长江口滨海湿地潮间带生态系统的多稳态特征

多稳态现象普遍存在于多种生态系统中,它与生态系统的健康和可持续发展密切相关,已成为生态学研究的热点与难点,但是目前有关滨海湿地生态系统多稳态的形成机制还缺乏深入研究.本文以崇明东滩鸟类自然保护区的潮间带生态系统为研究对象,通过以下内容,开展滨海湿地多稳态研究: 1)通过验证多稳态的判定依据“双峰”和“阈值”特征,证实长江口潮间带生态系统存在多稳态,并确定其稳态类型;2)通过监测潮间带生态系统水动力过程、沉积动力过程以及盐沼植物生长和扩散情况,分析盐沼植被与沉积地貌之间的正反馈作用,进而探讨潮间带生态系统多稳态的形成机制.结果表明: 1)潮间带生态系统的归一化植被指数(NDVI)频度分布存在明显的双峰特征,且盐沼植物成活存在生物量阈值效应,均证实潮间带生态系统存在多稳态,“盐沼”和“光滩”是潮间带生态系统的两种相对稳定状态;2)崇明东滩盐沼前沿的沉积地貌表现出泥沙快速淤积的趋势,显著促进了盐沼植物的生长,盐沼植物与泥沙淤积之间的这种正反馈作用是潮间带生态系统形成多稳态的主要原因;3)盐沼植被扩散格局监测结果在景观尺度上也表明,泥沙淤积作用促进了潮间带生态系统“盐沼”和“光滩”多稳态的形成.本研究既丰富了滨海湿地稳态转换的机理研究,也为我国开展海岸带保护、修复和管理提供了科学依据,具有重要的理论和实践意义.     

Tree–Grass Coexistence in the Everglades Freshwater System

Mosaic freshwater landscapes exhibit tree-dominated patches —or tree islands—interspersed in a background of marshes and wet prairies. In the Florida Everglades, these patterned landscapes provide habitat for a variety of plant and animal species and are hotspots of biodiversity. Even though the emergence of patchy freshwater systems has been associated with climate histories, fluctuating hydrologic conditions, and internal feedbacks, a process-based quantitative understanding of the underlying dynamics is still missing. Here, we develop a mechanistic framework that relates the dynamics of vegetation, nutrients and soil accretion/loss through ecogeomorphic feedbacks and interactions with hydrologic drivers. We show that the stable coexistence of tree islands and marshes results as an effect of their both being (meta-) stable states of the system. However, tree islands are found to have only a limited resilience, in that changes in hydrologic conditions or vegetation cover may cause an abrupt shift to a stable marsh state. The inherent non-linear and discontinuous dynamics determining the stability and resilience of tree islands should be accounted for in efforts aiming at the management, conservation and restoration of these features.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.     

Assessing the ecological risk from secondary salinity: A framework addressing questions of scale and threshold responses

Abstract Assessment of the ecological risk posed to native vegetation from the development of shallow and saline water tables is considered an urgent task in southern Australia. Ecological risk can be defined as the product of the likelihood of an ecological effect and the consequences of that effect. At present, the likelihood of the development of a shallow water table is determined by hydrological modelling at the catchment scale, and this in itself is often equated to ecological risk. In contrast, the ecological consequences of secondary salinity are generally investigated at the patch scale. Translating ecological likelihood and the ecological consequences of risk across these scales has proved problematic, both conceptually and quantitatively. Here we argue that the consideration of ecological risk within the context of the patch‐ or catchment‐scale is based upon human perceptions of spatial units, rather than the ecological scales at which various processes determine vegetation dynamics. By focusing on the processes that determine vegetation dynamics, and the dimensions of these processes, both spatial and temporal scales can be built into conceptual frameworks that aim to understand vegetation change, such as frameworks of alternative stable states. We present an alternative conceptual framework of the ecological risk from secondary salinity based around using the rates of processes, such as salt accumulation in the soil or the frequency of waterlogging, at the scales at which these processes occur. This framework also integrates concepts of vegetation states and transitions between meta‐stable states and alternative stable states.     

