Below- and Aboveground Biomass of <Emphasis Type="Italic">Spartina alterniflora</Emphasis>: Response to Nutrient Addition in a Louisiana Salt Marsh

The responses of Spartina alterniflora above- and belowground biomass to various combinations of N, P, and Fe were documented in a 1-year field experiment in a Louisiana salt marsh. Five levels of N additions to 0.25 m² plots resulted in 18% to 138% more live aboveground biomass compared to the control plots and higher stem densities, but had no effect on the amount of live belowground biomass (roots and rhizomes; R&R). There was no change in the aboveground biomass when P or Fe was added as part of a factorial experiment of +P, +N, and +Fe additions, but there was a 40% to 60% decrease in the live belowground biomass, which reduced the average R&R:S ratio by 50%. The addition of various combinations of nutrients had a significant affect on the belowground biomass indicating that the addition of P, not N, eased the need for root foraging activity. The end-of-the-growing-season N:P molar ratios in the live above- and belowground tissues of the control plot was 16.4 and 32.7, respectively. The relative size of the belowground standing stocks of N and P was higher than in the aboveground live tissues, but shifted downwards to about half that in fertilized plots. We conclude that the aboveground biomass was directly related to N availability, but not P, and that the accumulation of belowground biomass was not limited by N. We suggest that the reduction in belowground biomass with increased P availability, and the lower absolute and relative belowground standing stocks of P as plant tissue N:P ratios increased, is related to competition with soil microbes for P. One implication for wetland management and restoration is that eutrophication may be detrimental to long-term salt marsh maintenance and development, especially in organic-rich wetland soils.     

Responses of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> to Multiple Stressors: Changing Precipitation Patterns,Accelerated Sea Level Rise,and Nutrient Enrichment

Coastal wetlands, well recognized for their ecosystem services, have faced many threats throughout the USA and elsewhere. While managers require good information on the net impact of these combined stressors on wetlands, little such information exists. We conducted a 4-month mesocosm study to analyze the multiple stressor effects of precipitation changes, sea level rise, and eutrophication on the salt marsh plant Spartina alterniflora. Pots containing plants in an organic soil matrix were positioned in tanks and received Narragansett Bay (RI, USA) water. The study simulated three precipitation levels (ambient daily rain, biweekly storm, and drought), three levels of tidal inundations (high (15 cm below mean high water (MHW)), mean (MHW), and low (15 cm above MHW)), and two nutrient enrichment levels (unenriched and nutrient-enriched bay water). Our results demonstrate that storm and drought stressors led to significantly less above- and belowground biomass than those in ambient rain conditions. Plants that were flooded at high inundation had less belowground biomass, fine roots, and shoots. Nutrients had no detectable effect on aboveground biomass, but the enriched pots had higher stem counts and more fine roots than unenriched pots, in addition to greater CO₂ emission rates; however, the unenriched pots had significantly more coarse roots and rhizomes, which help to build peat in organogenic marshes. These results suggest that multiple stressors of altered precipitation, sea level rise, and nutrient enrichment would lead to reduced marsh sustainability.     

Relationship between aboveground and belowground biomass ofSpartina alterniflora (Smooth Cordgrass)

Aboveground and belowground biomass ofSpartina alterniflora were harvested during the period of peak aerial biomass from six sites along a latitudinal gradient ranging from Georgia to Nova Scotia. An equation relating live aboveground to live belowground biomass for short-form plants was formulated, using data collected in Delaware marshes. When data from the other sites were substituted into the equation, the mean live belowground biomass it predicted was within 15% of the value determined by harvesting at four of the five sites. At all sites, short-form plant live belowground biomass was concentrated in the upper 10 cm. Dead belowground biomass was located mostly in the top 15 cm in southern marshes, but was more evenly distributed with depth in northern marshes. Results were more ambiguous for tall-form plants, probably because of greater spatial variability in biomass distribution, and greater seasonal biomass dynamics.     

