Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations

1993 Evaluation of techniques for establishing sedge meadow vegetation Tony Lee Bremholm Iowa State University

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This Thesis is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Evaluation of techniques for establishing

sedge meadow vegetation

by

Tony Lee Bremholm

A Thesis Submitted to the

Graduate Faculty in Partial Fulfillment of the

Requirements for the Degree of

MASTER OF SCIENCE

Department: Botany Interdepartmental Major: Water Resources

Signatures have been redacted for privacy

Iowa State University Ames, Iowa

1993 ii

TABLE OF CONTENTS

Page GENERAL INTRODUCTION ...... 1

PART 1: COMPARISON OF NATURAL AND CREATED SEDGE MEADOWS ...... 3

INTRODUCTION ...... 4 SITE DESCRIPTIONS ...... 6 Natural sedge meadows ...... 6 Site selection ...... 6 Vegetation ...... 6 Soils ...... 7 Created sedge meadows ...... 10 Site description ...... 10 Soils ...... 10 METHODS ...... 13 Water table levels ...... 13 Natural sedge meadows ...... 13 Created sedge meadows ...... 14 Stratigraphy ...... 14 Natural sedge meadows ...... 14 Vegetation sampling ...... 14 Natural sedge meadows ...... 14 Created sedge meadows ...... 15 RESULTS ...... 17 Water table levels ...... 17 Natural sedge meadows ...... 17 Created sedge meadows ...... 17 Stratigraphy ...... 30 Natural sedge meadows ...... 30 Vegetation sampling ...... 30 Natural sedge meadows ...... 30 iii

Created sedge meadows ...... 30

PART II: SEDGE ESTABLISHMENT STUDIES ...... - ...... 42

INTRODUCTION ...... 43 METHODS ...... 45 Seed studies ...... 45 Seed germination ...... 45 Seed storage ...... 46 Growth studies ...... 48 Greenhouse studies ...... 48 Field studies ...... 49 RESULTS ...... 50 Seed studies ...... 50 Seed germination ...... 50 Seed storage ...... 51 Growth studies ...... 57 Greenhouse studies ...... 57 Field studies ...... 59

GENERAL DISCUSSION ...... , ...... 67 Comparison of natural and created sedge meadows ...... 67 Sedge establishment studies ...... 76 Conclusions and recommendations ...... 84

LITERATURE CITED ...... 85

ACKNOWLEDGEMENTS ...... 93

APPENDIX: MAPS SHOWING WELL PLACEMENTS AT THE FIVE NATURAL SEDGE MEADOW SITES ...... 94 1

GENERAL INTRODUCTION

Recent legislation regulating activities impacting on wetlands was designed to slow or reverse trends in wetland acreage lost (Kentula et al., 1992a; Owen and Jacobs, 1992).

However, reliance on compensatory mitigation to offset acreage losses may be degrading the quality and diversity of the nation's remaining wetland resources. Evaluations of created and restored wetlands indicate that our ability to create and restore wetland functions has not

progressed far enough to warrant the present reliance on mitigation to offset wetland losses

(Kusler and Kentula, 1989; Owen and Jacobs, 1992; Zedler and Weller, 1989).

In evaluating of the results of wetland mitigation projects in Florida, Washington and

Oregon, Kentula et al. (1992a) and Kentula et al. (1992b) noted the trend in created and restored freshwater wetlands offered as mitigation seemed to be toward hydrologically

isolated small ponds with steep banks and a ring of vegetation composed of volunteer species

around the shore. The types of wetlands resulting from mitigation were often different from the types of wetlands impacted.

The situation described has generated interest in developing methods to create and restore wetlands other than ponds and cattail marshes. Particularly lacking are methods to create and

restore wet meadows. These are seasonally flooded wetlands, often dominated by sedges

(sedge meadows), that occupy the transition zone between marshes and non-wetland areas

such as wet prairies.

This is a report on research conducted for the wet meadow creation project at the Des

Plaines River Wetland Demonstration Project (DPRWDP) in Lake County, Illinois. The work

was done between May, 1991 and June, 1993. The goal of the research was to evaluate

techniques available for establishing sedges of the genus Carex in created wet meadows. The

format of this report reflects the two main objectives of the research project. Part I is a

comparison of the hydrology, vegetation and soils of natural and created sedge meadows. 2

Part II reports on a series of studies conducted to evaluate sedge establishment techniques using seeds and seedlings. 3

PART I.

COMPARISON OF NATURAL AND CREATED SEDGE MEADOWS 4

INTRODUCTION

Information about the vegetation, soils and hydrology of natural wetlands should be used to guide the planning and design of created or restored wetlands to improve the chances of success (Garbisch, 1989; Kentula et a!., 1992a). Following project completion, the natural wetlands provide a standard of comparison for evaluating the success of the created or restored wetlands (Brown, 1991; Confer and Niering, 1992; D'Avanzo, 1989). The present study collected information about the vegetation, soils and hydrology of naturally occurring sedge meadows in the vicinity of the Des Plaines River Wetlands Demonstration Project

(DPRWDP) site in northeastern Illinois. This was done to guide the design of created sedge meadows constructed in experimental wetlands at the site and to provide baseline information for evaluating the success of the project.

Studies of the soils, hydrology, vegetation and ecology of wetlands containing sedge meadows have been concentrated in just a few areas. One area of study has been the hydrology, nutrient dynamics and restoration of floating fens in The Netherlands (Barendregt et ai., 1992; Verhoeven et a!., 1983; Verhoeven and Arts, 1987; Koerselman, 1989; Konings eta!., 1989; van der Valk and Verhoeven, 1988). Another area of study has been the hydrology and vegetation ecology of sedge meadows of Wick en Fen in Great Britain (Godwin and Bharucha, 1932). The effects of water level disturbance on sedge meadows have been studied in Europe and in the United States (Wilcox et a!., 1985; Sjoberg and Danell, 1983).

In the United States, ecological studies have focused on nutrient cycling and hydrology of peatlands (Heinselman, 1970; Siegal, 1983; Richardson et aI., 1978; Boelter, 1966; Verry and

Boelter). A significant body of work has begun to develop in the area oflife histories, nutrient dynamics and primary productivity of Carex species (Auclair, 1982; Bernard, 1973, 1975,

1990; Bernard and Gorham, 1978; Bernard and MacDonald, 1974; Bernard and Solsky,

1977; Bernard etai.,1988; Gorham, 1974; Grootjans and van Tooren, 1984; Schmid, 1984; 5

Verhoeven et aI., 1988).

Little has been published about the ecology of sedge meadows in northeastern Illinois.

What has been written tends to originate from Wisconsin, where sedge meadows are more abundant than in Illinois. Sedge meadows have, historically, received little attention aside from the forage value of the hay they produced and their productivity when drained (Stout, 1912).

This may be due in part to their ubiquity in the region, their often unremarkable, nearly monotypic floras, and to the difficult conditions encountered in trying to work in them.

Bernard and Gorham (1978) attributed the paucity of published information on sedge meadow ecology to the difficult taxonomy of the genus Carex and to the perception that sedges have little forage value. More recent research in these areas has generally focused on unique or endangered bog and calcareous fen ecosystems (Carpenter, 1990; Moran, 1981), and the information has not been readily available outside of agency reports.

Studies of the plant communities of sedge meadows are more readily available than are hydrologic studies. Ebinger (1977) and Hughes (1977) produced general flora and community descriptions of the Chain Q'Lakes region of Lake County, Illinois. Costello's (1936) analysis oftussock meadows in southeastern Wisconsin provided a thorough description of the flora and community ecology of Carex stricta meadows, but was lacking in quantitative hydrological information. Kelsey and Hootman (1992) monitored water table levels and hydric soils in a wet meadow/ wet prairie site in DuPage County, Illinois. Schennum (1990) emphasized the close association between sedge meadows and wet prairies in northeastern

Illinois.

Part I of this report describes the vegetation, soils and hydrology of natural sedge meadows and of the experimental wetlands at the DPRWDP. This information can be used to evaluate the experimental wetlands at the DPRWDP and to guide future management decisions in created sedge meadows. 6

SITE DESCRIPTIONS

Natural sedge meadolVs

Site selection Five sedge meadows in Lake and McHenry Counties in northeastern

Illinois were selected for study after consultation with personnel from state and county natural resource agencies and using data from the Illinois Natural Areas Inventory. The criteria used for selecting suitable sites were: classification of the site as a sedge meadow by the Illinois

Natural Areas Inventory, proximity to the DPRWDP, ease of access, dominance by tussock sedge (Carex stricta) and quality of the vegetation. Sites with exotic species problems were not included, nor were sites that were likely to be disturbed by visitors. Four of the sites

(Lyons Prairie and Marsh, Spring Grove Fen, Turner Lake Fen, and Wadsworth Prairie) were state nature preserves. The fifth, Des Plaines River trail, is in a Lake County Forest Preserve.

Work at all sites required prior approval and issuance of permits by the agencies involved. The

5 sites and their locations are listed in Table 1.

Vegetation Costello (1936) produced the definitive description of the type of sedge meadows (which he referred to as "tussock meadows") observed in the present study.

Table 1. Names and locations of natural sedge meadows sampled

Site County Location

Des Plaines River Trail Lake T46N RIlE sec 27 Wadsworth Prairiea Lake T46N RIlE sec 27 Turner Lake F ena Lake T46N R9E sec 21 Spring Grove Fena McHenry T46N R9E sec 30 Lyons Prairiea McHenryILake T43N R9E sec 4

aState Nature Preserves 7

Although Costello was working in southeastern Wisconsin, his descriptions are accurate for sedge meadows in northeastern Illinois. These sedge meadows are dominated by Carex stricta, or tussock sedge, which often forms mono dominant stands. In some places,

Calamagrostis canadensis is abundant, but never as abundant as Carex stricta. Carex lacllstris may form thick stands in the sedge meadows as well. However, Carex lacustris excludes other species where it grows, while Calamagrostis canadensis usually grows in

Carex stricta stands. Depending on the quality of the sedge meadow, a diverse flora offorbs and other sedges may be found among the grasses and sedges. Lower quality sedge meadows may have Typha sp., Phalaris arundinaceae, Lythrum salicaria or Rhanmllsfrangula

InvaSion.

Carex stricta's cespitose growth form results in the formation of pedestal-shaped tussocks that dominate the landscape in sedge meadows where they develop. Tussock size is a rough indicator of age, and Costello (1936) observed some tussocks he estimated to be 50 years old.

Tussocks can grow to well over a foot tall and to a similar diameter. This allows Carex stricta to maintain itself above potential competitors and above high water levels. Costello (1936) attributed the dominance of Carex stricta to the advantage gained through the height of the tussocks and the height of the plant itself. He believed Carex stricta's strategy was to permanently occupy a place through steady, vegetative growth, rather than through dispersal to new sites.

Soils Sedge meadows are found in a diversity of landscape positions in northeastern

Illinois. They may be found associated with fens, bog margins, lake shores, abandoned flood plain terraces, morainal upland depressions, former basins of glacial-age lakebeds, at the foot of morainal bluffs and in the dune-swale topography of ancient Lake Michigan beaches. What these sites all have in common are the proper hydrologic regime and soils to support sedge meadow vegetation. 8

Soils in the natural sedge meadows studied are typical of low, wet areas in northeastern

Illinois. They are, primarily, muck soils (highly decomposed organic soils of the Saprist suborder of Histosols), or Aquolls (Mollisois with aquic moisture regimes). Table 2, derived from Lake County soil survey maps (Soil Conservation Service, 1970), describes the major soils found at the 5 natural sedge meadow sites. Table 3 lists major soil series present at each site. These are not complete listings of all soils that support sedge meadows in northeastern

Illinois. Other soils were probably also present in the sites studied, but were too small to be included on soil survey maps. For example, Kelsey and Hootman (1992) studied a sedge meadow/wet prairie site in northeastern Illinois that occurred on a Typic Haplaquoll. This is probably a common mineral soil in sedge meadows, and may have occurred at the 5 sites studied, but was too small to be included on soil maps.

Houghton mucks are level or depressional Histosols (organic soils) composed of the remains of sedges, cattails and grasses. They are very poorly drained, with the water table at or near the surface all year. The soil organic component ranges from undecomposed litter to accumulations of peat and muck. They contain 20-50% organic matter. Soils of the Saprist suborder are the most highly decayed of Histosols and have the greatest bulk densities. Muck is more decomposed than peat and, therefore, has a finer texture and a greater mineral fraction than peat. Although they have a very high soil moisture holding capacity, soils of the Saprist suborder have the lowest water-holding capacities of Histosols (Brady, 1984).

Sawmill silty clay loams are level, poorly drained to very poorly drained mineral soils formed in alluvial material, under sedges and grasses (Mollisols). Water tables are high in the spring and depend on the water level in adjacent streams the rest of the year. These soils are frequently flooded. Organic matter content is high. Available moisture capacity is very high.

Sawmill soils may include small areas of Peotone silty clay loams and Houghton mucks.

