草原模式植物群落监测方法应用样例

模式植物群落是指能够反映某种植被分类单元基本特征, 并可作为准确描述该植被类型“标准”的典型植物群落。草原模式植物群落监测是中国生物多样性监测与研究网络——草原/荒漠植物多样性监测网的核心内容, 已初步确定了一套监测方法。本文应用这套方法分别建立了克氏针茅(Stipa krylovii)群系和百里香(Thymus mongolicus)群系模式植物群落监测固定样地, 作为中国生物多样性监测与研究网络——草原/荒漠植物多样性监测网的样例。介绍了这两个模式植物群落监测样地选取与设置的过程、群落调查方法和数据分析结果。通过实际观测和案例分析, 验证了“中国草原/荒漠植物多样性监测网模式植物群落监测方案”中监测方法的可行性, 同时根据监测中出现的问题, 提出了模式植物群落样地、监测频率和监测日期的选择原则, 以及监测过程中的操作规范等, 完善了模式植物群落监测内容、方法和指标体系的建议, 为开展草原模式植物群落监测提供了可借鉴的实例与技术储备。     

Plant diversity,species turnover and shifts in functional traits in coastal dune vegetation: Results from permanent plots over a 52‐year period

Question: What is the relationship between plant diversity and species turnover in coastal dune vegetation plots? How is the long‐term change in species composition of vegetation plots related to shifts in functional traits, and what does it tell us about the dominant processes? Location: Coastal dunes, the Netherlands. Methods: Our data set comprised 52 years of vegetation data from 35 permanent plots in grassland/scrub/woodland vegetation. Vegetation dynamics were described in terms of changes in species composition and abundance, and shifts in 13 functional traits related to resources capture and forage quality, regeneration and dispersal. Results: Species turnover in the plots was high, because of local extinction and colonization. Species‐rich plots were more stable in terms of species abundance and composition compared with species‐poor plots. Over time, the plots converged with respect to their abiotic conditions, as reflected by Ellenberg indicator values – indicating that the prevailing process was succession. The high species turnover reflected high invasibility: accordingly, the relative importance of annuals increased. Most newcomer annuals, however, were competitive generalists of little conservation value. The functional trait analysis allowed us to unravel the complexity of effects of disturbances and succession, and yielded information on the processes driving the observed vegetation dynamics. Conclusions: In this study, small‐scale species turnover was negatively related to species diversity, indicating more stability in species‐rich communities. Regarding shifts in trait diversity, unifying filters appeared to be more dominant than diversifying filters. Counteracting this homogenization process poses a challenge for nature management.     

金华北山南坡主要植被类型的群落特征

浙江省金华北山地处中亚热带北部, 在植被区划上属于中亚热带常绿阔叶林北部亚地带。该区植被是遭砍伐后恢复的次生林, 目前正处于快速的正向演替进程中, 这为研究常绿阔叶林植物群落动态演替机制及受损生态系统的恢复提供了平台。为进一步了解金华北山地区主要植被类型的群落特征, 该文以国际上通用的大样地调查方法, 采用固定样方对其南坡主要植物群落进行了调查, 其中森林样方面积30 m × 30 m, 灌丛样方面积为20 m × 10 m、30 m × 10 m。调查群落物种组成及其数量特征, 记录群落生境信息, 计算木本植物重要值, 分析群落的类型及特征, 并制作每个样方主要木本植物的空间分布图。论文提供了24组详细的群落样方数据(包括21个森林样方和3个灌丛样方), 含有11个群系。     

Local above‐ground persistence of vascular plants: Life‐history trade‐offs and environmental constraints

