盐城滨海滩涂湿地典型植物群落土壤微生物组成与结构特征

土壤微生物是生态系统维持正常结构与功能的重要组成部分,为探究盐城滩涂典型湿地土壤微生物群落结构特征,以江苏盐城滩涂互花米草、藨草、盐地碱蓬、芦苇及淤泥质光滩5种典型群落为对象,采用16S rRNA高通量测序技术分析0—10 cm(表层)、10—30 cm(中层)、30—60 cm(深层)土壤微生物多样性及群落结构。结果表明：(1)几种植物群落间,土壤微生物群落结构差异较大,主要体现在细菌群落结构的差异性,古菌群落结构差异相对较小。光滩与植物群落间,在土壤细菌种类及相对丰度上差异相对较大,互花米草群落与本土植物群落间,在微生物群落的细菌种类组成上存在较大差异;藨草群落土壤表层微生物群落结构与互花米草群落相似,深层与盐地碱蓬、芦苇群落相似。(2)同一群落不同层次土壤微生物群落结构相似,差异小于不同群落间土壤微生物群落的结构差异性;不同群落对应层次间,表深层土壤中五种群落土壤微生物多样性存在显著差异,中层土壤中五种群落微生物多样性差异不显著。总体上,植物群落类型对土壤微生物群落结构的影响大于土壤深度;与本土植物群落相比,互花米草群落土壤主要优势门微生物种类差异较小,但部分优势门微生物相对丰度...     

江苏滨海湿地芦苇和互花米草光合特性对模拟增温的响应

滨海湿地生态系统具有较高的初级生产力,是地球生态系统主要的碳库之一。然而,气候变暖和外来物种入侵通过改变植物光合特征性能使这一碳库的稳定性存在诸多的不确定性。利用在江苏盐城芦苇湿地和互花米草湿地建设的两个增温观测站,采用便携式光合荧光测量系统研究了本土植物芦苇（Phragmites australis）和入侵物种互花米草（Spartina alterniflora）光合特性对模拟增温的响应特征和机制。光合作用日调查变化曲线显示,增温使芦苇的净光合速率（P_n）、气孔导度（G_s）、蒸腾速率（T_r）和水分利用效率（WUE）都发生了显著的下降,但互花米草的各同名参数却表现出相反的变化特征,表明增温使互花米草的生理机能增强,促进了光合作用;根据P_n和胞间CO₂浓度（C_i）的变化趋势推断互花米草和芦苇光合速率变化均为非气孔限制因素驱动。利用直角双曲线修正模型拟合的光响应曲线结果显示,芦苇增温组的光响应曲线位于对照组下方,互花米草增温组的光响应曲线位于对照组上方;增温降低了所研究植物的光补偿点（LCP）,表明增温可提高两种植物利用弱光的能力;增温增加了互花米草光补偿点（LCP）和光饱和点（LSP）间值域范围,从而揭示了增温可有效提高互花米草利用光合有效辐射的能力;增温降低了芦苇的暗呼吸速率（R_d）,芦苇受到增温的胁迫,在减缓新陈代谢的同时也减弱了光合作用,而互花米草表现出相反特征。由此推测增温条件下入侵植物互花米草同化大气CO₂的能力（即碳汇能力）优于本土植物芦苇,这也是互花米草成为入侵物种的主要原因之一。     

Misting and nitrogen fertilization of shoots of a saltmarsh grass: effects upon fungal decay of leaf blades

We conducted a 12-week field manipulation experiment in which we raised the nitrogen availability (ammonium sulfate fertilization to roots) and/or water potential (freshwater misting) of decaying leaf blades of a saltmarsh grass (smooth cordgrass, Spartina alterniflora) in triplicate 11-m² plots, and compared the manipulated plots to unmanipulated, control plots. The ascomycetous fungi that dominate cordgrass leaf decomposition processes under natural conditions exhibited a boosting (>2-fold) of living standing crop (ergosterol content) by misting at the 1 st week after tagging of senescent leaves, but afterwards, living-fungal standing crop on misted blades was equivalent to that on control blades, confirming prior evidence that Spartina fungi are well adapted to natural, irregular wetting. Misting also caused 2-fold sharper temporal declines than control in instantaneous rates of fungal production (ergosterol synthesis), 5-fold declines in density of sexual reproductive structures that were not shown by controls, and 2-fold higher rates of loss of plant organic mass. Extra nitrogen gave a long-term boost to living-fungal standing crop (about 2-fold at 12 weeks), which was also reflected in rates of fungal production at 4 weeks, suggesting that saltmarsh fungal production is nitrogen-limited. Although bacterial and green-microalgal crops were boosted by manipulations of nitrogen and/or water, their maximal crops remained 0.3 or 2% (bacteria or green microalgae, respectively) of contemporaneous living-fungal crop. The fungal carbon-productivity values obtained, in conjunction with rates of loss of plant carbon, hinted that fungal yield can be high (>50%), and that it is boosted by high availability of nitrogen. We speculate that one partial cause of high fungal yield could be subsidy of fungal growth by dissolved organic carbon from outside decomposing leaves.     