Peotone soils are Cumulic Haplaquolls. They formed under grasses and sedges and have 9

Table 2. Major soil series in five natural sedge meadows in northeastern Illinois

Soil Classification

107 Sawmill silty clay loama Cumulic Haplaquolls 103 Houghton mucka Typic Medisaprists WI03 Houghton muck, weta Typic Medisaprists 82 Millington loama Cumulic Haplaquolls 327A Fox loam 0-4% slope Typic Hapludalfs 67 Harpster silty clay loama Typic Calciaquolls

ahydric soils (Environmental Laboratory, 1987)

Table 3. Major soil series at the five natural sedge meadow sites

Site Soil

Lyons Prairie WI03 Houghton muck, wet 67 Harpster silty clay loam Turner Lake WI03 Houghton muck, wet Spring Grove 82 Millington loam Des Plaines River Trail 103 Houghton muck 107 Sawmill silty clay loam Wadsworth Prairie 107 Sawmill silty clay loam

high organic matter content. They are similar to Sawmill soils, but may be more depressional and have more clay, subjecting them to prolonged ponding and high water tables rather than flooding as with Sawmill soils.

Millington loams are poorly developed floodplain soils formed in recently deposited alluvium. Texture is variable, from loam to fine sandy loam or silt loam. The soils may be 10 mucky at the surface. Sand and gravel may be found 30-40 inches down. Millington loam is high in organic matter, calcareous, and subject to high water table.

Harpster silty clay loams are level to depressional, poorly drained to very poorly drained soils, subject to ponding and high water tables. They formed in alluvial material under grasses and sedges, and may continue to receive eroded material washed down from upslope. Organic matter content is high. These soils are calcareous throughout, and snail shells are present in the upper 1-2 feet. Harpster soils may contain areas of Peotone silty clay loams and Houghton muck.

Created sedge meadows

Site description The created sedge meadows were located at the Des Plaines River

Wetlands Demonstration Project, west of Wadsworth, in Lake County, Illinois. The site was located on the west side of the Des Plaines River at the intersection of Wadsworth Road and

Highway 41. The sedge meadow complex consisted of 10 experimental wetlands (EWs), six of which were included in the present study (Figure 1). Each EW was surrounded by a berm.

The effective area within the berm in each EW was about 1.66 acres. The design allowed a gradual water level gradient to develop on the gentle slopes between channel and berm

(Figure 2). Water was pumped from the Des Plaines River into channels running the length of each EW. Water levels were controlled by the elevation of stoplogs at the outlet of each EW.

Soils Soils at the DPRWDP site (Table 4) are typical of soils that developed in flood plains and adjacent uplands in the region (Soil Conservation Service, 1970). Soils present reflect the changing environment of erosion and alluviation from past flooding and river channel meandering.

Sawmill soils formed in water-deposited silt and clay, under hydrophytic grasses and sedges. These soils are found in depositional environments, receiving material from upland erosion and stream alluviation. 11

Des plaines River

EW1D EW2D

EW1C EW2C

EW1B EW2B

Figure 1. Diagrammatic view of experimental wetland arrangement and water flow

EFFECTIVE AREA

32' VARIES 12~'1

1.5' min.

l' not to scale

Figure 2. Cross section of experimental wetland, DPRWDP 12

Table 4. Major soils present at DPRWDP experimental wetlands site

Soil Classification

107 SawmiII silty clay loam Cumulic Hapiaquolls 325A Dresden loam 0-2% slope MoIIic Hapludalfs 696A Zurich silt loam 0-2% slope Typic Hapludalfs 696B Zurich silt loam 2-4% slope Typic Hapludalfs

Dresden and Zurich soils formed higher in the landscape than Sawmill soils. Dresden soils formed on alluvial terraces, in silt and calcareous gravel, under Oak savannah. Zurich soils formed in glacial outwash, under deciduous forest, in silt and calcareous, stratified silt and sand. Dresden and Zurich soils are level to gently sloping, well drained to moderately well drained.

During construction of the sedge meadows, topsoil was removed to reach desired elevations, leaving subsoil horizons exposed at the surface. Because of the natural sloping of the site down toward the the river, a deeper excavation was required to achieve the desired elevations further from the river. The material exposed was more mineral in nature in the EWs distant from the river, and more organic closer to the river. The surface substrates that remained were a mosaic of previously subsurface horizons of the soils present. The surface substrates in the EWs were, therefore, quite variable in texture, both vertically and horizontally, ranging from silty clay to sand and gravel, often within a very short distance. 13

METHODS

Water table levels

Natural sedge meadows The accepted method for monitoring depth to shallow water tables is the unlined borehole, lined with perforated PVC pipe to prevent the hole from collapsing (Faulkner et al., 1989). It is considered adequate for most porous soils and is recommended for peat soils. That was the well design used in the present study.

In August, 1991, wells were installed in the five natural sedge meadows. Wells were 2 inch diameter PVC pipe, 8 feet long. Eighth-inch diameter holes were drilled 2 inches apart at alternating right angles in the lower 4 112 feet of pipe. Pipes were capped to prevent precipitation entering the top. Holes were drilled below the cap to allow equalization of atmospheric pressure in the wells.

Holes of smaller diameter than the pipes were augered into the ground and the wells inserted by force to a depth of 5 feet, leaving 3 feet above the surface. The pipes were cleared out with the auger and soil packed around the pipes at ground level as needed to seal any gaps between the soil and pipes. Because the sites were in nature preserves, impacts on the sites during installation were minimized by doing all of the installations by hand, and by foregoing the use of sand reservoirs at the bottom of the wells and bentonite clay sealant at the surface as is often done in less sensitive sites (Faulkner et al., 1989). At each site, four to six wells were placed in a line along the water depth gradient from upland to shallow marsh.(It is apparent from water table level records which wells were in the adjacent upland.) Maps showing well placement in the five natural sedge meadows are in the Appendix.

Water level in the wells was measured every two weeks from March 9, 1992 through

November 12, 1992. Standing water depth was measured from the soil surface. Level of the water table was measured with a battery-powered water level indicator, in which contact with water by a probe caused a needle to jump. If water tables were high enough, a tape measure 14 and flashlight was used, as that was more accurate than the probe, which had a resolution of several centimeters.

Created sedge meadows Two-inch diameter PVC pipe was prepared and installed as in the natural sedge meadows. Filter fabric was wrapped around the lower half of the pipes to prevent their filling with sediment. However, some filter fabric ripped during installation.

(Well #2 in EW2B filled with sand and remained so despite repeated attempts to clear it . This well was out of service throughout the study.) Four wells were placed along a transect from the top of the berm to the edge of the channel in EW IB, lC, ID, 2B, 2C and 2D. A fifth well was placed in each EW directly across the channel from the group of four wells. Wells were numbered starting with 1 at the top of the berm, to 4 nearest the water, and 5 across the channel by itself (EWID had a sixth well placed near well #5). Well transects were located where they would cross the soil amendment plots used in sedge establishment studies. Water level in the wells was measured manually every two weeks from mid-June through 12

November, 1992.

Stratigraphy

Natural sedge meadows During installation of the wells, depths to stratigraphic features were noted to allow description of stratigraphy at each well location. No distinction was made between peat and muck based on degree of decomposition. Organic material was simply referred to as peat. Therefore the term peat used in the stratigraphic descriptions includes muck as well.

Vegetation sampling

Natural sedge meadows The vegetation in the natural sedge meadows was sampled using 1- m2 square quadrats placed along the water table well transects. The abundances of species present were expressed as the percentage of the quadrat occupied by aerial parts of the species, as estimated by eye (Causton, 1988). A quadrat was placed at each well and at 15 every consecutive meter between wells except where vegetation composition was uniform for several consecutive meters. In that case, quadrats were sampled every second or third meter.

Created sedge meadows The experimental wetlands were seeded May 19-21, 1992. A seed mix of forbs and sedges found in native sedge meadows was prepared using seed purchased from wetland nurseries in northeastern Illinois and southeastern Wisconsin .. Three matrix grasses were used: oats (Avena sativa), barnyard grass (Echinochloa crllsgalli) and redtop (Agrostis alba). The matrix grasses were meant to provide a quick cover crop to reduce soil erosion and weed invasion. A study was done to evaluate the effect of different matrix grasses on vegetation establishment in the EWs. Both sides of each EW were divided into eight approximately equal sections. The eight matrix treatments, named after their matrix grasses (Table 5), were applied to randomly selected sections on each side. The "oat" treatment had no forb and sedge seed mix, and the "sedge" treatment had no matrix grass.

Seeds were sown by scattering by hand as evenly as possible.

In mid-August, 1992, cover estimates of species present in the EWs were made. Within areas planted with a seed mix, a 13 x 13 meter sampling area was delineated starting at a

Table 5. Eight seed matrix treatments evaluated for their effect on vegetation establishment (sedge = forb and sedge seed mix)

oats sedge oats + sedge redtop + sedge redtop + oats + sedge barnyard grass + sedge barnyard grass + oats + sedge barnyard grass + redtop + sedge 16 randomly selected location next to the berm. Nine I-m:! quadrats were sampled within the

sampling area. Quadrats were 2 meters apart. This arrangement allowed sampling of vegetation across a large part of the water depth gradient, from berm to channel. Because each matrix treatment was replicated 12 times, 108 quadrats should have been sampled from areas where each of the 8 seed mixes were sown. Other activities, particularly planting of plugs of Carex spp. within areas sown with the barnyard grass+oats+sedges mix in most of the EWs, made it impossible to sample areas sown with this mix in most of the EWs.

Consequently, only 18 quadrats were sampled in areas sown with this mix.

Monthly precipitation data was obtained from NOAA records of mean monthly

precipitation from all stations reporting in northeastern Illinois. 17

RESULTS

Water table levels

Natural sedge meadows Figures 3-7 show the water table levels observed for each of the natural sedge meadow sites from 9 March, 1992 to 12 November, 1992. All of the sedge meadows had standing water when observations began in March, 1992, indicating water tables above the surface. Water levels then fell steadily from early May until about the second week of July. Water levels increased in July after heavy rains that month. Water levels then dropped until heavy rains in September caused them to rise through the end of the observation period.

Records of monthly precipitation (Figure 8) indicate October and November of 1991 were wetter than normal, particularly due to heavy precipitation in October. The first six months of

1992 were drier than normal. May and June, 1992 were particularly dry. July and September were wetter than normal. July, 1992 was the second wettest on record, with an average 144 mm recorded over northern Illinois (NOAA, 1991 and 1992).

Created sedge meadows Figures 9-14 show the water table level measurements in the

EWs in from 15 June to 12 November, 1992. The significant changes in water table elevations seen are the result water level manipulations in the EWs. These included the initial filling of the EWs and subsequent changes in pumping rates or changes in the elevation of stoplogs at

EW outlets. Although little in the way of a consistent pattern can be seen, there is evidence of an effect due to distance from inlet pipes. EW IB, I C and ID are nearest the inlets, and responded sooner as a group to pumping than did the more distant EW 2B, 2C and 2D.

(There was a problem with pumping in EW 2 which prevented its filling with the others.) The southern tier ofEWs simply took longer to fill than did the northern tier. The remaining variability in water table levels is likely due to substrate heterogeneity and to the influence of other external factors. 60 --- well#1 ....+ ..... well#2 --*-_. well#3 ---8--- well#4 40

'-1L __ *---*--~ 20 *----=- " ~/+ , .. -+. ' / ...- + ...... + ...... +...... +.: ~ _- -- ./ ,0 B E G- ---e--B------5:\.'...... " '* I ----::..&.C-- . 0 0 , -'. ""--" +-/ '~'".\ +.······'-T/ W{\ ; I .•••. )LJ . , ,+... ," ,. -L: '...... '...... +. , +-' ,-., , , : 1Zf 0... -20 'm . , ,i?---Er-- ""'I) J o', ill ,. : t , : f -0 , , ,',: I 1, ,',," 00 \ :, - '\ :, -40 \ :r \ ~",,~m··..!i \ / ...... :.:... it' \ / ~~"" :1 , -60 ~... ",,.~. ',"*' ...... ""•. ~ ttl

-80 I I I I I I I I I I I I I I I I M10 A4 M2 M28 J25 J25 A19 S20 017 N12 M21 A18 M16 J15 J10 A6 S6 04 N1

Figure 3. Water table levels at Wadsworth Prairie, 10 March to 12 November, 1992 • well#1 ...... + ..... well#2 ---*--- well#3

---G--' well#4 )( well#5 ...... " well#6 20.0

0.0

,...... E -20.0 ... 0 '--'" -40.0 -\0 ...... Ji...... ,'-: ..c...... , ~ / ".41(' -60.0 \., ...... : 0... \. ~ .. (]) A ...»., ! LJ -80.0

-100.0 \~."l

-120.0 I I I I I I iii I I I 1 1 I I I Iii I M9 A4 M2 M28 J25 J25 A19 820 017 N12 M21 A18 M16 J15 J10 A6 86 04 N1

Figure 4. Water table levels at Des Plaines River Trail, 10 March to 12 November, 1992 10-1-*------==~li "'*---*--'*- /-* 0 , --- ,/ well#1 ...... L. , _~_*GJ' " ... rr. " '* ...... __"* " ?', ...+ .....+ .... '7' ~, -10 '. + " , +", < b - .+...... * *' " B. , . I • well #2 ..0" ~ ",.... '. }*, I '. : '{ /' 8' ',... '. I '. , { -20 /', 1Sl,, +'" "'+ '.\, / * +: ....., '. I...... " + ....+..... ,tP ~ --*-- *' ". "'" ' '...".., well #3 -E -30 lsJ._";\/~\/10("\\.-.-*;/ -1'5 0 ---B--

\+ -- ~"o=i"',+. :' " -40 >-.t' 1-.:.0 ~" +, -..c t..::J + ~,...+ ...... " well #4 ...... , , ,f 0.. S--EJ Q) -50 N "'0 o -60

-70

-80

-90 I I I I i I I I I I .,. I I I I I I M 10 A4 M 2 M 28 J 25 J 25 A 18 S 20 017 N 1 2 M21 A18 M17 J15 J10 A6 86 04 N2

Figure 5. Water table levels at Turner Lake, 10 March to 12 November, 1992 60 ' ,..------,

---well 1 40 '+'" .....+- ....

