How will warming affect the salt marsh foundation species Spartina patens and its ecological role?

Foundation species structure environments and create refuge from environmental stress. In New England high salt marsh, the grass Spartina patens is a foundation species that reduces salinity, anoxia, desiccation, and thermal stresses through canopy shading and root proliferation. In a factorial S. patens-removal and warming field experiment, foundation species removal strongly impacted every aspect of the community, reiterating the important role of the foundation species S. patens in the high marsh. Given this central role, we hypothesized that facilitation by the foundation species would be even more important under warmer conditions by ameliorating more severe thermal stress. However, the ecological role of S. patens was unaffected by experimental warming, and, independent of the presence of the foundation species, warming had only weak effects on the salt marsh ecological community. Only the foundation species itself responded strongly to warming, by significantly increasing aboveground production in warmed plots. Apparently, amelioration of thermal stress is not as important for salt marsh ecosystem function as S. patens’ moderation of salinity and desiccation stresses. From these experimental results, we anticipate that climate change-associated thermal stress will not greatly affect S. patens-dominated high marsh communities. In contrast, foundation species loss, an emergent conservation issue in Atlantic salt marshes, represents a critical threat to salt marsh ecosystem function.     

Effects of non-native <Emphasis Type="Italic">Spartina patens</Emphasis> on plant and sediment organic matter carbon incorporation into the local invertebrate community

The cycling of organic matter through food webs is a fundamental process that may be altered by the invasion of non-indigenous plants. We explored consequences of the invasion of non-indigenous Spartina patens to the composition of soil organic matter (SOM) and to detritivore and herbivore diets in the upper salt marsh within Corrubedo National Park, northwest Spain. We tested for the incorporation of S. patens carbon (C) into SOM and by detritivores and herbivores using stable isotope analysis, focusing primarily on detritivorous enchytraeid oligochaetes and herbivorous insects. Stable isotope results indicated that C derived from S. patens has been incorporated into SOM. Elevated densities of enchytraeids in stands of S. patens, and their incorporation of C derived from this plant, suggested that dense patches of S. patens may facilitate detritivore populations. In contrast, although insect herbivores used S. patens as habitat, there was little isotopic evidence for the widespread incorporation of S. patens-derived C by these consumers. The population and dietary response of enchytraeids to S. patens suggests that S. patens invasion could indirectly influence soil processes and pathways mediated by detritivore activity (e.g., soil respiration rates, nutrient retention and transformation, energy flow). The loss of food resources to insect herbivores alters local food webs. However, insect herbivores may move and feed on native plants elsewhere. As a result, insect populations may be less immediately impacted than soil detritivore populations by S. patens. Our study suggests that the influences of S. patens invasion extend beyond the more obvious changes in native plant abundance, to include differing responses in the cycling of organic matter between detritivore and insect herbivore food web pathways.     

A comparative study of above-ground productivity of dominant U.S. Gulf Coast marsh species

Above-ground productivity of dominant freshwater, brackish, and salt-marsh species from the U.S. Gulf Coast was evaluated using both gas exchange techniques and harvest methods. Both techniques showed significant differences in productivity among the study species which represent major components of their respective communities. Estimates of net aerial primary productivity using the harvest method yielded 3683 g dw (dry weight) m^?2yr^?1 for Spartina alterniflora (tall), 2008 g dw m^?2yr^?1 for S. alterniflora (short), 3677 g dw m^?yr^?1 for S. patens and 1641 g dwm^?yr^?1 for Panicum hemitomon. Carbon balance estimated from gas exchange calculation yielded values approximately equivalent to a biomass accumulation of 6024 g dw m^?2yr^?1 for S. alterniflora (tall), 3047 g dw m^?yr^?1 for S. alterniflora (short), 5702 g dw m^?yr^?1 for S. patens, and 2912 g dm^?yr^?1 for P. hemitomon. The net aerial primary production was estimated to be approximately 61% of total productivity in S. alterniflora (tall-form) and 66%o of total productivity in short-form, 64% in S. patens and 56%) in P. hemitomon. The assimilation data also indicated that Spartina alterniflora and S. patens continue carbon fixation throughout the year while assimilation in Panicum hemitomon is absent due to lack of live leaves during the winter. Various aspects of harvest and gas exchange techniques are discussed.     