A Landscape-Scale Assessment of Above- and Belowground Primary Production in Coastal Wetlands: Implications for Climate Change-Induced Community Shifts

Above- and belowground production in coastal wetlands are important contributors to carbon accumulation and ecosystem sustainability. As sea level rises, we can expect shifts to more salt-tolerant communities, which may alter these ecosystem functions and services. Although the direct influence of salinity on species-level primary production has been documented, we lack an understanding of the landscape-level response of coastal wetlands to increasing salinity. What are the indirect effects of sea-level rise, i.e., how does primary production vary across a landscape gradient of increasing salinity that incorporates changes in wetland type? This is the first study to measure both above- and belowground production in four wetland types that span an entire coastal gradient from fresh to saline wetlands. We hypothesized that increasing salinity would limit rates of primary production, and saline marshes would have lower rates of above- and belowground production than fresher marshes. However, along the Northern Gulf of Mexico Coast in Louisiana, USA, we found that aboveground production was highest in brackish marshes, compared with fresh, intermediate, and saline marshes, and belowground production was similar among all wetland types along the salinity gradient. Multiple regression analysis indicated that salinity was the only significant predictor of production, and its influence was dependent upon wetland type. We concluded that (1) salinity had a negative effect on production within wetland type, and this relationship was strongest in the fresh marsh (0–2 PSU) and (2) along the overall landscape gradient, production was maintained by mechanisms at the scale of wetland type, which were likely related to plant energetics. Regardless of wetland type, we found that belowground production was significantly greater than aboveground production. Additionally, inter-annual variation, associated with severe drought conditions, was observed exclusively for belowground production, which may be a more sensitive indicator of ecosystem health than aboveground production.     

<Emphasis Type="Italic">Spartina alterniflora</Emphasis> Biomass Allocation and Temperature: Implications for Salt Marsh Persistence with Sea-Level Rise

To predict the impacts of climate change, a better understanding is needed of the foundation species that build and maintain biogenic ecosystems. Spartina alterniflora Loisel (smooth cordgrass) is the dominant salt marsh-building plant along the US Atlantic coast. It maintains salt marsh elevation relative to sea level by the accumulation of aboveground biomass, which promotes sediment deposition and belowground biomass, which accretes as peat. Peat accumulation is particularly important in elevation maintenance at high latitudes where sediment supply tends to be limited. Latitudinal variation in S. alterniflora growth was quantified in eight salt marshes from Massachusetts to South Carolina. The hypothesis that allocation to aboveground and belowground biomass is phenotypically plastic was tested with transplant experiments among a subset of salt marshes along this gradient. Reciprocal transplants revealed that northern S. alterniflora decreased allocation to belowground biomass when grown in the south. Some northern plants also died when moved south, suggesting that northern S. alterniflora may be stressed by future warming. Southern plants that were moved north showed phenotypic plasticity in biomass allocation, but no mortality. Belowground biomass also decomposed more quickly in southern marshes. Our results suggest that warming will lead northern S. alterniflora to decrease belowground allocation and that belowground biomass will decompose more quickly, thus decreasing peat accumulation. Gradual temperature increases may allow for adaptation and acclimation, but our results suggest that warming will lower the ability of salt marshes to withstand sea-level rise.     

Effects of Shoreline Development on Composition and Physical Structure of Plants in a South Carolina High Marsh

Increased freshwater and nutrient runoff associated with coastal development is implicated in dramatically altering estuarine communities along eastern US shorelines. We examined effects of three categories of shoreline development on high-marsh environments within Murrells Inlet, South Carolina, USA by measuring sediment nutrients, porewater salinity, plant species diversity, and above- and belowground plant biomass. Effects on new plant growth also were examined in plot clearing and transplantation experiments. Greater nutrient availability in sediments along developed shorelines was reflected in greater aboveground biomass and nitrogen storage in Juncus roemerianus plant tissue. Plant species composition was not significantly different among levels of shoreline development. Zinc concentrations were greater in sediments from developed shorelines and may represent an easily measured indicator of shoreline development. Recently accelerating shoreline development in the southeastern USA may alter plant production, nitrogen storage, and sediment metal content in salt marshes.     