Effects of Wind Waves versus Ship Waves on Tidal Marsh Plants: A Flume Study on Different Life Stages of Scirpus maritimus

Recent research indicates that many ecosystems, including intertidal marshes, follow the alternative stable states theory. This theory implies that thresholds of environmental factors can mark a limit between two opposing stable ecosystem states, e.g. vegetated marshes and bare mudflats. While elevation relative to mean sea level is considered as the overall threshold condition for colonization of mudflats by vegetation, little is known about the individual driving mechanisms, in particular the impact of waves, and more specifically of wave period. We studied the impact of different wave regimes on plants in a full scale flume experiment. Seedlings and adult shoots of the pioneer Scirpus maritimus were subjected to two wave periods at two water levels. Drag forces acting on, and sediment scouring occurring around the plants were quantified, as these are the two main mechanisms determining plant establishment and survival. Depending on life stage, two distinct survival strategies emerge: seedlings present a stress avoidance strategy by being extremely flexible, thus limiting the drag forces and thereby the risk of breaking. Adult shoots present a stress tolerance strategy by having stiffer stems, which gives them a higher resistance to breaking. These strategies work well under natural, short period wind wave conditions. For long period waves, however, caused e.g. by ships, these survival strategies have a high chance to fail as the flexibility of seedlings and stiffness of adults lead to plant tissue failure and extreme drag forces respectively. This results in both cases in strongly bent plant stems, potentially limiting their survival.     

Simulation of the effects of deer browsing on forest dynamics

I introduced forest dynamics to a deterministic herbivore-vegetation model to examine the effects of vertically stratified forest structure on the dynamics of the herbivore–vegetation system, the resilience and stable states of vegetation, and the interactions between deer populations and vegetation. I constructed a model based on data from field studies performed in Hokkaido, northern Japan. Three phases of state were identified for a given deer density: (1) understory vegetation is maintained with a equilibrium proportion of canopy gaps in the absence of deer; (2) if the equilibrium proportion of canopy gaps is greater than that in the equilibrium state in the absence of deer, the understory vegetation can be maintained; and (3) the understory vegetation cannot be maintained. At the boundary between phases 2 and 3, the herbivore population level had discontinuous effects on vegetation. When the deer density was held below the threshold, forest vegetation had resilience to recover to the equilibrium stable state at the given deer density, although the equilibrium canopy gap ratio and vegetation biomass differed with deer density. However, the forest vegetation–herbivore system could not be maintained in a stable state without artificial deer population management if food limitation was the only mechanism to keep the deer population at low levels. The deer population must be kept below the boundary between phases 1 and 2 to maintain the forest regeneration processes. The level cannot be determined by observing the deer population; careful observation of forest regeneration processes is required.     

Catastrophic vegetation shifts and soil degradation in terrestrial grazing systems

It has long been recognized that alternative vegetation states may occur in terrestrial grazing systems. This phenomenon may be of great importance as small environmental fluctuations may lead to relatively sudden and irreversible jumps between vegetation states. Early theoretical studies emphasized saturation of herbivore feeding to explain multiple stable states and catastrophic behaviour. Recent studies on semi-arid grasslands and arctic salt marshes, however, relate catastrophic events in these systems to plant-soil interactions.     

