Above- and belowground biomass and nutrient pools of Spartina alterniflora (smooth cordgrass) in a South American salt marsh

In order to examine the role of position in the tidal range on biomass production and nutrient pools in Spartina alterniflora in an Argentinian estuary, we estimated productivity, the concentration of C, N and P in tissues and pools (concentration×biomass) of these elements in low (LM) and high (HM) zones. Aboveground biomass of S. alterniflora was higher in HM than in LM. Aboveground primary productivity was 106 and 439 g dry wt m^?2 year^?1 in LM and HM, respectively. Belowground biomass was similar in LM and HM. Belowground primary productivity was 526 and 744 g dry wt m^?2 year^?1 for LM and HM, respectively. Nutrient pools were higher in HM than in LM. Biomass and productivity values were low, which makes nutrient pools low. The lower values of the parameters analysed in LM than in HM indicate that position in the tidal range is an important factor in this system, possibly due to the effect of flooding. Moreover, this pattern is opposite to the general one observed in the northern hemisphere, meaning that studying marshes from different environments is worth doing. Because pools were higher in HM, this zone would be more important for nutrient input to the estuary.     

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。     

Effects of nitrogen addition on the growth of the salt marsh grassElymus athericus

Effects of nitrogen addition on the growth of the salt marsh grassElymus athericus were studied under greenhouse conditions. The addition of inorganic nitrogen (in the form of nitrate or ammonium and ranging from 0–24 g N/m²) stimulated the growth ofElymus athericus at the highest addition. Addition of nitrogen led to an increase of the soil nitrate concentrations both in the nitrate and ammonium treated soil in the first period of the experiment, whereas no differences were present at the end of the experiment. Ammonium in the ammonium treatments was transformed to nitrate within 15 days. In another experiment the values of the stable isotope nitrogen-15—expressed as δ¹⁵N-in nitrogen compounds used as fertilizer, in salt marsh soil and ofElymus athericus were measured. The δ¹⁵N of the N-compounds added (between ?3.2 and +2.6‰) were lower than the soil (ca.+10‰) and plants (ca.+8‰). During growth in water culture the δ¹⁵N of the leaves, stems and roots ofElymus athericus decreased from +9‰ to ?1‰. The latter value was close to the °¹⁵N of the N-compounds used in the water solution. Addition of N-compounds in soil culture, however, did not lead to such a decrease of the °¹⁵N ofElymus athericus. The difference in δ¹⁵N between soil nitrogen and the N-compounds added may be too small to be used successfully in ecological studies of nitrogen fluxes in the salt marsh environment.     

Consequences of omnivory for trophic interactions on a salt marsh shrub

Although omnivory is common in nature, its impact on trophic interactions is variable. Predicting the food web consequences of omnivory is complicated because omnivores can simultaneously produce conflicting direct and indirect effects on the same species or trophic level. We conducted field and laboratory experiments testing the top-down impacts of an omnivorous salt marsh crab, Armases cinereum, on the shrub Iva frutescens and its herbivorous and predatory arthropod fauna. Armases is a "true omnivore," consuming both Iva and arthropods living on Iva. We hypothesized that Armases would benefit Iva through a top-down trophic cascade, and that this benefit would be stronger than the direct negative effect of Armases on Iva. A field experiment on Sapelo Island, Georgia (USA), supported this hypothesis. Although Armases suppressed predators (spiders), it also suppressed herbivores (aphids), and benefited Iva, increasing leaf number, and reducing the proportion of dead shoots. A one-month laboratory experiment, focusing on the most common species in the food web, also supported this hypothesis. Armases strongly suppressed aphids and consumed fewer Iva leaves if aphids were available as an alternate diet. Armases gained more body mass if they could feed on aphids as well as on Iva. Although Armases had a negative effect on Iva when aphids were not present, Armases benefited Iva if aphids were present, because Armases controlled aphid populations, releasing Iva from herbivory. Although Armases is an omnivore, it produced strong top-down forces and a trophic cascade because it fed preferentially on herbivores rather than plants when both were available. At the same time, the ability of Armases to subsist on a plant diet allows it to persist in the food web when animal food is not available. Because omnivores feed on multiple trophic levels, their effects on food webs may differ from those predicted by standard trophic models that assume that each species feeds only on a single trophic level. To better understand the complexity of real food webs, the variable feeding habits and feeding preferences of different omnivorous species must be taken into consideration.     

