Ecological processes shaping Central Patagonian salt marsh landscapes

Plant zonation is one of the most conspicuous ecological features of salt marshes worldwide. In this work we used a combination of field transplant and greenhouse experiments to evaluate the importance of interspecific interactions and physical stress in the determination of the major plant zonation patterns in Central Patagonian salt marshes. There, Spartina alterniflora dominates the low marsh, and Sarcocornia perennis the high marsh. We addressed two questions: (i) What prevents Spartina alterniflora from colonizing the Sarcocornia perennis‐dominated high marsh zone? and (ii) What prevents Sarcocornia perennis from colonizing the Spartina alterniflora‐dominated low marsh zone? Our experimental transplants combined with neighbour exclusion treatments showed that the presence of Sarcocornia perennis negatively affects Spartina alterniflora, preventing it from surviving and/or spreading. Complementary field transplant and greenhouse experiments showed that Sarcocornia perennis did not survive the frequent tidal submersion by approximately 1.5 m of turbid seawater in the Spartina alterniflora zone, but its survival was independent of the presence of Spartina neighbours, and of the strong soil anoxia as well. Our results suggest that Spartina alterniflora is excluded by Sarcocornia perennis towards the low marsh, where frequent and prolonged submersion limit the survival of the latter. We provide and discuss key baseline information to facilitate the future design of ecophysiological experiments designed to accurately identify the exact mechanisms acting in every situation.     

Does low temperature prevent <Emphasis Type="Italic">Spartina alterniflora</Emphasis> from expanding toward the austral-most salt marshes?

Along the Atlantic coast of South America, the northern salt marshes (lower than 43°S) are dominated by Spartina species while the southern salt marshes (greater than 43°S) are dominated by Sarcocornia perennis. The most abundant Spartina species are Spartina densiflora which is present in most coastal marshes, and Spartina alterniflora that was never recorded above the ~42°25′S. It is not clear why S. alterniflora has not succeeded in the southern marshes, in which the low marsh zone remains as an extensive bared mud flat. We address the hypothesis that the absence of S. alterniflora in the south is driven by the cold temperatures inversely related with increasing latitudes along the East coast of Patagonia. To evaluate this hypothesis, we carried out an experiment in which we manipulated the temperature in combination with frost formation and photoperiod. We found that cold temperature produced a negative effect on S. alterniflora, and this effect seems accentuated by the frost but not by the reduction in the photoperiod. Our results support the hypothesis that the absence of S. alterniflora in the southernmost salt marshes of Patagonia is a consequence of the frost as an outcome of the co-occurrence of low temperature and high humidity. The importance of our results are discussed in the context of the global warming and how Spartina species enlarge their distributional range toward higher latitudes.     

Salt marsh colonization by a rocky shore invader: Balanus glandula Darwin (1854) spreads along the Patagonian coast

Balanus glandula, an east Pacific acorn barnacle from rocky shores, was introduced to Mar del Plata, Argentina more than 40 years ago and has spread over 17 latitudinal degrees southward. Here we report the first record of this species living in a soft-bottom environment colonizing the salt marsh plant species Limonium brasiliense, Spartina densiflora, S. alterniflora and Sarcocornia perennis. In addition, we describe the size frequency distribution, density and spatial distribution of the barnacles colonizing the different plant species. The size frequency distribution of Balanus showed a bimodal pattern in all plants. Barnacles were mostly large in S. densiflora, but small in S. alterniflora, with more balanced distributions of small and large barnacles on S. perennis and L. brasiliense. The highest density of barnacles was observed on S. perennis (x = 35.8 ind/cm², SD = 40.5) and S. alterniflora (x = 33.8 ind/cm², SD = 23), while the lowest on L. brasiliense (x = 1.5 ind/cm², SD = 1.18) and S. densiflora (x = 0.17 ind/cm², SD = 0.09). More than 90% of the barnacles on any given plant were found living. While barnacles colonized only the first few centimeters above the soil surface level in S. alterniflora and L. brasiliense, they reached their highest point on S. perennis. The finding of a rocky shore species successfully colonizing soft-bottom marshes within an invaded region brings new perspectives to discussions in biological invasion ecology, and raises additional considerations for coastal environmental management.     

