Facilitation in subnival vegetation patches

Abstract. We examined spatial relationships among species in the subnival zone of the central Caucasus. The species composition of 300 vegetation patches was analysed. 144 of them contained only one species, whereas the other 156 contained 2.36 ± 1.31 species, with species numbers distributed as follows: 59 patches with two species, 41 with three, 39 with four and 17 with five species. In the multi-species patches, the 22 most frequent species were examined and 46 statistically significant species associations, 36 positive and 10 negative, were found. Ten of these 22 species were typical subnival plants very rarely occurring at lower altitudes. The other 12 species are ‘invaders’ as they have broader altitudinal ranges of occurrence and are common in alpine and even in subalpine belts. Contrary to the typical subnival species the invaders were found exclusively in the patches with more than one species. Invader species were significantly more associated with subnival species while they were significantly less associated with other species than expected by chance. Our interpretation is that typical subnival species nurse plants from lower altitudes and facilitate their invasion to more adverse subnival environments.     

Interspecific primate associations in Amazonian flooded and unflooded forests

Stable associations between two or more primate species are a prominent feature of neotropical forest vertebrate communities and many studies have addressed their prevalence, and their costs and benefits. However, little is known about the influence of different habitat types on the frequency, seasonality, and composition of mixed-species groups in Amazonian forest primates. Here we examine the features of interspecific primate groups in a large mosaic of flooded (várzea and igapó) and unflooded (terra firme) forest in central Amazonia. In total, 12 primate species occurred in the study area, nine of which were observed in mixed-species associations. Primates were more than twice as likely to form associations in várzea forest than in terra firme forest. Squirrel monkeys were most frequently found in mixed-species groups in all forest types, most commonly in association with brown capuchins. Another frequent member of interspecific associations was the buffy saki, which often formed mixed-species groups with tamarins or brown capuchins. There was no seasonality in the frequency of associations in terra firme forest whereas associations in várzea forest were twice as frequent during the late-dry and early-wet seasons than in the late-wet and early-dry seasons. Interspecific primate associations were common in all forest types, but the degrees to which different species associate varied between these environments. We suggest that the temporal variation of várzea forest associations is connected with seasonal changes in habitat structure and resource abundance. However, more work is needed to pinpoint the underlying causes of mixed-species associations in all forest types and their strong seasonality in várzea forest.     

川西亚高山次生林恢复过程中乔木层优势种变化及其影响因子

川西亚高山森林在20世纪下半叶遭受长时期连续砍伐,形成了采伐迹地上天然次生林恢复演替序列。该研究采用空间代替时间方法,以立地条件相似的暗针叶林作为对照,分析川西亚高山暗针叶林采伐迹地上次生林自然恢复过程中乔木层优势种组成及其影响因子。结果表明:(1)研究区亚高山天然次生林恢复演替过程中,不同演替阶段土壤各养分差异显著,其中土壤总氮、土壤总磷、土壤有效磷、土壤有效氮均在针阔混交林阶段最高,土壤pH值的变化在5.42～7.39范围波动,土壤有机碳随恢复的进行而增加,土壤C/N主要集中在15左右,只有针叶林达到18.62。(2)研究区次生林以16个乔木树种为主,针叶树种主要有粗枝云杉、岷江冷杉,阔叶树种主要有红桦、青榨槭等,且不同恢复阶段的群落乔木层优势种差异显著(R_(ANOSINM)=0.439,P0.001)。(3)多元回归树分析发现,次生林乔木层优势种组成主要受林龄、土壤有机碳和总氮的影响,其中在次生林恢复演替后期(80a后),乔木层组成可能主要由于种间竞争导致,在次生林恢复演替前、中期(80a前),环境选择可能是影响乔木层组成差异的主导因素。(4)反向选择筛选结果显示,土壤总氮、海拔高度、土壤有效磷和土壤有机碳是影响乔木层优势种差异的主要因子。     

Phylogenetic and functional structures of succession in plant communities on mounds of Marmota himalayana in alpine regions on the northeast edge of the Qinghai–Tibet Plateau

