WETLANDS, Vol. 25, No. 2, June 2005, pp. 279–288 q 2005, The Society of Wetland Scientists NEW ENGLAND SALT MARSH POOLS: A QUANTITATIVE ANALYSIS OF GEOMORPHIC AND GEOGRAPHIC FEATURES Susan C. Adamowicz1,3 and Charles T. Roman2,4 1 Graduate School of Oceanography University of Rhode Island South Ferry Road Narragansett, Rhode Island, USA 02882 2 3 USGS Patuxent Wildlife Research Center University of Rhode Island Narragansett, Rhode Island, USA 02882 Present address and corresponding author: U.S. Fish and Wildlife Service 321 Port Road Wells, Maine, USA 04090 E-mail: [email protected] 4 Present address: National Park Service University of Rhode Island Narragansett, Rhode Island, USA 02882 Abstract: New England salt marsh pools provide important wildlife habitat and are the object of on-going salt marsh restoration projects; however, they have not been quantified in terms of their basic geomorphic and geographic traits. An examination of 32 ditched and unditched salt marshes from the Connecticut shore of Long Island Sound to southern Maine, USA, revealed that pools from ditched and unditched marshes had similar average sizes of about 200 m2, averaged 29 cm in depth, and were located about 11 m from the nearest tidal flow. Unditched marshes had 3 times the density (13 pools/ha), 2.5 times the pool coverage (83 m pool/km transect), and 4 times the total pool surface area per hectare (913 m2 pool/ha salt marsh) of ditched sites. Linear regression analysis demonstrated that an increasing density of ditches (m ditch/ha salt marsh) was negatively correlated with pool density and total pool surface area per hectare. Creek density was positively correlated with these variables. Thus, it was not the mere presence of drainage channels that were associated with low numbers of pools, but their type (ditch versus creek) and abundance. Tidal range was not correlated with pool density or total pool surface area, while marsh latitude had only a weak relationship to total pool surface area per hectare. Pools should be incorporated into salt marsh restoration planning, and the parameters quantified here may be used as initial design targets. Key Words: marsh pools, ditching, salt marsh, New England INTRODUCTION Able 1994, Erwin 1996, Wolfe 1996) have concluded that pools provide important fish and wildlife habitat, no research has been conducted in New England that quantifies basic geomorphic and geographic pool features. Salt marsh pools are significant because they provide habitat for numerous bird species. In Massachusetts, for example, Clarke et al. (1984) noted that shorebirds, wading species, terns, swallows, and crows were strongly attracted to salt marsh pools. In southern New Jersey, Master (1992) documented that various Salt marsh pools are soft-bottomed depressions that hold water throughout a tidal cycle and do not dry as shallower pannes often do (Figure 1; Chapman 1960). Despite the extensive literature on salt marshes, research specific to marsh pools is quite limited. Harshberger (1916), Miller and Egler (1950), and Redfield (1972) have provided the most extensive descriptions of pools but no quantitative analyses. While several authors (Daiber 1982, Clarke et al. 1984, Smith and 279 280 WETLANDS, Volume 25, No. 2, 2005 Figure 1. Small salt marsh pool in southern Maine (Wells, ME). Pool is 3–4 m in diameter and is located within a Spartina patens-dominated high marsh. shore and wading birds aggregated at pools to feed. Erwin (1996) supported the conclusion that salt marsh pools are vital year-round habitat for dabbling ducks, shorebirds, and wading birds. Pools are valuable to birds because they provide food items including fish and macrophytes, specifically Ruppia maritima L. (Daiber 1982). Historically, New England salt marshes have been extensively ditched for mosquito control and salt hay farming, resulting in lowered water-table levels and drainage of the marsh surface, often including salt marsh pools (Miller and Egler 1950, Redfield 1972). Salt hay farming has been a fairly common practice since colonial times (Rozsa 1995), but the most extensive and widespread ditching occurred in the early-tomid 1900s for mosquito control (Daiber 1986). Bourn and Cottam (1950) stated that, by 1938, approximately 90% of coastal marshes from Maine to Virginia had been ditched in an effort to reduce breeding habitat for the salt marsh mosquito (Ochlerotatus sollicitans [Walker]). In New Jersey and Rhode Island, it is reported that pools were specifically targeted for drainage (Cottam 1938, Price 1938). Given their ecological