崇明东滩盐沼植被扩散格局及其形成机制

长江河口盐沼植被的形成和演化是生物与其生长环境相互作用的结果。以崇明东滩盐沼植被典型扩散前沿为研究对象,2011至2012年期间调查了盐沼植被扩散前沿实生苗扩散、定居以及形成的扩散格局,同时测定了盐沼植被扩散前沿的潮滩冲淤动态和水文动力条件。研究结果表明,崇明东滩盐沼植被在扩散前沿形成了互花米草-光滩(Spartina alterniflora-Mudflat,SM)和互花米草-海三棱藨草-光滩(Spartina alterniflora-Scirpus mariqueter-Mudflat,SSM)两种典型的扩散格局。冲淤动态和水文动力条件是影响盐沼植被扩散格局的重要因子,尤其是在4—6月盐沼植物实生苗传播和定居的关键阶段。在此基础上,分析了东滩盐沼植被扩散前沿的生物-物理相互作用以及盐沼植被扩散格局的形成机制。研究结果不仅有助于理解长江河口地区盐沼植被扩散的生物物理过程,并对全球气候变化和海平面上升条件下滨海生态系统动态预测与湿地保护与管理具有重要的意义。     

EFFECT OF AN AMMONIUM NITRATE PULSE ON THE GROWTH AND ELEMENTAL COMPOSITION OF NATURAL STANDS OF SPARTINA ALTERNIFLORA AND JUNCUS ROEMERIANUS

A nitrogen (ammonium nitrate) pulse of 200 kg ha“¹ was added to stands of tall (1.0–1.5 m) Spartina alterniflora, short (< 0.5 m) Spartina alterniflora, and Juncus roemerianus in a Georgia salt marsh in July. The major response ten weeks later was an increase in the aerial biomass and a sharp reduction in the C/N ratio in short Spartina alterniflora. One year after the treatment the difference between the biomass in enriched and control plots was greater than ten weeks after treatment, but the C/N ratio in the plants in the treated plots had risen to that of the controls. The availability of nitrogen appears to limit growth in the middle elevation Georgia salt marsh (short S. alterniflora), but not in the lower (tall S. alterniflora) or higher (J. roemerianus) portions.     

Mosquito distribution in a saltmarsh: determinants of eggs in a variable environment

Two saltmarsh mosquitoes dominate the transmission of Ross River virus (RRV, Togoviridae: Alphavirus), one of Australia's most prominent mosquito‐borne diseases. Ecologically, saltmarshes vary in their structure, including habitat types, hydrological regimes, and diversity of aquatic fauna, all of which drive mosquito oviposition behavior. Understanding the distribution of vector mosquitoes within saltmarshes can inform early warning systems, surveillance, and management of vector populations. The aim of this study was to identify the distribution of Ae. camptorhynchus, a known vector for RRV, across a saltmarsh and investigate the influence that other invertebrate assemblage might have on Ae. camptorhynchus egg dispersal. We demonstrate that vegetation is a strong indicator for Ae. camptorhynchus egg distribution, and this was not correlated with elevation or other invertebrates located at this saltmarsh. Also, habitats within this marsh are less frequently inundated, resulting in dryer conditions. We conclude that this information can be applied in vector surveillance and monitoring of temperate saltmarsh environments and also provides a baseline for future investigations into understanding mosquito vector habitat requirements.     

Photosynthetic responses to light intensity of <Emphasis Type="Italic">Sarcocornia</Emphasis> taxa (Chenopodiaceae)

Salt marshes show a characteristic zonation of species distribution, which is correlated with marsh elevation. Radiation intensity and photoperiod change throughout the tidal frame. Photosynthetic response to light regime, 90–2450 μmol/(m² s), was determined in the laboratory for four closely related halophytic taxa of the genus Sarcocornia (Chenopodiaceae), which inhabit different positions in the tidal frame. Sarcocornia fruticosa, which germinates below vegetation cover and is found at high levels in the tidal frame, had the lowest maximum net photosynthetic rate and stomata conductance values. The two S. perennis subspecies demonstrated intermediate maximum net photosynthetic rates but S. perennis ssp. alpini reached light saturation point at higher light intensities. S. perennis ssp. perennis, found in the lowest elevations of the marshes, spends a significant proportion of its time submerged and therefore needs to take full advantage of available light. S. perennis ssp. alpini is exposed to very high light intensities in open salt pans at high elevations. The hybrid, S. perennis x fruticosa, which is currently found at intermediate elevations with less frequent inundation, had the highest net photosynthetic rate and chlorophyll a content. The ability to cope with high light levels may help to explain one of the environmental parameters, which affects distribution of four taxa throughout the tidal frame and also raise intriguing questions about the future role of the hybrid in the successive development of these marsh systems.     