Few studies have examined the succession of plant communities in the alpine zone. Studying the succession of plant communities is helpful to understand how species diversity is formed and maintained. In this study, we used species inventories, a molecular phylogeny, and trait data to detect patterns of phylogenetic and functional community structure in successional plant communities growing on the mounds of Himalayan marmots (Marmota himalayana) on the southeast edge of the Qinghai-Tibet Plateau. We found that phylogenetic and functional diversities of plant communities on marmot mounds tended to cluster during the early to medium stages of succession, then trended toward overdispersion from medium to late stages. Alpine species in early and late stages of succession were phylogenetically and functionally overdispersed, suggesting that such communities were assembled mainly through species interactions, especially competition. At the medium and late stages of succession, alpine communities growing on marmot mounds were phylogenetically and functionally clustered, implying that the communities were primarily structured by environmental filtering. During the medium and late stages of succession the phylogenetic and functional structures of plant communities on marmot mounds differed significantly from those on neighboring sites. Our results indicate that environmental filtering and species interactions can change plant community composition at different successional stages. Assembly of plant communities on marmot mounds was promoted by a combination of traits that may provide advantages for survival and adaptation during periods of environmental change.     

缙云山森林次生演替序列群落的物种多样性动态

通过１９８６ 和１９９６ 年对缙云山森林次生演替系列群落的重复调查,运用构成指数对这些群落进行演替排序,研究缙云山森林次生演替过程中的物种多样性动态．结果表明,构成指数（ ＣＩ） 可适当改变空间代替时间的局限;在缙云山森林次生演替过程中,物种丰富度呈先变大后变小的趋势,物种均匀度没有明显的动态规律,这与种间（ 内） 关系密切相关,因而适当干扰有利于群落内物种多样性的增大     

广东黑石顶自然保护区森林次生演替过程中群落的种间联结性分析

 采用2×2列联表，通过方差分析、X2检验、Ochiai指数对黑石顶森林次生演替过程中各阶段群落优势种的种间联结性进行了定量分析。研究结果表明：群落的总体种间关联性在演替恢复初期表现为无相关性，针阔叶混交林阶段为总体的正关联，阳生性常绿阔叶林阶段呈总体负关联，中生性常绿阔叶林阶段总体呈显著正关联。在演替初期主要种对间尚未形成一定的种间关系，演替至混交林和阳生性常绿阔叶林阶段，种间关系开始形成，但大多表现为直接的竞争关系，中生性常绿阔叶林阶段形成了复杂多样的种间关系，表现为同层树种间的种间结合或竞争和不同层种间的结合关系。     

植物种间联结研究内容与方法评述

植物种间联结是植物群落重要的数量和结构特征之一,是植物群落结构形成和演化的基础。研究种间联结,对于植物群落的稳定与演替、植被的恢复与重建等具有重要的理论意义和应用价值。为了便于探讨种间联结的形成机制,本文评述了种间联结的研究内容、计算方法及其影响因素,同时指出了现研究存在的问题及今后的研究方向。植物种间联结受多方面因素影响,既有人为取样尺度的影响,又有环境与生物自身的因素。在群落演替的不同阶段,种间联结性往往有所不同。为此,可采用多尺度排序法分析不同尺度格局上复杂的种间联结关系,重点加强植物"利他作用"及入侵种种间联结的研究。在Fisher精确检验的基础上,结合Pearson相关系数和Spearman秩相关系数来检验种对间的相关性,并利用生态位理论或从植物生理、分子遗传的角度研究密度主导的种间联结的变化,可以准确揭示植物种间的内在关联性。     

