Correspondence of seed traits with niche position in glacier foreland succession

Changes in species composition during succession are driven by biotic and abiotic factors leading to a multitude of niches occupied by distinct species. Gradient analyses of plant communities provide opportunities to approximate the niche position of species along a successional gradient. Several plant traits have been used to explain mechanisms governing successional sequences, but generalising changes in species traits during primary succession is still controversial. This study examined whether the seed mass and the optimum temperature for germination could explain the niche position of several glacier foreland species along a primary successional gradient in the Austrian Central Alps. We hypothesised that pioneer species should possess lighter seeds and a lower optimum temperature for germination than late successional species. We found significant differences in the seed mass between species, but the seed mass did not correspond with the assigned niche position on the successional gradient. Germination responses to temperature also differed significantly between species. Pioneer species performed better at lower temperatures than late successional species, suggesting that the optimum temperature for germination is a driver of niche separation. We discuss the interactions between seed traits and environmental conditions along the primary successional gradient emphasising the importance of temperature requirements for the germination. Differences in the regeneration characteristics are a major cue governing species turnover in glacier foreland succession.     

Changes in the structure and heterogeneity of vegetation and microsite environments with the chronosequence of primary succession on a glacier foreland in Ellesmere Island, high arctic Canada

Primary plant succession was investigated on a well-vegetated glacier foreland on Ellesmere Island in high arctic Canada. A field survey was carried out on four glacier moraines differing in time after deglaciation to assess vegetation development and microsite modification in the chronosequence of succession. The results showed evidence of directional succession without species replacement, which is atypical in the high arctic, reflecting the exceptionally long time vegetation development. During this successional process, Salix arctica dominated throughout all moraines. The population structures of S. arctica on these moraines implied the population growth of this species with progressing succession. The population density of S. arctica reflected the abundance of vascular plants, suggesting that development of the plant community might be related to structural changes and the growth of constituting populations. Through such growths of the population and the whole community with progressing succession, the spatial heterogeneity of vegetation gradually declines. Moreover, this vegetation homogenization is accompanied by changes in the spatial heterogeneity of microsite environments, suggesting significant plant effects on the modification of microsite environments. Accordingly, it was concluded that the directional primary succession observed on this glacier foreland is characterized by the initial sporadic colonization of plants, subsequent population growths, and the community assembly of vascular plants, accompanied by microsite modification.     

Can CSR classification be generally applied outside Britain?

The general applicability of CSR classification (a methodology for applying CSR theory) has not been tested outside Britain. We hypothesised that principal axes of trait variation for plant species native to southern European continental, sub-alpine and alpine bioclimatic zones correlate with CSR classification scores. Functional traits were quantified in situ for 506 species of 57 families. Multivariate analysis revealed three main modes in which plants invested biomass: (1) architectural extension of individual ramets (tall canopies, large leaves), (2) durable tissues (high leaf dry matter contents and carbon contents, low specific leaf areas) or (3) regenerative development (early, extensive flowering with delicate, nitrogen-rich leaves). These were highly significantly correlated with the extent of competitive ability (C), stress-tolerance (S) and ruderalism (R), respectively, determined by CSR classification. Despite being calibrated using only 43 British species, CSR classification is consistent with trait variation in a phylogenetically broader sample of species native to a wider range of bioclimatic zones.     

Life in harsh environments: carabid and spider trait types and functional diversity on a debris‐covered glacier and along its foreland

1. Patterns of species richness and species assemblage composition of ground‐dwelling arthropods in primary successions along glacier forelands are traditionally described using a taxonomic approach. On the other hand, the functional trait approach could ensure a better characterisation of their colonisation strategies in these types of habitat. 2. The functional trait approach was applied to investigate patterns of functional diversity and life‐history traits of ground beetles and spiders on an alpine debris‐covered glacier and along its forefield in order to describe their colonisation strategies. 3. Ground beetles and spiders were sampled at different successional stages, representing five stages of deglaciation. 4. The results show that the studied glacier hosts ground beetle and spider assemblages that are mainly characterised by the following traits: walking colonisers, ground hunters and small‐sized species. These traits are typical of species living in cold, wet, and gravelly habitats. The diversity of functional traits in spiders increased along the succession, and in both carabids and spiders, life‐history traits follow the ‘addition and persistence model’. Accordingly, there is no turnover but there is an addition of new traits and a variation in their proportion within each species assemblage along the succession. The distribution of ground beetles and spiders along the glacier foreland and on the glacier seems to be driven by dispersal ability and foraging strategy. 5. The proposed functional approach improves knowledge of the adaptive strategies of ground‐dwelling arthropods colonising glacier surfaces and recently deglaciated terrains, which represent landforms quickly changing due to global warming.     