Morphological adaptation to coastal marshes in spite of limited genetic structure in the Neotropical passerine Spartonoica maluroides (Aves: Furnariidae)

Tidal marshes present profound adaptive challenges to terrestrial vertebrates. For example, North American sparrows have relatively longer and thinner bills and darker dorsal plumage in coastal saltmarshes than in interior marshes. Bay‐capped wren‐spinetail (Furnariidae; Spartonoica maluroides) show a strong association with South American saltmarshes. We hypothesized that bay‐capped wren‐spinetail have similar morphological adaptations to North American sparrows to the saltmarsh environment, which would be indicative of the generality of selection on these traits in the coastal saltmarsh ecosystem. We captured individuals of S. maluroides from coastal saltmarshes and interior marshes. Populations were compared based on morphology and molecular markers. We found significant phenotypic differences in bill shape and plumage coloration (melanism) between S. maluroides populations from coastal and inland marshes. The low levels of genetic variation, weak geographical structure and shallow divergences, based on mitochondrial DNA and microsatellite data, suggest that coastal populations had a recent demographic expansion. Our results are consistent with the pattern of morphological divergence found between North American Emberizids. The possibility of convergent evolutionary adaptations between saltmarsh North American Emberizids and South American Furnariids suggests that there are strong selective pressures associated with saltmarsh environments on the beak, leading to adaptations for food acquisition, and on plumage coloration for better camouflage for predator avoidance (melanism). © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 78–91.     

Mangrove expansion and salt marsh decline at mangrove poleward limits

Mangroves are species of halophytic intertidal trees and shrubs derived from tropical genera and are likely delimited in latitudinal range by varying sensitivity to cold. There is now sufficient evidence that mangrove species have proliferated at or near their poleward limits on at least five continents over the past half century, at the expense of salt marsh. Avicennia is the most cold‐tolerant genus worldwide, and is the subject of most of the observed changes. Avicennia germinans has extended in range along the USA Atlantic coast and expanded into salt marsh as a consequence of lower frost frequency and intensity in the southern USA. The genus has also expanded into salt marsh at its southern limit in Peru, and on the Pacific coast of Mexico. Mangroves of several species have expanded in extent and replaced salt marsh where protected within mangrove reserves in Guangdong Province, China. In south‐eastern Australia, the expansion of Avicennia marina into salt marshes is now well documented, and Rhizophora stylosa has extended its range southward, while showing strong population growth within estuaries along its southern limits in northern New South Wales. Avicennia marina has extended its range southwards in South Africa. The changes are consistent with the poleward extension of temperature thresholds coincident with sea‐level rise, although the specific mechanism of range extension might be complicated by limitations on dispersal or other factors. The shift from salt marsh to mangrove dominance on subtropical and temperate shorelines has important implications for ecological structure, function, and global change adaptation.     

Seasonal patterns of daily net photosynthesis,transpiration and net primary productivity of Juncus roemerianus and Spartina alterniflora in a Georgia salt marsh

Summary Studies of the seasonal CO₂ and water vapor exchange patterns of Juncus roemerianus and Spartina alterniflora were conducted in an undisturbed marsh community on Sapelo Island, Georgia. Daily patterns of net photosynthesis, transpiration, leaf diffusive conductance and water-use efficiency in response to ambient conditions were monitored on intact, in situ plants. Net primary productivity was calculated from the daytime CO₂ fixation totals, nighttime CO₂ loss, leaf standing stock and aboveground to belowground biomass ratios for each plant type.The tall form of S. alterniflora had higher rates of photosynthesis and higher water-use efficiency values which, in conjunction with low respiratory losses and large leaf standing crop, results in high values of net primary productivity. The environmental factors in the marsh which permit these physiological responses occur in less than 10% of the marsh. Overall, the physiological capabilities of the short form of S. alterniflora were reduced in comparison to the tall form, but the combination of environmental factors which determine the physiological responses of this form occur in a much greater portion of the marsh, and the short form of S. alterniflora dominates the Sapelo Island marshes.The response patterns of the C₃ species, J. roemerianus, differed somewhat from the height forms of S. alterniflora. A large, seasonally constant leaf standing crop coupled with moderate rates of photosynthesis resulted in a net primary productivity value which was between the tall and short height forms of S. alterniflora. However, as with the tall S. alterniflora, the environmental conditions under which this high productivity and high water loss rate can be sustained are restricted to specific regions of the environmental gradient in the marsh.Contribution No. 435 from the University of Georgia Marine Institute     