Spatially explicit population responses of crayfish Procambarus alleni to potential shifts in vegetation distribution in the marl marshes of Everglades National Park,USA

Hydropattern disturbance has had wide-ranging impacts on wetland communities of the Florida Everglades, especially on the habitats and the aquatic biota of the seasonally flooded marl marshes. We used the Everglades crayfish Procambarus alleni as a model to study the associations among hydrology, vegetation distribution, and population dynamics to assess the potential impacts of hydrological changes on the aquatic faunal community in Everglades National Park. To classify benthic habitats as sources or sinks for the crayfish population, we quantified vegetation community structure using GIS maps in which dominant vegetation types were weighted by local hydroperiod (length of inundation). Regression analysis showed that this habitat classification was associated with crayfish density distribution. We then used a spatially explicit, stage-structured population model to describe crayfish population fluctuations under current environmental conditions and to simulate the potential population-level responses to habitat changes that might occur following hydrological restoration. In habitat that was initially saturated with crayfish, the crayfish population size declined under current environmental conditions and then stabilized at about 13% of the initial density over a 50-year period. A 4-month increase in hydroperiod was then simulated by converting shorter-hydroperiod Muhlenbergia-dominated marsh habitat to longer-hydroperiod Cladium-dominated marshes. The model predicted a rapid 7-fold increase in crayfish density following the simulated habitat restoration. This indicated that several functional effects may result from the restoration of historical hydropatterns in marl marshes: (1) the areal extent of habitat sinks will be reduced to isolated patches, whereas the spatial distribution of aquatic source habitats will expand; (2) crayfish population size will increase and persist over time; (3) the minimum threshold needed to increase secondary aquatic productivity may be a 7-month hydroperiod over 90% of the marl marsh landscape. Restoration of historical hydropatterns could thus have cascading positive effects throughout the Everglades aquatic food web.     

Spatial heterogeneity and irreversible vegetation change in semiarid grazing systems

Recent theoretical studies have shown that spatial redistribution of surface water may explain the occurrence of patterns of alternating vegetated and degraded patches in semiarid grasslands. These results implied, however, that spatial redistribution processes cannot explain the collapse of production on coarser scales observed in these systems. We present a spatially explicit vegetation model to investigate possible mechanisms explaining irreversible vegetation collapse on coarse spatial scales. The model results indicate that the dynamics of vegetation on coarse scales are determined by the interaction of two spatial feedback processes. Loss of plant cover in a certain area results in increased availability of water in remaining vegetated patches through run-on of surface water, promoting within-patch plant production. Hence, spatial redistribution of surface water creates negative feedback between reduced plant cover and increased plant growth in remaining vegetation. Reduced plant cover, however, results in focusing of herbivore grazing in the remaining vegetation. Hence, redistribution of herbivores creates positive feedback between reduced plant cover and increased losses due to grazing in remaining vegetated patches, leading to collapse of the entire vegetation. This may explain irreversible vegetation shifts in semiarid grasslands on coarse spatial scales.     

Effects of environmental context on the susceptibility of Atriplex patula to attack by herbivorous beetles

The susceptibility of plants to attack by insect herbivores often depends on local environmental conditions. This study documents variation in herbivore damage by the chrysomelid beetle Erynephala maritima to the annual forb Atriplex patula in two microhabitats within New England salt marshes: bare patches and dense matrix vegetation. Environmental conditions within bare patches differ from those within matrix vegetation in a number of ways. Bare patches are characterized by the absence of perennial grasses and rushes (matrix vegetation) and greater levels of physical stress, and are rapidly colonized by the fugitive annual, Salicornia europaea, a second host plant of these beetles. Surveys of herbivore damage across three marshes revealed that A. patula in bare patches had a greater proportion of leaves damaged by beetles than those within matrix vegetation. Presence or absence of matrix vegetation and presence or absence of S. europaea were experimentally manipulated to determine the proximate cause of this pattern. The presence of S. europaea significantly increased the susceptibility of A. patula to herbivory in experimental plots. Both the extent of herbivore damage to plants and the proportion of plants damaged through time were greater in treatments with S. europaea than in controls, regardless of the presence or absence of matrix vegetation. Plants in S. europaea addition treatments were also less likely to survive to reproduction. Decreased survival appears to result from increased herbivory, suggesting that the negative effect of S. europaea on A. patula is mediated indirectly through shared insect herbivores. These results support the hypothesis that indirect interactions between alternative host plants, mediated by insect herbivores, can be important in natural communities. Received: 9 January 1999 / Accepted: 29 April 1999     

