隆植物的特性及其在群落中的作用。这些例子表明,克隆性强烈地影响和制约着植物群落的空间格局与竞争关系。然而克隆性也能通过权衡活动性与局部持久性来缓解对物种共存的抑制。克隆植物具有在时空尺度上分株间相互调节的机制,直接体现在群落中小尺度（个体与个体间）与大尺度（种群与种群间）间的相互作用上。丰富了传统的竞争和生态位划分理论,为群落中物种共存提供了合理解释。因此克隆植物在群落中的出现拓宽了群落系统潜在机制的范围。     

Responses of Clonal Reef Taxa to Environmental Change

SYNOPSIS. Many reef taxa are predominantly clonal, and theirclones can spread over tens of meters and live for hundredsof years. Colony growth, which produces large colonies, andvegetative propagation, which can generate many clonal replicatesof colonies, affect the response of clonal taxa to climate changethrough a variety of mechanisms, some of which mitigate andsome which amplify effects on individuals. The large numbersof replicate individuals generated among clonal taxa may enablesome individuals to survive catastrophic mortality events suchas storms and then expand following the perturbation. In thosecircumstances clonality buffers the effects of environmentalchange. Conversely, the genetic uniformity of populations dominatedby few genotypes may leave clonal taxa more susceptible to physiologicstress than aclonal taxa. Consequently, clonal species may bemore sensitive to climate change that has chronic and/or acuteeffects on survival. Chronic stresses that reduce recruitmentwill have less obvious effects on clonal taxa than aclonal taxa.Under conditions of reduced recruitment, clonality will allowsome species to persist as relict populations due to the longevityof genets. The presence of relict populations has the appearanceof resistance to climate change. In fact, these taxa are responding,but at a slower rate. The long generation time of genets willslow the pace of evolution among clonal species, making adaptationat projected rates of climate change unlikely. The differentialresponse of species to environmental change will lead to transitionsin community structure as climate changes.     

Roads as Ecological Edges for Rehabilitating Coastal Dune Assemblages in Northern KwaZulu-Natal, South Africa

Abstract Post‐mining coastal dune rehabilitation north of Richards Bay, KwaZulu‐Natal started 20 years before field work for the present study commenced. This resulted in the development of a known age sere of coastal dune forest succession. These rehabilitating areas are fragmented by roads that may act as ecological edges. To establish whether roads affect regenerating bird, millipede, and rodent assemblages, multivariate techniques were applied to test for existence of edge and core assemblages within seral stages representative of the coastal dune forest successional sere. Edge and core assemblages were identified for both the bird and millipede communities but not for the rodent community. Low rodent numbers may have concealed edge effects, but the absence of edge and core assemblages could also be ascribed to the absence of a forest core. In the bird community species composition, richness, density, and total number of species contributed to the identification of edge and core assemblages. Within seral stages the species composition of millipedes differed between the edge and core assemblages. However, if the site was the same age the number of species in edge and core assemblages was similar. The generality of the edge concept should be approached with caution when dealing with taxa comprising species with such diverse natural histories as in the present study. It should also be kept in mind that some species require forest interiors for survival.     

Mycorrhizal fungal growth responds to soil characteristics,but not host plant identity,during a primary lacustrine dune succession

Soil factors and host plant identity can both affect the growth and functioning of mycorrhizal fungi. Both components change during primary succession, but it is unknown if their relative importance to mycorrhizas also changes. This research tested how soil type and host plant differences among primary successional stages determine the growth and plant effects of arbuscular mycorrhizal (AM) fungal communities. Mycorrhizal fungal community, plant identity, and soil conditions were manipulated among three stages of a lacustrine sand dune successional series in a fully factorial greenhouse experiment. Late succession AM fungi produced more arbuscules and soil hyphae when grown in late succession soils, although the community was from the same narrow phylogenetic group as those in intermediate succession. AM fungal growth did not differ between host species, and plant growth was similarly unaffected by different AM fungal communities. These results indicate that though ecological filtering and/or adaptation of AM fungi occurs during this primary dune succession, it more strongly reflects matching between fungi and soils, rather than interactions between fungi and host plants. Thus, AM fungal performance during this succession may not depend directly on the sequence of plant community succession.     