Water potential of standing-dead shoots of an intertidal grass

Summary Dead stems and leaves of smooth cordgrass (Spartina alterniflora Loisel.) undergo substantial fungal decay in the standing position. We conducted a series of experiments to determine the probable range of water availabilities that cordgrass-fungal decomposers experience; we determined the effects of duration of wetting and drying, site on shoots, leaching of osmotica, and stage of decomposition on water potential () of cordgrass shoots. Dried pieces of cordgrass shoots took up water rapidly when submerged, rising from values which were probably less than –150 megapascals (MPa) water potential to about –3.5 (soaked in seawater) or –1.5 (soaked in tapwater) within 5 min. Air-drying resulted in a return to low (<–7.5 MPa) within an hour. Literature reports show that most litter and wood-decomposing fungi which have been tested cannot grow at more negative than –6 MPa. The more lignified stems showed a more negative matric than leaves at water contents greater than 25% fresh weight. As leaves decomposed in the marsh, their increased (from –1.7 to –0.5 MPa, under standard conditions of 30 min freshwater soaking and 30 min air-drying), similar to what other investigators have found for wheat and forest litter. The water content at which cordgrass leaves reached –6 MPa was about 23% fresh weight, within the range (15–32%) found for wheat straw and forest litter.     

Restoration Success of Backfilling Canals in Coastal Louisiana Marshes

The need for effective marsh restoration techniques in Louisiana is a pressing issue as the state continues to lose coastal wetlands. Returning spoil banks to canals, known as “backfilling,” is an attractive restoration option because it restores marsh, prevents future wetland loss, and is cost effective. The restoration of 30 canals backfilled 20 years ago was examined in this study and compared to restoration success 5 and 10 years after backfilling. Ultimately, the success of backfilling was controlled by the amount of spoil returned to the canal and the position of the canal in the marsh. Up to 95% of the spoil area was restored to marsh when the spoil banks were adequately removed, but only 5% of the spoil area was restored at sites where spoil removal was poor. Restoration of organic matter, bulk density, water content, and plant communities of the former spoil areas was also constrained by the adequacy of spoil removal. Backfilling restored up to 90% of the organic matter, 92% of the bulk density, and 93% of the water content after 20 years at sites where spoil was properly removed. Canals backfilled in areas of intact marsh showed greater restoration success than canals backfilled in highly degraded marshes. This study indicates that the benefits of backfilling continue to increase over time, although complete restoration will take longer than 20 years. Improving the completeness of spoil removal, coupled with appropriate site selection, could speed up the restoration process and enhance the success of future backfilling projects.     

Genetic diversity and population genetic structure of saltmarsh Spartina alterniflora from four coastal Louisiana basins

Seventy-two Spartina alterniflora accessions originating from four coastal Louisiana basins (18 accessions per basin) were used to evaluate the genetic structure of this native perennial low-intertidal plant species. The objective of this study was to determine the population genetic structure and diversity of S. alterniflora accessions originating from these four basins using amplified fragment length polymorphism (AFLP) markers. A total of 250 unambiguous and highly repeatable AFLP markers, 186 of which (74.4%) were polymorphic, were obtained using four primer combinations. Overall, pairwise similarity estimates between accessions ranged from 0.70 to 0.93 (average = 0.80) with only a small portion of alleles (0.54–1.08%) unique to each basin. The average H_s (genetic diversity within coastal basins) was 0.20 with an H_s values of 0.19, 0.20, 0.20, and 0.21 for Mermentau, Terrebonne, Calcasieu, and Barataria-Breton basin, respectively. AMOVA analysis showed no genetic structure among basins, with the majority of genetic variation, 96.6%, residing within the basins. There was no indication of isolation by distance. Our results suggest that maintaining high levels of genetic diversity can be accomplished through the use of an adequate number of S. alterniflora samples collected within any large basin. Choosing parental lines from several Louisiana coastal basins for breeding purposes may not significantly increase genetic variability among the progeny lines.     