""+:...... +...... ,.:..... '-'+ well 2 20 --*-- .-. ~\.. E well 3 o o I 'g. "'I!il\,( \ 7 LF-''l5 El --8-- ---...c wel14 ...... 0.. -20 Q) ~ N "'0 wellS ..... -40

-60

-80 I I I I I I I I , I I I M10 A4 M2 M28 J25 J25 A18 820 017 N12 M21 A18 M17 J15 J10 A6 86 04 N2

Figure 6. Water table levels at Spring Grove Fen, 10 March to 12 November, 1992 20

10 ~..+.....;::*"::~*'~'*'--*-~- *. ---well 1 !CI..,-= +...... '. *. /-*- ~-* -- ~~-E)..-_~.__ ~ __ I , ~-~-~-*~::~= G . *-'"*I , ,~;;7T'\;; '. '1:1:] 0 .....+ .... '+...... +\ \ \ \ wel12 -10 \\ \ \ --*-- .- \. \ _Q :! i E -20 ~W" " ,'I'l wel13 \ • ';>IE, 'til/ 0 -30 --8-- -...c \. "~f well 4 +.. i ...... -40 0...... ~... ! Q) N ""'0 -50 N ...... J -60

-70

-80 I * -90 I I I I I I i I I I I i I I I I M9 A4 M2 M29 J25 J25 A17 820 017 N12 M21 A18 M17 J15 J10 A6 86 04 N2

Figure 7. Water table levels at Lyons Prairie, 9 March to 12 November, 1992 _ recorded D normal 200~------~------.

180 160 -E 140 E --c 120 o +:i 100 ...... m "0.. 80 IV u t..J ~ 60 Q.. 40

2:1 UJ II JFMAMJJASONDJFMAMJJAS 1991 1992

Figure 8. Northeastern Illinois 1991-92 monthly precipitation data and monthly normals 20 ......

~.Q_,- ~ .;' '- well2e1 10 ~_,. /.c:J-,. ':'.~-----~ ~-,---tIl " --er~--EJ -f- rr"/,/\\ \ }~ well2e2 0 . " ,: \, ,.I . \/ -"'/., :'/"*<'~\--*-- I :\ " ',,*----* ~\, . ;' '>Ji-,-' x· ...... x ~ well2e3 • : /~ " -E -10 .I ... ' () • --G-- '-' well2e4 ..c -20 +-' ·····x···· a.. ~ .; Q) I well2wS N l:J -30 ~ -40 / x ...... - ...... _.' -50 -" /------...... /--_.: , / ~ '1Ii -60~~~--~--.--.--~--.--.---r--.--.--~ J15 J9 A5 86 04 030 J26 J25 A19 820 017 N12

Figure 9. Water table levels in EWIB, DPRWDP, 15 June to 12 November, 1992 20 IGJ 10 ---well3e1

.....+ ..... 0 , , ' ,, ( .,"'-----~" '" " ;fE.. wel13e2 -10 ti / '-'-~-----* "'* ( ..... '''', ..., , .- --*-- ..-...... +., ....+ ...... "* ...,...... + ...... + ...... + -20 well3e3 E ...+ ..... 0 -30 ---8-- -£ wel13e4 of-' 0.. -40 Q) ~ N lJ -50 well3w5 I VI -60

-70~V -80-1

-90~~~--~--~~--~--~--~~--~--~~~ J15 J9 A5 86 04 030 J26 J25 A19 820 017 N12

Figure 10. Water table levels in EWIC, DPRWDP, 15 June to 12 November, 1992 20 Gl _---8..-, -II- ., '-', 8- --@ ' ,". s- --s-----EJ-----5:l 121------8'"-- '- 0 .. , well4e1 ...... ,," " ::-, r:::i ...... ~ "",..*".," ',...,!!". ...'m ...... + ..... " ~..- .;-,,-" ...... -20 .+...... +. well4e2 ... ..,'A ". .+ ...... + .+ ...... • -4°1+-V/~/ '.-/ / II ~:I~e3 -E () 60 -- - 1 ...c II ~:I~~e4 -+-' I ~-1 0... -80 ~ Q) N -0 well4w5 I 0\ -100~ / ...... II well4w6 -120~/ -140l

-160 J15 J9 A5 86 04 030 J26 J25 A19 820 017 N12

Figure II. Water table levels in EWID, DPRWDP, 15 June to 12 November, 1992 20 ~,, . ' ...... , ' I. !x lSI, well6e1 I j ". " I ,;,: ". '... B_~,.. ~t:::I- r-:-1 ... : _', ,.~..{3--, O I !!'. .) X s- --8,,·--8- ! ! \ -"'53 -+- ,: '. . : '. \ well6e2 ,:. '. ' -20 --*-- ....-... "\ ,// '" \\ }<'\ ..., X, t!J! / '_ '. ,/ '" ,X.lIE \ well6e3 E ", !~. ,X""' ,/ ", ..... *"," , () ',,: """ ,,· ..·X· *.-.--*---~-----8-- * '. i,' * .....--~.: ' -..c -40 '- ·X " --', /" wel16e4 +-' * +~\". '*, 0.. \ , , , I ..·.. x ....· (J) , '. • , , I ...... • •• tv "'0 . well6w51 -..I -60 1!Il...... / ...... II, ! ''II! rill -80 """",.",,,,j\/ -100~~~--~--~~--~--~~--~--~~--~ J15 J9 A5 86 04 030 J26 J25 A19 820 017 N12

Figure 12. Water table levels in EW2D, DPRWDP, 15 June to 12 November, 1992 20,"------.

>;<'111.,. ~ "'x --- "', . )( .....~ ~ well7e1 '" '" ". .' . ~...... ·~ ..~, ..x "l,. /£1." __8-" .. ",_...... + ..... /."', , " ./ ',--- . .. o I; '\' ...... ,.'oox ...... '" , ~ "'." \,'" I "''P, '~'-''" , ...... , , ... well7e2 -20 " . ~ , ," ."! '+ ". .I" " --*-- \ I ._ .-. ~ \ f" .*. ! I!J well7e3 \ , " / " '!/ E ~ / ' ...... / """:'! • I "'" "'" : o -40 \, " i --8-- , : . : ..c---- well7e4 +-' lfi>~.J...... ,...... ,: tv ""0 well7w51 00 \+- -80

-100

-120~~~--~~--~--~~--~~~~--~~~ J15 J9 A5 86 04 030 J26 J25 A19 820 017 N12

Figure 13. Water table levels in EW2C, DPRWDP, 15 June to 12 November, 1992 20 63""* If\ \ II' \ ,1 \ \ 10 "~"\ \ --- ::1 '\(" \ .....~, .. well8w1 ," \ ". \ "'-- ---*<----,"'""--- -, I" \ +. _ --"'" ---*-- "" -* ,~:, ~ ..".. \ ..._-+----- , \... 0 , .. It \ .. ,r• 1 \ .. \ .r rn " I well8w3 \ " . "\ .1I' , " \ II r.:1 .. -10 .1 \+:\ • .1Itl " ,...... , ---?lE--- ,I' > .-. ,:, " •••• ,,/ \'" J I "" \ \3- ...... '\:of,,:"/-"-+--"'-~~B3\ ~ f:~: "l well8w4 , I:' -- __ cc... ____ 0 J'O"I '.. ,', E -20 .. ,:1 tD" ••••••••• ~ L,;..I \', "'" \ r: 1 ',.... 0 \ ,:, .... ---8-- +. \ I;" " "'. ,;1·' \" '.:t- -..c -30 Ul ...... *: ,, well8e5 +-' \ ':+: , J tJ D- , J \ J I W -40 , J , J N "'0 C!J \0 -50

-60

-70

-80 .. J15 J9 AS' 86 04 030 J26 J25 A19 820 017 N12

Figure 14. Water table levels in EW2B, DPRWDP, 15 June to 12 November, 1992 30

Stratigraphy

Natural sedge meadows Observations of stratigraphy of the substrates supporting natural sedge meadows are presented in Figures 15-19. All of the columns had peat or muck at the surface, ranging in thickness from 30 cm to beyond the length of soil probes used, over

175 cm. (The exception was under well 1 at Turner Lake, which was not in sedge meadow, but rather on an adjacent upland.) Under the organic layer, either silt, sand or blue clay was found. The heterogeneity of the deposits under the muck reflects the extreme diversity of materials deposited in former lake beds and past stream channel movements and flooding in these areas. The Des Plaines River Trail site (Figure 16) exhibited signs of extremely active past river activity. The well 2 column contained a layer of peat buried by silt, indicating a shift in the nearby Des Plaines River channel or a period of active erosion had buried a former wetland. A clay layer buried by silt under well 5 suggests a similar history there.

Vegetation sampling

Natural sedge meadows The most frequently found sedge species in the natural sedge meadows was Carex stricta (Tables 6-7 ). It was the only Carex species all 5 sites.

Calamagrostis canadensis was the only other graminoid found at all 5 sites. Common forbs in these sedge meadows were Aster spp., Convolvulus sepium, Eupatorium maClilatllm, Lycopus spp., Mentha anJensis, Pycnanthemum virginial7um, and Solidago spp. All of these are common species in sedge meadows and other seasonally flooded wetlands in the midwest.

Created sedge meadows Table 8 lists all species found in the created sedge meadows in

August, 1992. The most abundant species were Lolium perenne (EW 1B, 1C, 2B), Ambrosia artemisifolia (EW1D) and Cyperlls esclllentlls (EW2D,2C). Table 9 lists the most abundant species by percent cover in each EW. Table 10 summarizes species recruitment data for areas seeded with the 8 different seed mixes. em well 1 well 2 well3 well4 key o

-20 Wi

-40 marl

,~ .. ,. ,. ,. ,. ,. -60 ,. ,. ,. ,. ,. """,. ,.,. ,. ,. \. .,.--, "\. "\. "\. "\. \. , , " ,. ,. ,. ,. ;' \. . \. ,." \. ,. ,. \. ,." ,. , "" " "" ,. ~ \. ,." ,.\. ,.\. ,." ,. -80 " "" ," '"" "" , ~ \. \. \. " " ,.\. ,.,. \. ,. ,. 181 / ,. ,. " , " " " ..... \. ,.\. ,.", \. ,.\. ,. / / / w " " "" " " \. ,." \. ,. ,. \. ,. \. ,. \. ,." "" ,. \. " \...... \. "" ,.\. ,." " ,., \. ,." ,.\. ,.\. ,.\. ,. " ,." ,.\. ,." ,., ,. ,. / , , -100 ,. ,.,. ,. """,. ,. ,. ,. ,. \. " " \. ", \. ,.\. ,." ,.\. ,-, """,. ,. ,. ,. ,. \. \. \. fi' ,." ,.,. ,., "","'""",/ , ,. ,. ,. "","',. ,. ,. ,. ,. "",.,. ,. ". ,. ,. -120 ,. , ,. ,. ,. silt '/ """,. ,. ,. ,. ,. ,. ,. ,. ,. """ -....' .... ~ ~ "",.""",. ,. ,. ,.,. ,".... ," .... ," ...... -140 """ f"....L..L- sand r:::81 -160 Ld

Figure 15. Stratigraphy of substrates beneath wells at Wadsworth Prairie em well 1 well 2 well3 wel14 wellS well 6 o key -20 m -40 marl

-60

-80 ~

,. ;' ,. / ,. gravel w ,. ,. ,. ,. ;' tv I "'" -100 ;' ";' ";' "" " / " I;' "'" / ;' / ( 1/////~ett: ~ I;' / ;' ;' ;' "'" / -120 I"'";' / ,. ,. silt I" I" I" I" I" ~

-140 sand I:8:l -160 LD

Figure 16. Stratigraphy of substrates beneath wells at Des Plaines River Trail site em well 1 well2 well3 well4 key o -20 m -40 · ...... sano.lY SOl -60 '.', ...... D -80 w silt w ," ,/ ", " / "';' , , , , , ,/ , , " " " -100 " " '" " . ", ", ", , " ;' "" " , " ,,- " " " " . " " " ,"" ," " " ," ," " "' :::::;::: : ," , " , ", " / . ~, , , , , sand -120 .. ,',',' " " " " " , ," , ", ", " / . ," , ", " , " / . r:l ... ', .. ', . ,-s~~~t5 , , , , Lill · , ...... ,/ ", ", " , " ,. -140 . :-:-:-:-:. /" ," ," ,/ " '. ," , ", " / " , ~"" ," ," , " , , " , " I' ", " " , / . -160 · . , ...... ," "" , " , " ," , " , " ," , /////," " " " ' · ....