Woody plant encroachment impacts on soil carbon and microbial processes: results from a hierarchical Bayesian analysis of soil incubation data

Quantitative methods were used to examine soil properties and their spatial heterogeneity in a 0-year fenced mobile dune (MD0), an 11-year fenced mobile dune (MD11) and a 20-year fenced mobile dune (MD20) in Horqin Sandy Land, Northern China. The objective of the study was to assess the effect of vegetation restoration on heterogeneity of soil properties in sand dunes and to provide a concept model to describe the relationship between vegetation succession and spatial heterogeneity variation of soil properties in the dunes. The results showed that the average values of vegetation cover, species number and diversity, soil organic carbon (C), total nitrogen (N), and electrical conductivity (EC) increased with the increase in fenced age of mobile dunes, while soil water content (0–20 cm) showed the reverse trend. Geostatistical analysis revealed that the spatial heterogeneity of soil organic C, total N, EC, very fine sand content, and soil water content (0–20 cm) increased from MD0 to MD11 with succession from sand pioneer plant to shrub species then decreased from MD11 to MD20 due to continuous development of herbaceous plants. Canonical correspondence analysis (CCA) showed that there was a relatively high correspondence between vegetation and soil factors, suggesting that the major gradients relating soil organic C, total N, EC, pH, slope, very fine sand content, and soil water content are the main factors for the distribution of dune plants and account for 68.1% of the species-environment relationship among the three sites. In addition, the distribution of the sand pioneer plant was positively related to the relative height of the sampling site and soil water content, and that of most herbaceous plants were determined by soil organic C, total N, EC, pH, and very fine sand content in mobile dunes. The conceptual model of relationship between vegetation succession and spatial heterogeneity of soil properties in mobile dunes suggests spatial patterns of soil properties are most strongly related to plant-induced heterogeneity in dune ecosystems prone to wind erosion, and conversely, the magnitude and degree of spatial heterogeneity in soil properties can influence the plant distribution pattern and vegetation succession of mobile dunes.     

Millipede communities in rehabilitating coastal dune forests in northern KwaZulu/Natal, South Africa

The rehabilitation, after mining, of coastal sand dunes north of Richards Bay by Richards Bay Minerals began some 18 years ago, and resulted in the simultaneous availability of a known-aged series of stands representative of coastal dune forest succession.
A survey of the millipede community in this area revealed increases in species diversity and a decrease in their density, with an increase in stand age. Development of these communities is characterized by replacement and addition of species, typical of ecological succession. Colonization of areas disturbed by mining reflects on the species reservoir present in the surrounding unmined forests with apparent pioneer species being either replaced or complemented by the relatively slow invasion of secondary species.
Comparisons of age-specific millipede community variables on rehabilitating dunes with those recorded in relatively undisturbed dunes suggest that the development of communities results from autogenic succession initiated through habitat rehabilitation. With several community parameters in rehabilitating dune forests being similar to those recorded in undisturbed forests, it is concluded that the millipede community can be restored through management options based on principles relating to ecological succession.     

Richness, diversity, and rate of primary succession over 20?year in tropical coastal dunes

