Consumer Control of Salt Marshes Driven by Human Disturbance

Abstract:  Salt marsh ecosystems are widely considered to be controlled exclusively by bottom–up forces, but there is mounting evidence that human disturbances are triggering consumer control in western Atlantic salt marshes, often with catastrophic consequences. In other marine ecosystems, human disturbances routinely dampen (e.g., coral reefs, sea grass beds) and strengthen (e.g., kelps) consumer control, but current marsh theory predicts little potential interaction between humans and marsh consumers. Thus, human modification of top–down control in salt marshes was not anticipated and was even discounted in current marsh theory, despite loud warnings about the potential for cascading human impacts from work in other marine ecosystems. In spite of recent experiments that have challenged established marsh dogma and demonstrated consumer-driven die-off of salt marsh ecosystems, government agencies and nongovernmental organizations continue to manage marsh die-offs under the old theoretical framework and only consider bottom–up forces as causal agents. This intellectual dependency of many coastal ecologists and managers on system-specific theory (i.e., marsh bottom–up theory) has the potential to have grave repercussions for coastal ecosystem management and conservation in the face of increasing human threats. We stress that marine vascular plant communities (salt marshes, sea grass beds, mangroves) are likely more vulnerable to runaway grazing and consumer-driven collapse than is currently recognized by theory, particularly in low-diversity ecosystems like Atlantic salt marshes.     

盐沼植物群落研究进展：分布、演替及影响因子

盐沼是全球温带及亚热带地区的主要滨海湿地类型之一,在我国分布广泛。盐沼湿地生态系统敏感、脆弱且具有重要的生态系统服务功能。理解盐沼植物群落时空分布动态的一般规律与生态学机制,是开展盐沼生态系统研究的基础与关键。海陆交界的特殊环境特征是影响盐沼湿地植物群落的空间分布及演替过程的主要因素。在海洋潮汐作用下,盐沼湿地中的盐度、水淹强度、氧化还原电位等非生物因子往往呈梯度分布,这也导致了生物群落中种内、种间关系的变化。在非生物及生物因子的共同作用下,盐沼植物群落也往往沿高程梯度呈带状分布。环境变化是盐沼植物群落演替的驱动因素,在海岸线相对较为稳定的盐沼,植物群落的演替多属自发演替,而在靠近的大型河口的一些持续淤涨的盐沼,植物群落演替通常属于异发演替。沿海地区的水产业、流域上游及沿海地区的工程、污染及生物入侵等直接或间接的人类活动已对盐沼湿地植物群落的产生了深刻影响。经过数十年发展,国际上盐沼植物群落学研究的热点领域主要包括盐沼植物群落与其他生物群落的相互关系、植物群落在盐沼生态系统过程中的作用等。在全球变化背景下,盐沼植物群落对气候变化与海平面升高也日益成为盐沼植物群落学相关的热点。     

Nitrogen accumulation and plant species replacement in three salt marsh systems in the Wadden Sea

Salt marsh development on the coastal barrier island of Schiermonnikoog (The Netherlands) was compared with two other salt marsh systems in the Wadden Sea. Accretion rate, nitrogen accumulation and changes in plant species composition were investigated using chronosequences. The age of the marsh was estimated from aerial photographs and old maps. In 7230 plots, the elevation of the marsh surface, the thickness of the sediment layer (clay) and the presence of plant species was recorded. In addition, the nitrogen pool was measured at each successional stage. Accretion rates were similar in the three salt marshes. Higher accretion rates were found at younger marshes. A strong linear relationship between nitrogen pool size and thickness of the clay layer was found for the three marshes. The accumulation rate of nitrogen is therefore strongly related to the accretion rate. Thus, more nitrogen is present in the sediment of later successional stages where more clay has accumulated. On the high salt marsh (55 cm+MHT),Ameria maritima disappeared andArtemisia maritima, Juncus gerardi andElymus athericus established at sites with a thicker clay layer. On the low salt marsh (25 cm+MHT),Plantago maritima, Puccinellia maritima andLimonium vulgare disappeared andAtriplex (Halimione) portulacoides established. Apparently, with the accumulation of clay and therefore of nitrogen, tall growing species take over in salt marshes not grazed by livestock.     