黄河首曲-玛曲县高寒草甸沙化演替进程中群落结构及种群生态位特征

以甘肃省玛曲县高寒沙化草甸为对象,应用空间序列代替时间序列的方法,对其不同沙化演替阶段植物群落主要种群结构及其生态位特征进行了研究.结果表明:(1)随着沙化演替的进行,物种组成数呈单峰式曲线变化规律,在轻度沙化阶段物种数最多(43种),禾草类和豆科类物种的重要值和生态位宽度均呈逐渐减少规律,优势度逐渐减弱;在极度沙化阶段(除垂穗披碱草外)几乎完全消失,莎草类和杂类草一些物种的重要值和生态位宽度则均呈逐渐增大规律,优势度逐渐增加,并在重度或极度沙化阶段成为优势种、次优势种或重要伴生种;随沙化演替进程,蒲公英、狼毒和乳白香青等物种则表现出波动性,最后消失,物种组成和数量的变化导致沙化演替进程中高寒草甸草场质量逐渐降低.(2)相邻沙化演替阶段植物群落的物种组成相似性指数较高,而相距较远的沙化演替阶段植物群落的物种组成相似性指数较小,沙化演替系列两个极端阶段的植物群落的物种组成相似性指数最小.(3)垂穗披碱草在整个沙化演替过程中重要值和生态位宽度均逐渐减小,但其值都较大.(4)生态位总宽度占前5位的物种依次是:垂穗披碱草、莓叶委陵菜、矮嵩草、高山唐松草和苔草,其值分别为1.653、1.438、1.387、1.295和1.204.(5)种群之间生态位重叠系数的平均值随沙化程度加深呈单峰式曲线变化规律,轻度沙化阶段种群的生态位重叠程度较高,为0.824.     

Steady state phytoplankton assemblages during thermal stratification in deep alpine lakes. Do they occur?

Phytoplankton seasonal and long-term succession can be described and functionally classified by associations similar as in terrestrial vegetation studies. Such a concept has to include 'climax' into pelagic succession which in turn leads to the question to what extent steady state assemblages occur and if during periods of dynamic equilibrium can be identified. Here we explore the situation with respect to the above question for deep, nutrient poor, alpine lakes in Austria. We first track the long-term development of phytoplankton biomass, their taxonomic structure and their relation to total phosphorus and chlorophyll-a as predictors of trophic state over the past 35 years. We then analyse this data set for coherent algal associations which can be ascribed to trait separated functional groups according to Reynolds et al. (2002). A three year period of stable environmental conditions has then be extracted from the progression of trophic state indices, having similar dominating species each year. These years were finally analysed for steady state conditions according to definitions given in Sommer et al. (1993). During thermal stratification, achievement of an equilibrium could be ruled out although coexistence of several dominating species lasted for several weeks. Habitat templates were constructed from environmental variables prior to biomass peaks for two species important in summer assemblages, the dinoflagellate Ceratium hirundinella and the diatom Fragilaria crotonensis. In summary, functional groups proved to be a valid and useful concept to describe species succession of phytoplankton in deep alpine lakes while pelagic climax is much less clear and steady state conditions were never met.     

喀斯特常绿落叶阔叶林个体死亡对空间格局及种间关联性的影响

树木死亡作为森林生态系统中一个重要的生态学过程,研究其如何影响物种的空间分布格局及种间关联性,可以为揭示树木死亡机制及群落动态变化规律提供重要参考。本研究根据木论25 hm²喀斯特常绿落叶阔叶林动态监测样地两次木本植物调查数据,采用点格局分析方法,分析了树木死亡前后空间格局和种间关系的变化。结果表明: 样地内个体死亡前后主要表现为聚集分布,但死亡后聚集分布的物种比例较死亡前略微下降,小尺度上呈现随机分布的物种比例有所增加,个体的死亡呈非随机性死亡。在物种水平上,13个优势种的死树和活树之间在0～30 m各尺度上呈显著正关联,表明样地内优势物种个体之间种内和种间竞争作用不激烈;树木死亡前后种间关联性主要为正关联,且大部分物种种间关系在死亡前后并未发生变化,表明群落已发展到相对稳定的阶段;死亡后,在1～30 m尺度上表现为正关联的物种对比例有所增加,负关联和无关联的物种对在大部分尺度上有所减少,说明个体死亡发生后,各个物种之间的竞争压力在一定程度上有所缓解。     