Dissolved Inorganic Nitrogen Pools and Surface Flux under Different Brackish Marsh Vegetation Types,Common Reed (<Emphasis Type="Italic">Phragmites australis</Emphasis>) and Salt Hay (<Emphasis Type="Italic">Spartina patens</Emphasis>)

The current expansion of Phragmites australis into the high marsh shortgrass (Spartina patens, Distichlis spicata) communities of eastern U.S. salt marshes provided an opportunity to identify the influence of vegetation types on pools and fluxes of dissolved inorganic nitrogen (DIN). Two brackish tidal marshes of the National Estuarine Research Reserve system were examined, Piermont Marsh of the Hudson River NERR in New York and Hog Island in the Jacques Coustaeu NERR of New Jersey. Pools of DIN in porewater and rates of DIN surface flux were compared in replicated pairs of recently-expanded P. australis and neighboring S. patens-dominated patches on the high marsh surface. Both marshes generally imported nitrate (NO₃⁻) and exported ammonium (NH₄⁺), such that overall DIN was exported. No differences in surface exchange of NO₃⁻ or NH₄⁺ were observed between vegetation types. Depth-averaged porewater NH₄⁺ concentrations over the entire growing season were 56% lower under P. australis than under S. patens (average 1.4 vs. 3.2 mg NH₄⁺ L⁻¹) with the most profound differences in November. Porewater profiles showed an accumulation of NH₄⁺ at depth in S. patens and constant low concentrations in P. australis from the soil surface to 50 cm depth, with no significant differences in porewater salinity. Despite these profound differences in porewater, NH₄⁺ diffusion from soils of P. australis and S. patens were not measurably different, were similar to other published rates, and were well below estimated rates based on passive diffusion alone. Rapid adsorption and uptake by litter and microbes in surface soils of both communities may buffer NH₄⁺ loss to flooding tides in both communities, thereby reducing the impact of P. australis invasion on NH₄⁺ flux to flooding waters.     

Salinity adaptation of plasma membrane H+-ATPase in the salt marsh plant Spartina patens: ATP hydrolysis and enzyme kinetics

Spartina patens, an intertidal C4 grass, grows in the upper salt marsh and tolerates coastal seawater salinity. The regulation of ion movement across the plasma membrane (PM) for plant salt tolerance is thought to be achieved by an electrochemical gradient generated by plasma membrane H⁺-ATPase. In this study, the change of PM H⁺-ATPase in response to NaCl was characterized for S. patens callus. Callus was cultured for 10 weeks under salinity levels of 0 mM, 170 mM, 340 mM, and 510 mM NaCl. Plasma membrane was isolated from a Dextran/PEG aqueous polymer two-phase system and the purity was demonstrated with membrane enzyme markers. There was a significant increase (up to 2-3 fold) of PM H⁺-ATPase activity when callus was grown on media containing NaCl. The incremental activation of PM H⁺-ATPase activity would enable the cell to tolerate higher cytoplasmic NaCl concentrations. PM H⁺-ATPase appeared to have a higher Vmax and a lower substrate concentration (Km to reach Vmax. When growth medium salinity increased from 0 mM to 170 and 340 mM, the Vmax of H⁺-ATPase increased from 0.64 to 1.00 and 1.73, respectively, while the Km decreased from 3.58 to 2.07 and 2.44 mM, respectively. In vitro NaCl inhibition kinetic data revealed a pattern of non-competitive inhibition by NaCl on PM H⁺-ATPase. The response of PM H⁺-ATPase in S. patens callus suggests that this species has evolved mechanisms that can regulate this important enzyme when cells are exposed to NaCl.     

