Lisamarie Windham

A comparison of Phragmites australisin freshwater and brackish marsh environments in North America

This paper compares the available North Americanliterature and data concerning several ecologicalfactors affecting Phragmites australisin inlandfreshwater, tidal fresh, and tidal brackish marshsystems. We compare aboveground productivity, plantspecies diversity, and sediment biogeochemistry; andwe summarize Phragmiteseffects on faunalpopulations in these habitats. These data suggest thatPhragmitesaboveground biomass is higher thanthat of other plant species occurring in the samemarsh system. Available data do not indicate anysignificant difference in the aboveground Phragmitesbiomass between marsh types, nor doesthere appear to be an effect of salinity on height.However, Phragmitesstem density wassignificantly lower in inland non-tidal freshwatermarshes than in tidal marshes, whether fresh orbrackish. Studies of the effects of Phragmiteson plant species richness suggest that Phragmitesdominated sites have lower diversity.Furthermore, Phragmiteseradication infreshwater sites increased plant diversity in allcases. Phragmitesdominated communities appearto have different patterns of nitrogen cyclingcompared to adjacent plant communities. Abovegroundstanding stocks of nitrogen (N) were found to behigher in Phragmitessites compared to thosewithout Phragmites. Porewater ammonium(NH4+) did not differ among plant covertypes in the freshwater tidal wetlands, but inbrackish marshes NH4+was much higher inSpartinaspp. than in neighboring Phragmitesstands. Faunal uses of Phragmitesdominated sites in North America were found to vary bytaxa and in some cases equaled or exceeded use ofother robust emergent plant communities. In light ofthese findings, we make recommendations for futureresearch.

Effects of Phragmites australis (common reed) invasion on aboveground biomass and soil properties in brackish tidal marsh of the Mullica river, New Jersey

Phragmites australis (common reed) has been increasing in brackish tidal wetlands of the eastern United States coast over the last century. Whereas several researchers have documented changes in community structure, this research explores the effects of Phragmites expansion on aboveground biomass and soil properties. We used historical aerial photography and a global positioning system (GPS) to identify and age Phragmites patches within a high marsh dominated by shortgrasses (Spartina patens and Distichlis spicata). Plots along transects were established within the vegetation types to represent a gradient of species dominance and a variety of ages of the Phragmites plots. In comparison to neighboring shortgrass communities, Phragmites communities were found to have nearly 10 times the live aboveground biomass. They also had lower soil salinity at the surface, a lower water level, less pronounced microtopographic relief, and higher redox potentials. These soil factors were correlated with the age and biomass of Phragmites communities, were increasingly different with increasing Phragmites dominance along the transects, and were increasingly altered by the ages of Phragmites communities until the factors stabilized in plots of 8 yr to 15 yr of age. We propose that Phragmites expansion plays an important role in altering these soil properties and suggest a variety of mechanisms to explain these alterations.

Uptake and distribution of metals in two dominant salt marsh macrophytes, Spartina alterniflora (cordgrass) and Phragmites australis (common reed)

We examined patterns of biomass accumulation and tissue concentrations of five metals—mercury, copper, zinc, chromium, lead—and two elements—carbon and nitrogen—to determine differences in net metal accumulation and distribution between Phragmites australis (common reed) and Spartina alterniflora (cord grass) which were growing intermingled in a contaminated low marsh. Data were collected at 2-month intervals across a growing season (April–October, 1999). Although they comprise only 5–15% of whole plant biomass for both species, roots consistently contained 70–100% of the whole plant metal burdens for both S. alterniflora and P. australis (shoot:root ratio <0.42). Stems and rhizomes had low and similar concentrations between plant species throughout the summer. Leaves of S. alterniflora, however, had consistently greater concentrations of Hg and Cr than those of P. australis. In contrast, the micronutrients Cu and Zn were enriched in P. australis leaf tissue in October, compared to S. alterniflora. Pools of metal in aboveground biomass were similar between plant species, but throughout the season S. alterniflora allocated more of this burden to leaf tissue than P. australis, which allocated more of the aboveground burden to stem tissue, a recalcitrant tissue with lower concentrations but greater biomass. The consistently higher concentrations and total pools of Hg and Cr in S. alterniflora leaf tissue and higher Zn and Cu in P. australis may result from differences in leaf phenology, root-influenced metal availability, or transport of dissolved metals. Because S. alterniflora shifts more of its Hg and Cr load into highly decomposable leaf tissues (as opposed to recalcitrant stems, roots, and rhizomes) this pathway of metal bioavailability would be reduced when S. alterniflora is replaced by P. australis.

