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.     

The impact of tidal inundation on salt marsh vegetation after de-embankment on Langeoog Island, Germany—six years time series of permanent plots

Salt marsh succession after de-embankment was monitored on the East Frisian barrier island Langeoog by investigating permanent plots. Seventy years after embankment salt marsh plants were once again influenced mainly by the tidal regime. From 2002 to 2004 the former high marsh and glycophytic vegetation died out and was replaced by species of lower salt marsh zones. Nitrophytic halophytes like Suaeda maritima, Atriplex prostrata and Artemisia maritima established first because of the high nutrient content in the soil, a direct result of former vegetation decay. With decreasing nitrogen afterwards other species became more competitive. Until 2007 Atriplex portulacoides became more dominant in the lower marsh and Elymus athericus reached dominance in areas where grazing has been abandoned in the high marsh. The dynamics in the study area is much lower than in natural marshes due to the still existing drainage system. Therefore vegetation units with low species diversity are widespread.     

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。     

Short-term and long-term facilitation of goose grazing by livestock in the Dutch Wadden Sea area

We studied the impact of livestock grazing on the distribution ofBranta bernicla bernicla (Dark-bellied Brent goose) in the Dutch Wadden Sea during spring. It was hypothesized that livestock facilitate short-term (within-season) grazing for geese as well as long-term (over years). Therefore we measured grazing pressure by geese in salt marsh and polder areas that were either grazed (spring-grazed) or ungrazed during spring (summer-grazed). Additionally, we carried out a preference experiment with captive geese to test the preference between spring-grazed and summer-grazed polder swards. We furthermore compared patterns of use by geese between long-term ungrazed and grazed salt marshes. In May, there is a difference in grazing pressure by geese between polder pastures that are grazed or ungrazed during spring. In this month, the ungrazed polder pastures are abandoned and the geese shift to either the grazed polder pastures or to the salt marsh. Vegetation in the polder that had been spring-grazed had a lower canopy height and a higher tiller density than summer-grazed vegetation. The captive geese in the preference experiment showed a clear preference for vegetation that had been spring-grazed by sheep over ungrazed vegetation. Goose grazing pressure was negatively correlated to canopy height, both on the polder and on the salt marsh. Within the plant communities dominated byFestuca rubra andPuccinellia maritima, marshes that were intensively grazed by livestock generally had higher grazing pressure by geese than long-term ungrazed or lightly grazed salt marshes.     

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.     

Mechanisms mediating plant distributions across estuarine landscapes in a low-latitude tidal estuary

Understanding of how plant communities are organized and will respond to global changes requires an understanding of how plant species respond to multiple environmental gradients. We examined the mechanisms mediating the distribution patterns of tidal marsh plants along an estuarine gradient in Georgia (USA) using a combination of field transplant experiments and monitoring. Our results could not be fully explained by the "competition-to-stress hypothesis" (the current paradigm explaining plant distributions across estuarine landscapes). This hypothesis states that the upstream limits of plant distributions are determined by competition, and the downstream limits by abiotic stress. We found that competition was generally strong in freshwater and brackish marshes, and that conditions in brackish and salt marshes were stressful to freshwater marsh plants, results consistent with the competition-to-stress hypothesis. Four other aspects of our results, however, were not explained by the competition-to-stress hypothesis. First, several halophytes found the freshwater habitat stressful and performed best (in the absence of competition) in brackish or salt marshes. Second, the upstream distribution of one species was determined by the combination of both abiotic and biotic (competition) factors. Third, marsh productivity (estimated by standing biomass) was a better predictor of relative biotic interaction intensity (RII) than was salinity or flooding, suggesting that productivity is a better indicator of plant stress than salinity or flooding gradients. Fourth, facilitation played a role in mediating the distribution patterns of some plants. Our results illustrate that even apparently simple abiotic gradients can encompass surprisingly complex processes mediating plant distributions.     

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.     

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。     

Hemiparasites generate environmental heterogeneity and enhance species coexistence in salt marshes.

Tidal inundation and salinity are considered to be controlling factors in salt marsh species distributions. Parasitic plants may also influence community organization as parasite-host interactions may play a functional role in stress amelioration due to physiological mechanisms for salinity tolerance and resource acquisition. Endangered root hemiparasites (Cordylanthus maritimus ssp. palustris and Cordylanthus mollis ssp. mollis) occupy unique habitat within fragmented northern California tidal wetlands. My objective was to examine the effects of these root hemiparasites on soil salinity, aeration, and community composition. I compared experimentally established bare patches, shaded and unshaded, and parasite removal patches to controls with hemiparasites across intertidal elevation gradients. Plant community composition, soil salinity, and redox potential were measured as response variables. In this field removal experiment, I demonstrated that parasite-host associations can enhance the amelioration of physical stress conditions in the salt marsh exceeding the passive role of shading by vegetation. Consumer-driven reduction of physical stress resulted in increased plant species richness, and the effect was most pronounced with elevated salinity and hypoxia stress. Although previous studies have demonstrated that removal of dominant plant biomass by herbivores can increase physical stress in salt marshes, this is one of the first examples of a positive indirect effect of a consumer on community diversity through physical stress relief. Greater understanding of biological interactions coupled with abiotic factors may improve rare plant conservation and salt marsh restoration success.     