The tropical coastal dunes in central Gulf of Mexico have been stabilizing over the last decades resulting in reduced substrate mobility, and promoting primary succession. We describe changes in species richness and diversity in dune vegetation during 20?years. Our questions: (a) Do species richness and diversity increase over time as predicted by models of ecological succession or do they show a hump-backed manner similar to the observations in temperate coastal dunes?, (b) What is the interaction between vegetation cover and diversity and species richness?, (c) Is there a relationship between species diversity and succession rate and does succession rate change over time?, and (d) How do plant functional types change during succession? In order to answer these questions, we set 140 4?×?4?m permanent plots in a mobile dune area and monitored vegetation cover and species richness from 1991 to 2011. In time, diversity increased in a logistic manner toward an asymptotic value once vegetation cover surpassed 60?%. Species richness increased in a humped-back shape, also reaching a maximum peak at 60?% vegetation cover. The succession rate of diversity was measured by the Euclidean distance, and showed a significant humped-back relation, meaning that it was slower in early and late successional stages. The study supports the intermediate disturbance theory. The conservation of coastal dunes vegetation should focus on all, species-poor and species-rich habitats that help to maintain the ecological integrity of these ecosystems. The understanding of community dynamics and diversity patterns becomes an essential component of coastal dune management and conservation.     

Arbuscular mycorrhizal fungal communities change among three stages of primary sand dune succession but do not alter plant growth

Plant interactions with soil biota could have a significant impact on plant successional trajectory by benefiting plants in a particular successional stage over others. The influence of soil mutualists such as mycorrhizal fungi is thought to be an important feedback component, yet they have shown benefits to both early and late successional plants that could either retard or accelerate succession. Here we first determine if arbuscular mycorrhizal (AM) fungi differ among three stages of primary sand dune succession and then if they alter growth of plants from particular successional stages. We isolated AM fungal inoculum from early, intermediate or late stages of a primary dune succession and compared them using cloning and sequencing. We then grew eight plant species that dominate within each of these successional stages with each AM fungal inoculum. We measured fungal growth to assess potential AM functional differences and plant growth to determine if AM fungi positively or negatively affect plants. AM fungi isolated from early succession were more phylogenetically diverse relative to intermediate and late succession while late successional fungi consistently produced more soil hyphae and arbuscules. Despite these differences, inocula from different successional stages had similar effects on the growth of all plant species. Host plant biomass was not affected by mycorrhizal inoculation relative to un‐inoculated controls. Although mycorrhizal communities differ among primary dune successional stages and formed different fungal structures, these differences did not directly affect the growth of plants from different dune successional stages in our experiment and therefore may be less likely to directly contribute to plant succession in sand dunes.     

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.     

亚高山草甸弃耕地植物群落演替的数量研究I.群落组成分析

本文采用由植被的空间序列推断时间上演替系列的方法对甘南亚高山草甸弃耕地上的植物群落组成进行了分析。结果表明：(1)弃耕初期,莎草,禾草类在群落中的地位从开始时的71%急剧地下降到弃耕十二年时的6%;以后随着密丛型的莎草,禾草的侵入,它们地位又开始上升而恢复到30%一40%：杂类草的地位在开始时上升较快,而到了后期又有所下降;毒害草在群落中的地位基本稳定,保持在10%左右。 (2)群落的种丰富度、均匀度、Simpson多样性指数随弃耕年限增加而增大,而群落优势度随弃耕年限而减小。 (3)优势度—多样性曲线在弃耕初期为几何分布型,中后期演变为对数正态分布型,而对放牧顶极群落又近似于MacArthur分布型。     

Restoration of riverine inland sand dune complexes: implications for the conservation of wild bees

1.  The evaluation of restoration measures is an important task of conservation biology. Inland sand dunes and dry, oligotrophic grasslands have become rare habitat types in large parts of Central Europe and their restoration and management is of major importance for the preservation of many endangered plant and insect species. Within such habitats, it is important to restore key ecosystem services, such as pollination networks. As wild bees are the most important pollinators in many ecosystems, they represent a suitable key group to evaluate restoration measures. Furthermore, the recent decline of many bee species and the potential ecological and economic consequences are currently topics of strong scientific interest.
2.  We studied the succession of bee communities in response to restoration measures of sand dunes and sand grasslands and compared these communities with those of old sand dune complexes.
3.  Our results show that wild bees respond rapidly to restoration measures indicated by a high species richness and abundance. The community structure of bees at restoration sites converged only slightly to those of the target sites. A higher similarity was found between bee communities at the restoration sites (sand dunes and grasslands), indicating that their close proximity was an important determinant of species overlap. Environmental factors such as the number of entomophilous plant species and moisture had a strong influence on wild bee species composition.
4.   Synthesis and applications . The restoration of inland sand dune complexes provides opportunities for colonization by a diverse wild bee community. Although it is difficult to establish a given target community, restoration measures gave rise to a high pollinator diversity and abundance, suggesting that community function can be re-established.     