Secondary succession dynamics in estuarine marshes across landscape-scale salinity gradients

Secondary succession plays a critical role in driving community structure in natural communities, yet how succession dynamics vary with environmental context is generally unknown. We examined the importance of seedling and vegetative recruitment in the secondary succession of coastal marsh vegetation across a landscape-scale environmental stress gradient. Replicate bare patches were initiated in salt, brackish, and oligohaline marshes in Narragansett Bay, Rhode Island, USA, and allowed to recover unmanipulated or with colonizing seedlings or vegetative runners removed for three years. Seed dispersal and seed bank studies were conducted at the same sites. We found that rates of recovery were 3-10 times faster in brackish and oligohaline marshes than in salt marshes. The fast pace of recovery in oligohaline marshes was driven by seedling colonization, while recovery was dominated by vegetative runners in brackish marshes and by both seedlings and runners in salt marshes. Seed and seedling availability was much greater in oligohaline marshes with up to 24 times the seed bank density compared with salt marshes. In contrast to the facilitated succession generally found in salt marshes, oligohaline marshes follow the tolerance model of succession where numerous species colonize from seed and are slowly displaced by clonal grasses whose recovery is slowed by preemptive competition from seedlings, contributing to the higher species diversity of oligohaline marshes. These findings reveal fundamental differences in the dynamics and assembly of marsh plant communities along estuarine salinity gradients that are important for conceptually understanding wetlands and for guiding the management and restoration of various types of coastal marshes.     

Die Umsetzung der FFH-Richtlinie in der Agrarlandschaft

The EU Habitats Directive (92/43/EEC) does include provisions for setting up the Natura 2000-network of protected areas based on listed species and habitats, and in addition specific regulations on species protection. Three Quarters of all designated sites (SCI’s) do not only include natural habitat types in a strict sense like forests or water-bodies, but also agricultural land. 18?% of the SCI’s even include between 25 and 50?% agricultural land and 24?% above 50?%. 48 species and three habitat types listed under the Habitats Directive have a clear focus in agriculture. Another eleven habitat types are dependant from a nature-friendly low intensity use or management. A large proportion of these habitats and species are actually in an unfavourable conservation status. The paper analyses the impact of EU nature conservation on agriculture based on the species and habitats falling under the Habitats Directive. On the other hand indirect negative influences of agriculture are discussed, that may have considerable impact on the future development of the conservation status of endangered habitats and species.     

滨海盐沼及其植物群落的分布与多样性

滨海盐沼是广泛存在于世界中、高纬度地区的一种湿地生态系统,具有抵御风暴潮灾害、净化污染物和为珍稀濒危生物提供适宜生境等重要的生态和经济价值。滨海盐沼因随高程变化而急剧变化的环境梯度和植物带状分布现象而为生态学者阐释自然界物种的分布机制提供了理想系统。主要概述了滨海盐沼的定义、特点、类型、全球分布以及潮汐作用、土壤盐度等环境因子特征;阐述了不同尺度下滨海盐沼的植物群落分布和多样性特征。在滨海盐沼植物群落的分布特征上,重点阐述了中尺度下的植物带状分布,即植物群落往往在白海向陆渐高的不同高程梯度上表现出显著的分带分布,不同植物各自占据该梯度上的一定区域。通常认为,带状分布是植物竞争和物理性胁迫共同调控的结果,但其在不同地理区域的普适性仍存争议。滨海植物群落多样性往往较低,在中、小尺度上盐沼植物多样性受控于盐度、潮汐等物理性胁迫、植物间相互作用等因子的作用;在大尺度上盐沼植物多样性可能随纬度增大而增加。系统深入地认识滨海盐沼植物群落生态格局和过程,将为气候变化、生物入侵等人类影响下的滨海盐沼生态系统的管理和恢复提供有益经验。     