Aboveground and belowground productivity ofSpartina alterniflora (Smooth Cordgrass) in natural and created Louisiana salt marshes

In Louisiana, salt marshes are being created in an effort to offset the large loss of such habitat that has occurred over the last 50 yr. Primary productivity is an important function and indicator of success for salt marsh creation and restoration projects. The aim of this study was to determine whether the aboveground and belowground productivity of the dominant salt marsh grassSpartina alterniflora in created marshes in southwest Louisiana began to approximate productivity levels in natural marshes, over time. Net annual aboveground primary productivity (NAPP) was measured by a harvest technique, while the ingrowth core method was used to estimate net annual belowground primary productivity (NBPP). NAPP levels were similar to those found in other, Louisiana salt marshes, while NBPP levels were similar to or higher than the reported range forS. alterniflora studied along the Atlantic and Gulf of Mexico coasts. NAPP tended to decrease as the created marshes aged, but the levels in the oldest, 19 year old, created marsh were still well above values measured in the, natural marshes. It was estimated that it would take 35 yr after marsh creation for NAPP in the created marshes to become equivalent to that in natural marshes. NBPP in the created marshes became equivalent to levels found in the natural marshes after 6–8 yr, but then belowground production increased with marsh age, reaching an asymptote that surpassed natural marsh levels. Equivalency in primary productivity has not been reached in these marshes. Elevation also affected productivity, as higher elevational sites with greater topographic heterogeneity had significantly lower aboveground and belowground biomass levels than those with elevations closer to mean sea level. This underscores the need to construct marshes so that their mean elevation and degree of topographic heterogeneity are similar to natural marshes.     

Primary production and decomposition ofSarcocornia fruticosa (L.) scott andPhragmites australis Trin. Ex Steudel in the Po Delta, Italy

From September 1994 through October 1995 aboveground and belowground production ofSarcocornia fruticosa andPhragmites australis was studied at two sites in the Po Delta. In 1995, aboveground production forS. fruticosa in an intertidal site was 678 g dw m⁻² yr⁻¹ with a peak live biomass of 1,008 g m⁻²; belowground production was 1,260 g m⁻² with a peak live biomass of 3,735 g m⁻². A litter bag decomposition study showed that after 69 wk there were 3.7%, 64.3%, and 66.6% of the original mass of leafy stems, woody stems, and roots, respectively. In a reed bed, which experiences brackish conditions,P. australis aboveground production was 876 g m⁻² with a peak live biomass of 780 g m⁻²; belowground production was 2,263 g m⁻² with a peak live biomass of 4,087 g m⁻². After 65 wk, there was 45.4%, 50.4%, and 29.3%, respectively, of leaves, stems, and rhizomes remaining of the initial biomass. At both sites, regular submersion by salt water probably leads to lower aboveground biomass and higher belowground biomass than reported for other Mediterranean coastal sites. The high belowground biomass can contribute to accretion to offset rising sea level.     

Net primary production and decomposition of salt marshes of the Ebre delta (Catalonia,Spain)

Net primary production was measured in three characteristic salt marshes of the Ebre delta: anArthrocnemum macrostachyum salt marsh,A. macrostachyum-Sarcocornia fruticosa mixed salt marsh andS. fruticosa salt marsh. Above-ground and belowground biomass were harvested every 3 mo for 1 yr. Surface litter was also collected from each plot. Aboveground biomass was estimated from an indirect non-destructive method, based on the relationship between standing biomass and height of the vegetation. Decomposition of aboveground and belowground components was studied by the disappearance of plant material from litter bags in theS. fruticosa plot. Net primary production (aboveground and belowground) was calculated using the Smalley method. Standing biomass, litter, and primary production increased as soil salinity decreased. The annual average total aboveground plus belowground biomass was 872 g m⁻² in theA. macrostachyum marsh, 1,198 g m⁻² in theA. macrostachyum-S. fruticosa mixed marsh, and 3,766 g m⁻² in theS. fruticosa biomass (aboveground plus belowground) was 226, 445, and 1,094 g m⁻², respectively. Total aboveground plus below-ground net primary production was 240, 1,172, and 1,531 g m⁻² yr⁻¹. There was an exponential loss of weight during decomposition. Woody stems and roots, the most recalcitrant material, had 70% and 83% of the original material remaining after one year. Only 20–22% of leafy stem weight remained after one year. When results from the Mediterranean are compared to other salt marshes dominated by shrubbyChenopodiaceae in Mediterranean-type climates, a number of similarities emerge. There are similar zonation patterns, with elevation and maximum aboveground biomass and primary production occurring in the middle marsh. This is probably because of stress produced by waterlogging in the low marsh and by hypersalinity in the upper marsh.     