Pattern analysis in successional communities – An approach for studying shifts in ecological interactions

Abstract. Many ecological studies have addressed issues of vegetation spatial patterns in attempts to understand the processes generating them. We investigated changes in ecological processes during succession via the analysis of shrubs’ spatial patterns in a system of linear sand dunes, an arid ecosystem located in the Negev Desert in Israel during three consecutive years. We hypothesized that spatial patterns change from clustered to regular as succession progresses due to changes in the relative importance of facilitation and competition in this environment. In this ecosystem communities of early successional stages are frequently disturbed by high rates of sand movement, whereas in later successional stages sand stability is high. We mapped in the field individual shrubs on high‐resolution aerial photographs, and converted the digital images to a GIS data set. Using Ripley's K‐function we analysed spatial patterns at three levels: the single‐species level, among species and at the individual level, in three communities characterizing different successional stages. In the early successional communities we found clustered spatial patterns, in comparison with stable habitats where spatial patterns tended to be regular. We argue that these shifts in spatial patterns are indicative of the assumption that in this sand‐dune system ecological interactions change from facilitation to competition as succession progresses. Further, we argue that these interactions operate in different spatial scales at the different successional stages, and that the study of these processes should be conducted at the spatial scales specific to each community.     

Changes in life history trait composition during undisturbed old-field succession

This paper describes the patterns of vegetation change by spontaneous succession on former agricultural land. In 1968, initiated by Heinz Ellenberg, an experimental study site was set up on an arable field in the new botanical garden of Göttingen University (Germany). The undisturbed successions on four plots of a long sere (since 1968/69) and on four additional plots of a shorter sere (since 1982) were evaluated. The seres were classified into stages by cluster analysis, which yield to four subsequent stages for the long sere and to three subsequent stages for the short sere. The early succession is characterized by a high proportion of species invasion, whereas with developing time species extinction increased. On all study plots undisturbed succession directed to the development of pioneer forests. Altogether 247 vascular plant species were recorded. The total species richness shows a fluctuating course during the successional seres, with significantly increased numbers shortly after succession has started and a decline in the pioneer-forest stage, when stands age and close down. The primary aim was to compare the characteristics (particular traits) of species occurring at different stages of succession. The majority of the traits concerned exhibited some clear trends in the course of succession. Exclusive reproduction by seeds decreased and the ability for additional vegetative reproduction increased. Anemochorous dispersal significantly decreased, whereas the importance of dispersal by animals, especially endochorous, dominates in the pioneer-forest stage. The mean seed weight significantly increased. During early succession, plant species staying green over winter dominated, but are of none relevance in the pioneer-forest stage, where summergreen plant species dominate. The change in strategy type features a significant increase of competitive species. The mean indicator value for light in the herb layer decreased significantly in the pioneer-forest stage.     