NET IMPACT OF A PLANT INVASION ON NITROGEN‐CYCLING PROCESSES WITHIN A BRACKISH TIDAL MARSH

Using comparative analysis of the rates of key processes, we have documented the net effect of a shift in plant species composition on nitrogen cycles with the example of the rapid expansion of Phragmites australis (common reed) and its replacement of short grasses (e.g., Spartina patens) in coastal marshes of the eastern United States. In this study, we measured nitrogen (N) uptake by marsh plants, N adsorption from the water column by litter, changes in N content of litter, sediment N mineralization, nitrification, and nitrate consumption in adjacent plots dominated either by P. australis or by historically dominant S. patens. Rates of individual processes were generally greater in P. australis than in S. patens, but the magnitude of difference varied greatly among processes. Seasonal measurements of standing stock nitrogen in plant tissue indicate that P. australis took up ∼60% more N than did S. patens, and annual rates of N immobilization were nearly 300% greater in P. australis litter than in S. pat...

COMPARISON OF BIOMASS PRODUCTION AND DECOMPOSITION BETWEEN PHRAGMITES AUSTRALIS (COMMON REED) AND SPARTINA PATENS (SALT HAY GRASS) IN BRACKISH TIDAL MARSHES OF NEW JERSEY, USA

The recent expansion of Phragmites australis (common reed) from the marsh-upland interface into high marsh zones provides an opportunity to assess the impact of individual plant species on biomass production and decomposition in salt marshes. Seasonal harvests of aboveground and belowground biomass demonstrate that annual production of P. australis is approximately three times greater for aboveground biomass, two times greater for belowground biomass, and 30% lower in root: shoot ratio than neighboring populations of S. patens. Whole-plant litter (stems and leaves) also decomposes at a much slower annual rate for P. australis (k=0.25) than S. patents litter (k=0.57). By crossing litter type with site of litter decomposition, I found these plant species to influence decay rates through litter type and not through their effects on marsh surface conditions (e.g., temperature, sedimentation rates). Based on these calculations, annual rates of carbon accumulation in the peat of high marshes are likely to increase 5-fold once P. australis becomes established due to its greater rates of biomass production and residence time in infrequently flooded brackish marshes.

Effects of Common Reed (Phragmites australis) Expansions on Nitrogen Dynamics of Tidal Marshes of the Northeastern U.S.

Several recent studies indicate that the replacement of extant species withPhragmites australis can alter the size of nitrogen (N) pools and fluxes within tidal marshes. Some common effects ofP. australis expansion are increased standing stocks of N, greater differentiation of N concentrations between plant tissues (high N leaves and low N stems), and slower whole-plant decay rates than competing species (e.g.,Spartina, Typha spp.). Some of the greater differences between marsh types involveP. australis effects on extractable and porewater pools of dissolved inorganic nitrogen (DIN) and N mineralization rates. Brackish and salt marshes show higher concentrations of DIN in porewater beneathSpartina spp. relative toP. australis, but this is not observed in freshwater tidal marshes whenP. australis is compared withTypha spp. or mixed plant assemblages. With few studies of concurrent N fluxes, the net effect ofP. australis on marsh N budgets is difficult to quantify for single sites and even more so between sites. The magnitude and direction of impacts ofP. australis on N cycles appears to be system-specific, driven more by the system and species being invaded than byP. australis itself. WhereP. australis is found to affect N pools and fluxes, we suggest these alterations result from increased biomass (both aboveground and belowground) and increased allocation of that biomass to recalcitrant stems. Because N pools are commonly greater inP. australis than in most other communities (due to plant and litter uptake), one of the most critical questions remaining is “From where is the extra N inP. australis communities coming?” It is important to determine if the source of the new N is imported (e.g., anthropogenic) or internallyproduced (e.g., fixed, remineralized organic matter). In order to estimate net impacts ofP. australis on marsh N budgets, we suggest that further research be focused on the N source that supports high standing stocks of N inP. australis biomass (external input versus internal cycling) and the relative rates of N loss from different marshes (burial versus subsurface flow versus denitrification).

Does Phragmites Expansion Alter the Structure and Function of Marsh Landscapes? Patterns and Processes Revisited

We assess the probability and importance of different spatial distributions ofPhragmites australis (Trin Ex Steud) within brackish tidal marshes of the mid-Atlantic United States coast. The comparative impact ofPhragmites expansion on the larger coupled marsh-estuary system may partially be a function of the landscape area dominated byPhragmites, the landscape position occupied byPhragmites, the landscape pattern created byPhragmites expansions, and the resulting impact on tidal drainage networks. We find evidence thatPhragmites establishment can occur at many landscape positions, and thatPhragmites spread within a marsh can occur via colonization (new patches), linear clonal growth (along a preferred axis), or circular clonal growth (non-directional, random spread). Early intervals ofPhragmites spread were dominated by colonization for all sites except for Piermont Marsh (which appeared to be dominated by linear clonal growth) and Lang Tract (which appeared to be dominated by circular clonal growth). Although 46–100% of new patches ofPhragmites occurred within 5 m of drainages, at only one site (Piermont Marsh, New York) didPhragmites populations remain concentrated along creek banks. Except for Iona Island, New York, which appears to be in an early stage ofPhragmites invasion, patch dynamics at all sites showed an increase followed by a decrease in patch number, as independent patches became established, expanded, and coalesced. We also found some evidence for a loss of first order streams at later stages ofPhragmites invasions in several sites (Hog Island, Lang Tract, Silver Run).

