American Journal of Botany 86(5): 697702. 1999.

EFFECTS

OF DRAINAGE AND SOIL ORGANIC CONTENT

ON GROWTH OF

SPARTINA ALTERNIFLORA (POACEAE)

AND

IN AN ARTIFICIAL SALT MARSH MESOCOSM1

DAVID E. PADGETT2

JEFFREY L. BROWN

Department of Biological Sciences, University of North Carolina, Wilmington, North Carolina 28403; and Department of Mathematical Sciences, University of North Carolina, Wilmington, North Carolina 28403 An articial salt marsh mesocosm was constructed using 680-L polypropylene tanks to determine the effect of soil drainage depth and organic content on growth and rhizome proliferation of the salt marsh smooth cordgrass Spartina alterniora. Soil drainage depth had no effect on accumulation of aerial or subsurface plant tissue, but tanks that had 2.5% soil organic content supported enhanced aerial tissue and rhizome growth compared to tanks that contained sand alone. We propose a mathematical model for predicting the mass of photosynthetically signicant leaf tissue without cutting and drying leaves. Implications of these ndings for salt marsh creation projects are discussed. Key words: biomass estimation; drainage; marsh creation; rhizome; soil organics, Spartina alterniora, Poaceae.

Numerous published reports have documented the importance of the salt marsh cordgrass Spartina alterniora Loisel. to tidal ecosystems along the U.S. Atlantic and Gulf coasts. In recent years, however, there has been increasing interest in this plant in connection with marsh creation projects (Broome, Seneca, and Woodhouse, 1986, 1988; Moy and Levin, 1991; Matthews and Minello, 1994; Padgett, Rogerson, and Hackney, 1998). This interest largely relates to the extensive, subsurface network of roots and rhizomes that cordgrass produces and its role in soil stabilization. Indeed, reports indicate that 60% or more of cordgrass biomass is found below the marsh surface (Good, Good, and Frasco, 1982; Broome, Seneca, and Woodhouse, 1986). To be sure, aerial tissues of this plant are important as habitat and ultimate food source for marsh animals (Teal, 1962; Odum and de la Cruz, 1967; Boesch and Turner, 1984; Posey, Alphin, and Powell, 1997). Their principal value to soil stabilization, however, is as a carbohydrate source that fuels proliferation of subsurface tissues. It necessarily follows that soil conditions conducive to growth of cordgrass stems and leaves likely would be those that promote rapid soil stabilization. In light of recent federal policy mandating no net loss of wetlands, the U.S. Army Corps of Engineers routinely requires replacement of an acre of marsh land for every acre impacted by construction projects. Despite this policy, the Corps currently has no regulation governing the type of soil that can be used to construct new marshes (Padgett, Rogerson, and Hackney, 1998). Soil organic content and the drainage characteristics it
1 Manuscript received 9 June 1998; revision accepted 6 October 1998. The authors thank UNC Wilmington undergraduate students Eric Honeycutt, Heather Altemus, Amy Dixon, and Amy Weaver for assistance with mesocosm construction, maintenance, and data gathering; Dr. Martin Posey (UNCW Biology) for assisting with ANOVA analyses; and Dr. Daniel B. Plyler (UNCW Biology) for critically reading a nal manuscript draft. The UNCW Center for Marine Sciences Research provided logistical assistance and nancial support for this study. This is contribution number 201 from the UNCW Center for Marine Sciences Research. 2 Author for correspondence.