石羊河下游白刺灌丛演替过程中群落结构及数量特征

白刺灌丛(Nitraria tangutorum Bobr.)是在石羊河下游存活面积最大的天然植被类型,研究不同演替阶段白刺灌丛群落的结构和数量特征可为该区白刺群落保护与恢复提供科学依据.根据白刺灌丛生长及其生境状况,在石羊河下游选择了初期发育、稳定、衰退、严重衰退的群落,研究了各演替过程中各阶段的群落物种组成、生活型谱,群落盖度、群落物种多样性及各演替阶段群落的相似性特征.结果表明,从初期发育阶段到稳定阶段的群落物种数量明显减少,而从稳定阶段至严重衰退阶段的演替过程中群落物种是逐渐增加的.在整个演替过程中,1年生植物所占比例逐渐增加,至严重衰退阶段时1年生植物所占的比重高达80％.不同演替阶段群落盖度最高、最低值出现的月份也不相同,5-10月群落盖度的平均值为稳定阶段＞初期发育阶段＞衰败阶段＞严重衰败阶段.各演替阶段多样性Simpson、Shannon-Wiener指数以及Pielou均匀度指数5-10月均呈现波动变化特征,Simpson、Shannon-Wiener指数大小顺序均表现为初期发育阶段＞严重衰退阶段＞衰退阶段＞稳定阶段,而均匀度Pielou指数为衰退阶段＞严重衰退阶段＞稳定阶段＞初期发育阶段.各演替阶段群落之间的相似性均较低,演替序列越相近,相似性系数越高.     

Importance of clonal plants and plant species diversity in the Northeast China Transect

In plant communities, the internal (genet-level) control mechanisms on a spatio-temporal scale of clonal plants impose strong constraints on spatial pattern as well as on competitive relations and, thus, species coexistence. Therefore, the presence of clonal species within a plant community affects spatio-temporal dynamics and plant species diversity. We examined the distribution of plants with different clonal growth forms in the Northeast China Transect (NECT) and correlated plant species diversity with the importance of clonal plants, and the importance of phalanx and guerilla clonal plants. Phalanx clonal plants were more abundant in western communities where the altitude was higher and both the soil nitrogen contents and precipitation were relatively low. Whereas guerilla clonal plants were more abundant in the middle of the NECT where the precipitation, mean annual temperature and photosynthetically active radiation were relatively high. In the relatively productive temperate typical steppe, plant species diversity was negatively correlated with the importance of phalanx clonal plants and positively correlated with the importance of guerilla clonal plants. In relatively unproductive temperate desert steppe, plant species diversity was positively correlated with the importance of both phalanx and guerilla clonal plants.     

The Association of Dispersal and Persistence Traits of Plants with Different Stages of Succession in Central European Man-Made Habitats

Traits related to seed dispersal, clonality and bud bank affect the success or failure of plant species. Using data from 13 successional seres in various human-made habitats the spectra of traits associated with dispersal and persistence were compared to determine the traits that can be used to predict the occurrence of particular plant species at each stage in a succession and how the importance of these traits changes over time. Differences in the traits of species associated with primary and secondary successions were also studied. Species with seeds that are dispersed by water (hydrochory) decreased in abundance during the course of succession. Species with a splitting main root, monocyclic and dicyclic shoots also decreased in abundance. Species capable of forming a potential below-ground bud bank, hypogeogenous rhizome and retaining a long-term connection with clonal offspring increased in abundance. The results indicate that seed dispersal is more important in determining the species composition in the early stages of succession whereas bud banks and clonal traits are more important in the later stages and for colonizing a locality. Primary and secondary seres did not remarkably differ in the trait spectra of the species present indicating that these trends occur in both types of succession.     

科尔沁沙地不同演替阶段冷蒿群落的结构特征

采用空间序列替代时间序列的方法,对科尔沁沙地不同演替阶段3个冷蒿群落的物种组成和物种多样性进行研究.结果表明:处于3个演替阶段的冷蒿群落为杠柳-冷蒿群落、杠柳-冷蒿+差巴嘎蒿群落及冷蒿-糙隐子草群落.3个冷蒿群落均以菊科、禾本科和藜科植物占优势.随着群落的演替,群落植物种数和科数逐渐减小,禾本科植物的种数逐渐减小,但菊科和禾本科植物种数所占比例逐渐增大.3个冷蒿群落中物种多样性顺序为:杠柳-冷蒿+差巴嘎蒿群落>杠柳-冷蒿群落>冷蒿-糙隐子草群落.3个冷蒿群落间的物种组成差异不大,群落间物种相似度较高.3个群落中冷蒿的盖度最大,其盖度随着群落的演替逐渐增大.     