Lead uptake, distribution, and effects in two dominant salt marsh macrophytes, Spartina alterniflora (Cordgrass) and Phragmites australis (Common reed)

We examined biomass accumulation, tissue concentrations of lead (Pb), and net uptake of Pb in Phragmites australis (common reed) and Spartina alterniflora (salt cord grass) grown under greenhouse conditions in sediment of different Pb concentrations. Sediment and newly emerged ramets of each plant species were collected in April 1999 from Tuckerton, NJ, a relatively clean salt marsh. One-gallon pots were filled with either control sediment (29 microg g(-1) Pb) or Pb-added sediment (68 microg g(-1) Pb), and the sediment moisture was kept saturated along with controlled additions of additional nutrients. At harvest in October, whole plant biomass was 60-85% greater for pots with P. australis than pots with S. alterniflora and a 40-70% reduction in biomass in response to the addition of Pb was observed for both species. In the high Pb treatments, both concentrations and pools of Pb were greater in the leaves of S. alterniflora than in leaves of P. australis at the end of the growing season. In both species, Pb concentrations were higher in lower leaves than upper leaves. The addition of Pb into experimental pots led to over an 800% increase in Pb standing stock for both species. In S. alterniflora, however, significantly more of this pool was allocated to aboveground biomass (leaves and stems) than to belowground biomass (roots and rhizomes). This difference in allocation was more profound at the higher sediment Pb concentration (Pb-added pots). This fundamental difference between the species in response to Pb contamination indicates that metal export into food webs or the water column should be greater in stands of S. alterniflora than in P. australis. These results suggest that in Pb-contaminated, and possibly all metal-contaminated sediments, the replacement of S. alterniflora with P. australis may reduce metal bioavailability by sequestering a greater proportion of its metal burden in belowground tissues which are likely to be permanently buried.

Patterns and processes of mercury release from leaves of two dominant salt marsh macrophytes,Phragmites australis andSpartina alterniflora

The release of mercury (Hg) from leaf tissue was compared between two dominant salt marsh macrophytes,Spartina alterniflora andPhragmites australis. Rates of Hg release were measured for individual leaves from late May to late July, along with concentrations of Hg in leaf tissue, rates of sodium (Na) release, and rates of transpiration. Leaves ofS. alterniflora consistently releasd 2–3 times more Hg than leaves ofP. australis. Leaves ofS. alterniflora also contained greater concentrations of Hg during these months. In contrast toP. australis leaves, rates of Na release were high forS. alterniflora and were correlated with rate of Hg release. Transpiration rates averaged 2.2 times greater forPhragmites as compared toS. alterniflora, and were not correlated with the other variables at the leaf level for either species. Leaf Hg concentration was highly correlated with Hg release for both species, but the slope was significantly greater forS. alterniflora. Monthly differences were profound for all climate and physiological variables measured, with high measurements in May, and lower measurements in June and July. For both species, the highest Hg content was found in lower leaves in May, followed by upper leaves in May. Hg accumulation in leaf tissue and release from both species appear to be greatest in the spring, although differences between the species persist throughout these peak months of the growing season.

Release into the environment of metals by two vascular salt marsh plants

Metals in contaminated salt marshes are mainly locked in the anaerobic layer of sediments, where they are tightly bound as sulfides and organic complexes. Vascular plants survive in saturated soils in part by pumping O2 into their root zones, changing their microenvironment to an oxic one. This, along with chelating exudates, mobilizes metals, allowing uptake by the roots. We compared the common reed Phragmites australis and cordgrass Spartina alterniflora in lab and field studies for ways in which they handle trace metals. Both plants store most of their metal burden in their roots, but some is transported to aboveground tissues. Spartina leaves contain approximately 2-3 x more Cr, Pb, and Hg than Phragmites leaves, but equivalent Cu and Zn. Furthermore, Spartina leaves have salt glands, so leaf excretion of all metals is twice that of Phragmites. In-depth studies with Hg indicate that Hg excretion correlates with Na release but not with transpiration, which is 2.2 x higher in Phragmites; and that more Hg accumulates in early-appearing leaves than in upper (i.e. later) leaves in both species. Spartina thus makes more metals available to salt marsh ecosystems than Phragmites by direct excretion and via dead leaves which will enter the food web as detritus.