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.     

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.     

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。     

Tidal and diurnal influence on food consumption of a salt marsh killifish Fundulus heteroclitus

Feeding patterns during four 24-h periods, sampled at 3-h intervals, were investigated for the mummichog Fundulus heteroclitus, in a Delaware, USA tidal marsh. Two factors potentially influencing feeding patterns, time of day and tide height, were examined. On 2 of the sampling periods a low tide occurred in the morning, while on the other 2 sampling periods a high tide occurred in the morning. Results are reported as g-dry wt. of food per g-dry wt. of fish. F. heteroclitus is primarily a daytime feeder that most actively feeds at high tide, regardless of whether or not the high tide inundates marsh surface areas. When tide height was sufficient to inundate the marsh surface, fish invaded these areas and consumed prey characteristic of the marsh surface. F. heteroclitus is an important link in energy transfers between the marsh surface and subtidal systems, enhancing its own energy supplies by consuming marsh surface prey whenever available.     

Sellafield waste radionuclides in Irish sea intertidal and salt marsh sediments

Low level liquid radioactive waste discharges from the Sellafield nuclear fuel reprocessing plant in north west England had generated environmental inventories of about 3 × 10¹⁶ Bq of¹³⁷Cs, 6.8 × 10¹⁴ Bq of^239,240Pu and 8.9 × 10¹⁴ Bq of²⁴¹Am by 1990. Most of the^239,240Pu and²⁴¹Am and about 10% of the¹³⁷Cs has been retained in a deposit of fine marine sediment close to the discharge point. The quantities of radionuclides discharged annually from Sellafield decreased by two orders of magnitude from the mid-1970s to 1990 but estimated critical group internal and external exposure decreased by less than one order of magnitude over this period. This indicates that during the period of reduced discharges, radionuclides already in the environment from previous releases continued to contribute to the critical group exposure and highlights the need to understand processes controlling the environmental distribution of the radionuclides. Redistribution of the contaminated marine sediment is potentially of major significance in this context, in particular if it results in transport of radionuclides to intertidal areas, where contact with the human population is relatively likely. A review is presented of published work relating to Sellafield waste radionuclides in Irish Sea sediments. Data on temporal and spatial trends in radionuclide concentrations and activity ratios are collated from a number of sources to show that the dominant mechanism of radionuclide supply to intertidal areas is by redistribution of the contaminated marine sediment. The implications of this mechanism of supply for trends in critical group radiation exposure are considered.     

Nitrogen exchange between a southeastern USA salt marsh ecosystem and the coastal ocean

The salt marsh ecosystem at North Inlet, South Carolina, USA consistently exported dissolved inorganic nitrogen via tidal exchange with the coastal Atlantic Ocean. Concentrations centrations of NH ₄ ⁺ and NO ₃ ^- +NO ₂ ^- displayed distinct tidal patterns with rising values during ebb flow. These patterns suggest the importance of biogeochemical processes in the flux of material from the salt marsh. NH ₄ ⁺ export peaked during the summer (15 to 20 mg m^-2 tide^-1) during a net balance of tidal water exchange. Remineralization of NH ₄ ⁺ within the salt marsh system appears to be contributing to the estimated annual net export of bout 4.7 g NH ₄ ⁺ -N m^-2 yr^-1. NO ₃ ^- +NO ₂ ^- exports were higher in the fall and winter of 1979 (2 to 4 mg N m^-2 tide^-1). The winter export coincided with a considerable net export of water with no distinctive concentration patterns, suggesting a simple advective export. However, the fall peak of NO ₃ ^- +NO ₂ ^- export occurred during a period of net water balance in tidal exchange and an insignificant freshwater input from the western, forested boundary. During the summer and fall, tidal concentration patterns were particularly apparent, suggesting that nitrification within the salt marsh system was contributing to the estimated annual net export of ca 0.6 g NO ₃ ^- +NO ₂ ^- -N m^-2 yr^-1.Contribution No. 637 from the Belle W. Baruch Institute of Marine Biology and Coastal Research     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