Microscale Spatial Distribution of Phragmites australis (Common Reed) Invasion into Spartina patens (Salt Hay)-dominated Communities in Brackish Tidal Marsh

Over the last century, Phragmites australis (common reed) has been expanding rapidly from the marsh–upland boundary into Spartina patens (salt hay)-dominated high marsh communities of the eastern US coast. Whereas direct and indirect human disturbances and changes in hydrology or salinity are likely to influence rates of spread at the landscape scale, the susceptibility of specific plant communities to invasion also influence rates of Phragmites expansion at the local scale. I measured microscale (0.25 m²) spatial patterns of culms (emerging buds and mature stems) in October 1993 at both expanding and stable boundaries of Phragmites populations within a S. patens-dominant matrix. In both expanding and stable plots, Phragmites culms were observed more frequently than expected on hummocks that were created by S. patens tussock-forming root structure. Culm density within a plot was correlated with the percent hummock cover within a plot. Further, Phragmites culms, particularly mature stems, were concentrated along the perimeter of the hummocks. Because the culms were not evenly distributed between hummocks and hollows, I suggest that invasion rates of Phragmites are limited in S. patens communities by microscale differences in hummock availability. The pattern of emergence suggests that expanding rhizomes of Phragmites encounter both competition with S. patens roots on the hummocks and physiological stressors (salinity, anoxia, sulfide concentrations) in the hollows.     

Effects of salinity on chlorophyll fluorescence and CO<Subscript>2</Subscript> fixation in C<Subscript>4</Subscript> estuarine grasses

The effects of salinity (sea water at 0 ‰ versus 30 ‰) on gross rates of O₂ evolution (J _O2) and net rates of CO₂ uptake (P _N) were measured in the halotolerant estuarine C₄ grasses Spartina patens, S. alterniflora, S. densiflora, and Distichlis spicata in controlled growth environments. Under high irradiance, salinity had no significant effect on the intercellular to ambient CO₂ concentration ratio (C _i/C _a). However, during photosynthesis under limiting irradiance, the maximum quantum efficiency of CO₂ fixation decreased under salinity across species, suggesting there is increased leakage of the CO₂ delivered to the bundle sheath cells by the C₄ pump. Growth under salinity did not affect the maximum intrinsic efficiency of photosystem 2, PS2 (F_V/F_M) in these species, suggesting salinity had no effect on photosynthesis by inactivation of PS2 reaction centers. Under saline conditions and high irradiance, P _N was reduced by 75 % in Spartina patens and S. alterniflora, whereas salinity had no effect on P _N in S. densiflora or D. spicata. This inhibition of P _N in S. patens and S. alterniflora was not due to an effect on stomatal conductance since the ratio of C _i/C _a did not decrease under saline conditions. In growth with and without salt, P _N was saturated at ∼500 μmol(quantum) m⁻² s⁻¹ while J _O2 continued to increase up to full sunlight, indicating that carbon assimilation was not tightly coupled to photochemistry in these halophytic species. This increase in alternative electron flow under high irradiance might be an inherent function in these halophytes for dissipating excess energy.     

Edaphic diatom communities associated with Spartina alterniflora and S. patens in New Jersey

Edaphic diatoms inhabiting the sediments beneath dwarf Spartina alterniflora Loisel. and S. patens (Ait.) Muhl. in Great Bay salt marsh, Tuckerton, New Jersey were collected from 24 September 1974 through 20 August 1975. Of the 91 taxa encountered, 8 were endemic to the dwarf S. alterniflora habitat and 42 endemic to the S. patens habitat. The edaphic diatom community associated with S. patens was comprised of a much greater number of taxa and possessed higher values for species diversity (H') and evenness (J') than the community associated with dwarf S. alterniflora. The salinity of the marsh surface showed a completely opposite trend, being greatly reduced at the S. patens habitat. A highly significant relationship (P < 0.001) between the number of diatom taxa and marsh surface salinity at the S. patens habitat was demonstrated by a least squares regression. This finding led to the conclusion that the dissimilarity in the structure of the two edaphic diatom communities was primarily due to the very low marsh surface salinities at the S. patens habitat from January through June, and that this sustained. low-salinity regime allowed a very large number of taxa to coexist only in the S. patens community. Comparison of the diatom flora of Great Bay salt marsh with that of a Delaware marsh studied previously by the author showed that 67.0% of the 91 taxa encountered in New Jersey also occur on the Delaware marsh.     