Secondary dune succession on Inhaca Island,Mozambique

Old field succession was studied on coastal dunes supporting tropical evergreen forest on Inhaca Island, Mozambique. Plots of 10×10 m were sited in three early successional stages and in relatively undisturbed forest. Woody species increased in number during succession; leptophylls were most frequent in younger vegetation, whereas microphylls and mesophylls were most frequent in forest. Grasses, shrubs and forbs dominated initially following abandonment, and shrubs persisted as dominants in the three early successional stages. The initial floristic composition model was generally supported by the pattern of species sequences, with many forest species entering early in the succession. Of the few species conforming to the relay floristic model, many were grasses and forbs of the forest understorey. Similarity between plots of equivalent vegetation age indicated that, at least in early succession, there was linearity in the successional pathway; there was no evidence for divergence or multiple pathways. In early succession, no accumulation was detected in either soil organic matter or extractable nutrients, thus providing little support for the facilitation model of succession. It is stressed that the findings are probably scale-dependent.     

Small-scale patch dynamics after disturbance in litter ant communities

The dynamics of re-colonisation of disturbed patches may aid in the understanding of spatial variation of species richness. The present study experimentally tested the hypothesis that the variation of litter ant local species richness and composition is caused by the dynamics of re-colonisation after disturbances. We were particularly interested in whether the re-colonisation was by pre-existent species or species new to the patches, and whether the succession of species evidences the existence of dominance-controlled or founder-controlled communities. Litter patches of a forest remnant in Southeast Brazil were disturbed by removing most animals through litter drying, and litter samples were returned to the same sites from where they were removed. Ant species richness and composition were compared before and 2 months after the disturbance. Dissimilarity among disturbed and non-disturbed samples was compared to infer the succession model occurring after disturbance. Ant species richness did not recover after 2 months, and species composition of the disturbed samples showed more new colonisers than pre-existent species. Dissimilarity among samples in the disturbed plots was smaller than in the control plots, indicating a directional, or dominance-controlled, succession. The changes in species composition observed were caused by a decrease of some species, particularly predators, and an increase of species that are possibly opportunistic. Patches of litter are naturally disturbed in time and space, and evidence from the present paper indicates that succession occurring in these patches would lead to different species richness and compositions. Thus the dynamics of re-colonisation contributes to explaining the diversity of litter-dwelling ant communities at larger spatial and temporal scales. In each patch the succession seems to be directional, with opportunist species re-colonising preferentially empty plots. Therefore, these communities may attain a high diversity due to a small-scale patch dynamics model.     

Structural changes of vegetation and its association with microclimate in a successional gradient of low thorn forest in northeastern Mexico

Understanding how microclimate and vegetation are associated during secondary succession is of primary importance for plant conservation in the face of the increasing land cover modification. However, these patterns are still unstudied for many plant communities. This study aimed to evaluate the structure (species richness, Shannon's diversity index, Simpson´s dominance index, abundance of each species, average height of species, species cover (%), species composition, and indicator values) of a low thorn forest fragment and to analyze its relation with microclimate along a successional gradient. Four stages of succession were delimited by the analysis of Landsat images, in the state of Tamaulipas, northeast Mexico. Statistical models incorporated species richness, diversity indices, abundance, height, and cover, as variables for searching differences between stages, or to evaluate microclimate associations. A total of 70 species, 54 genera, and 27 families were determined. Height of tree layer was the most important variable for discrimination of the successional stages. Conserved areas differed floristically from other stages, associated mainly with the lowest values of wind speed originated by tree layer characteristics. A significant association between species and microclimate was found, being wind speed and relative humidity the most important variables. Some species, due to their high importance values and their patterns of association with microclimate, may be considered as key taxa for low thorn forest, which is a threatened semitropical community in northeast Mexico. Conserved and late successional areas account for climatic regulation of this plant community, and the importance of these forest patches may be considered when establishing biodiversity protection areas.