inuences are two major factors among a myriad of other abiotic conditions affecting establishment of cordgrass outplants, thus it seems reasonable that they should be optimized during marsh creation efforts. The purpose of this paper is to present results from the rst year of a long-term salt marsh mesocosm study whose ultimate objective is to assess the inuence of these two important variables on production of aerial and subsurface tissues by S. alterniiora seedlings. Our particular interest is in monitoring rhizome proliferation. We view these horizontal, underground stems as critically important to soil stabilization because they serve as plant generating centers analogous to stolons of strawberries. Soil conditions favoring horizontal spread of rhizomes from an initial seedling thus would produce clonal offspring more rapidly, thereby generating more roots to anchor soil against erosion (Broome, Seneca, and Woodhouse, 1986, 1988). MATERIALS AND METHODS
Initial design and plantingTwelve, 680-L polypropylene tanks (stock number SST-4422 Polytank, Inc., Litcheld, Minnesota) were engineered with adjustable drains to control soil drainage depth and lled either with washed masonry sand (0.15% mean organic content, w/w) or masonry sand amended with canadian peat moss (2.5% mean soil organic content) to serve as a substrate for cordgrass seedlings (Figs. 13). Our experimental design involved ooding all tanks with unltered, natural seawater (35 ppt salinity), and then allowing them to drain to a prescribed depth below the soil surface on a continuous, cyclic basis that corresponded with the natural tidal cycle in the same geographic area. Within the experimental plot we constructed triplicate tanks of the following treatments: (1) sand substrate, ebbtide drainage to 5 cm below soil surface, (2) sand substrate, drainage to 15 cm, (3) sand peat substrate, drainage to 5 cm, and (4) sand peat, drainage to 15 cm. Tanks were installed outdoors in full sunlight at the UNC Wilmington Aquaculture facility at Wrightsville Beach, North Carolina (Fig. 3) in August 1996 and the seawater ood/ebb cycles allowed to precondition the soil for 9 mo before planting cordgrass seedlings. Seawater was drawn from Banks Channel, Wrightsville Beach, North Carolina, and held in two, 113 000-L storage tanks from which it fed our system on each 6-h ood tide cycle. All tanks were ooded synchronously through

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Figs. 13. Salt marsh mesocosm tanks. 1. Close-up view showing vertically adjustable ebb-tide drain (ed). Note overlow drain (od) on adjacent tank. 2. Schematic, cross-sectional diagram of mesocosm tank indicating ooding water inlet (fw). Fixed-position overow drain governs maximum water depth (mwd) on each ood cycle. Force exerted by hydraulic head pressure (hh) between mwd and ebb-tide drain (ed) governs velocity of drainage to achieve maximum drainage depth (mdd) beneath soil level (sl). Small arrows within tanks indicate path of ebb-tide water percolation through soil and gravel lter (gf). 3. Large-scale view of mesocosm experimental plot. a 2.5-cm diameter PVC (polyvinyl chloride) piping system controlled by an in-line electrical solenoid valve with a 24-h interval timer. We adjusted the timer setting daily so that ood/ebb cycles corresponded exactly to local tidal cycles. When the solenoid interrupted seawater ow, tanks drained by gravity over the 6-h ebb tide phase to the prescribed depth below the soil surface before the next ood cycle began. Soil drainage rate was calibrated for each tank by adjusting the vertical position of the external ebb-drain (Fig. 2) such that the target drainage depth was reached at approximately the same time in all treatments. Cordgrass seeds were collected in early October 1996 from tall-form plants in the nearby Shell Island salt marsh at the northern end of Wrightsville Beach and stored in the dark in 20 ppt seawater at 4 C until early March 1997 (Broome, Seneca, and Woodhouse, 1986). Seeds that germinated during cold storage were transplanted individually into 4 4 6 cm peat pots lled with commercial potting soil on 10 March and propagated in a greenhouse under natural light regimes for 9 wk. Seedlings were fertilized with Miracid commercial plant food every other week during this period. We outplanted seedlings into our mesocosm tanks on 21 May 1997, at which time the average seedling height was 8 cm. Nine seedlings were planted in each tank in a 3 3plant grid design with each 28 cm from any adjacent plant and all plants in outside rows 28 cm from the tank edge. At the time of planting, a bamboo shishkabob skewer was used to anchor each peat pot until subsequent root growth rmly established the seedling in its surrounding soil. These skewers were left in place throughout the growing season and served as points of reference for assessing horizontal rhizome growth (discussed below). Monitoring plant growthThe emergence of each new aerial shoot was recorded about every 2 wk throughout the growing season by measuring its linear distance to the closest seedling (seedling of presumed origin). These measurements were considered to be conservative indications of the horizontal rate of rhizome growth beneath the soil surface and were compiled for each treatment to assess the effect of soil organic content and drainage depth on rhizome proliferation. We quantied total production of aerial and subsurface tissues in each tank during the rst growing season in early October 1997. Aerial mass was harvested by cutting all plants at the soil surface and separating live from dead leaves as well as stems from leaves. Prior to harvesting living leaves, we measured their width and total length. These data were used to derive a model to predict the mass of standing-live leaves based on width/length measurments alone (rationale discussed below). Harvested tissues were washed free of supercial sediment, oven dried at 90 C to constant mass, and weighed. Subsurface biomass was estimated by taking duplicate soil cores to a depth of 30 cm using a sharpened 8.3-cm internal-diameter stainless steel cylinder. Coring was done midway between adjacent original plants, thus including only roots and rhizomes that had extended out a minimum distance 9.85 cm from each seedling. Each core was partitioned horizontally into six, 5-cm thick slabs using a hacksaw. The subsurface tissues in each were washed, separated into live vs. dead root and root vs. rhizome categories, oven dried, and weighed. Leaf biomass prediction modelOur rationale for developing a model to predict leaf biomass without actually cutting a plant was driven by the fact that, at this latitude, many plants overwinter in the standing-green condition and resume growth the following spring. We recognize, therefore, that yearly, destructive harvests would have an impact