Figure 17. Stratigraphy of substrates beneath wells at Turner Lake site em well 1 well2 well3 well4 wellS key o -20 m -40 marl :.:.:.:-;.:.:.;.:.:.:.;.;.;.:.:.:.:.;.;.;.-

~rttIttItII~ -60 --

-80 ~ •••••••••• ~:~:>~<:. gravel w .. .' .j:::. -100 < ~ -120 sand W.... -140 ••••••••••

. . -160 . . IlL1LLI

Figure 18. Stratigraphy of substrates beneath wells at Spring Grove site em well 1 well 2 well3 well 4 o key peat -20 Wij -40 silt -60 / , / , / , / , / , / , / / /

-80 sand w ... Vl -100 ..... '. '. m

-120

-140

-160

Figure 19. Stratigraphy of substrates beneath wells at Lyons Prairie site 36

Table 6. Mean percent cover of Carex stricta in natural sedge meadows in quadrats where it occurred

Site Mean % cover

Des Plaines River Trail 33.2 Wadsworth Prairie 35.3 Spring Grove Fen 29.5 Turner Lake Fen 3l.9 Lyons Prairie and Marsh 37.0

Recruitment was very sparse regardless of the seed mix used. Species were grouped into three classes for the purposes of statistical analyses: ruderals, i.e., species not in a seed mix; seed mix species; and all species (total). No Carex spp. were found in any of the quadrats sampled. Likewise, one of the grass matrix species, redtop, was not found in any quadrat.

There was no significant difference in mean total cover of ruderal species among the areas sown with different seed matrix treatments. Percent cover of seed mix species was also not significantly different between areas sown with different matrix treatments. Total percent cover was not significantly different in areas sown with different matrix treatments. There is a suggestion that the cover ofruderals was reduced in areas where oats (Avena sativa) or barnyard grass (Echinochloa crllsgalli) or both were matrix species. In summary, none of the seed mixes used was effective in establishing Carex spp., although scattered individuals of a few sedge meadow forbs were found. The vegetation in the created sedge meadows at the end ofl992 was a sparse cover (5 to 15%) of primarily ruderals (4 to 12%) and matrix grasses (1 to 3%). 37

Table 7. Percent frequency of occurrence of species found in natural sedge meadows. Only species that occurred with a frequency of 10% or higher at one or more sites are included. Numbers in parentheses are the number of quadrats sampled at a site (Des Plaines River Trail (DPRT), Wadsworth Prairie (Wads), Spring Grove (Spr), Turner Lake (Tur), and Lyons prairie (Lyo))

DPRT Wads Spr Tur Lyo Speciesa (154) (~1) (40) (27) (45)

Amaranthlls sp. 0 0 15 0 0 Angelica atropurpurea 0 0 10 7 0 Aster sp. 41 19 50 22 98 Bidens coronata 0 0 43 19 11 Boehmeria cylindrica 13 0 40 0 2 Calamagrostis canadensis 1 47 23 100 76 Campanliia sp. 9 6 3 4 0 Carex atherodes 0 25 0 0 0 Carex lacllstris 16 0 48 0 0 Carex rostrata 4 16 0 0 0 Carex sartwellii 6 0 0 0 2 Carex stipata 15 0 13 0 0 Carex stricta 75 79 68 100 91

~ Cirsillm sp. 5 0 ..) 0 0 Convolvulus sepium 2 20 0 48 9 Eleocharis palustris 10 5 5 0 2 Epilobillm sp. 0 0 0 19 4 Eupatorium maculatllm 2 1 40 19 69 Eupatorium per/oliatllm 22 0 3 0 2 Galillm sp. 15 5 18 0 4

~ Helenium autllmnale 5 1 .) 0 0 Impatiens capensis 12 0 58 0 2 Iris versicolor 2 2 18 0 0 a Species present at only one site with a frequency ofless than 10%: Acorus calamus, Caltha palustris, Chelone glabra, Eqllisell1m palus/re, Heliamhus grosseserra/lIs, Potentilla palus/ris, Sium suave. Solanum dlllcamera, Sparganium elllycalpUl11, Spirea alba, and Typha angllstifolia 38

Table 7. Continued

DPRT Wads Spr Tur Lyo Speciesa (154) (81) (40) (27) (45)

Lathyms palustris 0 1 0 4 11 Lycopus sp. 59 17 20 33 44 Lythrum alatum 2 0 0 0 9 Mentha anensis 40 9 5 11 9 Panicllm sp. 12 0 0 0 0 Phalaris arllndillaceae 43 0 15 0 9 Polygol1l1m sp. 67 64 45 19 0 Pycnanthemllm virginial1l1m 0 1 10 7 11 Ranlll1Cullls sp. 11 1 0 0 0 Rosa sp. 14 0 0 0 0 Rumex sp. 1 4 8 0 2 Sagittaria lati/olia 23 0 0 0 0 Salix sp. 1 6 0 0 2 Scirpus aClltlis 0 0 18 0 0 Scirplls atrovirells 30 0 0 0 2 SClltellaria galericulata 8 2 8 7 22 Solidago sp. 13 0 25 26 22 Spartina pectinata 12 0 30 0 Thelypteris sp. 0 0 0 37 0 Typha lali/olia 41 0 0 4 24 Verbena hastata 2 0 15 0 0 Veronicastrum virgilliclll1l 2 0 0 0 39

Table 8. Species found in 1992 in created sedge meadows at DPRWDP

Abutilon theophrasti Phleul1l pratense I Agropyron spp. Physostegia virginianaG Amaranthlls hybridlls Plantain major Ambrosia artemisifolia Plantain ruge/ii Avena sativaG Poa pratensis Barbarea vulgaris Polygonllm aviclilare Bidens spp. Polygonum erectllm Chenopodium album Polygonllm lapathafolium Cicuta bulbifera Polygonum pennsylvanicllm Cirsium spp. Polygonum persicaria Convolvulus sepium Populus deltoides Cyperus esculentlls Rumex crispus Dallclls carota Rumex verticillatlls Echinochloa crusgallia Salix spp. Eqllisetum spp. Setaria spp. Lolillm perel1ne Silene noctijlora Medicago luplilina Trifolium pratense Melilotlls officinales Trifolium repens Paniclfm dicotomijlorum Verbena hastataG Phalaris arundinaceae

a Species in seed mix 40

Table 9. Species with> 0.5% cover in experimental wetlands

EW Species % cover

IB Lolium perenne 3.2 Ambrpsia artemisifolia 1.9 Echinochloa crllsgalli 0.8 Avena sativa 0.7

lC Lolillm perenne 4.2 Ambrosia artemisifolia 4.2 Echinochloa crusgalli 2.4 Cyperus eSClllentlis 2.3 Avena sativa 0.6

ID Ambrosia artemisifolia 3.7 Echinochloa crusgalli 1.7 Lolium perenne 1.5 Cyperus esclilentlls 1.4 Salix spp. 1.3

2D Cyperus esclilentlls 3.2 Lolium perenne 1.7 Ambrosia artemisifolia 1.0 Echinochloa crusgalli 0.7

2C Cyperlls esclIlentlis 1.5 Echil1ochloa crllsgalli 0.7 Lolillm perenne 0.6

2B Lali1lm perenne 3.3 A vena sativa 1.7 Polygol1l1m pennsylvanicllm 0.9 Ambrosia artemisifolia 0.8 Table 10. Mean percent cover ofruderal species, seed mix species and total species in areas sown with one of8 seed mixes in the experimental wetlands at the DPRWDP

Seed Mix Ruderals Seed Mix Total Cover

N sum mean ± s.d. sum mean ± s.d. sum mean ± s.d. oats 108 828 7.67 ± 10.8 205 1.90 ± 2.8 1033 9.56 ± 12.9

oats+sedge 108 1033 9.56 ± 11.3 203 1.88 ± 2.2 1236 11.44 ± 12.9 .to...... barnyard+redtop+sedge 108 1067 9.88 ± 15.5 355 3.29 ± 6.6 1422 13.17± 17.7 barnyard+oats+sedge 18 76 4.22 ± 8.7 19 1.06 ± 2.4 95 5.28 ± 9.8 barnyard+sedge 108 730 6.76 ± 12.1 204 1.89 ± 2.2 934 8.65 ± 12.6 redtop+oats+sedge 117 1342 11.47±17.9 179 1.58 ± 2.0 1521 13.00 ± 18.7 redtop+sedge 108 1341 12.42 ± 16.3 248 2.30 ± 4.2 1589 14.71 ± 17.0 sedge 108 1169 10.82 ± 13.8 159 1.51 ± 2.1 1328 12.30 ± 12.3 42

PART II.

SEDGE ESTABLISHMENT STUDIES 43

INTRODUCTION

Very little information is available on germination of Carex species occurring in northeastern Illinois sedge meadows. The expansion of wetland restoration efforts in this area should encourage more studies, similar to Gillespie (1987) who studied germination of Care x stricta in southeastern Wisconsin, and Larso'n and Stearns (1990) who studied Carex scoparia germination and storage, for wetland restoration purposes. Published studies of Carex seed germination and viability have focused on four major areas: arctic species, rangeland species, weed seed ecology, and general wetland seed bank studies.

Chapin and Chapin (1980), Gartner et al. (1983), and Wein and MacLean (1973) worked with seed of Eriophonlm vaginatlln1, Carex bigelowii and other graminoids in the field of tundra reclamation in Alaska. Seed germination characteristics of Carex species have been

studied in the western United States in relation to forage value of Carex species in alpine

rangelands, where they comprise a significant part of the palatable forage (Amen and Bonde,

1964; Johnson et ai., 1965; Wiesner et ai., 1967), and for mine reclamation purposes in the

same areas (Haggas et ai., 1987). Comes el ai. (1978) evaluated the effect of water storage on germination of seeds of Carex lal1l1ginosa and other "weeds" to study the survival of weed

seeds in irrigation water. Finally, Carex species have appeared as components of general seed

bank studies for the restoration of prairie wetlands (Schmid, 1986; Galinato and van der Valk,

1986; van der Valk and Davis, 1978, 1979; van der Valk and Pederson, 1989; Welling et ai.,

1988).

Studies of vegetation establishment techniques relevant to the present study have been

done in the fields of wetland creation and restoration, mine reclamation, and prairie

restoration. Information about techniques for establishing vegetation in freshwater wetland

creation and restoration projects is becoming more readily available due to attention placed

on this area by wetlands regulatory legislation and the interest in using wetlands for 44 wastewater treatment. As a result, much of the information appears in government publications (Kusler and Kentula, 1989; Kentula et al., 1992a) and in engineering-oriented how-to manuals (Hammer, 1989; Hammer, 1992). However, the methods described in those sources are primarily for emergent and mud-flat vegetation, not wet meadows.

Coastal wetland restoration has been practiced for a much longer time and has received

Army Corps of Engineers attention and funding. As a result, techniques have progressed far beyond those available for freshwater wetlands restoration. Standard methods are established for revegetating salt marshes (Broome et al., 1988). Similarly, reclamation of mined lands has been practiced for many years, and proven techniques have been developed (Soil Conservation

Service, 1988; Lyle, 1987). In the field of prairie restoration, standard techniques have been developed (Schramm, 1976, 1990), many of which may be adaptable to sedge meadow creation and restoration, because sedge meadows are, in some ways, more similar to prairies than they are to marshes.

Finally, studies of the effects on soils of organic amendments have been done in the fields of agriculture (MacRae and Mehuys, 1985; Tester, 1990) and in mine reclamation (Lyle,

1987; Roberts et aI., 1988; Pietz et aI., 1989; Khaleel et aI, 1981).

Part II of this report describes a series of studies done to evaluate the use of seeds and seedlings to vegetate the created sedge meadows in the experimental wetlands at the

DPRWDP. This includes studies of Carex seed germination and viability and the effect of storage conditions on Carex seed germination and viabillity. It also includes greenhouse and field studies of the effect of soil amendments on the growth of Carex stricta seedlings. 45

METHODS

Seed studies

Seed germination Initial tests of seed viability and germinability were done in June,

1991, on seeds collected during the 1990 season. One ounce bags of seed obtained from wetlands nurseries were sampled by passing the seed through a divider in the Iowa State

University seed testing labs. An air column was used to separate the seed from the inert matter in a small, 1-5 gram sample. Further picking of the sample by hand removed any remaining inert matter. To test germination, 4 replicate batches of 100 seeds were placed on moist blotters in incubators with an alternating temperature regime, 15°C for 16 hours per day alternating with 25°C for 8 hours, for 3 weeks. Seedlings were counted and removed weekly as they germinated.

Seed viability was tested using methods adapted from those used by Iowa State University

Seed Testing Laboratory (Grabe, 1970). A 1% solution of tetrazolium dye (TZ) was prepared by mixing 3 liters distilled water, 32.19 g of sodium phosphate dibasic, 10.89 g of potassium phosphate monobasic and 30 g of2,3,5- triphenyl tetrazolium chloride.

Seeds were first soaked in distilled water for 24 hours at 35°C. Seed coats were then pierced with a needle or razor blade to allow penetration of the TZ dye, and soaked for an additional 24 hours in 1% TZ at 35°C. Embryos were observed by slicing the seeds in half. An embryo was considered to be alive if it had a red color indicating that the embryo was respmng.

A second series of comparable germinability and viability tests was done in December,

1991, on seeds collected during the 1990 and 1991 seasons. To test germination, 100 seeds of each species-year-source lot were placed on moist filter paper in petri dishes on a greenhouse bench for up to 8 weeks. Eighteen hour days were maintained with supplemental lighting.

Seeds were watered as needed to keep the filter paper moist. Seedlings were counted every 46 two to three days, and seedlings were removed as they germinated. The TZ viability test also was performed on these seed lots.

A third series of tests was done in July, 1992, on seed harvested in 1992. The seed had been air dried for several weeks after harvest and prior to testing. Only filled achenes were tested. This differed from previous tests in which no prior determination was made regarding achene fill status. Otherwise, methods used were the same as those used in the December,

1991 tests.