Landscape ecology of the Dutch coast

This paper is a summary and elaboration of an earlier publication in Dutch on the compilation of a landscape-ecological map, scale 1 : 50 000, of the Dutch coast. It is argued that such an integrated map is the best basis for the conservation and management of the coastal dunes and salt marshes. It may be combined with local more detailed vegetation maps, some examples of which are mentioned in the context of management. The Dutch North Sea coast is a ca. 350 km long chain of sandy beaches and sand dunes, from only 100 m to more than 10 km wide. On sheltered stretches of dune coasts along estuaries in the Southwest and on the Wadden Sea islands, salt marshes have developed. The small-scale gradient structure of the beach-dune-salt marsh complex is emphasized.     

‘Coastal squeeze’— an historical perspective

The use of salt marsh for agricultural use has probably been going on for thousands of years. This paper will look at the situation in southeast England, particularly the Wash. Here enclosure for extended grazing and over the last several hundred years, for arable cultivation, may have taken place since Roman Times. An enclosure of Freiston Shore salt marsh (1979) and a proposed further enclosure at Gedney Drove End at about the same time raised concerns about the cumulative effect of these developments on nature conservation interests. These concerns prompted the nature conservation agencies to oppose the Gedney Drove End enclosure. Though the conservation argument did not persuade the Government that no further enclosure should take place, economic circumstances changed such that the pressure for the creation of new agricultural land diminished. This marked the end of ‘reclamation’ in the Wash. Since then, in the UK at least, there have been no further enclosures of salt marsh for agriculture. What were the arguments that lead to this change? Up to this point the perceived wisdom, in the Wash at least, was that as enclosure took place new inter-tidal land was created to seaward— with no net loss of inter-tidal land. Today we accept that this is not the case and a policy of managed re-alignment has increasingly been adopted in England, at sites ranging from the Porlock shingle ridge in north Devon to the salt marshes of Freiston in the Wash. It is argued that recognition of ‘coastal squeeze’ probably began in the Wash some 20 years ago. It would appear that we are now witnessing a reversal of this trend. How far will it take us? Will we see a return of the large expanses of tidal swamp around the Wash and elsewhere along the southern North Sea coast? Is this an inevitable consequence of global warming? This paper provides an historical perspective of the issues and arguments that have led us to recognize ‘coastal squeeze’ and the importance of dynamic coast, including the European Commission’s ‘EURosion Project’, which reported in 2004 (Salman et al. 2004).     

Conserving international coastal habitat networks on migratory waterfowl flyways

Migratory waterfowl depend on habitat networks at local, national and international scales for their survival. Coastal habitats are key areas for many waterfowl. Different species use different biotopes and in different places, so overall many parts of the coastal resource are important. As well as national conservation efforts, waterfowl conservation is increasingly focussed worldwide on collaborative international conservation, catalysed by several measures e.g. the Ramsar Convention, the EC Conservation of Wild Birds Directive, and the Bonn Convention Agreement on the Conservation of African/Eurasian Migratory Waterbirds. Several international conservation plans are under development for single species, but a more effective approach may be to develop plans, for assemblages of migratory birds with similar habitat requirements. All such plans must incorporate future sustainable use of the habitats on which the birds depend. Yet migratory bird and coastal habitat conservation is still often approached separately, despite the two being now closely linked to the development of the Natura 2000 site network in the European Community. Implementing the 1992 EC Habitats Directive requires the selection of coastal habitat sites for designation, set in national and international contexts of resource distribution. International coastal habitat inventories are needed to underpin this process. Combining such inventories with assessment of the flyway habitat requirements of waterfowl species and assemblages offers great potential for identifying international coastal habitat networks that meet the objectives of both habitat and migratory waterfowl conservation.     