ANATOMICAL AND METABOLIC RESPONSES TO WATERLOGGING AND SALINITY IN SPARTINA ALTERNIFLORA AND S. PATENS (POACEAE)

The effects of waterlogging and salinity (25 or 325 mol m ³ NaCl) stressors on the anatomy and metabolism of the marsh grasses 5. alterniflora Loisel. and S. patens Aiton (Muhl.) were investigated in a V factorial greenhouse experiment over 30 d. Waterlogging and salinity in combination resulted in anatomical and metabolic responses in both species. Waterlogging reduced soil redox potential and decreased root-specific gravity significantly in both species. The inadequacy of aerenchyma development under hypoxia to support aerobic root respiration in S. patens was indicated by significant increases in root alcohol dehydrogenase (ADH) activity of 1,752% and 420%, respectively, in the low and high salinity treatments. ADH activity was not increased significantly by flooding of S. alterniflora. Proline concentrations in roots and leaves were low at low salinities and increased significantly at high salinities in both species, but only under drained conditions. Decrease in leaf elongation by high salinity occurred in drained, but not flooded treatments in both species. Under flooded conditions, leaf elongation was significantly greater in S. alterniflora than S. patens. Greatest leaf elongation occurred in flooded low salinity S. alterniflora plants that had the least proline. Although both species are adapted to waterlogging and salinity, S. alterniflora appears to be more tolerant of reducing soil conditions and less responsive to higher salinity than S. patens.     

Implication of nutrient and salinity interaction on the productivity of <Emphasis Type="Italic">Spartina patens</Emphasis>

Reintroduction of fresh water to coastal systems with altered hydrologic regimes is a management option for restoring degraded wetland habitats. Plant production in these systems is believed to be enhanced by increased nutrient availability and reduced salinity. Although studies have documented nutrient limitation and salinity stress in coastal marshes, interpreting the effects of freshwater reintroduction on plant production is difficult because high nutrient availability often is confounded with low salinity. We tested the hypothesis that plant growth response to nutrients does not vary with salinity in a greenhouse study. Treatments consisted of four nutrient concentrations and four non-lethal salinity levels; plant response was measured as biomass accumulation after 144 days of exposure. The significant interaction between salinity and nutrient concentrations indicates that response of Spartina patens marshes to freshwater inflows would vary by site-specific soil conditions. Biomass decreased with increased salinity at all four nutrient concentrations with variation among the nutrient concentrations decreasing as salinity increased. We demonstrate the importance of considering ambient salinity and nutrient soil conditions in restoration planning involving freshwater inflow. We propose salinity should remain a primary concern in restoration plans targeted at improving degraded S. patens-dominated marsh habitat.     

Evapotranspiration and water use efficiency in a Chesapeake Bay wetland under carbon dioxide enrichment

Wetlands evapotranspire more water than other ecosystems, including agricultural, forest and grassland ecosystems. However, the effects of elevated atmospheric carbon dioxide (CO₂) concentration (C_a) on wetland evapotranspiration (ET) are largely unknown. Here, we present data on 12 years of measurements of ET, net ecosystem CO₂ exchange (NEE), and ecosystem water use efficiency (EWUE, i.e. NEE/ET) at 13:00–15:00 hours in July and August for a Scirpus olneyi (C3 sedge) community and a Spartina patens (C4 grass) community exposed to ambient and elevated (ambient+340 μmol mol^?1) C_a in a Chesapeake Bay wetland. Although a decrease in stomatal conductance at elevated C_a in the S. olneyi community was counteracted by an increase in leaf area index (LAI) to some extend, ET was still reduced by 19% on average over 12 years. In the S. patens community, LAI was not affected by elevated C_a and the reduction of ET was 34%, larger than in the S. olneyi community. For both communities, the relative reduction in ET by elevated C_a was directly proportional to precipitation due to a larger reduction in stomatal conductance in the control plants as precipitation decreased. NEE was stimulated about 36% at elevated C_a in the S. olneyi community but was not significantly affected by elevated C_a in S. patens community. A negative correlation between salinity and precipitation observed in the field indicated that precipitation affected ET through altered salinity and interacted with growth C_a. This proposed mechanism was supported by a greenhouse study that showed a greater C_a effect on ET in controlled low salinity conditions compared with high salinity. In spite of the differences between the two communities in their responses to elevated C_a, EWUE was increased about 83% by elevated C_a in both the S. olneyi and S. patens communities. These findings suggest that rising C_a could have significant impacts on the hydrologic cycles of coastal wetlands.     