室内模拟波浪对2种典型滨海湿地挺水植物生物力学特征的影响

选取本地物种芦苇和入侵物种互花米草为研究对象,利用波浪装置对生长期的植物进行波浪处理后对其形态特征,根、茎、叶的拉力、弯曲特性进行研究,以探讨湿地植物对波浪作用的生物力学响应。结果表明,芦苇、互花米草经波浪处理后的茎直径均显著高于对照组（P=0.008,P=0.03）;互花米草总体呈现拉力或者弯曲载荷高于芦苇的现象（如茎秆）,但应力（根系）或静曲强度（茎秆）却要低于芦苇,即对应了互花米草比芦苇茎秆更粗壮的现象,但芦苇自身的强度会更强,其木质素和纤维素的结构成分显示出显著高于互花米草的现象（P<0.001,P<0.001）。总体来看,通过短时间的波浪模拟,植株在形态特征上发生了显著的变化,而植物自身的生物力学特性与形态、结构成分都息息相关。研究结果对于评价芦苇和互花米草在应对海浪影响的潜力、从植物生物力学角度探究滨海湿地的保护和修复都提供了一定的参考。     

Oligochaeta in Spartina stems: the microdistribution of Enchytraeidae and Tubificidae in a salt marsh,Sapelo Island,USA

The distribution and abundance of Enchytraeidae and Tubificidae in and around Spartina alterniflora plants in a tidal salt marsh on Sapelo Island, Georgia, USA were studied using two different sampling techniques: wet funnel extraction and stem dissection. At least 80% of all worms inhabited leaf sheaths at the bases of S. alterniflora plants, and densities were low in sediment, root and surface debris samples. Oligochaete densities were dependent on the position within the marsh, the height on stems and the stage of sheath decay. Six predominant species were identified and included Marionina appendiculata, Marionina spartinae, Marionina waltersi, Marionina paludis, and Monopylephorus parvus. Individual species were distributed differently on stems and enchytraeids were more common than tubificids on standing-dead and further up S. alterniflora stems. Estimates of oligochaete densities in salt marsh habitats are increased dramatically when the numbers of worms on stems are considered. Possible advantages of the stem microhabitat are discussed in relation to the biology and ecology of oligochaetes.     

Sediment addition enhances transpiration and growth of Spartina alterniflora in deteriorating Louisiana Gulf Coast salt marshes

Transpiration, leaf conductance, net photosynthesis, leaf growth, above-ground biomass and regeneration of new culms were studied in a rapidly subsiding Spartina alterniflora Lois. salt marsh following the addition at 47 and 94 Kg m^–2 of new sediment. Plant growth was enhanced in response to sediment addition as was evident by a significant increase in leaf area, above-ground biomass production and regeneration of new culms (p 0.05). Leaf conductance and transpiration rates were significantly greater in sediment treated plants than in control plants (p 0.05). Enhanced production of culms per unit area of marsh resulted in increased leaf area which allowed a greater capacity for net photosynthesis and contributed to increases in above-ground biomass of sediment treated plots.     

互花米草生物量变化对盐沼沉积物有机碳的影响

以江苏王港典型互花米草(Spartina alterniflora)盐沼湿地为研究对象,分析光滩及互花米草滩沉积物中有机碳的水平和垂向分布特征,了解互花米草生物量的季节动态变化,探讨二者之间的相互关系,在此基础上研究互花米草生物量分布和季节变化对沉积物中有机碳(TOC)含量的影响。结果表明,互花米草枯落物中的有机碳数量在两个月内衰减了40%,而表层沉积物中TOC含量及其中互花米草来源TOC所占比例的变化,均与互花米草地表枯落物量的季节变化存在两个月的"相位差",这与枯落物快速分解时间大致吻合,说明枯落物是表层沉积物中TOC的重要来源。高达60%的互花米草地下生物量分布在0—20cm深度内,该深度范围内沉积物中TOC含量较高,且TOC主要来源于互花米草。此外,不同深度TOC含量与地下生物量之间存在良好的正相关关系,说明地下生物量是影响沉积物TOC含量的重要因子。研究区互花米草年固碳能力为2274g m-2a-1,盐沼沉积物中TOC埋藏速率达到了470 g m-2a-1,是地表一个重要的碳汇;同时研究区每年向近岸水域输出大量的TOC,是近岸海域生态系统的一个重要碳源。     