Questions: 1. Which plant traits and habitat characteristics best explain local above‐ground persistence of vascular plant species and 2. Is there a trade‐off between local above‐ground persistence and the ability for seed dispersal and below‐ground persistence in the soil seed bank? Locations: 845 long‐term permanent plots in terrestrial habitats across the Netherlands. Methods: We analysed the local above‐ground persistence of vascular plants in permanent plots (monitored once a year for ca. 16 year) with respect to functional traits and habitat preferences using survival statistics (Kaplan‐Meier analysis and Cox’ regression). These methods account for censored data and are rarely used in vegetation ecology. Results: Local above‐ground persistence is determined by both functional traits (especially the ability to form long‐lived clonal connections) and habitat preferences (especially nutrient requirements). Above‐ground persistence is negatively related to the ability for dispersal by wind and to the ability to accumulate a long‐term persistent soil seed bank (‘dispersal through time’) and is positively related to the ability for dispersal by water. Conclusions: Most species have a half‐life expectation over 15 years, which may contribute to time lags after changes in habitat quality or ‐configuration (‘extinction debt’). There is evidence for a trade‐off relationship between local above‐ground persistence and below‐ground seed persistence, while the relationship with dispersal in space is vector specific. The rate of species turnover increases with productivity.     

Multiple environmental changes drive forest floor vegetation in a temperate mountain forest

Human‐induced changes of the environment and their possible impacts on temperate forest understory plant communities have been examined in many studies. However, the relative contribution of individual environmental factors to these changes in the herb layer is still unclear. In this study, we used vegetation survey data covering a time period of 21 years and collected from 143 permanent plots in the Northern Limestone Alps, Austria. Data on soil chemistry (49 plots), light condition (51 plots), soil temperature and moisture (four and six plots), disturbance (all plots), climate (one station in a clearing area), and airborne sulfur (S) and nitrogen (N) deposition (two forest stands) were available for analyses. We used these data together with plot mean Ellenberg indicator values in a path analysis to attribute their relative contributions to observed vegetation changes. Our analysis reveals a strong directional shift of the forest understory plant community. We found strong evidence for a recovery of the ground‐layer vegetation from acidification as response to decreased S deposition. We did not observe a community response to atmospheric N deposition, but we found a response to altered climatic conditions (thermophilization and drying). The path analysis revealed that changes in the light regime, which were related to small‐scale disturbances, had most influence on herb layer community shifts. Thermophilization and drying were identified as drivers of understory community changes independent of disturbance events.     

A study on long-term salt-marsh succession using permanent plots

Succession of plant communities on a salt marsh at the Boschplaat, Terschelling (The Netherlands) is described on the basis of relevés from permanent plots in three transects, covering a period of almost 30 years. All relevés were clustered, using a hierarchical fusion technique. Succession has been quantified in terms of transitions between clusters over time in all plots. Analysis of the results showed four clearly distinct successional trends, each restricted to a particular altitudinal zone of the salt marsh. Between zones there were only few interconnections. This implies that zonation does not represent succession, at least not on the time scale of this study. It is suggested that initial differences between zones in the processes that change the abiotic environment, form the basis of vegetation zonation at the Boschplaat.The study was financially supported by the Netherlands Organization for the Advancement of Pure Research (Z.W.O.).     

Restoration success of low‐production plant communities on former agricultural soils after top‐soil removal

Abstract. The success inrestoring seven low‐production vegetation types on former agricultural soil after top‐soil removal was investigated. The colonization and establishment of target species in permanent plots was recorded during the first nine years after restoration measures were taken. For each permanent plot abiotic site conditions were used to determine which of the vegetation types could persist there. A comparison of the actual vegetation in the permanent plots with reference releves of the selected vegetation type revealed a gradually increasing similarity during consecutive years for five vegetation types. This was due to the occurrence of an increasing number of target species and the number of permanent plots they occurred in. However, nine years after top‐soil removal a large number of the target species were still lacking from the vegetation in the permanent plots, although most were present in the local species pool. Seed dispersal therefore seems to be a major limiting factor for restoration of these low‐production vegetation types on formerly agricultural soils.     