Habitat utilization and alteration by the invasive burrowing isopod, Sphaeroma quoyanum, in California salt marshes

In recent years the pace of exotic species introduction and invasion has accelerated, particularly in estuaries and wetlands. Species invasions may affect coastal ecosystems in many ways. Alteration of sedimentary environments, through structure formation and burrowing, has particularly dramatic effects on coastal habitats. This study examines modification of channel bank and marsh edge habitat by the burrowing Australasian isopod Sphaeroma quoyanum Milne Edwards, in created and natural salt marshes of San Diego Bay and San Francisco Bay. Abundance and distribution patterns of this isopod species, its relationships with habitat characteristics, and its effects on sediment properties and bank erosion were examined seasonally, and in several marsh microhabitats. Mean isopod densities were 1541 and 2936 individuals per 0.25 m² in San Francisco Bay, and 361 and 1153 individuals per 0.25 m² in San Diego Bay study sites during December and July 1998, respectively. This isopod forms dense, anastomosing burrow networks. S. quoyanum densities did not differ as a function of location within creeks or location in natural versus created marshes. Burrows, which are on average 6 mm wide and 2 cm long, were associated with firm sediments containing high detrital biomass. Although erosion is a natural process along salt marsh banks, enclosure experiments demonstrated that isopod activities can enhance sediment loss from banks. In areas infested with S. quoyanum, losses may exceed 100 cm of marsh edge per year. The effects of habitat alteration by this invading species are likely to increase in severity in the coastal zone as these ecosystems become degraded. Received: 30 March 2000 / Accepted: 21 September 2000     

Behaviour of horses and cattle at two stocking densities in a coastal salt marsh

Livestock grazing has been practiced in salt marshes in the Wadden Sea area since 600 B.C. Currently livestock grazing is also applied for conservation management. However, effects of such grazing management on salt marshes are likely to vary depending on the species of livestock and stocking density due to differences in the behaviour of the animals. Yet, little is known about the behaviour of different livestock species and stocking densities grazing in salt marshes. We studied the grazing behaviour of horses and cattle by focal observation in an experiment with four different grazing treatments on a coastal salt marsh. In all treatments we recorded diet choice, movement and grazing activity, and spatial distribution. Livestock species shared an overlap in diet choice. Yet, horses more often foraged on the short grass Puccinellia maritima, while the cattle diet contained a higher amount of Aster tripolium. Horses travelled longer distances per day and spent more time grazing than cattle. Spatial distribution of cattle was significantly clustered, while horses showed a random distribution utilizing the whole area. Animal behaviour differs between livestock species and stocking densities with respect to diet choice, activity and spatial distribution.     

Coastal dynamics and adaptation to uncertain sea level rise: Optimal portfolios for salt marsh migration

The sustainability of dynamic natural systems often depends on their capacity to adapt to uncertain climate-related changes, where different management options may be combined to facilitate this adaptation. Salt marshes exemplify such a system. Marsh sustainability under rapid sea level rise requires the preservation of transgression zones - undeveloped uplands onto which marshes migrate. Whether these uplands eventually become marsh depends on uncertain sea level rise and natural dynamics that determine migration onto different land types. Under conditions such as these, systematically diversified management actions generally outperform ad hoc or non-diversified alternatives. This paper develops the first adaptation portfolio model designed to optimize the benefits of a migrating coastal system. Results are illustrated using a case study of marsh conservation in Virginia, USA. Results suggest that models of this type can enhance adaptation benefits beyond those available through current approaches.     

Causes and consequences of salt-marsh erosion in an Atlantic estuary in SW Spain

This study reports on the quantification of horizontal erosion by undermining of slopes in the atlantic mesotidal salt marshes of Odiel, SW Spain, and analyses its causes and consequences. Horizontal erosion has produced considerable losses of salt marsh area, including zones of mature salt marsh. Human pressure, such as from water-borne traffic or the exploitation of the slopes for the capture of bait, increases the natural erosion processes. The role of vegetation in protecting the slopes against erosion is studied. Channel banks covered with plants, many of which belong to species with long-living, above-ground creeping stems, were less eroded than those without vegetation cover. The enormous volume of sediments moved (ca. 7000m³ in one year) could contribute to the silting-up of the navigable channels of the estuary, so that continual dredging is necessary to allow access to shipping. These sediments are highly contaminated, and dredging exposes them more directly to the trophic network of the estuary. There is a considerable loss of natural resources. Finally, the integrated management of this coastal ecosystem is discussed.     