A technique for distinguishing between bacterial and non-bacterial respiration in decomposing Spartina alterniflora

A number of antibiotics were used to suppress bacterial activity in decomposing Spartina alterniflora. The effectiveness of each treatment was quantified using INT formazan vital staining and epifluorescent microscopy. Bacterial suppression of selected treatments was verified using standard plate count procedures. Chloramphenicol treated samples (exhibiting 87–90% bacterial suppression) were analyzed respirometrically and found to consume only 30% less O₂ than controls. Non-bacterial respiration (probably fungal) accounted for 70% of the respiration.     

Effects of saltmarsh invasion by Spartina alterniflora on arthropod community structure and diets

Invasive plants strongly affect physical and biotic environments of native ecosystems. Insects and other arthropods as one of the major components of many ecosystems are very sensitive to subtle changes in abiotic and biotic environments. We examined the effects of exotic Spartina alterniflora invasion on community structure and diets of arthropods in a saltmarsh previously dominated by native Phragmites australis in Yangtze River estuary through net sweeping and plant harvesting methods and stable isotope analysis. Our results showed that diversity indices were not significantly different between exotic and native plant communities, but the total abundance of insects estimated through plant harvesting method was found to be lower in Spartina monoculture than that in Phragmites monoculture. Community structure of insects in Spartina monoculture was dissimilar to that in Phragmites monoculture and Phragmites–Spartina mixture. Moreover, stable carbon isotope patterns of arthropods were significantly different between Phragmites and Spartina monocultures. Although some native arthropods (perhaps generalists) shifted their diets, many native taxa did prefer Phragmites to Spartina even in Spartina monoculture. Spartina invasions resulted in reduced abundances of some arthropds, and increased dominance of others feeding preferably on Spartina. This study provides evidence that invasive plants can change the community structure and diets of native arthropods, which will eventually alter the arthropod food web, and affect the integrity and functioning of native ecosystems within a nature reserve that has been set aside for conserving the native biodiversity and maintaining the ecosystem integrity. In this sense, Spartina invasions in the Yangtze River estuary need to be managed appropriately.     

互花米草入侵对鸟类的生态影响

生物入侵威胁本地物种生存,破坏生态系统的结构和功能,是导致全球生物多样性丧失的重要原因之一。外来植物是入侵生物中的重要一类,可以显著改变本地植被群落,并影响其他生物类群。鸟类作为生态系统中的较高营养级,对由入侵植物引起的栖息地变化十分敏感。互花米草自引入中国沿海以来,其分布区域不断扩散,多数研究认为互花米草入侵造成本地生物多样性降低和生态系统退化。系统梳理了互花米草入侵对鸟类栖息地、食物资源、繁殖、群落等方面的生态影响。主要负面影响有:(1)植被群落结构不利于鸟类栖息、筑巢、觅食;(2)鸟类食物资源丰度和多样性下降;(3)本地鸟类种群数量和物种多样性显著下降。在我国东部沿海湿地,互花米草入侵已经显著改变了植被与鸟类分布格局。但随着入侵历史的增长,少数小型雀形目鸟类却可以逐渐适应互花米草生境。互花米草入侵为某些非本地鸟类提供了空白生态位,在一定程度上丰富了本地物种多样性,对互花米草的快速清除反而可能不利于已适应并依赖互花米草生境的鸟类。综上,认为互花米草入侵对鸟类群落甚至整个生态系统的影响可能需要更多研究进行综合评价,应开展长期、大尺度、多因子的监测研究和多物种比较研究,建立生态评价模型并制定科学有效的互花米草管理对策。     