Belowground productivity of roots and rhizomes in a giant cordgrass marsh

Belowground production of roots and rhizomes in the top 20 cm of soil was 2.2 kg m^?2 yr^?1 based on a maximum minus minimum estimation procedure in a giant cordgrass (Spartina cynosuroides (L.) Roth) marsh in Mississippi. Approximately 1.9 kg m^?2 (86%) of this production occurred in late spring-summer and 0.3 kg m^?2 in late fall. This estimate ignores any production below 20 cm depth and is thus an underestimate. Production values increased to 4.0 kg m^?2 yr^?1 using Smalley’s technique and accounting for decomposition. Aboveground tissues (leaves and stems) were depleted in nitrogen in July which corresponded to peaks in both above- and belowground biomass. The low root/shoot ratio (2.6) on this marsh does not suggest that growth is nutrient limited. Indeed, total productivity (above- and belowground) for this marsh was high (between 4.4 and 6.2 kg m^?2 yr^?1).     

Aboveground Biomass Patterns of Dominant Spartina Species and Their Relationship with Selected Abiotic Variables in Argentinean SW Atlantic Marshes

Salt marsh zonation patterns generate different abiotic and biotic conditions that can accentuate species inherent differences in primary production and biomass. In South West Atlantic marshes, there are two Spartina species: Spartina alterniflora in the low intertidal and Spartina densiflora in the high intertidal. These two species are generally found in all marshes but with different dominance: In some marshes, the S. densiflora zone occupies higher extents, and in others, the S. alterniflora zone is the one that prevails. We found through field sampling that, in six studied marshes, there is greater S. densiflora live and total (i.e., dead+live) aboveground biomass (g m^?2) in the marshes dominated by S. densiflora than in the ones dominated by S. alterniflora. Spartina alterniflora had similar aboveground biomass in the six marshes, regardless of the dominance of each species. When comparing the two Spartina species within each marsh, S. densiflora had greater live and total biomass in the marshes it dominates. In the marshes dominated by S. alterniflora, both species had similar live and total biomass. In all marshes, there was greater dead S. densiflora biomass. A multivariate analysis using selected abiotic factors (i.e., salinity, latitude, and tidal amplitude) showed that S. alterniflora aboveground biomass patterns are mainly correlated with salinity, while S. densiflora live biomass is mainly correlated with salinity and latitude, dead biomass with salinity and tidal amplitude, and total biomass with salinity alone. We conclude that in S. densiflora dominated marshes, the main processes of that species zone (i.e., nutrient accumulation) will be accentuated because of its higher biomass. We also conclude that climatic conditions, in combination with specific Spartina biotic and ambient abiotic parameters, can affect marsh ecological functions.     

C3 and C4 Biomass Allocation Responses to Elevated CO2 and Nitrogen: Contrasting Resource Capture Strategies