Sediment manganese and biogenic silica as geochemical indicators in estuarine salt marshes of coastal Georgia, USA

Salt-marsh estuarine sediments are not homogeneous. It is obvious that a steady state cannot be assumed if the depositional environments under which salt marshes accumulate change from subtidal through non-vegetated intertidal to vegetated intertidal state during their formation. In addition to these, the supply of sedimentary material depends on the tidal prism which changes as salt marshes are formed. Based on the study of cores collected in the estuarine region of Georgia, USA, it was found that the Mn/Al ratios in sediments of marsh cores change from subtidal through non-vegetated intertidal to vegetated intertidal. The relative variation of biogenic silica preserved in sediments of marsh core reflects the relative supply rate of organic carbon (phytoplankton) produced in the water column to the sediment surface at the time of deposition. In this paper it is proposed that sediment manganese and biogenic silica may be applied as geochemical indicators of changing depositional environments and organic carbon originating from phytoplankton in the estuarine salt marsh sediments.     

Secondary succession on a high salt marsh at different grazing intensities

Succession occurs on a large part of German salt marshes following abandonment or reduction of grazing. Its speed and effect on the biodiversity of salt marshes has been discussed in the literature. Permanent plot studies show site-dependent differences in successional outcome. If grazing is to be continued, there is uncertainty about the stocking rates that are optimal for the conservation of plant diversity. We present an eight year permanent plot study with different grazing treatments on a high salt marsh at the marsh island of Hallig Langeness, Germany, which is enclosed by a summer dike. The study site is owned by WWF. Successional outcomes on the permanent plots depend on grazing intensity and depth of the groundwater table which is correlated with soil salinity. Intermediate stocking rates support a mixture of halophytes and glycophytes, the relative proportion of which depends on the depth of the groundwater table. Cattle grazing at stocking rates of 0.6 livestock units per ha were found to be optimal for plant diversity conservation. At sites with deeper groundwater tables and no grazing, dominantElymus spp. stands develop and diversity is strongly reduced. If the groundwater table is high, succession following grazing abandonment is retarded and small halophytes prevail.     

Phytoplankton in a temperate-zone salt marsh: Net production and exchanges with coastal waters

Phytoplankton production and associated variables were measured in Flax Pond, a 52 ha salt marsh on the north shore of Long Island, New York, from July 1972 to October 1973. Measurements made up to five times per day, once per week, yielded a mean annual net primary production, determined by the ¹⁴C technique, of 20.5 mg C/m³/h; daily means were as high as 60.0 mg C/m³/h. However, when productivity was calculated for the entire marsh ecosystem, the shallow water in the salt marsh produced only 11.7 g C/m² of marsh/year. There was a net flux of phytoplankton from the coastal waters into the marsh; during the summer up to 0.2 g chlorophy 11/m² of marsh was carried in with the tides daily and remained in the marsh. Analysis of the productivity data, as well as variables associated with productivity (pH, standing crop, nutrients, extinction coefficient), indicated that the aquatic portion of the marsh behaved more as a net consumer rather than a net producer of phytoplankton.Research carried out at Brookhaven National Laboratory, supported by the U.S. Atomic Energy Commission and the National Science Foundation under Grant No. AG-375.     