互花米草幼苗对重金属镉胁迫的生理响应

以互花米草为实验材料,通过模拟海水河沙培养实验,研究不同浓度镉胁迫对互花米草生理生化的影响。结果表明:随着镉浓度的增大,互花米草叶、根生物量逐渐降低,膜透性、丙二醛、SOD、POD酶活性随着镉浓度的增加而增加,其酶抗性也发挥到最大的程度。随着镉浓度的增加互花米草的光合特性发生较大变化,净光合速率、胞间CO₂和气孔导度都下降和减少。     

A simple approach to optimal control of invasive species

The problem of invasive species and their control is one of the most pressing applied issues in ecology today. We developed simple approaches based on linear programming for determining the optimal removal strategies of different stage or age classes for control of invasive species that are still in a density-independent phase of growth. We illustrate the application of this method to the specific example of invasive Spartina alterniflora in Willapa Bay, WA. For all such systems, linear programming shows in general that the optimal strategy in any time step is to prioritize removal of a single age or stage class. The optimal strategy adjusts which class is the focus of control through time and can be much more cost effective than prioritizing removal of the same stage class each year.     

Diel methane emissions in stands of Spartina alterniflora and Suaeda salsa from a coastal salt marsh

In this study, we aimed to understand the influence of plant type on the monthly variations of diel CH₄ fluxes from Spartina alterniflora and Suaeda salsa of coastal salt marshes at three growth stages (July, August and September). Dissolved CH₄ concentrations in porewater and sediment redox potentials were monitored, as were aboveground plant biomass and stem densities. CH₄ fluxes exhibited clear monthly variations and peaked in September in the S. alterniflora and S. salsa mesocosms. However, no discernible diel variation was observed in the CH₄ flux in the S. salsa mesocosm, probably due to its weak gas transport capacity. By contrast, notable diel variations of CH₄ flux with the peak of 1.42 and 3.67 mg CH₄ m⁻² h⁻¹ at 12:00 and the lowest of 0.75 and 2.11 mg CH₄ m⁻² h⁻¹ at 3:00 or 6:00 were observed in the S. alterniflora mesocosm on 11 August and 11 September, respectively, but not in July mainly due to low plant biomass masking diel variations in the porewater CH₄ concentration. The ratios of the maximum flux to minimum flux over the course of the day in the S. alterniflora mesocosm on 10 July, 11 August and 11 September were 1.28, 1.89 and 1.76, respectively, and corresponding values for porewater CH₄ concentration were 1.31, 1.39 and 1.17, respectively. CH₄ flux significantly correlated with CH₄ concentration in porewater, and both were significantly related to air temperature. These findings indicate that CH₄ production and CH₄ flux at the middle growth stage (August) exhibited greater responses to changes in air temperature, which in turn induced the higher diel variation. The higher diel cycle for CH₄ flux in August than in September was likely due to the higher proportion of CH₄ oxidized during diffusion within the aerenchyma system. Our results suggest that the extent of diel variations in CH₄ flux may have depended on the gas transport capacity of plants, and the highest diel variation occurred at the middle growth stage.     

Hydrologic,vegetation, and substrate characteristics of floating marshes in sediment-rich wetlands of the Mississippi river delta plain,Louisiana, USA

Floating marshes occur over 70% of the western Terrebonne Basin, Louisiana, USA, freshwater coastal wetlands. They are of several types: A free-floating thick-mat (45–60 cm) marsh dominated by Panicum hemitomon and Sagittaria lancifolia; a thick mat marsh dominated by Panicum hemitomon and Sagittaria lancifolia that floats part of the year, but whose vertical floating range is damped compared to adjacent water; and an irregularly-floating thin mat (< 30 cm) dominated by Eleocharis spp. in the spring and Ludwigia leptocarpa and Bidens laevis in the summer and fall. Floating mats must be almost entirely organic in order to be buoyant enough to float. The western Terrebonne wetlands receive large winter/spring supplies of suspended sediments from the Atchafalaya River. Even though sediment concentrations in the adjacent bayou are as high as 100 mg l^–1, the Panicum hemitomon/Sagittaria lancifolia free-floating marsh probably receives no over-surface sediments since it floats continuously. The bulk density data of the damped-floating marsh, however, suggest some mineral sediment input, probably during winter when this marsh is submerged. These two types of floating marsh could not have developed in the present sediment regime of the Atchafalaya River, but as long as they remain floating can continue to exist. Thin floating mats are found in areas receiving the least sediment (<20 mg 1^–1 suspended sediment concentration in adjacent bayous). This low sediment environment probably made possible their formation within the past 20 years. They may represent a transitional stage in mat succession from (1) existing thick-mat floating marsh to a degrading floating marsh, or (2) a floating marsh developing in shallow open water.Corresponding editor: D. Whigham     