Effects of exotic <Emphasis Type="Italic">Spartina alterniflora</Emphasis> on the habitat patch associations of breeding saltmarsh birds at Chongming Dongtan in the Yangtze River estuary,China

Smooth cordgrass (Spartina alterniflora) is one of the most invasive exotic plants of saltmarshes worldwide. To understand the effects of smooth cordgrass invasion on the habitat use and selection by breeding saltmarsh birds, we compared species number and abundance of breeding birds in native reed (Phragmites australis) and smooth cordgrass-invaded habitats (reed-cordgrass mixed habitats and cordgrass monocultures) at Chongming Dongtan in the Yangtze River estuary, China. We further examined the similarity of bird communities in different habitats and habitat selection by dominant bird species. For saltmarsh generalists, species number and abundance did not differ among the habitats. For saltmarsh specialists, species number and abundance did not differ in reed monocultures and reed-cordgrass mixed habitats but were significantly lower in cordgrass monocultures than in reed monocultures and reed-cordgrass mixed habitats. ANOSIM indicated that the difference in bird communities was larger between cordgrass monocultures and the habitats with reed than between the habitats with reed. The saltmarsh specialists preferred reed monocultures, while saltmarsh generalists avoided reed monocultures. Most species indicated no selection (neither preferred nor avoided) on reed-cordgrass mixed habitats, and no species preferred the cordgrass monocultures. The use of cordgrass monocultures by the common saltmarsh birds was negatively related to their body size. This study suggests that the spread of exotic smooth cordgrass has greatly affected the species composition and structure of local bird communities and has been especially disadvantageous to the saltmarsh specialists.     

Thermal stratification influences maturation and timing of spawning in a local Clupea harengus population

Maturation and timing of spawning in relation to temperature were studied in a local Atlantic herring Clupea harengus population inhabiting a small semi‐enclosed ecosystem (7 km²) separated from the larger outer fjord system by narrow sills on the west coast of Norway. Ambient temperatures varied annually up to 4° C during both the pre‐spawning and spawning periods from February to April, but without affecting the spawning time. Instead, the timing of spawning was found to be related to thermal stratification in response to spring warming, which occurred about the same time every year regardless of initial temperatures.     

闽江河口春季互花米草入侵过程对短叶茳芏沼泽土壤碳氮分布特征的影响

2014年4月,选择闽江口鳝鱼滩湿地中未被入侵的短叶茳芏群落(A)、互花米草入侵斑块边缘(B)以及互花米草入侵斑块中央(C)为研究对象,基于时空互代研究方法,探讨了互花米草入侵序列下湿地土壤碳氮空间分布特征的差异。结果表明,互花米草入侵显著降低了土壤的NO_3~--N含量(P0.05),但整体增加了NH_4~+-N含量,这与其入侵后导致湿地土壤颗粒组成发生显著变化(砂砾含量增加33.81%),进而促进了土壤的矿化作用和硝化作用,并有助于硝态氮的垂直淋失有关。互花米草入侵整体增加了土壤的碳氮含量和C/N比,与入侵进程和入侵前相比,互花米草入侵后湿地土壤的碳储量分别增加了8.73%和24.37%,氮储量则分别增加了10.22%和17.87%,这主要与其对闽江口湿地植物群落格局、养分生物循环以及强促淤作用引起的土壤颗粒组成等显著改变有关。研究发现,闽江口互花米草入侵对短叶茳芏湿地土壤碳氮含量的影响相对于江苏盐城、长江口以及杭州湾湿地的影响可能更为显著,其互花米草入侵较大改变了土壤中陆源和海源有机质的来源比例,使得入侵后湿地土壤养分的自源性增强。     

Effects of environmental gradients on the performances of four dominant plants in a Chinese saltmarsh: implications for plant zonation