Plants alter biomass allocation to optimize resource capture. Plant strategy for resource capture may have important implications in intertidal marshes, where soil nitrogen (N) levels and atmospheric carbon dioxide (CO₂) are changing. We conducted a factorial manipulation of atmospheric CO₂ (ambient and ambient?+?340?ppm) and soil N (ambient and ambient?+?25?g?m^?2?year^?1) in an intertidal marsh composed of common North Atlantic C₃ and C₄ species. Estimation of C₃ stem turnover was used to adjust aboveground C₃ productivity, and fine root productivity was partitioned into C₃?CC₄ functional groups by isotopic analysis. The results suggest that the plants follow resource capture theory. The C₃ species increased aboveground productivity under the added N and elevated CO₂ treatment (P?<?0.0001), but did not under either added N or elevated CO₂ alone. C₃ fine root production decreased with added N (P?<?0.0001), but fine roots increased under elevated CO₂ (P?=?0.0481). The C₄ species increased growth under high N availability both above- and belowground, but that stimulation was diminished under elevated CO₂. The results suggest that the marsh vegetation allocates biomass according to resource capture at the individual plant level rather than for optimal ecosystem viability in regards to biomass influence over the processes that maintain soil surface elevation in equilibrium with sea level.     

Phosphorus biological cycle in the different Suaeda salsa marshes of the Yellow River estuary, China

Much uncertainty exists in the phosphorus (P) cycle in the marshes of the intertidal zone. This study explored the P cycling in the two Suaeda salsa marshes [middle S. salsa marsh (MSM) and low S. salsa marsh (LSM)] of the Yellow River estuary during April 2008 to November 2009. Results showed seasonal fluctuations and vertical distributions of P in different S. salsa marsh soils, and variations in P content in different parts of plants due to water and salinity status. The N/P ratios of the different S. salsa were 9.87 ± 1.23 and 15.73 ± 1.77, respectively, indicating that plant growth in MSM was limited by N, while that in LSM was limited by both N and P. The S. salsa litter in MSM released P to the environment throughout the year, while that in LSM immobilized P from the environment at all times. The P absorption coefficients of S. salsa in MSM and LSM were very low (0.0010 and 0.0001, respectively), while the biological cycle coefficients were high (0.739 and 0.812, respectively). The P turnovers among compartments of MSM and LSM showed that the uptake amounts of roots were 0.4275 and 0.0469 g m^?2 year^?1 and the values of aboveground parts were 1.1702 and 0.1833 g m^?2 year^?1, the re-translocation quantities from aboveground parts to roots were 0.8544 and 0.1452 g m^?2 year^?1, the translocation amounts from roots to soil were 0.0137 and 0.0012 g m^?2 year^?1, the translocation quantities from aboveground living bodies to litter were 0.3157 and 0.0381 g m^?2 year^?1, and the annual return quantities from litter to soil were less than 0.0626 and ?0.0728 g m^?2 year^?1 (minus represented immobilization), respectively. P was an important limiting factor in S. salsa marshes, especially in LSM. S. salsa was seemingly well adapted to the low-nutrient condition and the vulnerable habitat, and the nutrient enrichment due to the import of N and P from the Yellow River estuary would be a potential threat to the S. salsa marshes.     

Effects of grazing by feral horses,clipping, trampling,and burning on a Georgia salt marsh

Responses ofSpartina alterniflora marsh to combinations of feral horse grazing, clipping, simulated trampling, and a late winter burn were studied on Cumberland Island National Seashore, Georgia. Replicated 200-m² plots were established and sampled bimonthly from July 1983 to November 1984. Clipping and trampling each reduced peak aboveground biomass by 20% in 1983 and 50% (clipping) and 55% (trampling) in 1984. A March burn reduced peak aboveground biomass by 35% in 1984. Trampling and burning earch reduced net aboveground primary production (NAPP) by 35%, but clipping did not reduce NAPP. Standing stocks of live rhizomes were correlated with aboveground biomass and were reduced with experimental treatments. Abundance of the periwinkle snail (Littorina irrorata) was also reduced. Horse grazing had a substantial impact on standing stocks and NAPP ofSpartina, but grazing was not uniform throughout the marsh. Moderately grazed plots had NAPP reduced by 25% compared to ungrazed plots. Heavily grazed plots had extremely low NAPP, and abovegroundSpartina never exceeded 40 g m^?2 dry mass compared to 360 g m^?2 within exclosures.     