Effectiveness of Turf Stripping as a Measure for Restoring Species-Rich Fen Meadows in Suboptimal Hydrological Conditions

Most species‐rich fen meadows in nature reserves in The Netherlands are acidified due to weaker upwelling of base‐rich groundwater. The present study investigated whether and why turf stripping combined with superficial drainage might promote the long‐term recovery of such meadows and restore the nutrient‐poor, buffered conditions they require. In a field experiment, we analyzed changes in vegetation composition, soil parameters, and soil water chemistry in stripped plots of degraded Cirsio‐Molinietum vegetation over 12 years. After the first five years, many species from the target communities occurred in stripped plots. Both vegetation and soil data showed positive effects of turf stripping on the acid‐buffering capacity. Because sulfate concentration in the soil water decreased over time, whereas the bicarbonate concentration increased, we inferred that there was internal alkalinization driven by sulfate reduction in low‐lying stripped plots. However, the succession toward more acidophilus plant communities, in both control and stripped plots, indicated gradual acidification. This may be caused by a continuing weakening of the upward seepage of base‐rich groundwater as shown by declining calcium concentrations in the soil water. Though turf stripping exposed a nutrient‐poor soil layer with a greater acid‐buffering capacity, these positive effects might not be sufficient to combat the ongoing acidification in the long term.     

Land use designation and vegetation community structure in the Adirondack uplands (New York,USA)

Question: Does forest vegetation community structure reflect legislative land use designations? Location: Adirondack Park, New York, USA. Methods: The Adirondack Park, located in northern New York State, is a mixture of public and private lands, with state‐owned Forest Preserve lands comprising ca. 42% of the 2.4 million ha, on which timber harvesting and many other forms of anthropogenic disturbance are prohibited. A survey of vegetation communities was conducted in eighteen upland catchments with differing land use history (managed and Forest Preserve), including overstory, understory, and dead wood (snags and downed woody debris) using randomly placed plots. Results: Mean overstory density and basal area were not significantly different between land uses, although mean overstory tree size was greater in Preserve catchments. Sapling densities were greater in managed catchments, while mean herb/shrub coverage was not affected by land use. Densities of 25% of common species were affected by land use, determined by GIS coverages constructed using an Inverse Distance Weighted estimation procedure. Discriminant Analysis of per‐plot plant community data correctly classified 89% of both managed and Preserve plots. Conclusion: The success of the Discriminant Analysis in classifying land uses based on vegetation communities indicates its potential utility of this method in comparing forest vegetation to a reference condition in this and other areas. The analysis suggests that at least 85 years is required for Adirondack up‐land catchments to recover following harvesting. Uncertainty in classification was related to heterogenous management and disturbance patterns within catchments.     

Oscillating vegetation dynamics in a wet heathland

Abstract. Question: The significance of disturbances caused by periodical inundation was investigated with respect to its effects on vegetation dynamics, species richness and fluctuations, and to the relevance of certain plant properties. Location and Method: At a sod‐cut stand within nutrient‐poor inland sand dunes, permanent plots along a transect were surveyed over a period of up to nine years after sod cutting. Results: In contrast to never inundated plots, periodically inundated plots were characterized by low vegetation cover and by high numbers of species belonging to many different communities, each of them with a low cover. Periodical inundations favoured the presence of pioneers, species tolerant of disturbances, species adapted to wet conditions and stoloni‐ferous species. Furthermore, annual fluctuations of species within each plot were higher and most species occurred only sporadically. Discussion: A comprehensive model is presented describing the relevant processes identified in the littoral zone. Changing water tables result in the creation of gaps. The re‐colonization of these gaps follows mainly from vegetative regeneration and less to the dispersion of diaspores. Highest species numbers in the zone of moderate disturbances result from a high rate of re‐colonization in spite of local extinctions following each disturbance event. It is suggested that colonization abilities are among the most important features for species occurrence at a site rich in disturbances (more important than competitive abilities and more important than a slow rate of displacement). For nature conservation such sites are very important, because they allow (rare) pioneer species to survive for longer periods of time.     