     

流动沙丘区和固定沙丘区丘间低地植物种间关联关系

丘间低地植物种间关联研究对揭示植物对生境变化的响应及植被演替机制具有重要意义。通过χ2检验和AC关联系数对科尔沁沙地22个不同面积的流动沙丘区和固定沙丘区丘间低地植物种间关联进行了比较分析。结果表明:在固定沙丘区,当丘间低地面积<0.5hm2时,|AC|≤0.3的种对数占所有种对数的百分比和|AC|≥0.7的种对数占所有种对数的百分比略大于流动沙丘区,当丘间低地面积>0.5hm2时,|AC|≤0.3的种对数占所有种对数的百分比显著小于流动沙丘区,|AC|≥0.7的种对数占所有种对数的百分比显著大于流动沙丘区;当面积<0.5hm2时,沙丘固定未引起丘间低地植物种间关系发生明显变化;当面积>0.5hm2时,沙丘固定导致丘间低地植物种间关系更加紧密。     

Nature and Age of Neighbours Matter: Interspecific Associations among Tree Species Exist and Vary across Life Stages in Tropical Forests

Detailed information about interspecific spatial associations among tropical tree species is scarce, and hence the ecological importance of those associations may have been underestimated. However, they can play a role in community assembly and species diversity maintenance. This study investigated the spatial dependence between pairs of species. First, the spatial associations (spatial attraction and spatial repulsion) that arose between species were examined. Second, different sizes of trees were considered in order to evaluate whether the spatial relationships between species are constant or vary during the lifetime of individuals. Third, the consistency of those spatial associations with the species-habitat associations found in previous studies was assessed. Two different tropical ecosystems were investigated: a montane cloud forest and a lowland moist forest. The results showed that spatial associations among species exist, and these vary among life stages and species. The rarity of negative spatial interactions suggested that exclusive competition was not common in the studied forests. On the other hand, positive interactions were common, and the results of this study strongly suggested that habitat associations were not the only cause of spatial attraction among species. If this is true, habitat associations and density dependence are not the only mechanisms that explain species distribution and diversity; other ecological interactions, such as facilitation among species, may also play a role. These spatial associations could be important in the assembly of tropical tree communities and forest succession, and should be taken into account in future studies.     

Rain forest composition and patterns of secondary succession in the Vava'u Island Group,Tonga

Abstract. The Vava'u island group, Tonga, comprises ca. 60 limestone islands on a single submarine platform overlain with rich soils derived from tephra deposits from nearby volcanic islands. The island group has moderate topographic relief (215 m) and is characterized by plateaus and steep cliffs. Humans settled in Tonga ca. 3000 yr ago and have exploited the flatter terrain for agriculture since that time. We conducted the first survey of forest composition in Vava'u, sampling remnant patches of late-successional forest as well as stands in various stages of secondary succession following agricultural abandonment. Plant species composition did not vary greatly with elevation over this short gradient, in contrast with patterns found on‘Eua, a higher island in Tonga. The most significant environmental gradient affecting species composition was coastal or maritime influence. However, the greatest variation in species composition and structure appeared to be related to species turnover during secondary succession, and we hypothesize a sequence of species replacements. Secondary forest begins to resemble late-successional forest in 30–50 yr in terms of structure and native species richness and therefore is of significant conservation value.     

Seasonal comparison of bacterial communities in rhizosphere of alpine cushion plants in the Himalayan Hengduan Mountains

Positive associations between alpine cushion plants and other species have been extensively studied. However, almost all studies have focused on the associations between macrofauna. Studies that have investigated positive associations between alpine cushion plants and rhizospheric microbes have been limited to the vegetation growing season. Here, we asked whether the positive effects that alpine cushion plants confer on rhizospheric microbe communities vary with seasons. We assessed seasonal variations in the bacterial diversity and composition in rhizosphere of two alpine cushion plants and surrounding bare ground by employing a high throughput sequencing method targeting the V3 region of bacterial 16S rRNA genes. Soil properties of the rhizosphere and the bare ground were also examined. We found that cushion rhizospheres harbored significantly more C, N, S, ammonia nitrogen, and soil moisture than the bare ground. Soil properties in cushion rhizospheres were not notably different, except for soil pH. Bacterial diversities within the same microhabitats did not vary significantly with seasons. We concluded that alpine cushion plants had positive effects on the rhizospheric bacterial communities, even though the strength of the effect varied in different cushion species. Cushion species and the soil sulfur content were probably the major factors driving the spatial distribution and structure of soil bacterial communities in the alpine communities dominated by cushion plants.     