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upon the production not only of aerial, but possibly also subsurface biomass in subsequent growing seasons. Our ultimate objective, pending verication of our predictive model, was to assess leaf biomass at the end of year 2 through the termination of the project by coupling standard harvests of dead and dying leaves with predictions of standinglive leaf mass in each tank without cutting them. Procedures used to develop and test this model are discussed below. Preliminary observations indicated that the bases of cordgrass leaves tightly ensheathed the stem and overlapped those of younger leaves produced successively closer to the plants longitudinal axis (centripetal leaf origin). We refer to these basal, overlapping leaf segments hereafter as leaf sheaths (LS). The blade (BL), however, represented the majority of the total length of each leaf and diverged from the stem such that both upper and lower epidermal surfaces were freely exposed to the atmosphere. The BLs of all leaves were uniformly dark green, but LS regions contained progressively less chlorophyll as a function of how deeply they were buried by the series of older leaves to their exterior. We inferred from this that BLs represented the most photosynthetically signicant aerial tissue and should serve as the focal point for our model. Observations of a large number of leaves revealed that BLs closely conformed to the shape of a gently tapering triangle (lanceolate) of uniform thickenss. We, therefore, hypothesized that a model could be developed to predict the total area of photosynthetically signicant leaf surface using basal width of each BL (determined at the point where the LS transitioned into the BL) and the BL length (from basal-width measurement point to the BL apex) alone. To test this, we cut the BLs from a number of living cordgrass leaves of various sizes from three natural salt marshes in the area. Each (now truncated lanceloate in shape) was measured to determine basal width and total length. These were then attened and photocopied before drying could cause any distortion. The actual area of each photocopied BL was determined by scanning the surface three times with a digital planimeter (Lasico Inc., Los Angeles, CA; Model 42P) and calculating the mean value. Upon closer examination we noted that BLs change shape slightly along their entire length as they mature. Margins of young leaves exhibit a smoothly convex taper from base to apex, older ones taper linearly, and fully mature leaves are slightly concave. We adjusted our model to accommodate this morphological variation by including an exponential factor that alters the equation as a function of leaf length (i.e., age). Our derived equation to predict area thus was predicted area PA Qe (0.098460.01681Q)

TABLE 1. Predicted biomass of sample cohorts of cordgrass leaves (arranged by size class) for determining accuracy of biomass prediction model.
Leaf length size No. of leaves class (cm) sampled Total biomass (g) Actual Predicted Relative error (%)a

015 1530 3045 4560 Total

a

19 149 118 19 305 actual

0.605 18.509 29.019 7.545 55.6738 predicted actual

0.599 19.080 28.284 6.730 54.693 100.

1.0 3.1 2.5 10.8 1.7

Relative error

yearly sampling to document changes in the infaunal community through time that might correlate with plant growth. We collected duplicate soil cores for analysis from comparable locations in each tank on 19 September 1997 after rst draining all tanks to a depth of 20 cm. Our coring tube was a 23.7-mm internal diameter plastic pipe, which was pushed vertically into the sediment to a depth of 84.2 mm. Recovered soil samples were xed in 10% formalin containing rose bengal stain and infauna extracted by ltration through a 75- m mesh sieve. Samples then were preserved in 70% ethanol and refrigerated until counts could be performed. Statistical analysesRaw data for each monitored parameter (discussed above) were grouped by treatment (N 3 tanks per treatment) and analyzed by two-way ANOVA (SAS Institute, Research Triangle Park, North Carolina) where soil organic content and drainage depth of the treatment were main effects.