基于功能性状的毛乌素沙地不同演替阶段适生植物筛选

沙地生态系统修复是恢复生态学研究的热点问题,适生植物筛选是修复的关键。植物功能性状反映了植物在不同环境中的生存策略,探究沙地植物功能性状及其与环境之间的关系,有助于筛选用于植被恢复的物种,为保护沙地生态系统提供理论依据。以毛乌素沙地为研究区,分析了1983-2015年间沙地典型飞播样地群落演替特征及其对环境因子的响应,建立基于10个植物功能性状的毛乌素沙地潜在种库,进一步筛选飞播恢复下沙地不同演替阶段的适生植物。研究表明:(1)飞播恢复下的毛乌素沙地植物群落分为三个演替阶段:固沙先锋物种群落、沙生植物为主的杂类草群落、中生植物为主的杂类草群落。(2)土壤因子是群落演替的主要驱动力,其中土壤全氮、土壤总有机碳、土壤硝态氮是影响群落演替的关键因素。(3)基于功能性状筛选出29种适生物种用于植被恢复,演替第一阶段可用雾冰藜、猪毛菜等,演替第二阶段可用拂子茅、无芒隐子草等,演替第三阶段可用草地风毛菊、猪毛蒿等。通过物种功能性状特征可以快速选择适合沙地退化生态系统修复的候选物种,为植被恢复提供了一定的理论支持。     

Patterns of Primary Succession on the Foreland of Coleman Glacier,Washington, USA

Patterns of community development vary among studied glacier forelands around the world. However, there have been few studies of primary succession on glacial forelands in temperate regions of North America. We described patterns in community composition, vegetation cover, diversity, and vegetation heterogeneity during primary succession on the foreland of Coleman Glacier, in Washington State, USA. Community composition changed rapidly with high turnover between age classes. Cover increased through succession as expected. Species richness and diversity were highest in early succession at small scales and in late succession at larger scales. At small scales, heterogeneity decreased in early succession but increased in mature sites. At larger scales, heterogeneity reached its lowest point earlier in succession. These scale-dependent patterns in diversity and heterogeneity differ from results of other studies of glacier forelands. We hypothesize that these patterns arise due to the development of a dense canopy of the deciduous shrub Alnus viridis followed by a dense canopy of Abies amabilis, Tsuga heterophylla, and Pseudotsuga menziesii.     

Early recruitment equals long-term relative abundance in an alpine saxicolous lichen guild

The relative abundance within a guild of six species of the lichen-forming fungal genus Umbilicaria was studied during primary colonization of rock surfaces in a chronosequence of ca. 245 y in an alpine glacier foreland in southern Norway. Although the density of the guild grew from zero to more than 1000 thalli/m2 and the species differ substantially in life history traits such as initial growth rate, maximal size, maturation rate and propagule types, the relative abundance among the species remained almost unchanged through those years. The relative abundance of species is correlated with their life history parameters, such as initial growth rate and size-related maturation. The pattern of relative abundance was also similar in the saxicolous communities outside the foreland, which are potentially several thousand years old. Outside the foreland however the density of the guild is only 1/10 of that in the oldest parts of the foreland, due to soil formation and vegetation growth that have covered many of the low profile rock habitats. Thus the areas affected by the disturbance of glacier expansions and retreats provide temporary opportunities for large increases in the population sizes of the members of the saxicolous community. The observations support the view that pre-emption of habitat rather than competitive exclusion is common in saxicolous lichen communities and that "succession" consists in the addition but rarely the loss of species.     