Food patch structure and plant resource partitioning in interspecific associations of amazonian tamarins

ávila-Pires’ saddle-back tamarins (Saguinus fuscicollis avilapiresi) and red-cap moustached tamarins (S. mystax pileatus), coexisting in highly stable mixed-species groups, overlapped considerably in their use of plant food resources at an Amazonian terra firme forest site. Overlap between food types consumed by the two species was particularly high during periods of lowest fruit availability, when they resorted to a common food supply, primarily the pod exudates of two emergent species of legume trees (Parkia nitida andParkia pendula) and nectar ofSymphonia globulifera. Within-group interspecific competition did not covary with independent measures of resource availability, contrary to predictions based on resource partitioning models. A greater number of both saddle-back and moustached tamarins were able to feed for longer patch residence periods within larger and more productive food patches, whereas small and clumped patches could be monopolized by the socially and numerically dominant moustached tamarins to the physical exclusion of the smaller-bodied saddle-back tamarins. Overall rates of interspecific aggression were extremely low, however, partly because patches that could be monopolized contributed with a minor proportion of either species’ diet. Saddle-backs foraged at lower levels in the understory and encountered smaller food patches more often, whereas moustached tamarins foraged higher and encountered more larger patches in the middle canopy. Although the two species led one another to differently-sized patches, moustached tamarins initiated most feeding bouts and encountered significantly larger and more productive patches that tended to accommodate the entire mixed-species group. Disadvantages of exploitative and interference feeding competition over plant resources, and advantages of shared knowledge of food patches, are but one component of the overall cost-benefit relationship of interspecific associations in tamarins.     

Small-scale spatial variability in phylogenetic community structure during early plant succession depends on soil properties

During early plant succession, the phylogenetic structure of a community changes in response to important environmental filters and emerging species interactions. We traced the development of temperate-zone plant communities during the first 7 years of primary succession on catchment soils to explore patterns of initial species assembly. We found pronounced small-scale differences in the phylogenetic composition of neighbouring plant assemblages and a large-scale trend towards phylogenetic evenness. This small-scale variability appears to be mediated by soil properties, particularly carbonate content. Therefore, abiotic environmental conditions might counteract or even supersede the effects of interspecific competition among closely related species, which are usually predicted to exhibit patterns of phylogenetic evenness. We conclude that theories on phylogenetic community composition need to incorporate effects of small-scale variability of environmental factors.     

Plant Functional Group Classification of Herbaceous Species in Quercus wutaishanica Communities in the Taiyue Mountains,Shanxi Province of China

Quercus wutaishanica is warm temperate deciduous broad leaved forest region one of the typical vegetation types. Qwutaishanica vegetation types as a representative of Shanxi, is the main part of shanxi forest. Past research focuses on the subject of Qwutaishanica results, the study of the herb layer seldom reported. The understory layer dominant species of Qwutaishanica community on Qiliyu of Taiyue Moutains, whose abundances clearly vary along environmental(altitudinal) gradients.Herbaceous plants highly responsive to changes in environmental conditions that reflect the dynamic relations between vegetation and environment. In this paper, using the method of sample, investigated the community of Qwutaishanica on Qiliyu of Taiyue Moutains, Shanxi. We used the results of this investigation to calculate species importance values and frequency, which were used to identify 27 dominant species in herbages. χ2 tests, together with association coefficient and percentage cooccurrence, were used to measure interspecific associations of the dominant species of herbaceous. Plant functional types (PFTS) were defined according to interspecific associations and cluster analysis of the dominant species. In studies of forest understory plant assemblages, designating PFTs according to dominant herbaceous species is feasible. Dominant herbaceous species formed seven PFTs: each with its unique spatial distribution and morphological characteristics, are identifiedⅠ.amphigenous Ⅱ.pelouse edge Ⅲ.wet sparse Ⅳ.gap drought resistant Ⅴ.hill edge Ⅵ.alpine Ⅶ.dank. Plant functional types(PFTs) of herbaceous plants varies with altitude gradients, which reflect the relationship between Qwutaishanica and environment well.