Using environmental heterogeneity to plan for sea‐level rise

Environmental heterogeneity is increasingly being used to select conservation areas that will provide for future biodiversity under a variety of climate scenarios. This approach, termed conserving nature's stage (CNS), assumes environmental features respond to climate change more slowly than biological communities, but will CNS be effective if the stage were to change as rapidly as the climate? We tested the effectiveness of using CNS to select sites in salt marshes for conservation in coastal Georgia (U.S.A.), where environmental features will change rapidly as sea level rises. We calculated species diversity based on distributions of 7 bird species with a variety of niches in Georgia salt marshes. Environmental heterogeneity was assessed across six landscape gradients (e.g., elevation, salinity, and patch area). We used 2 approaches to select sites with high environmental heterogeneity: site complementarity (environmental diversity [ED]) and local environmental heterogeneity (environmental richness [ER]). Sites selected based on ER predicted present‐day species diversity better than randomly selected sites (up to an 8.1% improvement), were resilient to areal loss from SLR (1.0% average areal loss by 2050 compared with 0.9% loss of randomly selected sites), and provided habitat to a threatened species (0.63 average occupancy compared with 0.6 average occupancy of randomly selected sites). Sites selected based on ED predicted species diversity no better or worse than random and were not resilient to SLR (2.9% average areal loss by 2050). Despite the discrepancy between the 2 approaches, CNS is a viable strategy for conservation site selection in salt marshes because the ER approach was successful. It has potential for application in other coastal areas where SLR will affect environmental features, but its performance may depend on the magnitude of geological changes caused by SLR. Our results indicate that conservation planners that had heretofore excluded low‐lying coasts from CNS planning could include coastal ecosystems in regional conservation strategies.     

Predictors of specialist avifaunal decline in coastal marshes

Coastal marshes are one of the world's most productive ecosystems. Consequently, they have been heavily used by humans for centuries, resulting in ecosystem loss. Direct human modifications such as road crossings and ditches and climatic stressors such as sea‐level rise and extreme storm events have the potential to further degrade the quantity and quality of marsh along coastlines. We used an 18‐year marsh‐bird database to generate population trends for 5 avian species (Rallus crepitans, Tringa semipalmata semipalmata, Ammodramus nelsonii subvirgatus, Ammodramus caudacutus, and Ammodramus maritimus) that breed almost exclusively in tidal marshes, and are potentially vulnerable to marsh degradation and loss as a result of anthropogenic change. We generated community and species trends across 3 spatial scales and explored possible drivers of the changes we observed, including marsh ditching, tidal restriction through road crossings, local rates of sea‐level rise, and potential for extreme flooding events. The specialist community showed negative trends in tidally restricted marshes (?2.4% annually from 1998 to 2012) but was stable in unrestricted marshes across the same period. At the species level, we found negative population trends in 3 of the 5 specialist species, ranging from ?4.2% to 9.0% annually. We suggest that tidal restriction may accelerate degradation of tidal marsh resilience to sea‐level rise by limiting sediment supply necessary for marsh accretion, resulting in specialist habitat loss in tidally restricted marshes. Based on our findings, we predict a collapse of the global population of Saltmarsh Sparrows (A. caudacutus) within the next 50 years and suggest that immediate conservation action is needed to prevent extinction of this species. We also suggest mitigation actions to restore sediment supply to coastal marshes to help sustain this ecosystem into the future.     

Eutrophication and Consumer Control of New England Salt Marsh Primary Productivity

Abstract:  Although primary productivity in salt marshes is thought to be controlled by physical forces, recent evidence suggests that human disturbances can drive a switch to consumer control in these ecologically valuable ecosystems. We tested the hypothesis that nitrogen enrichment can trigger consumer control in salt marshes in Narragansett Bay, Rhode Island, with (1) a field experiment in which we manipulated nutrient availability (with nutrient additions) and insect herbivory (with insecticide application), (2) a survey of 20 salt marshes that examined the relationship between marsh nutrient status and herbivore pressure, and (3) insect herbivore removal at high and low nutrient input sites to directly test the hypothesis that nutrient enrichment is increasing insect herbivory in these marshes. Experimental nitrogen eutrophication initially increased plant productivity but eventually led to reduced plant biomass due to insect herbivory, and our surveys revealed that marsh nitrogen supply was a good predictor of herbivore damage to plants. Insects had minimal impacts on primary productivity in pristine marshes, but suppressed primary productivity in eutrophic salt marshes by 50–75%. Thus, eutrophication is currently triggering consumer suppression of primary productivity in New England salt marshes and may ultimately jeopardize the ecological and societal services these systems provide.