贡嘎山冰川退缩区植物-土壤反馈与植物间相互作用

植物-土壤反馈是揭示陆地生物群落动态变化的关键环节,为理解植物间相互作用及植被群落变化过程奠定基础。本研究以贡嘎山冰川退缩区原生演替早(5~10年)、中(30~40年)和晚期(80~100年)3个阶段典型土壤以及各阶段优势植物为对象,采用盆栽控制试验,比较优势植物在不同土壤条件下的生物量,并量化植物间相互作用以及植物-土壤反馈的方向与强度,为探究贡嘎山冰川退缩区植被群落演替规律提供依据。结果表明:(1)植物-土壤反馈作用显著影响植物在本土中的生物量,早期沙棘(Hippophae rhamnoides)在本土中生长最差,沙棘的植物-土壤反馈系数为负值;演替中期冬瓜杨(Populus purdomii)的反馈系数趋于零;晚期峨眉冷杉(Abies fabri)在本土中生长最好,峨眉冷杉的反馈系数为正值。(2)混种时,早期沙棘与演替中、晚期植物间相互作用指数为负值;中期冬瓜杨、川滇柳(Salix rehderiana)与演替早、晚期植物的相互作用指数接近于零,晚期植物峨眉冷杉、麦吊云杉(Picea brachytyla)与演替早、中期植物相互作用指数为正值。从植物-土壤反馈的方向来看,贡嘎山植被演替从早期负反馈,中期中性反馈,过渡到晚期正反馈。此外,演替早期沙棘促进演替中晚期植物生长,演替中期冬瓜杨、川滇柳对演替早晚期植物无显著影响,晚期峨眉冷杉、麦吊云杉更利于与演替早中期植物相互竞争。结果显示,植物-土壤反馈与植物间相互作用共同驱动了贡嘎山冰川退缩区植被快速演替,直至顶极群落。     

Microbial succession of nitrate-reducing bacteria in the rhizosphere of Poa alpina across a glacier foreland in the Central Alps

Changes in community structure and activity of the dissimilatory nitrate-reducing community were investigated across a glacier foreland in the Central Alps to gain insight into the successional pattern of this functional group and the driving environmental factors. Bulk soil and rhizosphere soil of Poa alpina was sampled in five replicates in August during the flowering stage and in September after the first snowfalls along a gradient from 25 to 129 years after deglaciation and at a reference site outside the glacier foreland (>2000 years deglaciated). In a laboratory-based assay, nitrate reductase activity was determined colorimetrically after 24 h of anaerobic incubation. In selected rhizosphere soil samples, the community structure of nitrate-reducing microorganisms was analysed by restriction fragment length polymorphism (RFLP) analysis using degenerate primers for the narG gene encoding the active site of the membrane-bound nitrate reductase. Clone libraries of the early (25 years) and late (129 years) succession were constructed and representative clones sequenced. The activity of the nitrate-reducing community increased significantly with age mainly due to higher carbon and nitrate availability in the late succession. The community structure, however, only showed a small shift over the 100 years of soil formation with pH explaining a major part (19%) of the observed variance. Clone library analysis of the early and late succession pointed to a trend of declining diversity with progressing age. Presumably, the pressure of competition on the nitrate reducers was relatively low in the early successional stage due to minor densities of microorganisms compared with the late stage; hence, a higher diversity could persist in this sparse environment. These results suggest that the nitrate reductase activity is regulated by environmental factors other than those shaping the genetic structure of the nitrate-reducing community.     

Ecosystem development in short‐term postglacial chronosequences: N and P limitation in glacier forelands from Santa Inés Island,Magellan Strait