Primary production and heterotrophy of a pennate and a centric salt marsh diatom

Radionuclide labeled sodium bicarbonate and glucose were used to measure the rates of primary production and glucose uptake by Nitzschia obtusa var. undulata and Cymbella pusilla. The presence of glucose in the medium enhanced primary production in both diatom species. Uptake of glucose in C. pusilla was photoinhibited. A maximum of 9.3 nmol C mg dry wt^-1 h^-1 of glucose was taken up in the light without the presence of NaHCO₃ in the medium and with NaHCO₃ present the value was 18 nmol C mg dry wt^-1 h^-1. The ratio of inorganic carbon fixed to the total carbon budget (autotrophy and heterotrophy) was 0.51 for N. obtusa var. undulata and 0.62 for C. pusilla.     

Biotic and abiotic stress effects on nitrogen chemistry in the salt marsh cordgrassSpartina alterniflora (Poaceae)

Summary Planthopper (Insecta: Homoptera) feeding stress induces a senesence-like response in the leaves ofSpartina alterniflora characterized by decreased soluble protein, an increased total amino acid pool, and elevated levels of 10 individual amino acids. Increased proline and tryptophan in response to planthopper feeding could not be fully explained by protein degradation. Low degrees of soil salinity stress resulted in an increased total free amino acid pool and elevated levels of 7 amino acids. Anaerobic soil stress resulted in decreased glutamic acid and increased asparagine. Low salinity and anaerobic stress had no effect on soluble protein levels. Glycinebetaine was not affected by the stresses examined in this study.     

Latitudinal variation in top-down and bottom-up control of a salt marsh food web

The shrub Iva frutescens, which occupies the terrestrial border of U.S. Atlantic Coast salt marshes, supports a food web that varies strongly across latitude. We tested whether latitudinal variation in plant quality (higher at high latitudes), consumption by omnivores (a crab, present only at low latitudes), consumption by mesopredators (ladybugs, present at all latitudes), or the life history stage of an herbivorous beetle could explain continental-scale field patterns of herbivore density. In a mesocosm experiment, crabs exerted strong top-down control on herbivorous beetles, ladybugs exerted strong top-down control on aphids, and both predators benefited plants through trophic cascades. Latitude of plant origin had no effect on consumers. Herbivorous beetle density was greater if mesocosms were stocked with beetle adults rather than larvae, and aphid densities were reduced in the "adult beetle" treatment. Treatment combinations representing high and low latitudes produced patterns of herbivore density similar to those in the field. We conclude that latitudinal variation in plant quality is less important than latitudinal variation in top consumers and competition in mediating food web structure. Climate may also play a strong role in structuring high-latitude salt marshes by limiting the number of herbivore generations per growing season and causing high overwintering mortality.     

Macroinfauna of a Southern New England salt marsh: seasonal dynamics and production

The animal-habitat relationships and seasonal dynamics of the benthic macroinfauna were investigated from November 1986 to October 1988 in the Great Sippe-wissett salt marsh (Massachusetts, USA). Total macrofaunal abundance varied seasonally, displaying a peak in late spring and early summer, then declining sharply during late summer and recovering briefly in fall before collapsing in winter. Three macroinfaunal assemblages were found in the marsh, distributed along gradients of environmental factors. These included a sandy non-organic sediment assemblage, a sandy organic sediment assemblage and a muddy sediment assemblage. The species groups characteristic of unstable sandy non-organic sediments included the polychaetes Leitoscoloplos fragilis, Aricidea jefreyssi, Magelona rosea and Streptosyllis verrilli, the oligochaete Paranais litoralis, and the crustacean Acanthohaustorius millsi. Sandy organic sediments were characterized by the polychaetes Marenzelleria viridis, Capitella capitata, Neanthes succinea, N. arenaceodonta, Polydora ligni and Heteromastus filiformis, the oligochaete Lumbricillus sp., and the mollusc Gemma gemma. In muddy sites, the polychaete Streblospio benedicti and the oligochaetes Paranais litoralis and Monopylephorus evertus were the dominant species. Secondary production of benthic macroinfauna in each of these habitats was estimated. The highest values of biomass and production were recorded in the sandy organic sediments. Secondary production was estimated to be 1850 kJ m^-2 yr^-1 in sandy organic areas, but only 281 kJ m^-2 yr^-1 in sandy non-organic areas and 113 kJ m^-2 yr^-1 in muddy areas. This results in an area-weighted average production of 505 kJ m^-2 yr^-1 for the unvegetated areas of the marsh. The Great Sippewissett salt marsh has an area of 483800 m², the total secondary production of the macroinfauna for the whole unvegetated area of the marsh was estimated as 4651 kg dry wt yr^-1, expressed as somatic growth. This production value seems consistent with production data obtained for other intertidal North Atlantic environments.     