Seeds of Carex stricta, C. aqllatilis, C. atherodes, C. iaclistris, C. comosa and C. iasiocarpa were obtained from wetland nurseries and personal collections. Seed availability varied from year to year, so not all species were available all years. Table 11 lists species, year of harvest, year obtained, and source of Carex seeds tested. As noted, seeds tested were collected during the prior year or the same year as they were tested. Seeds collected in the year prior to testing were obtained for testing after being in storage at the nurseries. Their storage methods consist of keeping seeds in sealed plastic tubs, in an unheated barn.

Seed storage The effect of storage conditions on fresh (collected and tested same year) and old (collected the previous year) seed germinability and viability was tested using seeds of

C. atherodes, C. aquatilis, C. iaclistris, and C. stricta. Seeds of these species were placed in storage from February, 1992 through July, 1992 under a combination of four temperature

(room temperature, 4°C, -4°C, and alternating 4°CI-4°C ) and two moisture conditions (wet, dry) for a total of 8 treatment combinations. The 4°CI-4°C is designated the aIt treatment.

Two hundred and ten (210) seeds of each seed lot were placed in sterile sand in plastic ice cube trays wrapped in cellophane. In the wet treatment, seeds were stored under standing water or ice. For the alternating -4°C/4°C treatment, seed trays were moved every two weeks from -4°C to 4°C to -4°C etc.

At the end of the study, half the seeds were tested for germinability and half for viability 47

Table 11. Carex species, year seed harvested, year seed obtained, and source of Carex seeds

Year Year Species harvested obtained Sourcea

Carex stricta 1990 1991 SEW 1990 1991 NIN 1991 1991 SEW 1991 1991 NIN 1991 1991 NEI 1992 1992 NE2 1992 1992 SEW

C. lasiocarpa 1991 1991 NEI

C. rostrata 1991 1991 NEI

C. aquatilis 1990 1991 NIN 1991 1991 NIN 1991 1991 SEW 1992 1992 NE2

C. lacllstris 1990 1991 NIN 1990 1991 SEW 1991 1991 NEI 1991 1991 NIN 1991 1991 SEW 1992 1992 SEW

C. comosa 1990 1991 SEW

C. atherodes 1990 1991 SEW 1990 1991 NIN 1991 1991 SEW 1991 1991 NIN

a NIN = northern Illinois nursery SEW = southeast Wisconsin nursery NE 1 = northeast Illinois source # 1 NE2 = northeast Illinois source #2 48

A second seed storage study was begun in July, 1992, using seeds harvested in June,

1992. Tested were seeds of C. stricta and C. aquatilis from the northeast Illinois source #2, and C. stricta and C. iaclistris from a southeastern Wisconsin nursery. The freshly harvested seed was air dried several weeks, then placed into storage. Two replicate lots of 100 seeds of each seed lot were wrapped in plastic mesh 'and placed in Nalgene bottles. These bottles were filled with either dry sand, distilled water, muck from natural sedge meadows, or goose droppings in distilled water. The bottles were stored in one of four temperature regimes

(room, 4°C, -4°C, and alternating 4°C/-4°C) from July 27, 1992 until November 27, 1992.

The bottles were then placed in a refrigerator until April 27, 1993, when testing began. To test germinability, seeds were placed on sterile sand on a greenhouse bench. Each replicate was placed in a separate container to prevent contamination with muck or goose droppings.

Containers were watered as often as needed to keep the sand saturated to the surface.

Seedlings were counted and removed as they germinated, every fourth day for 6 weeks.

Growth studies

Greenhouse studies The first greenhouse study was begun March 11, 1992. Six to eight week old Carex stricta seedlings were planted in four inch pots containing one of the soil amendment treatments. Treatments were topsoil/no topsoil, fertilizer/no fertilizer, and compost/no compost. These studies were arranged in a 2x2x2 factorial design with 18 replicates of each treatment combination. Subsoil from the DPRWDP sedge meadows was the control growth medium. Topsoil and compost were mixed with the subsoil in equal parts.

Topsoil was the topsoil that was stripped from the DPRWDP during construction. Compost was composted yard waste provided by Wetlands Research, Inc. Fertilizer was Greensweep liquid lawn food, 27-1-4, applied at a rate equivalent to 1 quart/SOOO square feet. Fertilizer was applied once, when seedlings were potted.

Pots were placed in plastic trays and watered as needed to keep the soil saturated 49 continuously. After three months in the greenhouse, the number of shoots per plant was counted. The plants were then harvested, soil was washed from below-ground parts and the above- and below-ground dry mass of the plants was determined. Leaf mortality was minimal in the three month duration of the study and any leaves that did die remained attached to the plant and were included in above-ground dry mass.

The second greenhouse study was begun in October, 1992. A second batch of composted yard waste from WRI, Inc. was mixed with sterile sand to produce growth media having 0,

10,20,33.3, 50,66.7, 80, 90, or 100% percent compost by volume. There were sixteen replicates of each treatment. Two to four week old Carex stricta seedlings were planted in 4 inch pots containing a compost treatment. The bottoms of the pots were set in plastic bags to help retain water. After three months in the greenhouse, the plants were harvested and their above- and below-ground dry mass was measured.

Field studies A study of the effects of soil amendments on the growth of Carex stricta seedlings was established in June, 1992 in the experimental wetlands at the DPRWDP. A concurrent study examined the effect of soil amendments on recruitment of species from a seed mix sown in all the plots. Sixteen one meter square plots were established on the east and west sides ofEW1B, 1C,lD, 2B, 2C and 2D. The overall experimental design was a 2x2x2 factorial, with treatments: compost/no compost, topsoil/no topsoil and fertilizer/no fertilizer.

Compost, topsoil and granular fertilizer were mixed into the substrate with a rototiller.

Fertilizer used was Scott's Starter Fertilizer (17-23-6). It was applied at a rate of20.5 g • m- 2.

Carex stricta seedlings were planted in half of the quadrats. After two weeks to allow establishment, the number of shoots per Carex slricta plug was counted. After two months, i.e., in August, the number of shoots was again counted for each plug.

In August, 1992, percent cover of species was estimated in quadrats without Carex stricta plugs to evaluate the effect of soil amendments on recruitment from the seed mix. 50

RESULTS

Seed studies

Seed germination Tests conducted in June, 1991 indicated very poor germination

«2%) and very low viability (1 to 7%) for the four Carex species tested (Table 12), with the exception of Care x comosa (58,5% viability), Low viability and poor germination were primarily due to the large number of unfilled achenes and the many perigynia that contained no achene, Fungal damage and infestation by cecidomyidae larvae were also common, Many of the achenes that were filled contained a dead embryo,

Table 12, Germination rate and percent viability of commercially available Carex seed in tests conducted in June, 1991, one year after harvest

Year Percent Percent Species Harvested Source Viability Germination

Carex stricta 1990 SEW 1.0 0,5 C. atherodes 1990 SEW 1.3 0,3 C. comosa 1990 SEW 58,S 1.5 C. laclistris 1990 SEW 7,0 0,0

In the second study, started in December, 1991 (Table 13) 20-27% of Care x stricta and

lA-25,5 % of C. aqllatilis were viable, but only 7,6 and 2 % germinated, respectively, No seeds harvested, in 1990 germinated in December, 1991. Tetrazolium tests showed that many of the seeds that did not germinate were viable, Percent viability was as high as 64 and 69% for Carex laclIstris and C. atherodes, respectively (Table 13),

In the third germination/viability study, fresh seeds collected during the 1992 growing season were tested in July, 1992, Germination was good for all species tested, This was due in 51

Table 13. Germination rate and percent viability of Carex seed, determined December, 1991

year Percent Percent Species harvested source Viability Germination

Carex stricta 1990 SEW 5 0 1990 NIN 2 0 1991 SEW 21 5 1991 NIN 27 7.6 1991 NEI 20 0 C. atherodes 1990 SEW 12 0 1990 NIN 11 0 1991 SEW 1 0 1991 NIN 69 0 C. /asiocarpa 1991 NEI 0.5 0 C. rostrata 1991 NEI 0 0 C. aquatilis 1990 NIN 3 0 1991 NIN 1.4 1.4 1991 SEW 25.5 2 C. /acllstris 1990 NIN 16 0 1990 SEW 23 0 1991 NIN 17 0 1991 SEW 64 0

part to the fact that only filled achenes were tested. Viability was also high, except for Carex lacllstris (Table .14). Percent germination exceeded percent viability for two species, indicating problems with either the tetrazolium test, interpretation of the results or significant variability in the viability of different lots of seeds.

Seed storage Results of the first seed storage study are presented in Tables 15-21.

Initial percents viability and germinability were significantly greater in 1991 seed than 1990 52

Table 14. Percentage of seeds of Care x spp. collected in 1992 that were viable and germinable as tested July, 1992

Year Percent Percent Species Harvested Source Viability Germination

Carex aqllatilis 1992 NE2 64 33 C. stricta 1992 SEW 57 77 C. laclistris 1992 SEW 10 44

seed (Table 15). Percent viability was significantly less after storage than it was before.

Viability was reduced by on half. Percent germinability was not significantly different after storage than before. There were no significant differences in percent viability or in percent germinability between temperature treatments (Table 16). Percent viability was significantly less after storage than before. Percent germinability was not significantly different after storage than it was before.

Percent viability after storage was not significantly different between wet and dry

Table 15. Initial (Ini) and final (Fin) percent viability and germinability (mean ± s.d.) of seeds of Carex spp. collected in 1990 and 1991 in the first seed storage study.

Viability Germination

Year Ini* Fin Ini* Fin

1990 15.5 ± 4.8 7.8 ± 7.0** O.O± 0.0 0.7±2.1 1991 32.1 ± 23.6 15.9 ± 15.4** 2.3 ± 2.9 2.5 ± 4.4

*means significantly different between years (p ~ 0.05) **significant difference between initial and final means (p ~ 0.05) 53 treatments (Table 17). Percent germinability was significantly greater after wet than after dry storage. Final percent viability and percent germinability were significantly different than initial percent viability and percent germinability, respectively.

Percent viability was significantly less after storage for Carex lacustris, C. aquatilis and C. atherodes, and was not significantly different for Carex stricta (Table 19). Percent germinability was not significantly different for any species tested, averaged over all treatments. Neither percent viability nor percent germinability were significantly different after storage due to temperature treatment (Table 20). There was no significant difference between percent viability before and after storage for Carex aquatilis and C. stricta due to moisture treatment (Table 21). The only significant difference in percent germinability before and after storage due to moisture was seen in the dry treatment in Carex aquatilis and C. stricta.

Overall, due to large variability, there were no significant differences between either percent viability or percent germinability before and after storage (Table 21).

Table 16. Initial (Ini) and final (Fin) percent viability and germinability (mean ± s.d.) of seeds of Carex spp. stored under four different temperature regimes in the first seed storage study (N=22)

Viability Germination

Temp. Ini Fin Ini Fin

Room 26.1 ± 2l.0 14.1 ± 15.3* l.5 ± 2.6 2.0 ± 3.4 4°C 26.1±2l.0 14.3 ± 12.6* l.5 ± 2.6 3.0 ± 5.8 -4°C 26.1 ± 2l.0 9.6 ± 8.4* l.5 ± 2.6 1.1±1.9 Alt. 26.1 ± 21.0 13.8 ± 16.9* l.5 ± 2.6 1.3 ± 2.9

* significant difference between initial and final means (p =s; 0.05) 54

Table 17. Initial (Ini) and final (Fin) percent viability and germination (mean ± s.d.) of seeds of Carex spp. stored wet and dry in the first storage study (N=44)

Viability Germination

Moisture Ini Fin Ini Fin**

Wet 26.1 ± 20.7 14.3 ± 15.9* 1.5 ± 2.6 3.1 ± 5.0* Dry 26.1±20.7 11.6 ± 10.8* 1.5 ± 2.6 0.6 ± 1.1 *

*significant difference between initial and final means (p ::; 0.05) **treatment means significantly different (p ::; 0.05)

Table 18. Initial (Ini) and final (Fin) percent viability and germinability (mean ± s.d.) of seeds of Carex spp. stored in different temperature and moisture regimes in the first storage study (N= 11)

Viability Germination

Temp Moist Ini Fin Ini Fin

Room Wet 26.1±21.5 18.5 ± 18.5 1.5 ± 2.7 3.5 ± 4.3 Dry 26.1±21.5 9.7± 10.4* 1.5±2.7 0.5 ± 1.0 4°C Wet 26.1 ± 21.5 15.7 ± 13.3 1.5 ± 2.7 5.6 ± 7.4 Dry 26.1 ± 21.5 12.8 ± 12.3* 1.5 ± 2.7 0.3 ± 0.6 _4°C Wet 26.1±21.5 8.9 ± 8.0* 1.5 ± 2.7 1.4 ± 2.2 Dry 26.1±21.5 10.3 ± 9.0* 1.5 ± 2.7 0.7 ± 1.6 Alt. Wet 26.1 ±21.5 13.9 ± 21.2* 1.5 ± 2.7 1.9 ± 4.0 Dry 26.1±21.5 13.7 ± 12.3 1.5 ± 2.7 0.7±1.1

* significant difference between initial and final means (p ::; 0.05)

Results of the second seed storage study are presented in Tables 23-28. There were no significant differences between percent germination treatment means for Carex iaclIstris