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.     

Impact of cattle grazing on the occupancy of a cryptic, threatened rail

Impacts of livestock grazing in arid and semiarid environments are often concentrated in and around wetlands where animals congregate for water, cooler temperatures, and green forage. We assessed the impacts of winter-spring (November-May) cattle grazing on marsh vegetation cover and occupancy of a highly secretive marsh bird that relies on dense vegetation cover, the California Black Rail (Laterallus jamaicensis coturniculus), in the northern Sierra Nevada foothills of California, U.S.A. Using detection-nondetection data collected during repeated call playback surveys at grazed vs. ungrazed marshes and a "random changes in occupancy" parameterization of a multi-season occupancy model, we examined relationships between occupancy and habitat covariates, while accounting for imperfect detection. Marsh vegetation cover was significantly lower at grazed marshes than at ungrazed marshes during the grazing season in 2007 but not in 2008. Winter-spring grazing had little effect on Black Rail occupancy at irrigated marshes. However, at nonirrigated marshes fed by natural springs and streams, grazed sites had lower occupancy than ungrazed sites. Black Rail occupancy was positively associated with marsh area, irrigation as a water source, and summer vegetation cover, and negatively associated with marsh isolation. Residual dry matter (RDM), a commonly used metric of grazing intensity, was significantly associated with summer marsh vegetation cover at grazed sites but not spring cover. Direct monitoring of marsh vegetation cover, particularly at natural spring- or stream-fed marshes, is recommended to prevent negative impacts to rails from overgrazing.     

Twenty years of salt marsh succession on a Dutch coastal barrier island

After a formerly grazed salt marsh was released from cattle grazing, changes in plant species composition were monitored for 20 yr, using vegetation maps and permanent plots. Three areas, differing in age and nutrient status were compared. The number of plant species and plant communities decreased.Elymus athericus (Elytrigia pungens) became dominant in most plant communities after 5–20 yr on the oldest and most productive salt marsh. In younger areas it took more time forE. athericus to become dominant. At least 7 cm of clay seemed to be a prerequisite for this plant species to increase in dominance. The results from monitoring over decades are discussed in view of the knowledge on succession over centuries as derived from a chronosequence.     

25 years of salt marsh erosion in Essex: Implications for coastal defence and nature conservation

This paper presents the results from a study which was undertaken to monitor, map and quantify salt marsh change along 440 km of shoreline within the county of Essex, south-east England, between 1973 and 1998. Results indicate that during this 25-yr period, 1000 ha of salt marsh has been lost in Essex, primarily due to coastal erosion. This figure represents ca. 25% of the total salt-marsh area originally present in Essex in 1973. The salt marshes of Essex are important nature conservation areas, with many sites designated as Special Protection Areas under the EC Birds Directive (79/409/EEC) and as Special Areas of Conservation under the EC Habitats Directive (92/43/EEC). Salt marshes are also natural features which significantly dissipate wave and tidal energy, thereby playing an important role in contributing to effective coastal defence. The large-scale loss of salt marsh in Essex has, therefore, implications for both nature conservation and flood defence. Potential hypotheses for, and implications of such losses are discussed in this paper, together with the identification of potential management approaches to alleviate the losses.     

Dendrochronology of Atriplex portulacoides and Artemisia maritima in Wadden Sea salt marshes

The study uses a rather unusual method, dendrochronology, to investigate the growth and survival of Atriplex portulacoides L. and Artemisia maritima L. on salt marshes at two field sites on the Dutch North Sea barrier islands of Terschelling and Ameland. By providing information on longevity of these typical salt-marsh shrubs, dendrochronology offers an indirect way to investigate the influence of management regime – grazing in this case – on marsh quality and areal extent. Diminishment of salt marshes is a continuing concern in the northern Netherlands. The two shrub species studied here, A. portulacoides and A. maritima, are common to salt marshes. With their extensive roots and branches, they facilitate sedimentation and stabilize salt marshes. Using dendrochronology, this study found that annual growth rings could be identified to determine shrub age and growth. In A. portulacoides these rings took the form of a narrow band of terminal parenchyma. In A. maritima they were made up of unlignified marginal parenchyma together with higher vessel density at the beginning of the growing season. Growth rings indicated that intense grazing was clearly detrimental to the survival of A. portulacoides at the Terschelling site. However, grazing facilitated survival of A. maritima at the Ameland site by reducing light and nutrient competition from grasses. No growth trends could be found, however, as the lifespan for both species is short and many other influences on shrub growth could be identified.     