Tidal marsh plant responses to elevated CO2, nitrogen fertilization,and sea level rise

Elevated CO₂ and nitrogen (N) addition directly affect plant productivity and the mechanisms that allow tidal marshes to maintain a constant elevation relative to sea level, but it remains unknown how these global change drivers modify marsh plant response to sea level rise. Here we manipulated factorial combinations of CO₂ concentration (two levels), N availability (two levels) and relative sea level (six levels) using in situ mesocosms containing a tidal marsh community composed of a sedge, Schoenoplectus americanus, and a grass, Spartina patens. Our objective is to determine, if elevated CO₂ and N alter the growth and persistence of these plants in coastal ecosystems facing rising sea levels. After two growing seasons, we found that N addition enhanced plant growth particularly at sea levels where plants were most stressed by flooding (114% stimulation in the + 10 cm treatment), and N effects were generally larger in combination with elevated CO₂ (288% stimulation). N fertilization shifted the optimal productivity of S. patens to a higher sea level, but did not confer S. patens an enhanced ability to tolerate sea level rise. S. americanus responded strongly to N only in the higher sea level treatments that excluded S. patens. Interestingly, addition of N, which has been suggested to accelerate marsh loss, may afford some marsh plants, such as the widespread sedge, S. americanus, the enhanced ability to tolerate inundation. However, if chronic N pollution reduces the availability of propagules of S. americanus or other flood‐tolerant species on the landscape scale, this shift in species dominance could render tidal marshes more susceptible to marsh collapse.     

GROWTH OF THE FILAMENTOUS GREEN ALGA CTENOCLADUS CIRCINNATUS (CHAETOPHORALES,CHLOROPHYCEAE) IN RELATION TO ENVIRONMENTAL SALINITY1

Clones of the filamentous green alga Ctenocladus circinnatus Borzi were isolated from algae collected at Abert Lake (Oregon) and Mono Lake (California). Stock cultures were exposed to varied salinities of natural lake water to examine the effects on growth rate, cell form, chlorophyll a, and water content. Growth rates were reduced in both clones with increased salinity over the range 25–100 g·L^?1 and were almost completely inhibited at 150 g·L^?1. Chlorophyll a increased between salinities of 25 and 100 g·L^?1, reflecting slower growth, higher proportions of akinetes, and smaller cell sizes as salinity increased. Tissue water content remained essentially constant from 25 to 100 g·L^?1 salinity. Shorter cell dimensions with increased salinity suggest that a lower surface-to-volume ratio may reduce the potential for passive loss of cell water. Prior acclimation of stock cultures to elevated salinity provided no enhancement of growth response at any salinity. The results indicate that environmental salinity can limit the productivity and distribution of Ctenocladus in nature.     