Abrupt shifts in African savanna tree cover along a climatic gradient

Aim To describe patterns of tree cover in savannas over a climatic gradient and a range of spatial scales and test if there are identifiable climate‐related mean structures, if tree cover always increases with water availability and if there is a continuous trend or a stepwise trend in tree cover. Location Central Tropical Africa. Methods We compared a new analysis of satellite tree cover data with botanical, phytogeographical and environmental data. Results Along the climatic transect, six vegetation structures were distinguished according to their average tree cover, which can co‐occur as mosaics. The resulting abrupt shifts in tree cover were not correlated to any shifts in either environmental variables or in tree species distributions. Main conclusions A strong contrast appears between fine‐scale variability in tree cover and coarse‐scale structural states that are stable over several degrees of latitude. While climate parameters and species pools display a continuous evolution along the climatic gradient, these stable structural states have discontinuous transitions, resulting in regions containing mosaics of alternative stable states. Soils appear to have little effect inside the climatic stable state domains but a strong action on the location of the transitions. This indicates that savannas are patch dynamics systems, prone to feedbacks stabilizing their coarse‐scale structure over wide ranges of environmental conditions.     

Intraspecific competition in models for vegetation patterns: Decrease in resilience to aridity and facilitation of species coexistence

Patterned vegetation is a characteristic feature of many dryland ecosystems. While plant densities on the ecosystem-wide scale are typically low, a spatial self-organisation principle leads to the occurrence of alternating patches of high biomass and patches of bare soil. Nevertheless, intraspecific competition dynamics other than competition for water over long spatial scales are commonly ignored in mathematical models for vegetation patterns. In this paper, I address the impact of local intraspecific competition on a modelling framework for banded vegetation patterns. Firstly, I show that in the context of a single-species model, neglecting local intraspecific competition leads to an overestimation of a patterned ecosystem’s resilience to increases in aridity. Secondly, in the context of a multispecies model, I argue that local intraspecific competition is a key element in the successful capture of species coexistence in model solutions representing a vegetation pattern. For both models, a detailed bifurcation analysis is presented to analyse the onset, existence and stability of patterns. Besides the strengths of local intraspecific competition, also the difference between two species has a significant impact on the bifurcation structure, providing crucial insights into the complex ecosystem dynamics. Predictions on future ecosystem dynamics presented in this paper, especially on pattern onset and pattern stability, can aid the development of conservation programs.     

Hydrology-Driven Regime Shifts in a Shallow Tropical Lake

Shifts between alternative stable states have become a focus of research in temperate shallow lakes. Here we show that sharp transitions between a clear, macrophyte-dominated state and a turbid state without submerged plants can also occur in tropical floodplain lakes, albeit driven by a largely different set of mechanisms. We show how a shallow lake in the Pantanal becomes covered by an exploding population of the submerged macrophyte Egeria najas Planchon as the water level rises during the annual high-water period. Water clarity increases spectacularly in this period due to flushing with river water that has lost most of its suspended matter during its slow flow over the flooded vegetated plains. A few months later when the water level drops again, the submerged plant beds die and decompose rapidly, triggering a phase of increasing turbidity. During this period an increase in dissolved organic matter, suspended matter, and phytoplankton biomass results in a sharp deterioration in water clarity. The concomitant water level decrease largely counteracts the effects on the underwater light climate, so that the amount of light at the bottom may not differ in comparison with the high-water period. Therefore, changes in light climate seem unlikely to be the sole driver of the vegetation shifts, and other mechanisms may prevent recovery of the submerged vegetation until the next high-water episode. Also, contrary to what is found in temperate lakes, there is no evidence for top-down control of phytoplankton biomass associated with the macrophyte-dominated state in our tropical lake. Author Contributions  Simoni Maria Loverde-Oliveira, Vera Lúcia Moraes Huszar—conceived the study, Simoni Maria Loverde-Oliveira—performed research and analyzed data, Simoni Maria Loverde-Oliveira, Vera Lúcia Moraes Huszar, Nestor Mazzeo, Marten Scheffer—wrote the paper.     