Physical conditions and biotic interactions are believed to be the determinants of plant zonation in saltmarshes. However, in rapidly developing estuarine marshes, succession is regarded as the primary process responsible for plant zonation and it is controlled mainly by environmental factors. Salinity and inundation are two important factors responsible for the distribution pattern of dominant plants in coastal saltmarshes. Here we conducted a common garden experiment as well as a field transplanting to examine the responses of four dominant saltmarsh plants (native Scirpus mariqueter, Scirpus triqueter and Phragmites australis, and exotic Spartina alterniflora) in the Yangtze River estuary to environmental gradients, which may help us understand their current and potential zonation. The results showed that Scirpus adapted to freshwater and less inundated habitats, Phragmites performed well in brackish or freshwater environments with less inundation, and Spartina tolerated the highest salinity and deepest inundation. In the harshest environments (the highest salinity and water level), only Spartina performed well. In the mild environments, however, there were only minor differences in the performances among the four species. The potential ranges of Phragmites and Spartina were predicted to be larger than their current ones, and their lower boundaries might be set by tidal scour rather than edaphic factors. With the saltmarsh succession, invasive Spartina in the Yangtze River estuary might ultimately replace Scirpus, and alter the zonal patterns of native saltmarsh plants, which will lead to severe ecosystem consequences. Thus, proper management measures (e.g., repeated mowing) need to be implemented to control this invasive exotic plant, and restore the vulnerable ecosystems invaded by Spartina in the Yangtze River estuary.     

The saltmarshes of Coll and Tiree

Summary

A survey was made of the saltmarshes of the Inner Hebridean islands of Coll and Tiree. Seventeen plant communities are described and classified using the National Vegetation Classification (NVC). The saltmarshes are compared with those described from elsewhere in the region, and from other parts of Britain, and their species composition and range of communities place them firmly within the Type C — ‘western Scottish’ — marsh type described by Adam (1978). The islands support several saltmarsh communities and species of considerable phytogeographical and nature conservation importance.     

Exotic Spartina alterniflora invasion alters ecosystem–atmosphere exchange of CH4 and N2O and carbon sequestration in a coastal salt marsh in China

Coastal salt marshes are sensitive to global climate change and may play an important role in mitigating global warming. To evaluate the impacts of Spartina alterniflora invasion on global warming potential (GWP) in Chinese coastal areas, we measured CH₄ and N₂O fluxes and soil organic carbon sequestration rates along a transect of coastal wetlands in Jiangsu province, China, including open water; bare tidal flat; and invasive S. alterniflora, native Suaeda salsa, and Phragmites australis marshes. Annual CH₄ emissions were estimated as 2.81, 4.16, 4.88, 10.79, and 16.98 kg CH₄ ha^?1 for open water, bare tidal flat, and P. australis, S. salsa, and S. alterniflora marshes, respectively, indicating that S. alterniflora invasion increased CH₄ emissions by 57–505%. In contrast, negative N₂O fluxes were found to be significantly and negatively correlated (P < 0.001) with net ecosystem CO₂ exchange during the growing season in S. alterniflora and P. australis marshes. Annual N₂O emissions were 0.24, 0.38, and 0.56 kg N₂O ha^?1 in open water, bare tidal flat and S. salsa marsh, respectively, compared with ‐0.51 kg N₂O ha^?1 for S. alterniflora marsh and ?0.25 kg N₂O ha^?1 for P. australis marsh. The carbon sequestration rate of S. alterniflora marsh amounted to 3.16 Mg C ha^?1 yr^?1 in the top 100 cm soil profile, a value that was 2.63‐ to 8.78‐fold higher than in native plant marshes. The estimated GWP was 1.78, ?0.60, ?4.09, and ?1.14 Mg CO₂eq ha^?1 yr^?1 in open water, bare tidal flat, P. australis marsh and S. salsa marsh, respectively, but dropped to ?11.30 Mg CO₂eq ha^?1 yr^?1 in S. alterniflora marsh. Our results indicate that although S. alterniflora invasion stimulates CH₄ emissions, it can efficiently mitigate increases in atmospheric CO₂ and N₂O along the coast of China.     