Sexual reproduction is light-limited as marsh grasses colonize maritime forest

Climate change is driving abiotic shifts that can threaten the conservation of foundation species and the habitats they support. Directional range shifting is one mechanism of escape, but requires the successful colonization of habitats where interspecific interactions may differ from those to which a species has adapted. For plants with multiple reproductive strategies, these range-edge interactions may alter the investment or allocation toward a given reproductive strategy. In this study, we quantified sexual reproduction of the clonal marsh grass Spartina patens across an inland colonization front into maritime forest being driven by sea-level rise. We find that flowering is variable across S. patens meadows, but consistently reduced in low light conditions like those of the forest understory. Observational surveys of S. patens flowering at four sites in the Delmarva Peninsula agreed with the results of two experimental manipulations of light availability (shading experiment in S. patens-dominated marsh and a forest dieback manipulation). These three approaches pinpointed light limitation as a principal control on S. patens flowering capacity, suggesting that light competition with taller upland species can suppress S. patens flowering along its upland migration front. Consequently, all propagation in shaded conditions must occur clonally or via seeds from the marsh, a reproductive restriction that could limit the potential for local adaptation and reduce genetic diversity. Future research is needed to determine whether the lack of flowering is the result of a trade-off between sexual and clonal reproduction or results from insufficient photosynthetic products needed to achieve either reproductive method.     

Relationship of soil hydrogen sulfide level to net carbon assimilation ofPanicum hemitomon andSpartina patens

Panicum hemitomon Schult andSpartina patens (Ait) Muhl. plants from Louisiana Gulf Coast fresh and brackish marshes were subjected to hydrogen sulfide under controlled sediment redox conditions. Net carbon assimilation responses of both species to the combined sediment anaerobiosis and hydrogen sulfide concentrations was measured.Panicum hemitomon was more sensitive to hydrogen sulfide as compared toSpartina patens. Initiation of reduction in net carbon assimilation inP. hemitomon began when H₂S concentrations of soil solution exceeded 0.22 mgl^-1. Reductions in net carbon assimilation inS. patens were also noted at H₂S concentrations exceeding 0.34 mgl^-1. The reduction in net carbon assimilation of both species measured at elevated H₂S concentrations suggests that extreme anaerobiosis and elevated sulfide could contribute to the growth reduction of these species under certain conditions. However based on H₂S concentration in fresh and brackish marsh soil profiles, levels were too low to cause any adverse effects ofPanicum hemitomon. In brackish marsh soils containing hydrogen sulfide of 3.4 mgl^-1 in soil solution, sulfide could be a major factor limiting growth ofS. patens.     

Role of ENA ATPase in Na<Superscript>+</Superscript> efflux at high pH in bryophytes

Potassium or Na⁺ efflux ATPases, ENA ATPases, are present in all fungi and play a central role in Na⁺ efflux and Na⁺ tolerance. Flowering plants lack ENA ATPases but two ENA ATPases have been identified in the moss Physcomitrella patens, PpENA1 and PpENA2. PpENA1 mediates Na⁺ efflux in Saccharomyces cerevisiae. To propose a general function of ENA ATPases in bryophytes it was necessary to demonstrate that these ATPases mediate Na⁺ efflux in planta and that they exist in more bryophytes than P. patens. For these demonstrations (1) we cloned a third ATPase from P. patens, PpENA3, and studied the expression pattern of the three PpENA genes; (2) we constructed and studied the single and double Δppena1 and Δppena2 mutants; and (3) we cloned two ENA ATPases from the liverwort Marchantia polymorpha, MpENA1 and MpENA2, and expressed them in S. cerevisiae. The results from the first two approaches revealed that the expression of ENA ATPases was greatly enhanced at high pH and that Na⁺ efflux at high pH depended on PpENA1. The ENA1 ATPase of M. polymorpha suppressed the defective growth of a S. cerevisiae mutant at high K⁺ or Na⁺ concentrations, especially at high K⁺.     

Population differentiation inSpartina patens: Water potential components and bulk modulus of elasticity