benefits, restoring pool habitats should be an important salt marsh restoration objective. To enhance the success of such work, welldocumented guidelines for pool creation are necessary. The objective of this paper is to begin addressing this need by providing a quantitative baseline analysis of geomorphic (e.g., pool depth and size) and geographic (e.g., density, spatial arrangement, and coverage) attributes of salt marsh pools for ditched and unditched salt marshes located throughout New England. By studying a gradient of marshes from unditched to intensely ditched, this study also evaluates the relationship between ditch density and geographic factors such as pool density and pool surface area. METHODS Salt marsh pools were assayed using both field investigations and aerial photographic surveys coupled with geographic information system (GIS) analysis. The complete effort extended from the Long Island Sound shoreline of Connecticut to mid-coast Maine, USA. A total of 32 sites were surveyed by either or both methods (Table 1). Several criteria were used to select marsh study areas. All sites were broad as opposed to fringing systems (e.g., average width over 150m). Impounded, filled, and tide-restricted marsh sites were not sampled in order to minimize variability. All surveyed locations were in public ownership. Unditched sites were very uncommon; only eight such locations were included in this study, mostly in Massachusetts and Maine. Some marshes had discrete areas of ditched and unditched marsh that were considered separate study areas for the purposes of this work. Adamowicz & Roman, NEW ENGLAND SALT MARSH POOLS 281 Table 1. Location of ditched and unditched marsh study sites, summary of survey method employed at each site, and determination of pool density (number of pools/km of transect). Latitude Longitude Survey Method1 Pool Density (no. of pools/km) DITCHED MARSHES Hammonasett State Park, Point, Clinton, Ct Hammonasett State Park, Rotary, Clinton, CT Grass Island 1, Guilford, CT Grass Island 2, Guilford, CT S. B. McKinney NWR, Hammonck Dock, West Site, CT S. B. McKinney NWR, Hammonck Dock, East Site, CT Great Island, Old Lyme, CT Unnamed Island, Old Lyme, CT Watts Island, Niantic, CT Barn Island Management Area (1), Stonington, CT Barn Island Management Area (3&4), Stonington, CT Sachuest Point NWR, Middletown, RI Sapowet Wildlife Management Area, Tiverton, RI Coggeshall Marsh, NBNERR, Portsmouth, RI 100 Acre Cove, Barrington, RI Felix Neck, Martha’s Vineyard, MA Water Street, Essex, MA Annisquam River Marshes, Gloucester, MA Corn Island, Essex, MA Moody Marsh, Wells, ME Wells NERR, North Area, Wells, ME Granite Point, Biddeford, ME Morse River, Phippsburg, ME Squirrel Point, Arrowsic, ME Average Ditched Marshes (SE) N41815.3859 N41815.3859 N41816.1009 N41816.3639 N41816.7309 N41816.7909 N41817.2139 N41817.1159 N41818.0259 N41820.3309 N41820.2499 N41828.6839 N41835.0129 N41839.0469 N41845.9859 N41824.9679 N42837.9529 N42837.9689 N42839.4359 N43815.8849 N43820.4109 N43824.9809 N43845.1309 N43849.4109 W72832.3979 W72832.3979 W72839.3269 W72839.4439 W72828.5709 W72828.2909 W72819.8109 W72819.3159 W72813.1939 W71852.3909 W71852.0139 W71814.6229 W71812.3199 W71820.6359 W71818.2449 W70833.5619 W70846.2519 W70841.0739 W70844.8029 W70835.5239 W70832.3409 W70823.3869 W69849.4709 W69847.4509 F/D F/D F/D/C D/C D D D D D F/D/C F/D/C F/GPS F/GPS F/GPS F/D D D D D F/C C F/AP D D 1.7 0.2 0.0 0.6 0.0 1.9 1.1 17.6 2.1 3.8 1.8 1.8 7.1 1.8 7.4 4.0 0.0 6.2 11.3 19.8 10.7 18.7 5.5 0.4 5 (6) UNDITCHED MARSHES S. B. McKinney NWR, Bridgeport, Ct Nauset Marsh, Eastham & Orleans, MA Annisquam River Marshes, Gloucester, MA Parker River NWR, Newbury, MA Wells NERR, Harbor Area, Wells, ME Scarborough Marsh, Scarborough, ME Spurwink River, Pleasant Hill, ME Reid State Park, Georgetown, ME Average Unditched Marshes (SE) N41809.3769 N41849.3489 N42838.6639 N42843.3239 N43819.0509 N43833.6939 N43835.2649 N43846.7279 W73808.2979 W69957.4709 W70841.0379 W70847.6569 W70934.2909 W70819.8289 W70815.7679 W69844.2679 D F/D D F/D C F F F/D 4.8 21.3 14.6 23.1 7.5 13.8 31.3 24.3 18 (9) Marsh Type and Location 1 Survey methods included georectified true color aerial photographs (AP), color infrared images (C), digital orthophotographic images (D), field survey (F), and field-generated GPS maps (GPS). Field Survey Field surveys were conducted at 16 salt marshes throughout New England during the 1999 and 2000 growing seasons (Table 1). At each salt marsh, a study area was identified as a discrete region bounded by major