Bias in vegetation databases? A comparison of stratified‐random and preferential sampling

Aim: Vegetation plots collected since the early 20th century and stored in large vegetation databases are an important source of ecological information. These databases are used for analyses of vegetation diversity and estimation of vegetation parameters, however such analyses can be biased due to preferential sampling of the original data. In contrast, modern vegetation survey increasingly uses stratified‐random instead of preferential sampling. To explore how these two sampling schemes affect vegetation analyses, we compare parameters of vegetation diversity based on preferentially sampled plots from a large vegetation database with those based on stratified‐random sampling. Location: Moravian Karst and Silesia, Czech Republic. Methods: We compared two parallel analyses of forest vegetation, one based on preferentially sampled plots taken from a national vegetation database and the other on plots sampled in the field according to a stratified‐random design. We repeated this comparison for two different regions in the Czech Republic. We focussed on vegetation properties commonly analysed using data from large vegetation databases, including alpha (within‐plot) diversity, cover and participation of different species groups, such as endangered and alien species within plots, total species richness of data sets, beta diversity and ordination patterns. Results: The preferentially sampled data sets obtained from the database contained more endangered species and had higher beta diversity, whereas estimates of alpha diversity and representation of alien species were not consistently different between preferentially and stratified‐randomly sampled data sets. In ordinations, plots from the preferential samples tended to be more common at margins of plot scatters. Conclusions: Vegetation data stored in large databases are influenced by researcher subjectivity in plot positioning, but we demonstrated that not all of their properties necessarily differ from data sets obtained by stratified‐random sampling. This indicates the value of vegetation databases for use in biodiversity studies; however, some analyses based on these databases are clearly biased and their results must be interpreted with caution.     

Post‐fire regeneration of Mediterranean plant communities at a regional scale is dependent on vegetation type and dryness

Question: We tested whether (1) the change in composition and structure of whole plant communities after fire is directly related to regeneration of the dominant tree species in the canopy; (2) the change in structure and composition of plant communities several years after fire decreases with the proportion of obligate seeders and (3) the proportion of obligate seeders in plant communities increases with the dryness gradient. Location: Catalonia (NE Spain) Methods: We measured floristic differences between burned and long‐since burned sites in eight vegetation types across a climate gradient. We compared 22 sites burnt in 1994 in paired plots with 22 sites that had not been burnt since the 1940s. In each site we placed plots in burned and long‐since burned areas, where we identified the presence and abundance of all plant species. Results: When the tree canopy recovers, structure and composition of the vegetation also return to the long‐since burned community; when tree canopy does not recover, composition of the post‐fire community varies compared to the long‐since burned one. A higher proportion of obligate seeders in the pre‐fire community promotes quicker regeneration of the original community. The proportion of obligate seeders increased along the dryness gradient. Conclusions: Regeneration of plant communities after fire depends on the vegetation type before the fire. Regeneration increases when the dominant tree or shrub species persists after fire and with a higher proportion of obligate seeders in the pre‐fire community. The proportion of obligate seeders varies along the dryness gradient, which suggests that vegetation in drier areas (when seeders are more abundant) recovers earlier than in moister areas.     

Change within and among forest communities: the influence of historic disturbance,environmental gradients,and community attributes

Understanding how ecological communities change over time is critical for biodiversity conservation, but few long‐term studies directly address decadal‐scale changes in both the within‐ and among‐community components of diversity. In this study, we use a network of permanent forest vegetation plots, established in Great Smoky Mountains National Park (USA) in 1978, to examine the factors that influence change in community composition within and among communities. In 2007, we resampled 15 plots that were logged in the late 1920s and 15 plots that had no documented history of intensive human disturbance. We found that understory species richness decreased by an average of 4.3 species over the 30‐yr study period in the logged plots, but remained relatively unchanged in the unlogged plots. In addition, tree density decreased by an average of 145 stems ha^?1 in the logged plots, but was relatively stable in the unlogged plots. However, we found that historic logging had no effect on within‐community understory or tree compositional turnover during this time period. Instead, sites at lower elevations and sites with lower understory biomass in 1978 had higher understory compositional turnover than did sites at higher elevations and sites with higher understory biomass. In addition, sites with lower soil cation exchange capacity (CEC) and with lower tree basal area in 1978 had higher tree compositional turnover than did sites with higher soil CEC and higher tree basal area. Among‐community similarity was unchanged from 1978 to 2007 for both the logged and unlogged plots. Overall, our results indicate that human disturbance can affect plant communities for decades, but the extent of temporal change in community composition may nevertheless depend more on environmental gradients and community attributes.     