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.     

Root respiration and oxygen flux in salt marsh grasses from different elevational zones

Plants growing in waterlogged environments are subjected to low oxygen levels around submerged tissues. While internal oxygen transport has been postulated as an important factor governing flooding tolerance, respiration rates and abilities to take up oxygen under hypoxic conditions have been largely ignored in plant studies. In this study, physiological characteristics related to internal oxygen transport, respiration, and oxygen affinity were studied in low intertidal marsh species (Spartina alterniflora and S. anglica) and middle to high intertidal species (S. densiflora, S. patens, S. foliosa, a S. alterniflora × S. foliosa hybrid, S. spartinae, and Distichlis spicata). These marsh plants were compared to the inland species S. pectinata and the crop species rice (Oryza sativa), corn (Zea mays), and oat (Avena sativa). Plants were grown in a greenhouse under simulated estuarine conditions. The low marsh species S. anglica was found to transport oxygen internally at rates up to 2.2 μmol O₂ g fresh root weight⁻¹ h⁻¹. In contrast, marsh species from higher zones and crop species were found to transport significantly less oxygen internally, although rice plants were able to transport 1.4 μmol g⁻¹ h⁻¹. Under hypoxic conditions, low marsh species were better able to remove dissolved oxygen from the medium compared to higher marsh species and crops. The oxygen concentration at which respiration rates declined due to limited oxygen (P _crit) was significantly lower in low marsh species compared to inland and crop species; P _crit ranged from <4 μM O₂ in the low marsh species S. anglica up to 20 μM in the inland species corn. Flooding-sensitive crop species had significantly higher aerobic respiration rates compared to flooding-tolerant species in this study. Crop species took up 3.6–6.7 μmol O₂ g⁻¹ h⁻¹ while all but one marsh species took up <3.5 μmol O₂ g⁻¹ h⁻¹. We conclude that oxygen transport, aerobic demand, and oxygen affinity all play important and interrelated roles in flood tolerance and salt marsh zonation.     

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.     

Diel flux of dissolved carbohydrate in a salt marsh and a simulated estuarine ecosystem

The concentrations of total dissolved carbohydrate (TCHO), monosaccharide (MCHO) and polysaccharide (PCHO) were followed over a total of ten diel cycles in a salt marsh and a 13 m³ seawater tank simulating an estuarine ecosystem. Their patterns are compared to those for total dissolved organic carbon (DOC), CO₂, pH, O₂, chlorophyll a, phaeopigments and solar radiation. During 5 of the 6 marsh studies, PCHO underwent periods of sustained accumulation starting in the late morning or early afternoon and continuing into the early evening. These periods possibly represent release of recently synthesized PCHO from phototrophs. similar patterns were not found in the tank although direct associations between TCHO and phaeopigment dynamics suggest that zooplankton excretion was an important source of dissolved carbohydrate. The numbers of planktonic bacteria determined in one tank study increased rapidly during a late morning PCHO pulse and varied inversely with PCHO throughout the afternoon and evening, indicating that they were able to respond rapidly and control natural substrate concentrations on a time scale of a few hours. MCHO fluctuated to a much lesser extent than PCHO at both locations with levels possibly maintained near the bacterial uptake threshold or in a closely regulated steady state. TCHO concentration changes over 2-to 3-h sampling intervals suggest very rapid net system release and uptake with summer rates frequently exceeding 30 g C l^-1h^-1 in the marsh and 20 g C l^-1 h^-1 in the tank.