The potential role of roots and rhizomes in structuring salt-marsh benthic communities

The density of the Carolina marsh clam,Polymesoda caroliniana (Bosc), was determined in three adjacent tidal marsh communities which differed only in plant species composition. Clam density was inversely related to the density (biomass) of plant roots and rhizomes in sediments and directly related to density of plant stems (numbers). Clam abundance was not related to the basal area of plant stems. Each plant community contained clams of various ages from juvenile to adult indicating continued recruitment and survival. These data suggest thatP. caroliniana is most abundant inJuncus roemerianus marshes because there are fewer roots and rhizomes (mean of 2.5 kg m^?2) to hamper burrwing as compared toSpartina alterniflora andcynosuroides (5.1 and 6.3 kg m^?2, respectively) dominated marshes. Salinity, floding frequency, distance from flooding water, and sediment type were essentially constant among the three plant communities. Root/rhizome density should be collected along with other environmental parameters during studies of benthic organisms on marshes because it potentially limits the occurrence or abundance of some species.     

Nitrogen Pools of Macrophyte Species in a Coastal Lagoon Salt Marsh: Implications for Seasonal Storage and Dispersal

High nitrogen (N) loading rates received by coastal bays can have deleterious effects on aquatic ecosystems. Salt marshes can intercept land-based N through seasonal plant uptake, denitrification, and burial. Salt marshes fringing Delaware’s Inland Bays are characterized by different plant species occurring in close proximity. To evaluate N pool retention and loss for the dominant plant species, we measured seasonal N concentration and pool size, N resorption efficiency, loss during decomposition, and soil N. Seasonal variation in N pools and fluxes differed among species. Seasonal differences in the total N pools of the herbaceous species were largely influenced by belowground fine root and dead macro-organic matter fluxes. N production rate estimates ranged from 18 g N m⁻² year⁻¹ aboveground for the high marsh shrub to 40.8 g N m⁻² year⁻¹ above- and belowground for the high marsh rush illustrating the importance of incorporating species-specific dynamics into ecosystem N budgets.     

Demography and biomass of the seagrass Zostera marina in a Mexican Coastal Lagoon

From January 1987 to February 1988 the annual biomass cycle and demography of the seagrass Zostera marina were assessed in San Quintin Bay, a shallow coastal lagoon on the Pacific coast of Baja California, Mexico. Shoot density and aboveground biomass were sampled monthly along two intertidal transects parallel to the shore. Belowground biomass was sampled every 2 mo. Shoot density differed between transects, ranging from 929±71 (SE) in July to 279 ±80 shoots m⁻² in December, at the deeper transect (I). At the shallow transect (II) there was not a significant difference through time, and a mean of 737 shoots m⁻² was calculated. Lateral shoots were present year round and represented between 1% and 30% of total density at transect I and between 3% and 25% at transect II. Reproductive shoots were present from March to September at both transects, with a density range of 77±28 shoots m⁻² (March) to 9±3 shoots m⁻² (September), and represented 5% of total shoot density. Neither aboveground biomass nor LAI (Leaf Area Index) differed between transects, with values ranging between 77±14.5 g dry wt m⁻² (October 1987) and 13±2.4 g dry wt m⁻² (February 1988) for aboveground biomass, and between 0.6±0.2 m² leves m⁻² substrate (January) and 2.7±0.3 m² leaves m⁻² substrate (September) for LAI. Neither root biomass nor rhizome biomass differed between transects, or as a function of time; the mean value for roots was 17 g dry wt m⁻² and for rhizomes 29 g dry wt m⁻². Belowground biomass represented 54% of total biomass. We found a significant correlation between aboveground biomass and LAI (r=0.949 for transect I, and 0.926 for transect II) as well as between total biomass (aboveground and belowground) and LAI (r=0.814), which allows us to consider using LAI as a predictor of these variables. Biomass changes were related to changes in shoot weight (r=0.676 at transect I; 0.582 at transect II), more than to changes in shoot number. Water temperature was found to be the driver of biomass changes in the aboveground compartment.     