Remnant Populations and Plant Functional Traits in Abandoned Semi-Natural Grasslands

Although semi-natural grasslands in Europe are declining there is often a time delay in the local extinction of grassland species due to development of remnant populations, i.e., populations with an extended persistence despite a negative growth rate. The objectives of this study were to examine the occurrence of remnant populations after abandonment of semi-natural grasslands and to examine functional traits of plants associated with the development of remnant populations. We surveyed six managed semi-natural grasslands and 20 former semi-natural grasslands where management ceased 60–100 years ago, and assessed species response to abandonment, assuming a space-for-time substitution. The response of species was related to nine traits representing life cycle, clonality, leaf traits, seed dispersal and seed mass. Of the 67 species for which data allowed analysis, 44 species declined after grassland abandonment but still occurred at the sites, probably as remnant populations. Five traits were associated with the response to abandonment. The declining but still occurring species were characterized by high plant height, a perennial life form, possession of a perennial bud bank, high clonal ability, and lack of dispersal attributes promoting long-distance dispersal. Traits allowing plants to maintain populations by utilizing only a part of their life cycle, such as clonal propagation, are most important for the capacity to develop remnant populations and delay local extinction. A considerable fraction of the species inhabiting semi-natural grasslands maintain what is most likely remnant populations after more than 60 years of spontaneous succession from managed semi-natural grasslands to forest.     

Species and functional trait re-assembly of ground beetle communities in restored grasslands

Ecosystem restoration provides unique opportunities to study community dynamics under succession and can reveal how consumer communities re-assemble and respond to successional changes. Studying community dynamics from both taxonomic and functional trait perspectives also may provide more robust assessments of restoration progress or success and allow cross-system comparisons. We studied ground beetle (Coleoptera: Carabidae) communities for three years in a restored grassland chronosequence with sites from 0 to 28 years old. We measured traditional community metrics (abundance, richness, Shannon diversity) and functional trait metrics based on species’ body length, wing morphology, activity time, phenology, and diet. Communities had high species richness and abundance in early successional stages, but these declined in later stages to low levels comparable to an adjacent grassland remnant. Species composition also shifted with time, converging with the remnant. Although functional richness, like species richness, declined as succession progressed, functional divergence quickly increased and was maintained over time, suggesting niche differentiation in established communities. Young sites were typified by small, macropterous, phytophagous species, while older sites contained larger species more likely to be flightless and carnivorous. Prescribed burns also affected traits, decreasing prevalence of larger species. This study demonstrates that functionally diverse consumer communities can self-assemble under restoration practices. In a relatively short amount of time both morphological and trophic level diversity are established. However, prescribed fire intended to control non-desirable plants may also shape beetle community functional composition, and restoration managers should consider if plant community benefits of fire outweigh potential declines in consumer function.     

Soil oxidases recovered faster than hydrolases in a 50-year chronosequence of desert revegetation

Aims

Desert characterized by alkaline soil with low organic matter and nutrients has a high soil oxidative potential. We hypothesized that oxidase activities would recover faster than hydrolases during the succession of sand-fixing community.

Methods

Sand dunes stabilized in different years, including a moving sand dune and a steppe at the southeastern fringe of the Tengger Desert, were selected to investigate restoration of extracellular enzyme activities (EEAs) in a 50-year chronosequence.

Results

Oxidases showed significantly higher activities than hydrolases at all ten studied sites and EEAs exibited a decreasing trend from catalase, phenol oxidase, sucrase, urease, alkaline phosphatase, α-Amylase to cellulase. After 50 years of revegetation, most EEAs in topsoil recovered to 50–83% of that of the steppe except for urease. Oxidase activities recovered earlier and faster than hydrolases, while hydrolases activities attained the fastest recovery at 19–25 years in the 50-year chronosequence.

Conclusions

Recovery of EEAs was modulated by the succession of the sand-fixed community: oxidases activities exhibited peak recovery rates at the stage when shrubs dominated the community, while recovery of hydrolases activities appeared to be mainly regulated by biological soil crusts and annual plants.