(Table 23). Significant differences were observed between treatment means for Carex aqllatilis and C. stricta (Tables 24,25). High variabilities make interpreting treatment effects difficult, but significant differences were observed between some treatments (Tables 26-28). 55

Table 19. Initial (Ini) and final (Fin) percent viability and germinability (mean ± s.d.) for seeds of four Carex species in the first storage study

Viability Germination

Species Ini Fin Ini Fin

C. lacustris 29.9 ± 19.8 16.2 ± 12.7* 0.2 ± 1.4 1.0 ± 2.3 C. aquatilis 13.0 ± 13.4 6.3 ± 8.0* l.5 ± 0.5 2.0 ± 5.0 C. atherodes 30.7 ± 27.7 8.1 ± 15.2* O.O±O.O 0.1 ± 0.5 C. stricta 24.3 ± 3.6 20.7 ± 12.2 6.4 ± 1.6 6.1±4.9

*significant difference between initial and final means (p ~ 0.05)

Table 20. Initial (Ini) and final (Fin) percent viability and germinability (mean ± s.d.) of seed of four Carex species stored in different temperature regimes in the first storage study

Viability Germination

Temp. Ini Fin Ini Fin

Carex lacllstris Room 30.0 ± 21.2 21.9 ± 20.5 0.0 ± 0.0 l.3 ± 2.8 4°C 30.0 ± 2l.2 16.9 ± 11.0 0.0 ± 0.0 l.9 ± 3.6 -4°C 30.0 ± 2l.2 10.0 ± 5.9 0.0 ± 0.0 0.0 ± 0.0 Alt. 30.0 ± 2l.2 16.0 ± 8.8 0.0 ± 0.0 0.6 ± 0.9 Carex aquatilis Room 13.0 ± 15.0 4.8 ± 9.5 1.5 ± 0.6 l.3 ± l.9 4°C 13.0 ± 15.0 7.5 ± 9.0 l.5 ± 0.6 5.5 ± 9.7 -4°C 13.0 ± 15.0 8.5 ± 9.8 l.5 ± 0.6 0.8 ± l.5 Alt. 13.0 ± 15.0 4.3 ± 5.7 l.5 ± 0.6 0.5 ± l.0 Carex atherodes Room 30.7 ± 29.7 8.5 ± 8.3 0.0 ±O.O 0.3 ± 0.8 4°C 30.7 ± 29.7 6.8 ± 9.9 0.0 ± 0.0 0.0 ± 0.0 -4°C 30.7 ± 29.7 3.5±2.1 0.0 ± 0.0 0.2 ± 0.4 Alt. 30.7 ± 29.7 13.5 ± 28.7 0.0 ± 0.0 0.0 ± 0.0 Carex stricta Room 24.5 ± 4.0 16.5 ± 10.0 6.5 ± l.7 6.5 ± 4.8 4°C 24.5 ± 4.0 27.0 ± 13.1 6.5±l.7 7.0 ± 7.8 -4°C 24.5 ± 4.0 19.0 ± 10.4 6.5 ± 1.7 4.8 ± 1.0 Alt. 24.5 ± 4.0 19.5 ± 15.4 6.5 ± 1.7 5.5 ± 5.3

* si&nificant difference between initial and final means (p ~ 0.05) 56

Table 21. Initial (Ini) and final (Fin) percent viability and germinability (mean ± s.d.) of seeds of Carex species when stored under two different moisture regimes in the first storage study

Viability Germination

Moisture Ini Fin Ini Fin

Carex iaclistris wet 30.0 ± 20.5 17.0 ± 16.6* 0.0 ±O.O 1.6±3.1 dry 30.0 ±20.5 15.4±8.1* 0.0 ±O.O 0.3 ±0.7 Carex aquatilis wet 13.0 ± 13.9 7.3 ± 8.8 1.5 ± 0.5 3.8 ±6.7 dry 13.0 ± 13.9 5.3 ±7.5 1.5 ± 0.5 0.3 ± 0.7* Carex atherodes wet 30.7 ± 28.3 12.1±20.7* 0.0 ± 0.0 0.3 ± 0.6 dry 30.7 ± 28.3 4.1 ± 4.0* 0.0 ±o.o 0.0 ± 0.0 Carex stricta wet 24.5 ± 3.7 19.1±9.8 6.5 ± 1.6 9.6 ± 4.1 dry 24.5 ± 3.7 21.9±14.1 6.5 ± l.6 2.3 ± 1.5*

* significant difference between initial and final means (p ~ 0.05)

Grovvth studies

Greenhouse studies Results of the first greenhouse soil amendment study indicate that

all amendments are beneficial to growth of C. stricta to some extent (Table 29). Greatest

number of shoots was found on plants grown in the topsoil+fertilizer (T+ F) treatment,

, 57

Table 22. Initial (In i) and final (Fin) percent viability and germinability (mean ± s.d.) of seeds of Carex species stored under various temperature and moisture regimes in the first storage study

Viability Germination

Treatment Ini Fin Ini Fin

Carex faclIstris wet/room 30.0 ± 22.9 28.3 ± 26.2 0.0 ±O.O 2.5±3.7 wet/4°C 30.0 ± 22.9 16.8 ± 14.2 0.0 ±O.O 3.8 ±4.5 wet/-4°C 30.0 ± 22.9 8.8 ± 8.6 0.0 ± 0.0 0.0 ± 0.0 wet/a It 30.0 ± 22.9 14.3 ± 12.0 0.0 ± 0.0 0.3 ± 0.5 dry/room 30.0 ± 22.9 15.5 ± 13.7 0.0 ± 0.0 0.0 ± 0.0 dry/4°C 30.0 ± 22.9 17.0 ± 9.0 0.0 ±O.O 0.0 ±O.O dry/-4°C 30.0 ± 22.9 11.3 ± 1. 7 0.0 ±O.O O.O±O.O dry/alt 30.0 ± 22.9 17.8 ± 5.1 0.0 ± 0.0 1.0 ± 1.2 Carex aqllatilis wet/room 13.0 ± 18.4 9.5±13.4 1.5 ± 0.7 2.5 ± 2.1 wet/4°C 13.0 ± 18.4 9.0 ± 12.7 1.5 ± 0.7 10.0 ± 14.1 wet/-4°C 13.0± 18.4 8.0±11.3 1.5 ± 0.7 1.5±2.1 wet/alt 13.0 ± 18.4 2.5 ± 3.5 1.5 ± 0.7 1.0±1.4 dry/room 13.0 ± 18.4 0.0 ±O.O 1.5±0.7 0.0 ±O.O dry/4°C 13.0 ± 18.4 6.0 ± 8.5 1.5 ± 0.7 1.0±1.4 dry/-4°C 13.0± 18.4 9.0±12.7 1.5±0.7 0.0 ± 0.0 dry/alt 13.0 ± 18.4 6.0 ± 8.5 1.5±0.7 0.0 ± 0.0 Carex atherodes wet/room 30.7 ± 33.2 10.7 ± 10.7 0.0 ± 0.0 0.7 ± 1.2 wet/4°C 30.7 ± 33.2 9.7 ± 14.2 0.0 ±O.O 0.0 ± 0.0 wet/-4°C 30.7 ± 33.2 3.3 ± 1.5 0.0 ±O.O 0.3 ±0.6 wet/alt 30.7 ± 33.2 24.7 ± 41.0 0.0 ±O.O 0.0 ±O.O dry/room 30.7 ± 33.2 6.3 ±6.7 0.0 ±O.O 0.0 ±O.O dry/4°C 30.7 ± 33.2 4.0 ± 4.4 0.0 ±O.O 0.0 ±O.O dry/-4°C 30.7 ± 33.2 3.7 ±2.9 0.0 ±O.O 0.0 ±O.O dry/alt 30.7 ± 33.2 2.3 ± 1.5 0.0 ±O.O 0.0 ±O.O Carex stricta wetlroom 24.5 ± 4.9 20.0 ± 14.1 6.S±2.1 10.5±2.1 wet/4°C 24.5 ± 4.9 29.5 ± 3.5 6.5 ± 2.1 13.5 ± 3.5 wet/-4°C 24.5 ± 4.9 18.5 ± 3.5 6.5 ± 2.1 5.5 ±0.7 wet/alt 24.5 ± 4.9 8.5 ± 2.1 6.5 ± 2.1 9.0 ± 5.7 dry/room 23.5 ± 4.9 13.0±7.1 6.5 ± 2.1 2.5 ± 0.7 dry/4°C 24.5 ± 4.9 24.5 ± 21.9 6.5 ± 2.1 0.5 ± 0.7 dry/-4°C 24.5 ± 4.9 19.5 ± 17.7 6.5±2.1 4.0 ± 0.0 dry/alt 24.5 ± 4.9 30.5 ± 14.8 6.5±2.1 2.0 ± 1.4

*significant difference between initial and final means (p s 0.05) 58

Table 23. Percent germinability (mean ± s.d.) offresh seeds of Carex laclistris stored under various temperature and moisture regimes in the 1992-1993 storage study and tests for significant differences between treatments (T-test) (Initial germinability was 44.0%)

Mean Treatment

da dc df dr da dc df dr rna mc mf mr wa wc wf wr

0.0 ±O.O dry/alt 1.5 ± 0.7 dry/cold 1.0 ± 1.4 dry/froz 10.0 ±4.2 dry/room 0.0 ±O.O dung/alt 0.0 ±O.O dung/cold 0.0 ±O.O dung/froz 0.5 ±0.7 dung/room 0.0 ±O.O muck/alt 0.5 ± 0.7 muck/cold 0.0 ± 0.0 muck/froz 5.5 ±0.7 muck/room 0.5 ±0.7 wet/alt 5.0 ± 0.0 wet/cold 0.0 ± 0.0 wet/froz 15.5 ± 7.8 wet/room

*means significantly different (p :::; 0.05)

followed by fertilizer + compost (F+C), topsoil (T) and topsoil+compost (T+C) treatments.

Above ground dry mass was highest in the topsoil+fertilizer+compost (T+F+C) and topsoil+compost (T+C) treatments. All the other soil amendments also significantly increased aboveground biomass. Belowground mass likewise was highest in treatments in which topsoil was added, but all treatments with amendments increased belowground mass significantly. The main treatment effects of the soil amendments (Table 30) are all statistically significant.

The second greenhouse study tested the effect of percent compost on growth of Carex stricta. Above ground, below ground and total dry mass increased linearly with percent compost (Tabtes 31, 32, 33). However, due to large variances, differences in total dry mass 59

Table 24. Percent germinability (mean ± s.d.) offresh seeds of Carex aquatilis stored under various temperature and moisture regimes in the 1992-1993 storage study and tests for significant differences between treatments (T-test) (Initial germinability was 33.0%)

Mean Treatment

da dc df dr da dc df dr rna mc mf mr wa wc wf wr 63.5 ± 4.9 dry/alt * * * * * * * 54.5 ± 19.1 dry/cold * 77.5 ± 17.7 dry/froz 22.0 ± 1.4 dry/room * * * * * * * O.O±O.O dung/alt * * * 1.5 ± 2.1 dung/cold * * 0.0 ± 0.0 dung/froz * * * 1.5 ± 0.7 dung/room * * * 0.0 ± 0.0 muck/alt * * * 37.5 ± 0.7 muck/cold * * 1.5 ± 2.1 muck/froz * 19.0 ± 14.1 muck/room 0.5 ± 0.7 wet/alt * * 58.0 ± 4.2 wet/cold * 5.0 ± 1.4 wet/froz * 53.0 ± 2.8 wet/room

*means significantly different (p ~ 0.05)

were not significant for treatments of33% percent compost or greater nor for above ground dry mass for 50% compost or greater.

Field studies Results of field tests of the effects of soil amendments on growth of Carex stricta are presented in Tables 34 and 35. Numbers of shoots were significantly greater than than in the control treatment in the compost, fertilizer, compost+fertilizer and topsoil+compost treatments.

The effect of soil amendments on establishment of species in the field soil amendment study is summarized in Table 36. Treatment effect means are contrasted in Table 37. The species recrutted were placed into two categories, ruderals and species that were in the seed -60

Table 25. Percent germinability (mean ± s.d.) offresh seeds of Carex stricta stored under various temperature and moisture regimes in the 1992-1993 storage study and tests for significant differences between treatments (T-test) (Initial germinability was 77.0%)

Mean Treatment

da dc df dr da dc df dr rna mc mf mr wa wc wf wr 11.5±6.4 dry/alt * 39.5 ± 12.0 dry/cold 26.5 ±4.9 dry/froz * 10.0 ± 1.4 dry/room * * * 4.0±2.8 dung/alt 11.5±0.7 dung/cold * 1.0 ± 1.4 dung/froz * 11.5 ±4.9 dung/room 0.5 ±0.7 muck/alt * 11.5 ± 4.9 muck/cold 3.5±2.1 muck/froz * 6.0±4.2 muck/room 8.0±2.8 wet/alt * 40.0 ± 17.0 wet/cold S.0±2.8 wet/froz * 35.5 ± 0.7 wet/room

*means significantly different (p ::; 0.05)

Table 26. Percent germinability (mean ± s.d.) of fresh seeds of Carex species stored under various moisture regimes in the 1992-1993 storage study and tests for significant differences between treatments (T-test)

Mean Treatment

Muck Dung Water Dry 7.2± 11.4 Muck * 2.4 ± 4.5 Dung * * 16.8 ± 21.2 Water 23.7 ± 26.4 Dry

*significant diffet"ences between means (p ::; 0.05) 61

Table 27. Percent germinability (mean ± s.d.) offresh seeds of Care x species stored under various temperature regimes in the 1992-1993 storage study and tests for significant differences between treatments (T-test)

Mean Treatment

AIt Cold Froz Room 7.3 ± 17.8 AIt * 23.0 ± 23.8 Cold * 10.1 ± 22.3 Froz 15.5 ± 15.6 Room

*significant differences between means (p ::;; 0.05)

Table 28. Percent germinability (mean ± s.d.) offresh seeds of Carex species stored under various moisture and temperature regimes in the 1992-1993 storage study and tests for significant differences between species (T -test)

Mean Treatment

C.aquatilis C. lacus/ris C. stricta 24.7 ± 28.0 C. aquatilis * * 2.5 ±4.7 C. lacus/ris * 15.0 ± 15.7 C. stricta

*significant differences between means (p ::;; 0.05)

mixes sown. The total cover of seed mix species was very low (1 to 10%) and increased only slightly in the soil amendments, particularly topsoil amendment treatments. The majority of the cover in seed mix species was due to Echinochloa crusgalli. Ruderal cover (5 to 40%) increased significantly in treatments with soil amendments, particularly those containing topsoil. This is prewmably due to ruderal seeds being added with the soil and better growing 62 conditions in these treatments. In general, percent cover of both volunteer and seed mix species was significantly greater in plots treated with compost and in plots treated with topsoil than in plots without those treatments (Table 37). Percent cover of volunteer species was greater in plots receiving fertilizer treatment than in plots receiving no fertilizer. There was not a significant difference in percent cover of seed mix species in plots treated with fertilizer and those without fertilizer.