Identifying Preferential Associates to Initiate Restoration Plantings

Ecological restoration typically aims to re‐establish dominant plant species and their native associates, despite the lack of guidance on which associates to introduce initially. Analysis of naturally occurring plant communities can provide criteria to shorten the list of species that are associated with dominants, in order to focus revegetation efforts on species likely to establish. Using the example of sedge meadows, we evaluated wetland vegetation data from Laurentian Great Lakes wetlands to identify “preferential associates,” that is, species that co‐occur more frequently than expected based on their overall abundance. A total of 176 taxa occurred within the two hundred and thirty‐nine 1 × 1 m² plots in 48 wetlands that contained Carex stricta, a widespread tussock‐forming sedge. Of 58 species that co‐occurred with C. stricta where it was dominant (≥50% plot cover), we identified 26 associates using Bray–Curtis similarities and we determined that 12 of the 26 were preferential using an electivity index. The electivity index identified preferential associates even when they occurred infrequently or had low mean cover per plot. We provide guidance on how to initiate restoration with preferential associates.     

The RAINFOR database: monitoring forest biomass and dynamics

Problem: Data from over 100 permanent sample plots which have been studied for 10–20 years need a suitable system for storage which allows simple data manipulation and retrieval for analysis. Methods: A relational database linking tree records, taxonomic nomenclature and corresponding environmental data has been built in MS Access as part of the RAINFOR project. Conclusion: The database allows flexible and long‐term use of a large amount of data: more than 100 tree plots across Amazonia, incorporating over 80 000 records of individual trees and over 300 000 total records of tree diameter measurements from successive censuses. The database is designed to enable linkages to existing soil, floristic or plant‐trait databases. This database will be a useful tool for exploring the impact of environmental factors on forest structure and dynamics at local to continental scales, and long term changes in forest ecology. As an early example of its potential, we explore the impact of different methodological assumptions on estimates of tropical forest biomass and carbon storage.     

Conversion of sagebrush shrublands to exotic annual grasslands negatively impacts small mammal communities

Aim The exotic annual cheatgrass (Bromus tectorum) is fast replacing sagebrush (Artemisia tridentata) communities throughout the Great Basin Desert and nearby regions in the Western United States, impacting native plant communities and altering fire regimes, which contributes to the long‐term persistence of this weedy species. The effect of this conversion on native faunal communities remains largely unexamined. We assess the impact of conversion from native perennial to exotic annual plant communities on desert rodent communities. Location Wyoming big sagebrush shrublands and nearby sites previously converted to cheatgrass‐dominated annual grasslands in the Great Basin Desert, Utah, USA. Methods At two sites in Tooele County, Utah, USA, we investigated with Sherman live trapping whether intact sagebrush vegetation and nearby converted Bromus tectorum‐dominated vegetation differed in rodent abundance, diversity and community composition. Results Rodent abundance and species richness were considerably greater in sagebrush plots than in cheatgrass‐dominated plots. Nine species were captured in sagebrush plots; five of these were also trapped in cheatgrass plots, all at lower abundances than in the sagebrush. In contrast, cheatgrass‐dominated plots had no species that were not found in sagebrush. In addition, the site that had been converted to cheatgrass longer had lower abundances of rodents than the site more recently converted to cheatgrass‐dominated plots. Despite large differences in abundances and species richness, Simpson’s D diversity and Shannon‐Wiener diversity and Brillouin evenness indices did not differ between sagebrush and cheatgrass‐dominated plots. Main conclusions This survey of rodent communities in native sagebrush and in converted cheatgrass‐dominated vegetation suggests that the abundances and community composition of rodents may be shifting, potentially at the larger spatial scale of the entire Great Basin, where cheatgrass continues to invade and dominate more landscape at a rapid rate.     