creeks, roads, upland, or similar features. Selected study areas did not include tidal channels greater than 10 m wide, while all natural channels less than 10 m wide (‘‘creeks’’) and all ditches were included. Large channels were excluded because they were representative of open water or aquatic habitat (Dame et al. 1992, Heck et al. 1995, Bertness 1999). Within each study area, line transects were used to identify sample pools for a number of measures as described below. The initial transect was randomly located along the main tidal channel or shoreline, with subsequent transects set at 100-m intervals. Each transect extended perpendicularly from the edge of the marsh to the upland border and was measured using a metric survey tape. Only pools in direct contact with transect lines were included in the field study. The number of pools located on the transects was used to calculate a measure of pool density (number of pools/ km transect). Adapting the standard line-intercept 282 method used in vegetation ecology (Mueller-Dombois and Ellenberg 1974), pool coverage was defined as the linear distance of pool falling directly on the transect line (m of pool/km transect). The total transect distance sampled ranged from 293 m at the Sapowet Marsh (RI) study area to 2,177 m at Hammonasett State Park (CT). For each individual pool encountered along the transects, pool surface area was estimated by measuring the major and minor axes of regularly shaped pools (Ingolfsson 1994) and applying the formula for the area of an ellipse. Irregular pools were subdivided into simple geometric shapes, with appropriate data taken to calculate surface area. For each pool, geographic parameters such as distance to nearest neighboring pool and distance to nearest tidal flow (either a ditch or creek) were also measured. Pool depth was based on an average of at least three measurements of the distance from the surface of the surrounding marsh surface to the pool bottom. Water depth (measured as surface of water to pool bottom) was recorded for use in comparison with other studies, but it is less consistent than pool depth since the depth of water alone is variable depending on tides, rainfall, and other factors. To categorize qualitatively the location of pools along a gradient from the frequently flooded low marsh zone to the less frequently flooded high marsh, the vegetation type adjacent to each pool was recorded based on visual inspection. Three such types were distinguished: one dominated by Spartina alterniflora Loisel (tall form), one dominated by S. patens, and a third dominated by S. alterniflora of intermediate height. Spartina alterniflora (tall form) characterized more frequently flooded (low) marsh areas, while the other vegetation types characterized less frequently flooded (high) marsh regions Aerial Photography and Geographic Information System Analysis Digital orthophotographic quadrangle (DOQ) images were used to increase the number of marshes studied. Black and white DOQ images (Multi-resolution Seamless Image Database ‘‘MrSID’’ compression) were downloaded from World Wide Web sites for Connecticut (0.5 m per pixel resolution) and Massachusetts (1.0 m per pixel resolution). DOQs at the same resolution were obtained from the University of Rhode Island Environmental Data Center and the United States Fish and Wildlife Service Gulf of Maine Program for Rhode Island and Maine, respectively. In addition, color infrared (CIR) images were available for a limited number of sites in Connecticut (1:12,000, August 1995) and Maine (Wells NERR 1:12,000, August 1998; Moody Marsh 1:8,000, June 1992). A true WETLANDS, Volume 25, No. 2, 2005 color photograph was available for Granite Point Marsh in Maine (1:8000, May 1986). A total of 21 ditched and 6 unditched marshes were surveyed with ArcView 3.2 geographic information system (GIS) software applied to the DOQs and CIR photographs. Site boundaries were delineated in the same manner as in the field surveys. Transects were established on the imagery following methods used in the field surveys, and density, distance measures, and coverage (m of pool/km of transect) were then calculated. For each individual pool encountered along the transects, surface area was calculated from digitized ArcView shape files. Additionally, within the study boundary for each site, all pools, creeks, and ditches were delineated with ArcView shape files to calculate pool density and total pool surface area per hectare of marsh study area. Comparison of Survey Methods The two survey methods (field survey transects and