Table 29. Growth of Care x stricta seedlings in soil (Control) from the DPRWDP amended with compost (C), fertilizer (F), topsoil (T) or some combination of them under greenhouse conditions. (All treatment means were significantly different from the control mean at the 0.05 level.)

Shoots/plant Biomass (g) ± s.d.

Treatment mean ± s.d. above below

Control 3.3 ± 1.3 0.6 ±0.3 0.7 ± 0.3 F 5.4 ± 1.9 1.3 ± 0.5 2.3 ± 1.3 C 6.4 ± 2.2 1.6 ± 0.5 1.6 ± 0.6 T 8.8 ±2.3 1.6 ± 0.6 3.0 ± 1.4 F+C 8.9 ±2.9 1.6±0.4 2.5 ± 0.7 T+F 9.6 ±4.3 1.9 ± 0.6 2.8 ± 1.0 T+C 8.3 ±2.6 2.1 ± 0.5 2.8 ± 1.0 T+F+C 7.7 ± 2.9 2.2 ±0.8 2.7±1.5 63

Table 30. Mean number of shoots per plant and mean above- and below-ground shoot biomass of Carex stricta in the greenhouse soil amendment treatments with (+) and without (-) topsoil, fertilizer, and compost.

Shoots/plant Biomass (g) ± s.d.

Treatment Mean ± s.d. above below

- topsoil 6.0± 2.9 1.3 ± 0.6 1.8 ± 1.0 + topsoil 8.6±3.1* 2.0 ± 0.7* 2.8 ± 1.2*

- fertilizer 6.7 ± 3.0 1.5±0.7 2.1 ± 1.3 + fertilizer 7.9 ± 3.4* 1.8 ± 0.7* 2.6 ± 1.2*

- compost 6.8 ± 3.6 1.3 ± 0.7 2.2 ± 1.4 + compost 7.8 ± 2.7 1.9 ± 0.6* 2.4 ± 1.1

* Means significantly different at 0.05 level.

Table 31. Mean final above-ground, below-ground, and total mass of Carex slricta in response to percent compost

Compost (%) Biomass (g) ± s.d.

Above Below Total

o 0.02 ± 0.01 0.02 ± 0.03 0.04 ± 0.04 10 0.12 ± 0.05 0.11 ± 0.07 0.23 ± 0.12 20 0.13 ± 0.07 0.10 ± 0.06 0.23 ± 0.13 33 0.26 ± 0.12 0.20 ± 0.12 0.45 ± 0.23 50 0.33 ± 0.15 0.25 ± 0.15 0.58 ± 0.28 67 0.32 ± 0.17 0.19 ± 0.11 0.52 ± 0.27 80 0.42 ± 0.23 0.24±0.15 0.67 ± 0.38 90 0.46±0.18 0.27 ± 0.15 0.73 ± 0.31 100 0.44 ± 0.28 0.29 ± 0.25 0.73 ± 0.52 64

Table 32. T-Statistics for significantly different treatment means at the 0.05 level from percent compost soil amendment study (see Table 25 for means)

% Compost 100 90 80 67 50 33 20 10 0

Aboveground Biomass 100 3.2 4.0 5.1 90 2.9 4.9 5.5 7.3 80 2.3 3.6 4.6 5.9 67 3.5 3.4 5.1 50 2.5 6.0 6.0 8.4 33 2.7 2.8 6.3 20 4.6 10 6.9 Belowground Biomass 100 3.9 90 4.2 3.1 4.8 80 2.8 3.3 4.6 67 -2.4 3.4 3.0 4.8 50 3.9 5.4 5.9 33 4.1 20 3.7 10 3.5 Total Biomass 100 2.8 3.3 4.8 90 4.8 4.4 6.3 80 3.4 4.3 5.5 67 3.6 3.3 5.0 50 5.4 6.7 8.0 33 2.6 2.4 5.4 20 4.4 10 5.1 65

Table 33. Linear regressions of dry mass to % compost

Equation

aboveground mass = 0.0042 (% compost) + 0.0677 R2= 0.94 belowground mass = 0.0023 (% compost) + 0.0695 R2= 0.84 total mass = 0.0065 (% compost) + 0.1372 R2= 0.92

Table 34. Initial and final number of shoots (mean ± s.d.) of Carex stricta plugs planted in the field in various soil amendment treatments (compost=C; fertilizer=F; topsoil=T)

Shoot Density Treatment Initial Final

Control 5.1 ± 2.3 11.8 ± 5.0 C 4.7 ± 2.9 15.2 ± 9.9* F 5.8 ± 2.8 14.0 ± 7.4* T 4.8 ± 2.8 12.2 ± 6.6 C+F 5.2 ± 3.2 16.9 ± 10.4* T+C 5.8 ± 3.3 15.5 ± 8.4* T+F 5.0±2.9 11.5± 6.0 T+C+F 4.7 ± 2.7 14.3 ± 9.7

* Mean significantly different from control mean at the 0.05 level.

Table 35. Treatment means of final shoot number of Carex stricta plugs in different soil amendment treatments in field tests

Treatment Mean± s.d. T -statistic

-topsoil 14.5 ± 8.7 +topsoil 13.4 ± 8.0 1.6

-compost 12.4 ± 6.4 +compost 15.5±9.7 4.5*

-fertilizer 13.7± 7.9 +fertilizer 14.2 ± 8.8 0.8

* t-test significantly different at p <0.05 66

Table 36. Total cover (mean % ± s.d.) ofruderals and species in the seed mix for various soil amendment treatments on plots without Carex stricta plugs (compost=C; fertilizer=F; topsoil=T)

Treatment Weeds Seed Mix Total

Control 3.4 ± 4.2 1.7±3.0 S.1 C 9.7 ± 10.7 3.2 ± 3.3 12.9 F 7.8 ± 10.6 1.2 ± 2.0 9.0 F+C 13.3± 7.1* 6.0 ± 6.6* 19.3 T 7.6± S.9 1.8±1.7 9.4 T+C 24.8 ± 10.7* 6.8 ± 4.2* 31.6 T+F 17.7± IS.7* 3.3 ± 2.6 21.0 T+F+C 34.2 ± 17.S* 6.1 ± 3.8* 40.3

* means significantly different from control mean at p $ 0.05

Table 37. Treatment means ± s.d. of percent cover of volunteer (weeds) and seed mix species response to soil amendments compost (C), fertilizer (F) and topsoil (T)

Treatment Weeds T -statistic Seed Mix T -statistic

+C 21.0 ± IS.2 4.9* S.6± 4.7 4.7* -C 9.1±11.2 2.0 ± 2.4

+F 18.5±16.4 2.8* 4.2±4.S 1.1 -F 11.S ± I1.S 3.4±3.7

+T 21.1 ± 16.2 4.8* 4.5 ± 3.7 2.1 * -T 8.S ± 8.9 3.0 ± 4.S

*t-test significantly different at p <0.05 67

GENERAL DISCUSSION

Comparison oj natural and created sedge meadows

Water table levels During the first growing season following their construction, water table levels in the DPRWDP experimental wetlands did not resemble water table levels observed in natural sedge meadows. The differences were likely due to differences in the source of water and in the nature of the substrates. In fact, water table level may not be the best indicator of functional hydrology in natural sedge meadows. Finally, monitoring wells may not have functioned properly in the experimental wetlands, producing erroneous indications of water table level.

Water tables in the natural sedge meadows showed signs of seasonal fluctuations that were controlled by the interaction of groundwater discharge, precipitation and evapotranspiration. The influence of above and below normal precipitation on water table levels depends on the status of the vegetation, which depends on time of year. Water tables in the experimental wetlands seemed to be independent of seasonal precipitation and vegetation influences, and were controlled by the rate of pumping, elevation of stoplogs at outlets and variability of infiltration rates of substrates.

Large fluctuations in water table levels in the experimental wetlands were the result of changes in pumping or changes in elevation of the stoplogs at EW outlets (Figures 14-19). In general, except for EWs Ie and 2D, water levels in the EWs do not reflect intense rains that fell in July and September. The outlet design causes water in the EWs to spill out when it reaches the height of the stoplogs, limiting the maximum depth of standing water possible.

Water tables depths observed in the wells were the product of these controlled factors combined with substrate differences, and other, less well known or unknown factors. The variability during the first weeks of record is due to the effect offirst introducing water into the EWs . As the EWs were first filled, an unknown quantity of water was absorbed initially 68 by soils, and by bank storage in the berms.

Distance from inlets also had an observable effect on water tables. The inlets pumping water into the complex are located in EW lA, IB and IC. These three EWs were the first to fill when pumping began in June. (There was some problem with pumping into EWIB which prevented its filling with the others.) The fluctuation patterns of these three EWs were fairly

similar through the rest of the season, reflecting their similar location nearest the inlets.

Differences between water tables in the three wetlands are likely due to substrate or elevation differences, or both.

EW 2B, 2C and 2D are located in the southern tier ofEWs. They, too, exhibited similar responses through the season. Again, this reflects their similar positions, i.e., distant from the water pumps. It simply took longer for water to reach those EWs, and weeks for them to fill.

Their water table responses generally lagged behind those of the northern tier ofEWs.

Differences between water tables in the three wetlands are likely due to substrate or elevation differences, or both.

Other factors interacted in unknown ways to contribute to the water table depths

observed. Sediments in the EW s are a mosaic of past deposition and erosion events, as the

Des Plaines River flooded and shifted its channel. Substrates vary both horizontally and vertically, ranging from coarse sand to clay in very short distances. This textural variability within and among EWs likely contributed to differences in infiltration and water retention and in lateral and vertical water movement that were reflected in water table measurements.

Sediment texture could also permit groundwater recharge or discharge in sandier areas and' cause water perching in clayey areas, with unknown effects on observed water tables. In some cases, water was observed standing on the surface at a well, while the water in the well was inches below the surface. In other places, the reverse was seen. A hole probed 10 inches deep remained dry indefinitely, despite its location next to a well with water at 6 inches below the 69 surface. Soil surfaces were often dry and cracked despite a relatively shallow water table.

Other factors potentially influencing observed water tables in unknown ways were precipitation, water levels in the Des Plaines River, and flow from surrounding uplands, including the deep-water experimental wetlands upslope from the sedge meadow EWs.

The well design itself may have created false indications of water table where it did not occur. Faulkner et al. (1989) attributed erroneous water table measurements in these types of wells to channelized flow downward in vertically oriented pores. Water accumulating in a well would indicate a water table when surrounding soil was unsaturated. In addition, with no seal around the wells at the soil surface, precipitation and surface runoff could enter the wells.

Regardless of the cause, water tables in the created sedge meadows did not resemble those of natural sedge meadows in 1992.

Interpreting the water table fluctuations in the natural sedge meadows is complicated by the lack of information about the components of the hydrologic budget for any of the sites.

Basic information about the source of water, groundwater or surface runoff, is not known. It was assumed that groundwater discharge was the primary source in sites known as fens

(Turner Lake Fen and Spring Grove Fen), but the relationship of areas of groundwater disharge to the locations of water table wells is unknown. The source of water in the other sites is also unknown. They may be groundwater or surface water fed slope or depressional wetlands (Novitzki, 1982) or a combination of these. A more thorough study made throughout the year, including use of piezometers and water chemistry observations, could have shed some light on this most basic wetland feature.

The hypothesized hydrologic regime includes fairly steady rates of groundwater discharge throughout the year, unpredictable periods of above and below normal precipitation and associated surface runoff, and a strong capability for drawdown by vegetation. Resultant water tables reflect the balance of these factors. During 1992, water tables in the natural sedge 70 meadows were above the surface (i.e. had standing water) from the time observations began in early March until mid- to late- May (Figures 3-7). This may have been due to heavy precipitation in October and November of the previous year (Figure 8) combined with steady groundwater discharge and low or non-existent evapotranspiration from late fall to early spring. The rapid decline in water tables observed from late May to early July corresponded to a period of rapid growth of vegetation and the cumulative effect of seven consecutive months of below normal precipitation (Figure 8), particularly in April, May and June. Short-lived rises in water table levels in July and September were associated with above normal precipitation during those months. Many of the sedge meadows are in a position to receive surface runoff from large areas of adjacent uplands in addition to the precipitation falling on the wetlands themselves. High evapotranspiration rates in July were more efficient at drawing down these pulses of water than was evapotranspiration in September.