Pressure-volume technique was utilized to evaluate salinity response among three populations ofSpartina patens (Ait.) Muhl. from Louisiana Gulf coast marshes. Plants were subjected to salinities of 85 and 425 mol m^?3 for 77 d in a greenhouse. Ψ_w and Ψ_π decreased in all populations in response to increases in salinity. There were 32% decrease in Ψ_sat, 42% decrease in Ψ_tlp in response to salinity changes from 85 to 425 mol m^?3 in the Ferblanc population. Similarly, there were 35% and 41% decrease in Ψ_sat in the Clovelly and Lake Tambour populations, respectively. All populations showed the ability to adapt to the increased salinity as was evidenced by osmotic adjustment. However, the Lake Tambour population appeared to have superior ability to adapt to high salinity through having a significantly lower osmotic potential at saturation (Ψ_sat), osmotic potential at turgor loss point (Ψ_tlp), and maximum turgor potential (Ψ_P(max)) compared to other populations. Ferblanc and Clovelly populations revealed the ability to adapt to saline environments to a lesser extent as compared to the Lake Tambour population. Results indicate that there is a potential for selection of superior strains ofSpartina patens for use in marsh restoration projects aiming at prevention of wetland loss in certain coastal areas.     

Nitrogen and carbon dynamics in C3 and C4 estuarine marsh plants grown under elevated CO2 in situ

Summary Carbon dioxide concentrations were elevated in three estuarine communities for an entire growing season. Open top chambers were used to raise CO₂ concentrations ca. 336 ppm above ambient in monospecific communities of Scirpus olneyi (C₃) and Spartina patens (C₄), and a mixed community of S. olneyi, S. patens and Distichlis spicata (C₄). Nitrogen and carbon concentration (% wt) of aboveground tissue was followed throughout growth and senescence. Green shoot %N was reduced and %C was unchanged under elevated CO₂ in S. olneyi. This resulted in a 20%–40% increase in tissue C/N ratio. There was no effect of CO₂ on either C₄ species. Maximum aboveground N (g/m²) was unchanged in S. olneyi, indicating that increased productivity under elevated CO₂ was dependent on reallocation of stored N. There was no change in the N recovery efficiency of S. olneyi in pure stand and a decrease in the mixed community. Litter C/N ratio was not affected by elevated CO₂ suggesting that decomposition and N mineralization rates will also remain unchanged. Continued growth responses to elevated CO₂ could, however, be limited by the ability of S. olneyi to increase the total aboveground N pool.     

Effects of regular salt marsh haying on marsh plants, algae, invertebrates and birds at Plum Island Sound, Massachusetts

The haying of salt marshes, a traditional activity since colonial times in New England, still occurs in about 400 ha of marsh in the Plum Island Sound estuary in northeastern Massachusetts. We took advantage of this haying activity to investigate how the periodic large-scale removal of aboveground biomass affects a number of marsh processes. Hayed marshes were no different from adjacent reference marshes in plant species density (species per area) and end-of-year aboveground biomass, but did differ in vegetation composition. Spartina patens was more abundant in hayed marshes than S. alterniflora, and the reverse was true in reference marshes. The differences in relative covers of these plant species were not associated with any differences between hayed and reference marshes in the elevations of the marsh platform. Instead it suggested that S. patens was more tolerant of haying than S. alterniflora. Spartina patens had higher stem densities in hayed marshes than it did in reference marshes, suggesting that periodic cutting stimulated tillering of this species. Although we predicted that haying would stimulate benthic chlorophyll production by opening up the canopy, we found differences to be inconsistent, possibly due to the relatively rapid regrowth of S. patens and to grazing by invertebrates on the algae. The pulmonate snail, Melampus bidendatus was depleted in its δ¹³C content in the hayed marsh compared to the reference, suggesting a diet shift to benthic algae in hayed marshes. The stable isotope ratios of a number of other consumer species were not affected by haying activity. Migratory shorebirds cue in to recently hayed marshes and may contribute to short term declines in some invertebrate species, however, the number of taxa per unit area of marsh surface invertebrates and their overall abundances were unaffected by haying over the long term. Haying had no impact on nutrient concentrations in creeks just downstream from hayed plots, but the sediments of hayed marshes were lower in total N and P compared to references. In sum, haying appeared to affect plant species composition but had only short-term affects on consumer organisms. This contrasts with many grassland ecosystems, where an intermediate level of disturbance, such as by grazing, increases species diversity and may stimulate productivity. From a management perspective, periodic mowing could be a way to maintain S. patens habitats and the suite of species with which they are associated.