RESULTS The accuracy of our live-leaf-BL-biomass prediction model is summarized in Table 1 wherein actual masses of leaf BL cohorts collected from area marshes are compared to predicted values within the same size class. Results indicate that the model predicted biomass within an average of 1.7% of actual mass for leaves with BLs up to 45 cm in length. Longer leaves were subject to greater predictive errors (10.8% for BLs between 45 and 60 cm length). In terms of predicting the mass of standing-live mesocosm leaf BLs (Table 2), our model proved accurate within 6.2% for the 5-cm drain/sand, 4.8% for 5-cm/sand peat, and 5.0% for 15-cm/sand treatment. The mean prediction error for the 15-cm/sand peat treatment was articially high (10.3%) due to a 25.2% error for one replicate tank that was measured by a student assistant. The remaining two replicates of this treatment were measured by DEP (who measured 11 of the 12 tanks) and were 2.0 and 3.7%, respectively. All seedlings outplanted into each of the 12 mesocosm tanks survived transplantation and continued growing throughout the rst season. Aerial tissue biomass accumulations determined during the October harvest are summarized in Table 2. Two-way ANOVA indicated that drainage depth had no signicant effect on accumulation of stem, live (actual mass), or dead leaf tissue. Treatments containing peat moss (2.5% soil organic content w/w), however, supported signicantly more stem (Pr F 0.0518, 1 df) and dead leaf mass (Pr F 0.0494, 1 df), but were not signicantly different from sand tanks (0.15% soil organic content) for live-leaf biomass.

where Q 0.667wL, w blade basal width, and L blade length base to tip. We tested the accuracy of our model by comparing the predicted area of a cohort of randomly collected BLs to their known areas as determined by planimetry. Regression analysis of this comparison yielded a coefcient of determination of 0.977. Relating BL area to biomass involved determining the average density per unit area of randomly collected cordgrass leaves and using these data to derive the following mass-prediction equation: Predicted mass PAe (
5.0110 0.04147 PA)

As with the area equation, we tested our mass model by comparing the predicted mass of 305 BLs collected from local salt marshes to their actual, oven-dry masses (Table 1). The application of this model to mesocosm leaves involved measuring BLs of all green leaves in each tank while they were still attached to stems then cutting them for determinations of actual dry mass. Invertebrate recruitmentPosey, Alphin, and Powell (1997) have shown that vegetational changes in created marsh soils are reected in recruitment patterns of small invertebrates that live within the soil matrix (infauna). Therefore, we thought it would be useful to conduct

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TABLE 2. Effects of treatment on production of aerial biomass.

Mean biomass per tank (g)a Live leaves Treatmentc Stemsb Dead leaves Actual Predictedd

5-cm 5-cm 15-cm 15-cm

a

drain/sand drain/sand drain/sand drain/sand

peat peat

0.488 1.544 1.015 1.960

(0.166) (0.573) (0.698) (1.208)

1.051 2.355 1.635 2.557

(0.398) (0.960) (0.238) (1.280)

0.430 1.310 0.952 2.007

(0.190) (0.560) (0.365) (1.932)

0.421 1.242 0.975 1.637

SD in parentheses. b Live dead. c N 3 tanks per treatment. d Based on mathematical model.

Soil coring to assess subsurface plant growth (Table 3) showed that limited root tissue extended outward farther than 9.85 cm from the original seedlings and that none had penetrated deeper than 20 cm below the surface. It is interesting to note that there were no dead roots at any soil depth in any of the 24 cores. Only one core contained a rhizome fragment (sand substrate only, 15-cm drainage depth; fragment contained in core slab that extended from 5 to 10 cm soil depth). Statistical analysis revealed no signicant effect of soil drainage depth or substrate organic content either on total root biomass accumulation, on the maximum soil depth to which roots penetrated, or on root preference for a particular depth stratum within the 30 cm long core prole. Table 4 summarizes compiled rhizome lengths as affected by treatment. Statistical analysis indicated that peat-containing tanks supported a highly signicant inF 0.0182, 1 df), crease in total rhizome length (Pr but that soil drainage depth had no effect. In addition, we noted that peat-containing tanks produced a total of 126 new shoots from rhizomes compared to only 58 for sand tanks. We inferred that a new shoot arose from a rhizome if it was at least 4 cm from its seedling of presumed origin. Results of invertebrate infaunal counts are summarized in Table 5. We noted a substantial predominance of oligochaetes in all cores regardless of treatment, but found the variability for all taxa between replicate cores as well as between replicate tanks to be too great to warrant statistical comparisons. This spatial variability appeared consistent with that of natural samples analyzed by Posey, Alphin, and Powell (1997). DISCUSSION Our leaf biomass equation predicted the mass of standing-live leaf BLs within 6.6% of the actual value for all 12). We consider this mesocosm tanks (Table 2, N
TABLE 3. Mean root mass contained in duplicate soil cores.