DOQ transects) were compared using paired t-tests for each pool variable (pool density, pool coverage, nearest neighbor distance, pool size, distance to tidal flow) to determine if the two data sets for transects could be combined. A total of nine sites (6 ditched, 3 unditched) were used in this analysis, but because some surveys (variously among the field and DOQ methods) did not encounter pools, sample size was either 7 or 9 sites for the 5 parameters of concern. It should be noted that, while the same study sites were used in each survey method, initial transects were given independent, random starts with the result that each survey provided a statistical representation of that site. Statistical Analyses Based on results from the method comparison noted above, pool densities (number of pools/km transect) for field and digital orthophotographic quadrangle (DOQ) transect studies were combined to test for differences attributable to marsh type (ditch vs. unditched) using a two-sample t-test. Relying on field data alone, ANOVA was used to determine if the proportion of the total number of pools differed among the three marsh zones [S. alterniflora (tall), S. patens, S. alterniflora (intermediate)] or between ditched and unditched marsh sites. Logistic regression was applied to determine the environmental variable(s) (distance to nearest neighboring pool, distance to nearest tidal flow, pool surface area, log of pool surface area, and pool depth) that best predicted pool location (SAS 1985). The effect of marsh type on a series of pool variables measured from DOQs was determined by two-sample t-test. While it was instructive to examine ditched versus Adamowicz & Roman, NEW ENGLAND SALT MARSH POOLS 283 Table 2. Average values (6 SE) for geographic measures of pools in ditched and unditched New England salt marshes based on digital orthophotographic images. Data generated from transects placed on the imagery and digitizing techniques. Geomorphic Measures Transect Data Pool Cover/Transect (m/km) Individual Pool Size (m2) Distance to Nearest Neighbor Pool (m) Distance to Tidal Flow (m) Area Data Density (number of pools/ha) Total Pool Surface Area (m2/ha) Ditch Length (m/ha) Creek Length (m/ha) Ditched Marshes Unditched Marshes t-value df p 31 6 37 227 6 305 83 6 47 205 6 113 22.85 20.47 24 21 ,0.01 0.32 15 6 14 12 6 15 966 10 6 5 1.07 0.65 21 21 0.15 0.26 23.12 22.78 9.70 23.52 24 24 19 6 ,0.001 ,0.01 ,0.0001 ,0.01 4 229 217 38 6 6 6 6 5 242 100 44 unditched marshes, some marshes have been ditched more intensively than others. Similarly, ditched and unditched marshes also varied in the amount of natural creek length that each possessed. Because ditches and creeks both convey water to and from the marsh, these parameters may be correlated with pool density and the amount of pool surface area per hectare of marsh. Therefore, separate regression analyses were used for ditches and creeks to determine the relationship between these factors and pool density and total pool surface area per hectare on data obtained from DOQ sources. In addition, tidal range and latitude were each regressed against pool density and total pool surface area. Paired Study In order to examine the effect of ditching on salt marsh pools more closely and to control for potential effects of tidal range and latitude, ten total areas (five paired sites) in five marsh systems were selected from the DOQ data set in large marshes where a ditched section was adjacent to an unditched portion of marsh but separated by a hydrologic feature (e.g., tidal creek) sufficient to consider the areas to be independent. These pairings included sites from Old Lyme, CT; Newbury, MA; Gloucester, MA; Essex, MA; and Wells, ME. The same procedures were followed for delineating study areas, ditches, creeks, and pools as previously noted. Linear regressions and paired t-tests were used to determine the effect of ditching on pool density (number of pools/ha salt marsh) and total pool surface area per hectare salt marsh (m2 pool/ha salt marsh). RESULTS Pool Density, Size, and Other Geographic Measures Transect data from field and DOQ evaluations showed that pool density in unditched marshes was 13 913 0 171 6 6 6 6 7 489 0 89 more than three times that of ditched marshes (Table 1, t29 5 24.25, p,0.0001). Pool density ranged from 0 to 19.8 pools/km of transect in ditched sites and from 4.8 to 31.3 pools/km of transect in unditched marshes. When calculated on an area basis from DOQ evaluations, pool density (number of pools/ha salt marsh) again was three times greater in