放牧和围封条件下羊草碱化草地中斑发布格局研究

应用分形方法对松嫩平原羊草碱化划地中斑块分布格局进行定量估计,分析了放牧干扰和围封条件下斑块分布特征及动态。结果表明：随放牧强度增加,羊草斑块的斑块化指数增大,斑块面积范围的上下限变小,分布趋于破碎化和不均匀,斑块公布格局趋于复杂,自相似规律发生转折的尺度增多。围封后,１９８９～１９９３年羊草斑块总面积增加,斑块数目减少,斑块化指数降低,１９９４年后羊草斑块分布格局基本稳定。羊草斑块分布的双对数曲     

Coastal salt-marshes plant communities of the Salicornietea fruticosae class in Apulia (Italy)

An overview of the salt-marsh herbland and scrub vegetation belonging to the class Salicornietea fruticosae Br.-Bl. et Tx. ex A. Bolòs y Vayreda 1950 in Apulia is presented. Data available from literature have been supplemented with original relevés performed in different locations of the Apulia region. On the basis of a total of 297 relevés, fifteen communities have been defined, according to the traditional phytosociological system based on dominant and/or diagnostic taxa. For comparison purposes, the salt-marsh vegetation has been classified using numerical methods. The results obtained show that most of the clusters correspond to specific associations, and confirm the division into vegetation alliances and orders. Numerical analysis also allowed us to assign the proper allocation of some associations and plant communities drawn from literature. Five alliances, with plant communities characterized by specific ecological features, have been discriminated: Sarcocornion alpini and Arthrocnemion glauci (lower marshes), Salicornion fruticosae (middle marshes), Inulion crithmoidis and Suaedion brevofoliae (upper marshes). In addition, during the field work, a population of Halocnemum strobilaceum (Arthrocnemo-Halocnemetum strobilacei), new record for the Apulia region, has been found.     

黄土高原天然次生林植被演替过程中土壤团聚体有机碳动态变化

土壤团聚体物理保护是促进有机碳积累主要机制之一。以黄土高原子午岭林区天然次生林植被演替群落为对象,研究从农田、草地（白羊草,Bothriochloa ischaemum）、灌木林（沙棘,Hippophae rhamnoides）、先锋林（山杨,Populus davidiana）到顶级林（辽东栎,Quercus liaotungensis）5个植被演替阶段0-20 cm土壤团聚体稳定性和团聚体有机碳的动态变化,并分析团聚体有机碳的影响因素。结果表明：土壤团聚体稳定性随着植被演替显著提高（P<0.05）,顶级林的团聚体稳定性最高;土壤有机碳含量和各粒径土壤团聚体（> 2 mm、2-0.25 mm、0.25-0.053 mm、<0.053 mm）有机碳含量均随着植被演替而增加。除草地0.25-0.053 mm团聚体有机碳含量最高外,其他演替阶段均为0.25-2 mm粒径最高。根系生物量、凋落物生物量、微生物生物量碳、团聚体稳定性均与团聚体有机碳含量呈显著正相关关系（P<0.05）。总体而言,长期植被演替有助于团聚体稳定性和团聚体有机碳累积。     

Vegetation patches improve the establishment of Salvia mexicana seedlings by modifying microclimatic conditions