Glacier foreland moraines provide an ideal model to examine the patterns of ecosystem development and the evolution of nitrogen and phosphorous limitation over successional time. In this paper, we focus on a 400‐year soil chronosequence in the glacier forelands of Santa Inés Island in the Magellan Strait, southern Chile by examining forest development on phosphorus (P)‐poor substrates in a uniquely unpolluted region of the world. Results show a steady increase in tree basal area and a humped trend in tree species richness over four centuries of stand development. The increase in basal area suggests that the late successional tree species were more efficient nutrient users than earlier successional ones. Total contents of carbon (C) and nitrogen (N) in soils increased during the chronosequence, reaching an asymptote in late succession. The net increases in soil C : N, C : P and N : P ratios observed over successional time suggest that nutrient limitation is maximal in 400‐year‐old substrates. Foliar C : N and C : P ratios also increased over time to reach an asymptote in old‐growth stages, following soil stoichiometric relationships; however the foliar N‐to‐P ratio remained constant throughout the chronosequence. Biological N fixation was greater in early postglacial succession, associated with the presence of the symbiotic N‐fixer Gunnera magellanica. Declining trends of δ¹⁵N in surface soils through the 400‐year chronosequence are evidence of decreasing N losses in old‐growth forests. In synthesis, glacier foreland chronosequences at this high South American latitude provide evidence for increasing efficiency of N and P use in the ecosystem, with the replacement of shade‐intolerant pioneers by more efficient, shade‐tolerant tree species. This pattern of ecosystem development produces a constant foliar N : P ratio, regardless of variation in soil N‐to‐P ratio over four centuries.     

海岛植物不同演替阶段植物功能性状与环境因子的变化规律

植物功能性状与环境之间的关系是功能性状研究的重点,环境因子驱使植物功能性状发生变化,进而推动群落发生演替。以平潭岛4个不同演替阶段的森林植被(灌草丛、针叶林、针阔混交林、常绿阔叶林)为研究对象,结合不同群落演替阶段的物种特征和群落结构,分析海岛不同演替阶段茎、叶功能性状的变化规律,以及功能性状与环境因子的关系。结果表明:(1)随着演替的进行,土壤养分和水分逐渐增加,土壤pH逐渐下降。比叶面积(SLA)、叶片氮含量(LNC)、叶片磷含量(LPC)、茎氮含量(SNC)、茎磷含量(SPC)下降后上升,叶厚度(LT)、叶片碳含量(LCC)、茎碳含量(SCC)与之相反,叶干物质含量(LDMC)、茎组织密度(STD)逐渐上升。(2)冗余分析表明,演替早期植物主要分布在土壤pH、容重高的贫瘠环境,拥有较高SLA、SNC、SPC、LPC的性状组合;演替后期植物主要分布在土壤养分和水分高的肥沃环境,拥有较高的STD、LDMC、LCC、LNC的性状组合。其中,土壤有机质和全氮含量是影响海岛植物演替过程中功能性状变化的关键环境因子。研究海岛植物功能性状与环境之间的关系随演替的变化规律,探讨各演替阶段功能性状和环境特征,以及功能性状如何响应环境变化。旨在为今后选择合适的树种进行海岛植被修复和重建提供依据。     

冰川消退带微生物群落演替及生物地球化学循环

冰川是生物圈重要组分之一。由于全球气候变化世界多地冰川加速消融,暴露原本被冰盖覆盖的区域,这些区域被称为冰川消退区域(glacier retreat area)或冰川前部区域(glacier foreland)。自暴露开始消退区随即发生初生演替,随着演替进行,物质循环逐步建立,生物量和土壤C、N总量逐步增加。生态系统C、N输入最初以矿化外来物为主,逐渐转变为以生物固C、固N为主。演替早期生态系统的发育主要受土壤C、N含量的限制,而演替后期的限制性营养物转变为P。演替区域土壤逐渐发育并促进生态位的分化,细菌、真菌、古菌,病毒及其他微生物群落的生物量和多样性不断增加直至达到该地区可承受的极值。随着生存条件的改善,不同生态策略物种的更替导致每个演替阶段微生物群落结构的差异。整体上,伴随演替进行微生物群落丰度、结构和活性呈现梯度性变化。气候变化对冰川消退带生态演替结果产生多方面的影响,而这些影响结果又综合反馈气候变化,因此目前难以准确估计气候变化对消退带生态演替的净效应。综述了近年冰川消退带微生物群落演替方面相关的研究结果,同时分别对该区域物质循环的建立、微生物群落演替和气候变化造成的影响这三个方面进行详细描述,并指出当前研究的不足。     