unditched marshes compared to ditched marshes (Table 2). Other parameters, including total pool surface area per hectare (m2 pool/ha salt marsh), pool coverage (m pool/km transect), and creek length (m creek/ha salt marsh), were all greater in unditched marshes (Table 2). Somewhat unexpectedly, several pool measures were the same between ditched and unditched marshes. Pools in ditched marshes were no further from the nearest neighboring pools or from nearest tidal flow than in unditched marshes. In both ditched and unditched marshes, pool size averaged about 200 m2 (Table 2). It should be noted that 100-Acre Cove, RI was eliminated as an outlier from all but pool density calculations since pools there were up to 35 ha in size— much larger than the next largest pool in the study (0.56 ha, Scarborough Marsh, ME). Influence of Ditching Intensity, Creeks, Tidal Range, and Latitude Along a gradient from unditched to highly ditched marshes, both pool density (number of pools/ha salt marsh) and total pool surface area (m2 pool/ha salt marsh) decreased significantly (Figure 2). The weak coefficients of determination reflect the high variability in pool density and total pool surface area for marshes with no ditches. There was also a negative relationship between ditch length per hectare and creek length per hectare (Figure 3), with the regression equation indicating that about 300 m of ditching per hectare of marsh study area would result in little or no creek hab- 284 WETLANDS, Volume 25, No. 2, 2005 Figure 2. Relationship of pool density and total pool surface area with ditching intensity (ditch length, m/ha). itat, at least from the study sites in this investigation. When pool parameters were regressed against natural creek length per hectare (Figure 4), there was a positive relationship in each case; as creek density increased, so too did pool density and total pool surface area. Mean tidal range (F1,24 5 3.25, p50.08, R250.02) and latitude (F1,24 5 2.04, p50.17, R250.08) were not Figure 3. related to pool density. However, there was a weak relationship between total pool surface area per hectare of marsh and both tidal range (F1,24 5 6.02, p50.02, R250.20) and latitude (F1,24 5 4.69, p50.04, R250.16). While there were no or only weak relationships between tidal range or latitude with pool density and total pool surface area, the fact that the marsh study areas were distributed along a continuum of tidal range Relationship of creek length and pool length per hectare of salt marsh. Adamowicz & Roman, NEW ENGLAND SALT MARSH POOLS Figure 4. 285 Relationship of pool density and total pool surface area with natural creek intensity (creek length, m/ha). and latitude could influence the significant relationship between ditching and pool density and total pool surface area (Figure 2). To eliminate possible confounding influences of tidal range and latitude on pool parameters, five pairs of DOQ salt marsh study areas (1 pair 5 1 ditched site adjacent to an unditched site), located along the continuum, were analyzed. Obviously, the ditched and unditched sites of each pair were located at the same latitude and were influenced by the same tidal regime. Once again, unditched marshes had greater pool density (paired t-test, t4 5 23.15, p 5 0.02) and total pool surface area (paired t-test, t4 5 22.97, p 5 0.02) than ditched sites. Ditched marshes had 4.3 pools per ha 6 2.5 (SD) and 249 m2 6 292 m2 total pool surface area per marsh hectare; unditched marshes had 27 pools per ha 6 20 and 1063 m2 6 270 m2 total pool surface area per marsh hectare. Similarly, pool density and total pool surface area per hectare of salt marsh decreased with increasing total ditch length per hectare of salt marsh (linear regression; n 5 10, R25 0.42, p,0.001; R25 0.78, p,0.05; respectively, Figure 5). Figure 5. Relationship of pool density and total pool surface area with ditching intensity based on paired ditched and unditched marshes. 