Human disturbance has disrupted the dynamics of plant communities. To restore these dynamics, we could take advantage of the microclimatic conditions generated by remaining patches of vegetation and plastic mulch. These microclimatic conditions might have great importance in restoring disturbed lava fields located south of Mexico City, where the rock is exposed and the soil is shallow. We evaluated the effects of both the shade projected by vegetation patches and plastic mulch on the mean monthly soil surface temperature (T _ss) and photosynthetic photon flux density (PPFD) and on the survival and growth of Salvia mexicana throughout the year. This species was used as a phytometer of microsite quality. Shade reduced the T _ss to a greater extent than mulch did. Both survival and growth were enhanced by shade and mulch, and the PPFD was related with seedling growth. During the dry season, plant biomass was lost, and there was a negative effect of PPFD on plant growth. At micro-meteorological scales, the use of shade projected by patches of vegetation and mulch significantly reduced the mortality of S. mexicana and enhanced its growth. Survival and growth of this plant depended on the environmental quality of microsites on a small scale, which was determined by the environmental heterogeneity of the patches and the landscape. For plant restoration, microsite quality must be evaluated on small scales, but on a large scale it may be enough to take advantage of landscape shade dynamics and the use of mulch to increase plant survival and growth.     

Gradual regime shifts in spatially extended ecosystems

Ecosystem regime shifts are regarded as abrupt global transitions from one stable state to an alternative stable state, induced by slow environmental changes or by global disturbances. Spatially extended ecosystems, however, can also respond to local disturbances by the formation of small domains of the alternative state. Such a response can lead to gradual regime shifts involving front propagation and the coalescence of alternative-state domains. When one of the states is spatially patterned, a multitude of intermediate stable states appears, giving rise to step-like gradual shifts with extended pauses at these states. Using a minimal model, we study gradual state transitions and show that they precede abrupt transitions. We propose indicators to probe gradual regime shifts, and suggest that a combination of abrupt-shift indicators and gradual-shift indicators might be needed to unambiguously identify regime shifts. Our results are particularly relevant to desertification in drylands where transitions to bare soil take place from spotted vegetation, and the degradation process appears to involve step-like events of local vegetation mortality caused by repeated droughts.     

The impact of grazing on plant communities,plant populations and soil conditions on salt marshes

Grazing an abandoned salt marsh causes retrogressive succession, since mid salt-marsh communities change into lower salt-marsh communities. Grazing and mowing are compared in detail. Both management practices enhance species diversity in an abandoned salt marsh. This can be attributed to the removal of litter. The finding that lower salt-marsh species appear more with grazing than with mowing or abandoning is not related to a higher soil salinity as compared to mowing or abandoning, but probably to locally baring of the soil by grazing animals. Only species of pioneer or unstable environments seem to have a persistent seed bank, for other species seed dispersal seems to be a limiting factor for their establishment.Nomenclature follows Heukels & van Ooststroom (1977) for species; Westhoff & den Held (1969) for syntaxa.Mrs R. Rusthoven analyzed the soil samples, Mr E. Leeuwinga drawed the figures, and Mrs J. O'Brien corrected the English text.     

Competition between Hardwood Hammocks and Mangroves

A bstract The boundaries between mangroves and freshwater hammocks in coastal ecotones of South Florida are sharp. Further, previous studies indicate that there is a discontinuity in plant predawn water potentials, with woody plants either showing predawn water potentials reflecting exposure to saline water or exposure to freshwater. This abrupt concurrent change in community type and plant water status suggests that there might be feedback dynamics between vegetation and salinity. A model examining the salinity of the aerated zone of soil overlying a saline body of water, known as the vadose layer, as a function of precipitation, evaporation and plant water uptake is presented here. The model predicts that mixtures of saline and freshwater vegetative species represent unstable states. Depending on the initial vegetation composition, subsequent vegetative change will lead either to patches of mangrove coverage having a high salinity vadose zone or to freshwater hammock coverage having a low salinity vadose zone. Complete or nearly complete coverage by either freshwater or saltwater vegetation represents two stable steady-state points. This model can explain many of the previous observations of vegetation patterns in coastal South Florida as well as observations on the dynamics of vegetation shifts caused by sea level rise and climate change.