Oxygen loss from Spartina alterniflora and its relationship to salt marsh oxygen balance

Spartina alterniflora has been reported to lose significant amounts of oxygen to its rhizosphere with potentially important effects on salt-marsh biogeochemical cycling and plant productivity. The potential significance of this oxidative pathway was evaluated using laboratory split-chamber experiments to quantify oxygen loss from intact root systems under a wide variety of pre-treatment and incubation conditions including antibiotics to inhibit microbial respiration. The aerenchyma system of S. alterniflora was found to transport O₂, N₂, Ar, and CH₄ from above-ground sources to its below-ground roots and rhizomes. While non-respiratory gases were observed to move from the lacunae to water bathing the root systems, net O₂ loss did not occur; instead oxygen present outside of the roots/rhizomes was consumed. Net oxygen loss was found when resistance to gas transport was reduced in the lacunae-rhizosphere pathway by placing the root systems in a gas phase and when plant respiration was significantly reduced. Root system respiration appeared to be the major variable in the plant oxygen balance. When root and rhizome respiration was inhibited using poisons or lowered by cooling, the oxygen deficit was greatly reduced and oxygen loss was indicated. The effect of seasonal temperature changes on root system oxygen deficit presents a possible explanation as to why Spartina produces root systems with respiration rates that cannot be supported by gas transport. Overall, while oxygen loss from individual plant roots is likely, integrating measured root system oxygen loss with geochemical data indicates that the mass amount of oxygen lost from S. alterniflora root systems is small compared to the total oxygen balance of vegetated salt marsh sediments.     

Spread of Exotic Cordgrasses and Hybrids (Spartina sp.) in the Tidal Marshes of San Francisco Bay, California, USA

Four species of exotic cordgrass (Spartina sp.) occur in the San Francisco estuary in addition to the California native Spartina foliosa. Our goal was to map the location and extent of all non-native Spartina in the estuary. Hybrids of S. alterniflora and S. foliosa are by far the most numerous exotic and are spreading rapidly. Radiating from sites of deliberate introduction, S. alterniflora and hybrids now cover ca. 190 ha, mainly in the South and Central Bay. Estimates of rate of aerial increase range from a constant value to an accelerating rate of increase. This could be due to the proliferation of hybrid clones capable of rapid expansion and having superior seed set and siring abilities. The total coverage of 195 ha by hybrids and other exotic cordgrass species is slightly less than 1% of the Bay's tidal mudflats and marshes. Spartina anglica has not spread beyond its original 1970s introduction site. Spartina densiflora has spread to cover over 5 ha at 3 sites in the Central Bay. Spartina patens has expanded from 2 plants in 1970 to 42 plants at one site in Suisun Bay. Spartina seed floats on the tide, giving it the potential to export this invasion throughout the San Francisco estuary, and to estuaries outside of the Golden Gate. We found isolated plants of S. alterniflora and S. densiflora in outer coast estuaries north of the Bay suggesting the likelihood for the San Francisco Bay populations to found others on the Pacific coast.     

Nitrogen cycling in Louisiana Gulf Coast brackish marshes

Nitrogen fixation and nitrogen accumulation were measured in a Louisiana Spartina patens brackish marsh. Using the acetylene reduction technique calibrated with direct ¹⁵N₂ assimilation, an equivalent of 90.0 µ g N g^–1 yr^–1 was fixed. Fixation was greater in the summer months and in the upper portion of the soil profile. Extractable ammonium increased with depth and was negatively correlated with ethylene production. Average ammonium concentration in the sediment was 39 µg NH₄ ⁺-N g^–1 sediment. Cesium-137 dating of the soil profile showed the marsh was vertically accreting at a rate of 0.60 cm yr^–1. Calculations using vertical accretion rate, bulk density, and total nitrogen content of sediment indicate that the marshes are accumumating 7.2 g Nm^–2 yr^–1 thus serving as a major nitrogen sink. Measured nitrogen fluxes were incorporated with existing flux measurement in developing a nitrogen budget for the marsh.