黄土高原植被不同演替阶段优势种的光合生理特性

在黄土高原子午岭林区,按照植被的演咎序列,分别选择植被次生演替不同阶段的优势种-白羊草、铁杆蒿、沙棘、狼牙刺、山杨和辽东栎,对其土壤与叶片氮素含量、叶片气体交换参数、叶绿素含量及叶绿素荧光参数进行了观测．结果表明：随着植被演替,0—20cm土壤全氮含量呈增大趋势,植被优势种叶片氮含量先升高后降低,叶绿素含量变化与优势种叶片氮含量相似．灌木群落优势种（沙棘、狼牙刺）叶片氮含量明显高于其它群落优势种（P〈0．05）,草本群落优势种（白羊草、铁杆蒿）和灌木群落优势种的光合速率（Pn）高于早期森林群落优势种（山杨）和顶级群落优势种（辽东栎）,叶片气孔导度（ga）与Pn变化规律相似．草本群落和灌木群落优势种叶片蒸腾速率（Tr）较高,随着演替的进行t逐步下降,顶级群落优势种辽东栎Tr较低．PSⅡ最大光化学量子效率（Fv／Fm）值呈缓慢增长趋势,辽东栎〉山杨〉沙棘〉狼牙刺〉铁杆蒿〉白羊草,而PSⅡ量子效率（ΦPSⅡ）值呈先增加后降低趋势;光化学淬灭系数（qp）在演替过程中呈整体增加趋势,而非光化学淬灭系数（qNP）呈先增加后降低趋势．这说明植被不同演替阶段优势种的光合生理特征以及对微生境的适应性存在明显差异．     

Plant succession and rhizosphere microbial communities in a recently deglaciated alpine terrain

This study describes how early and late successional plant species affect soil microorganisms in alpine ecosystems. We quantify the relative importance of plant species and soil properties as determinants of belowground microbial communities. Sixteen plant species were selected from six successional stages (4–14–20–43–75–135 years) within the foreland of the Rotmoosferner glacier, Austria, and at one (reference) site outside the foreland. The size, composition and function of the communities of microorganism in the bulk soil and the rhizosphere were characterized by ninhydrin-reactive nitrogen, phospholipid fatty acids and enzyme activities (β-glucosidase, β-xylosidase, N-acetyl-β-glucosaminidase, leucine aminopeptidase, acid phosphatase, sulphatase). The results show that the microbial data could be grouped according to early (up to 43 years) and late-colonizing plant species (75 or more years). In early succession, no plant species or soil age effect was detected on the microbial biomass, phospholipid fatty acids, or enzyme activity. The rhizosphere microbial community was similar to that in the bulk soil, which in turn was determined by the abiotic environmental conditions. In late succession, improved soil conditions probably mediated plant species effects on the belowground microbial community. The most pronounced rhizosphere effects were attributed to plant species of the 75- and 135-year-old sites. The microbial colonization (size, composition, activity) of the bulk soil predominantly followed changes in vegetation cover, plant life forms and soil organic matter. In summary, the observed successional pattern of the above- and belowground communities provides an example of the facilitation models of primary succession.     

青藏高原东缘海螺沟冰川退缩区植被原生演替的氮磷动态

在青藏高原东缘海螺沟冰川退缩区植被原生演替序列的典型地段依次设置7个采样点(冰川退缩0、10、30、40、50、80和127年),测定各演替阶段表层土壤与植被各层氮(N)、磷(P)含量及生态系统生物量,分析不同演替阶段N、P养分的积累与循环特征.结果表明:乔木层叶、枝、根的N、P含量随演替下降,而干的N、P含量在演替末...     

Environmental gradients and succession patterns of carabid beetles (Coleoptera: Carabidae) in an Alpine glacier retreat zone