286 Pool Location within Marshes Pools were unequally distributed among the three marsh zones (ditched and unditched marshes combined), with the greatest proportion of pools consistently found in the high marsh compared to low marsh or intermediate S. alterniflora zones (ANOVA, arcsine square-root transformation of proportion data, F3,38 5 10.89, p,0.0001; least squares means post-hoc test, p,0.0005). Marsh type (ditched vs. unditched) was not a significant factor (ANOVA, F1 5 0.07, p50.79), and there was no significant interaction between marsh type and pool distribution across zones. Distance to tidal flow was the best predictor for determining pool location along a flooding gradient when given the three vegetation types (S. patens, S. alterniflora (intermediate), S. alterniflora (tall); logistic regression, Wald X2 5 11.19, 1df, p50.0008). As would be expected, distance to tidal flow was greater for pools in S. patens areas, 31.4 m 6 33.6 m (SD, n5109), compared to tall S. alterniflora (6.9 m 6 10.3m, n514) or the intermediate S. alterniflora regions (13.9 m 6 14.8 m, n550) (least squares means post-hoc test, p,0.003). When discriminating between just low (S. alterniflora (tall)) and high marsh (S. patens) locations, distance to tidal flow and pool size both were significant variables in predicting pool location (logistic regression; distance to tidal flow, Wald X2 5 6.39, 1df, p50.01; pool size, Wald X2 5 4.92, 1df, p50.03). Pool size was greatest in the high marsh (310 m2 6 817 m2, n5109) compared to the low marsh zone (229 m2 6 794 m2, n514) (logistic regression, Wald X2 5 4.92, 1df, p50.03; least squares means post-hoc test, p,0.05). Pool Depth and Water Depth Pool depth was the same regardless of marsh type (ditched 5 26.0 cm 6 12.7 cm, n598; unditched 5 30.2 cm 6 21.1 cm, n580) or pool location (high marsh 5 27.3 cm 6 15.7 cm, n5109; intermediate S. alterniflora 5 27.9 cm 6 18.0 cm, n550; low marsh 5 31.4 cm 6 23.0 cm, n514; F3,131 5 0.74, p50.53). Similarly, water depth did not differ between marsh types or marsh vegetation zone and averaged 19.1 cm (range 1–65 cm) for ditched marshes and 18.1 cm (range 1–71 cm) for unditched sites (F5,140 5 0.73, p50.60). The shallowest depths were from pools that were breached and drained almost completely at low tide. The maximum water depth (71 cm) and maximum pool depth (92 cm) were both from pools in Nauset Marsh, MA (an unditched marsh). Methodology Comparison Information for this paper was generated from two methods, field survey transects and DOQ transects. WETLANDS, Volume 25, No. 2, 2005 Paired t-tests indicated no significant difference between the methods for pool density (number of pools/ km transect; t8 5 21.07, p50.16), pool coverage (m pool/km; t8 5 20.46, p50.33), nearest neighbor distance (t6 5 1.63, p50.08), or distance to tidal flow (t6 5 0.55, p50.30). However, there was a significant difference in average individual pool size (m2) (log transformed data, t6 5 2.71, p50.02). In general, pools sampled in DOQ transects were larger. This may have been due to the limited resolution of DOQs compared to field surveys or merely due to the fact that DOQ transects were located in slightly different positions compared to those in the field, resulting in a different set of pools actually being sampled. Throughout this paper, field and DOQ transect data were only combined for the comparison of pool density (number of pools/km transect) between ditched and unditched marshes. DISCUSSION New England salt marsh pools are remarkably similar between ditched and unditched sites. Average values for pool size, depth, nearest neighbor distance, and distance to tidal flow are similar (Table 2). Additionally, most pools are located in the high marsh zone of salt marshes. Conversely, a recent quantitative comparison of ditched and unditched salt marshes in New Jersey found marsh pools in an unditched marsh to be larger than those of a ditched marsh, and further, unditched marsh pools had a lower nearest neighbor distance (Lathrop et al. 2000). Except to mention the difference in geographic location, we offer no explanation for the divergence of our findings. While New England pools appear similar between marsh types, ditched marshes have less pool habitat than unditched marshes, a relationship that holds for both pool density and pool coverage (m pool/km transect; Table 2). In fact, whether based on the number of pools per kilometer of transect or the number of pools per hectare of salt marsh, ditched marshes have 70% fewer pools than unditched marshes. Similar to our findings, studies in New Jersey salt marshes found that unditched sites had dramatically more pool habitat than ditched sites (Lathrop et al. 2000, Lathrop and Bognar 2001). Collectively, these findings seem to support Redfield’s (1972) hypothesis that ditching results in decreased pool density. Further, our findings (Figure 3) and those of Lathrop et al. (2000) would support a hypothesis that ditching may influence the density of natural salt marsh creeks. The fact that marshes with an abundance of natural creeks also have a high density of pools (Figure 4), while intensively ditched marshes have few pools (Figure 2) or creeks (Figure 3), suggests that ditches Adamowicz & Roman, NEW ENGLAND SALT MARSH POOLS and creeks drain marshes differently. Further study is necessary to determine the precise hydrologic interactions among creeks, ditches, and pools. Of particular interest is the manner in which ditches, intertidal creeks, and subtidal creeks influence sub-surface marsh drainage. While it is clear that the unditched marsh sites in this study have a greater density of pools and more pool surface area than ditched marshes, there is considerable variation in these relationships pointing to heterogeneity in marsh morphology. As noted, some unditched marshes have many pools, while others have few (Figure 2, Table 2), thereby contributing to habitat variability. With ditching, this important variation in habitat is reduced and marshes are more uniform, at least with regard to pool geography. Heterogeneity of unditched marshes included in this study can be related to several factors. Our marsh study areas extended from Long Island Sound to the Gulf of Maine, an extensive geographic region with considerable variation in climate, tidal range, sediment supply and type, and other factors (Roman et al. 2000). It is likely that processes of pool formation and maintenance vary in response to these factors. When considering all of the unditched and ditched sites, there was a weak relationship between total pool surface area and latitude, as well as tidal range, suggesting that marshes to the north and marshes with a high tidal range (typically also to the northern portion of our study region) have a greater total pool surface area. It is also noted that our study marshes were classified as back-barrier, embayment, or river valley, and it is likely that pool formation and maintenance vary among these types. Classic papers on New England salt marsh geomorphology (Harshberger 1916, Miller and Egler 1950, Redfield 1972) offer hypotheses for pool formation, but additional studies are needed to describe the role of tidal range, climate (particularly ice), classification type, and other factors influencing marsh pool geography. Implications for Restoration The descriptive analysis of pools from unditched New England marshes may provide some design targets as efforts to re-establish pool habitat are considered. On average, 9% (913 m2/ha) of an unditched marsh is composed of pool habitat, with an average density of 13 pools/ha. The range of pool size was broad (3–3,786 m2), but large pools were rare, with only 6% of all pools in the entire DOQ survey being greater than 800 m2. Pool depth, measured as the average distance from marsh surface to pool bottom, was about 30cm, with actual water depth usually less (i.e., pools are often not flooded to the marsh surface). Most 287 pools are found within the high marsh zone. In the absence of site-specific, pre-alteration data to use as restoration targets, managers may want to consider these average pool characteristics, while incorporating variation in pool size, depth, and density into restoration plans. Restoration of marsh pool habitat is an important resource management objective but should proceed with caution. Pools are just one of many different habitat types integrated within the salt marsh mosaic, and all pool restoration plans must carefully consider the overall effects on marsh development processes, vegetation dynamics, fish and wildlife support functions, and other marsh processes and roles. ACKNOWLEDGMENTS This study was supported by the U. S. Geological Survey, administered through the USGS Patuxent Wildlife Research Center, Coastal Field Station at the University of Rhode Island. Brandi Bornt and Susan Wilson (URI Coastal Fellows), Mike Morrison, and interns from the Rachel Carson National Wildlife Refuge provided invaluable assistance with fieldwork. Ron Rozsa (Office of Long Island Sound Programs), Robert Houston, and Rick Schauffler (U. S. Fish and Wildlife Service) provided aerial imagery. Lynn Carlson (Brown University) assisted with geo-rectification of several aerial photographs. LITERATURE CITED Bertness, M. D. 1999. The Ecology of Atlantic Shorelines. Sinauer Associates, Inc., Sunderland, MA, USA. Bourn, W. S. and C. Cottam 1950. Some biological effects of ditching tidewater marshes. Fish and Wildlife Service, U.S. Department of Interior, Washington, DC, USA. Research Report 19. Chapman, V. J. 1960. Salt Marshes and Salt Deserts of the World. The University Press, Aberdeen, UK. Clarke, J., B. A. Harrington, T. 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