Patterns of Fish Use and Piscivore Abundance within a Reconnected Saltmarsh Impoundment in the Northern Indian River Lagoon, Florida

Nearly all saltmarshes in east-central, Florida were impounded for mosquito control during the 1960s. The majority of these marshes have since been reconnected to the estuary by culverts, providing an opportunity to effectively measure exchange of aquatic organisms. A multi-gear approach was used monthly to simultaneously estimate fish standing stock (cast net), fish exchange with the estuary (culvert traps), and piscivore abundance (gill nets and bird counts) to document patterns of fish use in a reconnected saltmarsh impoundment. Changes in saltmarsh fish abundance, and exchange of fish with the estuary reflected the seasonal pattern of marsh flooding in the northern Indian River Lagoon system. During a 6-month period of marsh flooding, resident fish had continuous access to the marsh surface. Large piscivorous fish regularly entered the impoundment via creeks and ditches to prey upon small resident fish, and piscivorous birds aggregated following major fish movements to the marsh surface or to deep habitats. As water levels receded in winter, saltmarsh fish concentrated into deep habitats and emigration to the estuary ensued (200% greater biomass left the impoundment than entered). Fish abundance and community structure along the estuary shoreline (although fringed with marsh vegetation) were not analogous to marsh creeks and ditches. Perimeter ditches provided deep-water habitat for large estuarine predators, and shallow creeks served as an alternative habitat for resident fish when the marsh surface was dry. Use of the impoundment as nursery by transients was limited to Mugil cephalus Linnaeus, but large juvenile and adult piscivorous fish used the impoundment for feeding. In conclusion, the saltmarsh impoundment was a feeding site for piscivorous fish and birds, and functioned as a net exporter of forage fish to adjacent estuarine waters.     

滩涂湿地入侵种互花米草植被覆盖度的高空间分辨率遥感估算

互花米草是沿海滩涂生态系统的重要入侵物种,其分布状况和覆盖度是湿地生态研究的重要参数和基础。以宁德三沙湾(三都澳)滩涂湿地为研究区,以SPOT6 6 m空间分辨率卫星影像为数据源,对互花米草分布和植被覆盖度进行研究,并与同期10 cm空间分辨率无人机影像进行比较验证。结果表明,影像覆盖区域内互花米草总面积为20.19 km~2,其中蕉城区互花米草分布较广,面积为9.63 km~2,占研究区互花米草总面积的47.70%。互花米草植被覆盖度整体上以40%-60%和60%-80%的中、高覆盖度分布为主,其分布面积分别为5.44 km~2和4.95 km~2,占互花米草总分布面积的26.92%和24.52%,而40%以下的低覆盖度和80%以上较高覆盖度分布相对较少。SPOT6遥感影像估算得到的互花米草植被覆盖度具有较好的精度,与无人机影像值之间的均方根误差RMSE为0.117,线性回归决定系数R~2为0.918,可用于滩涂湿地植被覆盖度分析。     

人工管理和自然驱动下盐城海滨湿地互花米草沼泽演变及空间差异

将盐城国家级自然保护区核心区划分为人工管理区和自然湿地区两种模式,根据2000年、2006年、2011年3个时相的ETM+影像,运用RS、GIS技术和景观生态学方法,分析不同驱动力下互花米草沼泽景观变化的差异.结果表明:(1)人工管理区,2000-2011年,互花米草沼泽面积由272hm2增加到781hm2;平均分维数呈下降的趋势,聚合度指数呈上升的趋势,表明互花米草沼泽在空间上趋于聚集,在形状上趋于规则;从互花米草沼泽增长速率看,2000-2011年,呈现先快后慢的特征.在景观空间演变上,互花米草沼泽的平均宽度从598.679m增加到1719.002m,表现出向海陆两个方向扩张的特征.(2)自然湿地区,2000-2011年互花米草沼泽面积由2023hm2增加到3974hm2;平均分维数和聚合度指数呈先上升后下降的趋势;米草沼泽增长稳定,年增长约160-180 hm2/a,单位长度岸线年增长面积约为15 hm2· km-1·a-1左右;比较发现,自然湿地区互花米草沼泽的增长速度高于人工管理区的增长速度.在景观空间演变上,互花米草沼泽平均宽度从1625.586m增加到3193.317m,2000-2006年表现出向海洋单方向扩张的特征,2006-2011年表现出向海陆两个方向扩张的性质.(3)在自然条件下互花米草沼泽扩张受到气候、地貌过程、水文过程、植被类型及种间竞争的影响,表现出明显的带状特征.在人工管理下,通过建设拦水堤坝等,阻止了潮汐作用下海水的扩散能力,生态过程发生改变,致使生态系统类型向淡水湿地方向演变,抑制了互花米草沼泽的扩张,表现出一定的镶嵌格局.