Accelerated by global warming, retreating glaciers leave behind spatially ordered moraines with underlying primary succession and disturbance. Current knowledge of primary succession comes mainly from studies of vegetation dynamics. Information about above-ground macroinvertebrates is still scarce. We used carabid beetles (Coleoptera; Carabidae) as indicator taxon to assess the effects of (1) terrain age (species turnover along the proglacial chronosequence) and (2) small-scale habitat architecture (vegetation cover, surface texture) on the carabid assembly. For this purpose, 33 sampling sites with pitfall traps were installed throughout the glacier foreland Morteratsch (Engadine, Switzerland), adjacent sparse forests serving as reference sites. With a total of 33 carabid species on the foreland and another 2 on the reference sites, the study area yielded a very high carabid species diversity compared to other glacier forelands. In general, the age of deglaciation proved to be a highly significant predictor for the carabid distribution, especially for particularly discriminant species. Observed species richness and activity densities showed bimodal patterns with a steep increase within the first ca. 40?years, a decline between around 40–90?years, and a further increase towards the terminal moraine. There was no evidence of dispersal-stochasticity: distinct clusters of sites with similar species composition were found. Microhabitat suitability proved to be a secondary effect, embedded in a temporal framework of primary succession. Surface cover with litter, herbs and dwarf-shrubs turned out to be the crucial habitat factors. Habitat loss as a result of climate warming will primarily affect cold-stenotopic carabids, but may potentially be absorbed by active selection for cooler microhabitats.     

茂兰喀斯特森林不同演替阶段植物叶片功能性状与土壤因子的关系

植物如何改变功能性状来适应环境一直是生态学的研究征点。为探究茂兰喀斯特森林不同演替阶段植物叶片的适应策略，该文以茂兰自然保护区5个不同演替阶段(草本、灌木、灌乔、乔木和顶极群落阶段)优势种为研究对象，测定不同演替阶段的优势植物叶片功能性状与土壤理化性质。结果表明：(1)随着植被正向演替的进行，土壤全氮(STN)含量、土壤有机质(SOM)含量、土壤含水量(SWC)逐渐增加，土壤全磷(STP)含量和土壤全钾(STK)含量先增加后减少，土壤pH值整体呈减小的趋势。(2)随着植被演替的进行，叶面积(LA)、叶干物质含量(LDMC)、叶厚度(LT)和叶片碳含量(LCC)逐渐上升，比叶面积(SLA)与叶片钾含量(LKC)则与之相反，叶片氮含量(LNC)呈先升后降的趋势，叶片磷含量(LPC)呈先降后升的趋势。(3)冗余分析表明，演替初期植物主要分布在土壤pH值高而STK、STP、SWC、SOM、STN相对低的环境中，群落内植物叶片采取高SLA、LNC、LPC,低LA、LDMC、LT、LWC的性状组合，演替晚期植物主要分布在土壤水分和养分含量较高的环境，LDMC、LT、LA、LWC与演替初期相比呈上升...     

Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats

Changes in traditional agricultural systems in Europe in recent decades have led to widespread abandonment and colonization of various habitats by shrubs and trees. We combined several vegetation databases to test whether patterns of changes in plant diversity after land abandonment in different habitats followed similar pathways. The impacts of land abandonment and subsequent woody colonization on vegetation composition and plant traits were studied in five semi-natural open habitats and two arable habitats in six regions of Europe. For each habitat, vegetation surveys were carried out in different stages of succession using either permanent or non-permanent plots. Consecutive stages of succession were defined on a physiognomic basis from initial open stages to late woody stages. Changes in vegetation composition, species richness, numbers of species on Red Lists, plant strategy types, Ellenberg indicator values of the vegetation, Grime CSR strategy types and seven ecological traits were assessed for each stage of the successional pathway. Abandonment of agro-pastoral land-use and subsequent woody colonization were associated with changes in floristic composition. Plant richness varied according to the different habitats and stages of succession, but semi-natural habitats differed from arable fields in several ecological traits and vegetation responses. Nevertheless, succession occurred along broadly predictable pathways. Vegetation in abandoned arable fields was characterized by a decreasing importance of R-strategists, annuals, seed plants with overwintering green leaves, insect-pollinated plants with hemi-rosette morphology and plants thriving in nutrient-rich conditions, but an increase in species considered as endangered according to the Red Lists. Conversely, changes in plant traits with succession within the initially-open semi-natural habitats showed an increase in plants thriving in nutrient-rich conditions, stress-tolerant plants and plants with sexual and vegetative reproduction, but a sharp decrease in protected species. In conclusion, our study showed a set of similarities in responses of the vegetation in plant traits after land abandonment, but we also highlighted differences between arable fields and semi-natural habitats, emphasizing the importance of land-use legacy.