Changes in forest understorey vegetation in Norway related to long‐term soil acidification and climatic change

Question: Does the understorey vegetation of Norwegian boreal forests change in relation to broad‐scale, long‐term changes? Location: Norway. Methods: Permanently marked 1‐m² vegetation plots from 17 monitoring reference areas in forests dominated by Picea abies (11 areas, 620 plots) and Betula spp. (six areas, 300 plots) were analysed twice, at the start in 1988–1997 and 5 yr later (1993–2002). Species subplot frequency data were analysed separately for each area by univariate and multivariate statistical methods; 5‐yr changes in single species abundances, species number per plot and species composition were tested. Results: Two distinct patterns of change were found: 1. Abundance of several vascular plant species decreased in SE Norwegian Picea forests, most noticeably of species with a preference for richer soils, such as Oxalis acetosella. 2. Abundance of many bryophyte species as well as bryophyte species number per plot increased in forests of both types over most of Norway. Conclusions: The pattern of vascular plant changes is probably a time‐delayed response of long‐lived, mainly clonal, populations to acidified soils resulting from deposition of long‐distance airborne pollutants. The pattern bryophyte changes, with reference to the close link between climatic conditions for growth and abundance changes for Hylocomium splendens established in previous demographic studies, is related to climatic conditions favourable for bryophyte growth. We conclude that many forest understorey plants are sensitive indicators of environmental change, and that the concept used for intensive monitoring of Norwegian forests enables early detection of changes in vegetation brought about by broad‐scale, regional, impact factors.     

Herbaceous vegetation change in variable rangeland environments: The relative contribution of grazing and climatic variability

Abstract. A 44‐yr record of herbaceous vegetation change was analysed for three contrasting grazing regimes within a semi‐arid savanna to evaluate the relative contribution of confined livestock grazing and climatic variability as agents of vegetation change. Grazing intensity had a significant, directional effect on the relative composition of short‐ and mid‐grass response groups; their composition was significantly correlated with time since the grazing regimes were established. Interannual precipitation was not significantly correlated with response group composition. However, interannual precipitation was significantly correlated with total plant basal area while time since imposition of grazing regimes was not, but both interannual precipitation and time since the grazing regimes were established were significantly correlated with total plant density. Vegetation change was reversible even though the herbaceous community had been maintained in an altered state for ca. 60 yr by intensive livestock grazing. However, ca. 25 yr were required for the mid‐grass response group to recover following the elimination of grazing and recovery occurred intermittently. The increase in mid‐grass composition was associated with a significant decrease in total plant density and an increase in mean individual plant basal area. Therefore, we failed to reject the hypotheses based on the proportional change in relative response group composition with grazing intensity and the distinct effects of grazing and climatic variability on response group composition, total basal area and plant density. Long‐term vegetation change indicates that grazing intensity established the long‐term directional change in response group composition, but that episodic climate events defined the short‐term rate and trajectory of this change and determines the upper limit on total basal area. The occurrence of both directional and non‐directional vegetation responses were largely a function of (1) the unique responses of the various community attributes monitored and (2) the distinct temporal responses of these community attributes to grazing and climatic variation. This interpretation supports previous conclusions that individual ecosystems may exist in equilibrial and non‐equilibrial states at various temporal and spatial scales.     

Plant colonization windows in a mesic old field succession

Abstract. Closed canopy vegetation often prevents the colonization of plant species. Therefore the majority of plant species are expected to appear at the initial phase of post‐agricultural succession in mesic forest environment with moderate levels of resources. This hypothesis was tested with data from the Buell‐Small Successional Study, NJ, USA, one of the longest continuous fine‐scale studies of old‐field succession. The study started in 1958, including old fields with different agricultural histories, landscape contexts, and times of abandonment. In each year of the study, the cover values of plant species were recorded in 48 permanent plots of 1 m² in each field. We analysed the temporal patterns of colonization at plot scale and related these to precipitation data and other community characteristics. The number of colonizing species decreased significantly after ca. 5 yr, coinciding with the development of a continuous canopy of perennial species. However, species turnover remained high throughout the whole successional sequence. The most remarkable phenomenon is the high inter‐annual variation of all studied characteristics. We found considerable temporal collapses of vegetation cover that were synchronized among fields despite their different developmental stages and distinctive species compositions. Declines of total cover were correlated with drought events. These events were associated with peaks of local species extinctions and were followed by increased colonization rates. The transitions of major successional stages were often connected to these events. We suggest that plant colonization windows opened by extreme weather events during succession offer optimum periods for intervention in restoration practice.