quite acceptable, particularly in light of its including the 25% sampling error associated with one of the 15 cm/ sand peat tanks (discussed above). It should be noted that this 6.6% error, although acceptable, was considerably higher than the 1.7% error for sample BLs used to derive the model (Table 1). We attribute this difference to the fact that it was more difcult to measure mesocosm leaves while they were still attached to parent plants and to the smaller size of many mesocosm BLs compared to those used to develop the model. Realistic application of our biomass model to mesocosm BLs in future years will necessitate measuring them in their standing-live condition. This requirement coupled with the fact that inaccurate measurements of small leaves created proportionally large estimation errors explains why mesocosm estimates were less accurate overall than the sample cohorts. Most long-term eld experiments require periodic, destructive harvests of aerial plant tissues to quantify production. These involve cutting live as well as dead plant tissue at the soil surface for drying and weighing (e.g., Broome, Seneca, and Woodhouse, 1986). This is a justiable practice without measurable, long-term consequences when an experimental plot is sufciently large to permit sampling in areas not previously impacted. Our mesocosms, however, presented only 1.25 m2 of actual marsh surface per tank; thus the impact of removing standing-live leaves, which could overwinter and resume growth the following year, would increase as plants grow larger and denser. Although validation of our model necessitated a destructive harvest of aerial tissue at the end of year 1, we conclude that the accuracy of our BL modeling will obviate this in subsequent years. We should note that our harvest of aerial mass included stems as well as leaves. We do not, at this time, have an adequate way of quantifying total production either of LS tissue or the standing-live stem mass that it surrounds

Mean root mass (mg) per core in indicated sampling incrementa Treatmentb 05 cm 510 cm 1015 cm 1520 cm

5-cm 5-cm 15-cm 15-cm

a

drain/sand drain/sand drain/sand drain/sand

peat peat

0 42.9 (42.8) 1.47 (2.54) 1.57 (2.14)

0.32 31.0 2.07 6.10

(0.46) (53.2) (1.89) (5.82)

0.28 1.17 0.77 1.67

(0.49) (1.85) (0.75) (1.60)

0.08 0.37 0.40 0.30

(0.01) (0.64) (0.69) (0.44)

Cores cut horizontally into 5 cm thick increments. SD in parentheses. b N 3 tanks per treatment.

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TABLE 4. Effect of treatment on rhizome growth rate as evidenced by linear distance between newly emergent shoots and closest original seedling.
Mean rhizome length (cm)a per tank per observation date Treatment
b

16 Jul

30 Jul

27 Aug

13 Sep

5-cm 5-cm 15-cm 15-cm

a

drain/sand drain/sand drain/sand drain/sand

peat peat

9.7 10.0 1.7 21.7

(10.6) (10.0) (2.9) (14.3)

40.0 60.7 29.0 68.0

(21.1) (27.6) (20.7) (50.5)

53.7 126.3 84.7 197.0

(28.9) (43.1) (28.9) (125.4)

53.7 151.7 90.7 235.3

(28.9) (38.3) (22.0) (131.9)

SD in parentheses. b N 3 tanks per treatment.