Relationship between waterfowl and the seagrassRuppia maritima in a Southwestern Atlantic coastal lagoon

We evaluated the distribution of waterfowl in relation to a seagrass (Ruppia maritima) patch, to infauna, and on its relationship with substrate characteristics. An experiment performed in the Mar Chiquita coastal lagoon (37°46′S, 57°27′W; Argentina) was used to evaluate the effect of herbivory on widgeon grass,R. maritima. Control plots of equal size, located between bird exclosures, were exposed to herbivory. The experiment ran for 31 d, starting on December 1, 1994. Censuses showed that black-necked swan ( $\bar x = 50$ birds ha^?1, SD = 37, n = 10) and coots ( $\bar x = 42$ birds ha^?1, SD = 28, n = 10) were the most important (always present) of the waterfowl seen feeding onR. maritima. The results of the experiment showed greater leaf lengths, lower belowground (rhizomes and roots) biomass, greater aboveground (leaves and shoots) biomass, and greater abundance of the polychaeteHeteromastus similis in exclosure plots. There were no exclosure effects on total biomass (belowground plus aboveground), reproductive parts (fruits and pre- and postpollination flowers), or abundance of most benthic infauna. Topographic surveys showed that sediment surface was higher within theR. maritima patches, but there were no significant differences in granulometric composition. Surveys also showed that bird distribution was strongly associated with the presence ofR. maritima.     

Comparison of giant cutgrass productivity in tidal and impounded marshes with special reference to tidal subsidy and waste assimilation

Aboveground live standing crop of giant cutgrass (Zizaniopsis miliacea) populations in similar freshwater tidal and impounded nontidal marshes were almost identical (peaking at 1,039 g per m² in each). The mortality, however, was greater in the tidal marsh resulting in significantly (95% level) greater annual production of aboveground cutgrass in the tidal (1,530±103 g per m² per yr) than the impounded (1,172±88 g per m² per yr) marsh, a 31% difference which we consider to be a measure of tidal subsidy. Belowground production also was found to average higher in the tidal marsh, but estimates were not as satisfactory as the aboveground results due to sampling difficulties. Combined annual above and belowground net production comes to an estimated 2,048 ±101 g per m² per yr for the tidal and 1,481±219 for the impounded cutgrass marsh. The potential of freshwater tidal marshes for tertiary treatment of wastes is briefly discussed.     

Rhizome growth dynamics of native and exotic haplotypes ofPhragmites australis (Common reed)

Phragmites australis (common reed), a clonal grass, has expanded from a minor component of the mid-Atlantic wetlands to a dominant species. It has been suggested that invasive populations ofPhragmites are an exotic haplotype responsible for the dramatic increase in the distribution of the species. We used field observations and measurements and a greenhouse assay to compare native (haplotype F) and exotic (haplotype M) populations, growing adjacent to one another in a brackish marsh near Odessa, Delaware. In the marsh, shoots of the exotic strain emerged from the rhizomes earlier than those of the native and by March there was an order of magnitude more new shoots of the exotic strain than the native. In August, the exotic strain was 30% taller than the native, had twice the leaf biomass, and twice the total biomass. Nine of ten morphological and biomass characteristics measured differed significantly between the native and exotic strains. A greenhouse assay was conducted by planting rhizomes collected in March in shallow trays and growing them for 70 d followed by shoot harvest (Harvest 1). Rhizomes were measured, replanted, and grown for 35 d after which time they were measured and shoots were harvested (Harvest 2). At Harvest 1, shoot height was approximately 80% greater in the exotic strain, shoot biomass was three times higher, aboveground to belowground biomass ratio was twice as high, and rhizome internode length was 50% greater in the exotic strain than the native. These traits, in addition to number of shoots, were also greater in the exotic strain at Harvest 2. The number of rhizome buds at Harvest 1 was three times greater in the native than in the exotic strain. The greater number of rhizome buds in the native would seem to be an advantage, but it did not result in more shoot production. Buds were maintained in an inactive state that does not allow this strain to compete well in a wetland environment inhabited by a more efficient spreader. The earlier emergence of new shoots from the rhizomes, the greater aboveground structure, the greater rhizome internode length, and the quick transition of rhizome buds to shoot or rhizome explain in part the exotic strain's advantage over the native and the mechanisms for its invasive nature.