Role of Crab Herbivory in Die-Off of New England Salt Marshes

Abstract:  Die-offs of cordgrass are pervasive throughout western Atlantic salt marshes, yet understanding of the mechanisms precipitating these events is limited. We tested whether herbivory by the native crab , Sesarma reticulatum , is generating die-offs of cordgrass that are currently occurring on Cape Cod, Massachusetts (U.S.A.), by manipulating crab access to cordgrass transplanted into die-off areas and healthy vegetation. We surveyed 12 Cape Cod marshes to investigate whether the extent of cordgrass die-off on creek banks, where die-offs are concentrated, was related to local Sesarma grazing intensity and crab density. We then used archived aerial images to examine whether creek bank die-off areas have expanded over the past 2 decades and tested the hypothesis that release from predation, leading to elevated Sesarma densities, is triggering cordgrass die-offs by tethering crabs where die-offs are pervasive and where die-offs have not yet been reported. Intensity of crab grazing on transplanted cordgrass was an order of magnitude higher in die-off areas than in adjacent vegetation. Surveys revealed that Sesarma herbivory has denuded nearly half the creek banks in Cape Cod marshes, and differences in crab-grazing intensity among marshes explained >80% of variation in the extent of the die-offs. Moreover, the rate of die-off expansion and area of marsh affected have more than doubled since 2000. Crab-tethering experiments suggest that release from predation has triggered elevated crab densities that are driving these die-offs, indicating that disruption of predator–prey interactions may be generating the collapse of marsh ecosystems previously thought to be exclusively under bottom-up control .     

A neighboring plant species creates associational refuge for consumer and host

Examples of plant-animal and plant-plant associational defenses are common across a variety of systems, yet the potential for them to occur in concert has not been explored. In salt marshes in the Gulf of Mexico, the marsh periwinkle (Littoraria irrorata) is an abundant and conspicuous member of the community, climbing up the stems of marsh plants to remain out of the water at high tide. Though Littoraria are thought to primarily utilize stems of marsh cordgrass Spartina alterniflora as a source of food and refuge, Littoraria were more abundant in mixed assemblages of Spartina and Juncus roemerianus than in Spartina-only areas at the same tidal height. Mesocosm experiments confirmed that Juncus provided a refuge for Littoraria, with predation by Callinectes sapidus (but not Melongena corona) reduced when Juncus was present. However, Littoraria's utilization of Juncus as well as the effectiveness of Juncus as a refuge depended strongly on plant height: when Juncus was experimentally clipped to a shorter height than Spartina, snail abundance on Spartina and snail predation by crabs increased. Interestingly, this plant animal refuge led to a corresponding refuge for Spartina from Littoraria: Spartina plants lost less biomass to snail grazing when growing with Juncus in mesocosm and field experiments, and Spartina plants in natural assemblages were taller when growing with Juncus than when growing alone, even in the presence of abundant snails. This example highlights the potential importance of plant plant and plant-animal associational refuges in competitive plant assemblages.     

‘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).     

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.     

Increased primary production shifts the structure and composition of a terrestrial arthropod community

Numerous studies have examined relationships between primary production and biodiversity at higher trophic levels. However, altered production in plant communities is often tightly linked with concomitant shifts in diversity and composition, and most studies have not disentangled the direct effects of production on consumers. Furthermore, when studies do examine the effects of plant production on animals in terrestrial systems, they are primarily confined to a subset of taxonomic or functional groups instead of investigating the responses of the entire community. Using natural monocultures of the salt marsh cordgrass Spartina alterniflora, we were able to examine the impacts of increased plant production, independent of changes in plant composition and/or diversity, on the trophic structure, composition, and diversity of the entire arthropod community. If arthropod species richness increased with greater plant production, we predicted that it would be driven by: (1) an increase in the number of rare species, and/or (2) an increase in arthropod abundance. Our results largely supported our predictions: species richness of herbivores, detritivores, predators, and parasitoids increased monotonically with increasing levels of plant production, and the diversity of rare species also increased with plant production. However, rare species that accounted for this difference were predators, parasitoids, and detritivores, not herbivores. Herbivore species richness could be simply explained by the relationship between abundance and diversity. Using nonmetric multidimensional scaling (NMDS) and analysis of similarity (ANOSIM), we also found significant changes in arthropod species composition with increasing levels of production. Our findings have important implications in the intertidal salt marsh, where human activities have increased nitrogen runoff into the marsh, and demonstrate that such nitrogen inputs cascade to affect community structure, diversity, and abundance in higher trophic levels.