without destructive harvest, but propose to develop a model to predict these during the second year of this project. Even though LS tissue and stems are much less signicant than BLs as sources of photosynthate (because of shading that results from the overlapping growth habit described above), they represent signicant carbohydrate sinks that could not be quantied except by destructive harvest, which would also kill standing-live BLs they subtend. All 108 cordgrass seedlings outplanted into our mesocosm tanks became successfully established regardless of treatment. Prior to outplanting, these seedlings were maintained in a greenhouse and watered only with tap water. Their surviving the osmotic shock associated with instantaneous exposure to full-strength seawater indicates a hardiness that underscores the value of this species for marsh restoration projects (Woodhouse, 1979). Statistical comparisons of aerial biomass as a function of treatment indicated no signicant effect of soil drainage depth on accumulation of leaves (live or dead) or stem tissue. Padgett, Rogerson, and Hackney (1998) also studied effects of soil drainage on cordgrass production but noted a direct correlation with aerial tissue accumulation during the rst growing season after transplantation. By the end of the second season, however, they found no differences regardless of soil drainage depth. It may be that their rst-year results differed from the present study due to use of buckets that were much smaller than our mesocosm tanks and outplanting of more mature plants directly from a natural marsh rather than greenhouse-grown seedlings. Mendelssohn and Seneca (1980) similarly compared effects of soil drainage on cordgrass growth in eld and greenhouse experiments and noted exactly opposite effects on aerial mass accumulation. They found that greater soil saturation reduced overall plant height, biomass, and stand density in the eld but caused increases in the smaller scale greenhouse trials. These perplexing differences coupled with those cited above suggest that the overall scale of nature-simulation experiments likely has a pronounced effect on their correlation with plant growth in natural marshes. Unlike the effect of drainage depth, we noted a statis-

tically signicant increase in stem and dead leaf mass in treatments that contained peat moss. Conversely, there was no signicant difference in living-leaf mass when peat tanks were compared to sand. Increased total aerial tissue production (stem and leaf masses combined) could result from greater adsorption of soil nutrients onto the smaller particles present in organic soils as compared to sand (Lindau and Hossner, 1981; Craft, Seneca, and Broome, 1991). It seems counterintuitive, however, that increased organics would stimulate overall plant production yet increase leaf mortality. We can neither explain this nor nd other literature that indicates why increased organic content resulted in more dead leaf mass compared to sand treatments. In general, we conclude that increased production of aerial tissue in peat-containing soils suggests that soil organic content could be a significant factor in early establishment and aerial growth of cordgrass in created salt marshes. Results of soil coring (Table 3) revealed that neither soil drainage nor organic content had a signicant effect on accumulation of subsurface tissue or on its vertical distribution. Our drainage depth effects are in agreement with ndings of Padgett, Rogerson, and Hackney (1998). It was interesting to note that there was very little tissue in any of our soil cores, and only one contained a segment of rhizome. Furthermore, we found that all roots and rhizomes were alive. Judging from the steady increase in size of stems and leaves over the growing season, we infer that there was corresponding production of new root mass. The fact that our cores were taken midway between adjacent plants means that only roots and rhizomes extending out (horizontally) farther than 9.85 cm from a seedling were included in the samples. This, in turn, suggests that the majority of root tissue must have been clumped within 9.85 cm of its associated seedling. We conrmed this by carefully excavating (then replanting) several seedlings and found the great majority of root mass within a 5-cm radius. It is well documented that cordgrass roots are fundamentally important in marsh soil stabilization (see review by Matthews and Minello, 1994). Accordingly, the limited horizontal root proliferation noted herein might sug-

TABLE 5. Invertebrate infauna recruited into mesocosm sediments from ooding water.
Individuals present (SD) Treatment Oligochaete Copepod Nematode Polychaete

5-cm 5-cm 15-cm 15-cm

drain/sand drain/sand drain/sand drain/sand

peat peat

212.7 107.7 76.7 145.0

(71.2) (32.8) (76.4) (86.4)

0 4.0 (1.8) 14.5 (16.4) 2.0 (3.5)

25.0 20.5 18.8 3.3

(18.5) (32.9) (31.8) (2.0)

2.8 0.8 2.0 1.5

(1.8) (1.0) (3.5) (1.8)

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gest that rapid soil stabilization in created marshes would mandate planting propagules very close together regardless of soil organic content. Indeed this would be desirable, but Woodhouse (1979) points out that close plant spacing adds dramatically both to labor and cost. Broome, Seneca, and Woodhouse (1986) experimented with cordgrass spacing intervals of 45, 60 and 90 cm in a coastal North Carolina project and found 60 cm to be the best compromise between cost efciency and soil stabilization. It seems likely that every marsh creation project would attempt to achieve a balance between cost efciency and rapid soil stabilization. In this connection, Broome, Seneca, and Woodhouse (1986) noted the importance of maximizing rhizome proliferation as an alternative to close propagule spacing. They implied that rhizome proliferation from original transplants can be an efcient means of vegetating barren soil between transplants without adding to cost. The present experiments showed a highly signicant increase in linear rhizome growth when the sand substrate was amended with peat moss (Table 5, compare 13 September data). Drainage depth, on the other hand, had no effect. Only one of our 24 soil cores contained even a single rhizome fragment regardless of substrate organic content. This could suggest that tanks containing peat produced longer, but still few rhizomes per original seedling. If rhizomes are important in soil stabilization, we suggest that the true measure of their worth is not their length, but rather the total number of aerial shoots (each with a subtending root network) they produce. We infer, therefore, that the peat-containing tanks encouraged greater soil stabilization in that their rhizomes generated 126 aerial shoots compared to a total of 58 shoots in sand tanks. The microinvertebrate fauna recruited into mesocosm tanks exhibited a great deal of spatial heterogeneity similar to that reported by Posey, Alphin, and Powell (1997) in a North Carolina created salt marsh. We, therefore, feel that these data, whose value will become apparent as a baseline for evaluating subsequent annual samples, should be viewed as preliminary. We note simply that oligochaetes were vastly predominant in all treatments and that the ranked order of predominance for all treatnematodes copepods ments was oligochaetes polychaetes. Our major inference from the present study is that soil organic content is fundamentally important to early establishment and proliferation of smooth cordgrass seedlings and may well determine the ultimate success of a

created marsh. Accordingly, we strongly recommend that regulatory agencies consider enacting rules mandating use of ll material with appropriate organic content for all salt marsh creation projects. Although this view might be considered unduely burdensome to developers, the likelihood that it may enhance chances for a successful project could save money in the long run. LITERATURE CITED
BOESCH, D. F., AND R. E. TURNER. 1984. Dependence of sheries species on salt marshes: the role of food and refuge. Estuaries 7: 460 468. BROOME, S. W., E. D. SENECA, AND W. W. WOODHOUSE, JR. 1986. Longterm growth and development of transplants of the salt-marsh grass Spartina alterniora. Estuaries 9: 6374. , , AND . 1988. Tidal salt marsh restoration. Aquatic Botany 32: 122. CRAFT, C. B., E. D. SENECA, AND S. W. BROOME. 1991. Porewater chemistry of natural and created marsh soils. Journal of Experimental Marine Biology and Ecology 152: 187200. GOOD, R. E., N. F. GOOD, AND B. R. FRASCO. 1982. A review of primary production and decomposition dynamics of the belowground component. In V. S. Kennedy [ed.], Estuarine comparisons, 139158. Academic Press, New York, NY. LINDAU, C. W., AND L. R. HOSSNER. 1981. Substrate characterization of an experimental marsh and three natural marshes. Soil Science Society of America Journal 45: 11711176. MATTHEWS, G. A., AND T. J. MINELLO. 1994. Technology and success in restoration, creation, and enhancement of Spartina alterniora marshes in the United States, vol. 1, Executive summary and annotated bibliography. NOAA Coastal Ocean Program Decision Analysis Series, Number 2. NOAA Coastal Ocean Ofce, Silver Spring, MD. MENDELSSOHN, I. A., AND E. D. SENECA. 1980. The inuence of soil drainage on the growth of salt marsh cordgrass Spartina alterniora in North Carolina. Estuarine and Coastal Marine Science 11: 27 40. MOY, L. D., AND L. A. LEVIN. 1991. Are Spartina marshes a replaceable resource? A functional approach to evaluation of marsh creation efforts. Estuaries 14: 116. ODUM, E. P., AND A. A. DE LA CRUZ. 1967. Particulate organic detritus in a Georgia salt marsh-estuarine ecosystem. In G. A. Lauff [ed.], Estuaries, 383388. American Association for the Advancement of Science Publication Number 83. Washington, DC. PADGETT, D. E., C. B. ROGERSON, AND C. T. HACKNEY. 1998. Effects of soil drainage on vertical distribution of subsurface tissues in the salt marsh macrophyte Spartina alterniora Lois. Wetlands 18: 35 41. POSEY, M. H., T. D. ALPHIN, AND C. M. POWELL. 1997. Plant and infaunal communities associated with a created marsh. Estuaries 20: 4247. TEAL, J. M. 1962. Energy ow in the salt marsh ecosystem of Georgia. Ecology 43: 614624. WOODHOUSE, W. W. 1979. Building salt marshes along the coasts of the continental United States. U.S. Army, Corps of Engineers Coastal Engineering Research Center Special Report Number 4. U.S. Army, Corps of Engineers, Fort Belvoir, VA.