Carter et al. (1978) reported that water loss by evapotranspiration of sedges and willows was 10-30% higher than open water evaporation in Minnesota peatlands. Other researchers have noted the ability of wetland vegetation to draw down water table levels measurably.

Williams (1968) observed that larger, vegetated wetland basins could draw down water tables in adjacent uplands during dry periods to the extent that when it did finally rain, a groundwater mound formed under the wetland, reversing the hydraulic gradient and causing groundwater recharge from the wetland. Owen et al. (1989a,1989b) described the drawdown of water tables by evapotranspiration as "strong" in Wisconsin wetlands during the growing season. They found that the water table was drawn down sooner in the spring and rose more slowly in the fall in wetlands populated by well-established sedges and other vegetation than in restored wetlands with less well established vegetation. Carpenter (1990) observed a diurnal pattern of drawdown in Wisconsin fens. Daily drawdowns of up to 8cm were recorded in sedge meadows at the base of the fens. Water tables returned to their originalleve\s after 71 sunset. It is unknown whether time of day water table measurements were made had any influence on the resultant water table record in the present study.

In the only comparable modern study of sedge meadow hydrology in northeastern Illinois,

Kelsey and Hootman (1992) observed water tables in a wet prairie/sedge meadow complex in

DuPage County, Illinois, in 1990 and 1991. They used lined boreholes of PVC pipe (not perforated as they were in the present study) to observe water tables. During 1990, a year with above average precipitation, they observed very stable water tables punctuated by short duration rises in response to rainy periods. During 1991, a year with an extended period of below average precipitation, they observed a rapid decline in water tables during the growing

season, followed by a return to near-surface levels in the autumn. Their observations during the dry year are similar in shape to the depth/time graphs of 1992 in the five sedge meadow

sites in the present study. Their site's steep water table decline occurred about five weeks

later, however, than it did in the 1992 observations of the present study.

Interestingly, in their study, water tables only rose above the surface in their sedge

meadows on two occasions, for only a few days at a time. Both occurred during the spring of

1991. This indicates differences in hydrology at the two groups of sites, differences that are

reflected in soils present. The DuPage County sites are on Typic Haplaquoll soils, not

Histosols as were most of the five sedge meadow sites in the present study. Although their

sites have an aquic moisture regime, they are not as saturated as long or as often as would be

indicated by the presence of Histosols. It may be possible to use information about the soils

present to predict hydrologic regimes at a site or to identify sites suitable for sedge meadows.

The sedge meadows in the present study were predominantly on muck, primarily

Houghton Muck, a Typic Medisaprist. A muck soil contains 20-50% organic matter. The

Saprist (= muck) suborder of Histosols includes the most decomposed of Histosols, with plant

parts not recognizable. They have a fine texture, which can be powdery when dry (Brady, 72

1984). The fine texture of muck provides a greater proportion of capillary pores in a given volume of soil than in coarser textured soils or less well decomposed peats. This capillarity is

sufficient to create a very wide tension-saturated zone, or capillary fringe above the water table (Paivanen, 1973). This zone is defined by Heliotis and DeWitt (1987) as" the region

immediately above the water table where pores are fully saturated but the pressure head is

negative, indicating that water is held in place by surface-tension forces." The height of the water column, or thickness of the capillary fringe, is inversely proportional to the pore size,

which in muck soils is quite small.

Few data are available on reported thicknesses of capillary fringes in muck soils. Paivanen

(1973) reported that capillary rise in peat increased with bulk densitiy. He reported capillary

rise in undecomposed peats ranged from 15-30 cm. The greatest bulk density he gave figures

for was 0.156 g. m- 3, with a capillary rise of 70 cm. Brady (1984) reports that Saprists have

the greatest bulk densities of Histosols, 0.2 g • m- 3 or more. Extrapolating from Paivanen

(1973) gives a capillary rise of about 89 cm for a soil with a bulk density of 0.2 g • m- 3 . This

would be sufficient to keep soils in the sedge meadows saturated to the surface when water

tables fell as low as 89 cm below the surface. While water tables on adjacent uplands (e.g.

Turner Lake # 1, and Des Plaines River Trail #6) were deeper than this at times, none of the

water table depths in sedge meadow vegetation ever fell below 75-80 cm in 1992. Owen

(1989a, 1989b) estimated capillary fringes in the Wisconsin wetlands they studied to be

"several decimeters" thick. This was sufficient to keep the rooting zone saturated, via its

connection with the water table, through the summer.

It is this capillary fringe that keeps sedge meadow soils saturated to the surface, even

when water tables are 80-90 cm deep. In fact, water table depth is not as relevant an indicator

of the effective hydrology of sedge meadows as is depth to saturated soils. Replicating natural

sedge meadow water table depths in created sedge meadows will not replicate natural sedge 73 meadow hydrology, unless the created sedge meadows have a comparable substrate, i.e. muck. In addition, the hydrologic regime must account for differential rates of drawdown by evapotranspiration through the year.

Despite their disparate locations, landscape settings and potential water sources, water table depths in the natural sedge meadows were surprisingly similar. Data from Wadsworth

Prairie (Figure 3) and Spring Grove Fen (Figure 6) are almost identical. Their water table maxima and minima are similar, as are the timing of fluctuations. These two sites occupy very similar positions in the lanscape, on broad, flat alluvial terraces along the Des Plaines River

(Wadsworth Praie) and Nippersink Creek (Spring Grove Fen). They are protected from all but the most extreme flooding by natural levees. It is possible these two sites represent a broad category of sites suitable for sedge meadow development in the region: the abandoned alluvial terrace, often on the inner bend of stream meanders, in former oxbow lakes or backswamps.

High morainal bluffs on either side of the flood plain could be the recharge area for local groundwater flows discharging as springs near the sedge meadows. These may be classified as groundwater depression or surface water depression wetlands (Novitzki, 1982).

A second class of sites suitable for sedge meadow formation may be represented by the

Des Plaines River Trail and Lyons Prairie sites (Figures 4 and 7). They, too, had similar water table maxima and minima, and timing of water tables rising and falling above and below the surface. These sites are both located at the bases of morainal bluffs, at the margin of flood plains of the Des Plaines (Des Plaines River Trail) and Fox Rivers (Lyons Prairie). These might be groundwater slope or surface water slope wetlands (Novitzki, 1982).

A third class of sites may be represented by Turner Lake Fen. These would be surface water slope wetlands (Novitzki, 1982) associated with lake margins. The water tables at the site seem to be linked to water levels in Turner Lake (Figure 5). Fluctuations are buffered by the large lake volume. Drainage ditches dug near the wells connect the sedge meadow to the 74 lake, assuring continuity in their fluctuations.

The sedge meadow sites in different landscape positions share the feature of being low energy wetlands, hydrodynamically. The highly organic soils indicate that sedge meadows are not exposed to high energy flooding. Accumulation of organic matter is an indicator of hydrologic energy status. Organic matter was observed to increase in freshwater marshes in

New Jersey on a gradient from actively flooded stream banks to less actively flooded inland high marshes. In coastal marshes, flooding frequency was seen to be directly related to organic export (Gosselink and Turner, 1978).

Stratigraphy Analysis of stratigraphy of substrates (Figures 9-13) at the natural sedge meadows reveals that depth of peat or muck is quite variable, ranging from 30 cm

(Wadsworth Prairie #2) to over 170 cm (Spring Grove #5). Underlying the muck is either sand, silt or blue clay. These are similar to stratigraphies described by Owen et al.

(1989a, 1989b). Those authors emphasized the role stratigraphy plays in maintenance of hydrology in natural wetlands, and the problems caused by improper stratigraphy in restored wetlands. Particularly problematic were areas with sand intermixed with peat. This interrupted the capillarity of the peat, causing surface soils to be too dry.

The substrates in the created sedge meadow EWs at DPRWDP in 1992 did not resemble those observed in the natural sedge meadows. Natural sedge meadow substrates were organic to the surface. Those in the created sedge meadows were generally mineral to the surface. Soil textures ranged from clay to silt, sand and gravel, with an organic matter gradient increasing toward the river, representing the remains of" A" horizons of soils not stripped during construction. When not flooded or saturated to the surface, substrates were best described as hard and dry. In sandy areas, substrates became unconsolidated when wet, as evidenced by the filling of well #2 in EW 2B with wet sand that flowed into the well as rapidly as it was cleaned out. Owen et al. (1989a,1989b) noted substrate hardness was 2-3 times greater in the created 75

wetlands they evaluated than in natural wetlands. They stressed that improper substrates

caused problems for vegetation due to surface conditions that were too dry and too hard.

O'Brien (1977) described the effect of substrate organic content on soil temperatures. He

noted that saturated, highly organic soils would be slower to warm up in the spring, remain

cooler in the summer and stay warm longer in the fall than unsaturated, inorganic mineral

substrates, due to the specific heat of water in the muck.

Amending the soil in the created sedge meadows with organic matter may be essential, or

at least highly beneficial, for developing the proper soil moisture, temperature and textural

conditions necessary to establish sedge meadow vegetation. Mulching with straw, hay or even

sawdust is a standard practice in mine reclamation to control erosion, modify soil

temperatures, prevent soil crusting and increase water availability (Lyle, 1987). Roberts et at.

(1988) working in the area of reclamation of surface mined lands, report that sludges, paper

mill wastes, sawdust and other organic amendments have been used as alternatives to standard

fertilizers. They all increased water holding capacity, cation exchange capacity, soil

aggregation and organic matter content of mine soils.

Vegetation sampling A comparison of the vegetation in natural sedge meadows with

that in the created sedge meadows in 1992 reveals that only two species, Verbena hastata and

Physostegia virginiana, were present in both. The vegetation in the natural and created sedge

meadows could be described as dissimilar. Vegetation cover in the created sedge meadows

was a sparse mixture of ruderals and matrix species (Tables 8-10). The site presented itself as

highly disturbed, and was invaded by opportunistic species common to disturbed areas,

. including Cyperus esclIlelltllS, Ambrosia artemisifolia and Lolium perellne. The coarse, hard,

dry, inorganic nature of the substrates undoubtedly played a role in the composition of the

vegetation in the created sedge meadows in 1992. This was combined with a lack of

recruitment from seed mix species. In addition, most of the EW s were flooded soon after seed 76 was sown. This may have washed away seed of mix species, drowned seedlings or prevented germination altogether. The failure of seed mix species to have established was either lack of germination or failure of seedlings to survive until the vegetation survey in August. Personal observations suggest that lack of establishment was due to low or nonexistent germination rates of seed mix species, particularly of Carex species.

Schramm (1978, 1990), working in the field of prairie restoration, identified two major variables that influence the success of plantings: weed seed present at the site and the degree of prompt germination of seed planted. These are the the very same factors that caused the failure to establish sedge meadow species in the experimental wetlands. Schennum (1990) has pointed out that many prairie species find refuge in sedge meadows in northeastern Illinois.

This emphasizes that sedge meadows have both wetland and upland characteristics. Perhaps, from the standpoint of the vegetation, sedge meadows should be seen as more like prairie than marsh. Vegetation establishment in created and restored sedge meadows may benefit from the knowledge gained and techniques developed in three decades of prairie restorations. The field of sedge meadow creation and restoration should not be making the same mistakes the prairie restorationists made thirty years ago.

For weed control, Schramm (1978, 1990) recommended allowing weed seeds to germinate, followed by shallow cultivation or herbicide application two or more times from

April through June, before planting. This could also be accomplished in the experimental wetlands by shallow flooding to kill weed seedlings (Fredrickson and Taylor, 1982). Schramm

(1978, 1990) also recommends annual spring burning, beginning the first spring after planting.

More optimistically, Schramm (1978, 1990) reported that in prairie restorations, apparently ungerminated seed has often shown up as plants years after planting.

Sedge establishment studies

Seed germination It is likely that low germination rates of seeds of Carex species 77 hampered efforts to establish sedge meadow vegetation using seed. The low germination rates of Carex seeds may be due to improper germination conditions, dormancy, low rates of viable seed production, loss of viability of seeds in storage, or a combination of these. The present study did not attempt to determine the most appropriate conditions for germination of the seeds evaluated. Results of germination and viability studies do not suggest that low germination rates were due to improper germination conditions. Gillespie (1989) and Kerans

(1990) note the highest germination rates of Carex stricta seeds were obtained on saturated soils at 10c C or higher in sunlight. These are similar to conditions in the greenhouse during germination tests in the present study.

Relatively high germination rates in freshly harvested Carex seeds in 1992 indicates seeds of these species were not dormant when mature. Some of them may require several weeks of afterripening, however. Seeds continued to germination until about eight weeks after seeds were sown, suggesting some time was needed foHowing harvest for seeds to mature fully.

The Carex species studied are perennials, reproducing almost exclusively vegetatively in nature. It should not be surprising that seeds of these species are unreliable as propagules.

Many Carex species have been observed to have very little sexual reproduction. Verhoeven et af. (1988), working at a site in The Netherlands, reported flowering shoot production by

Carex rostrata of about 20%, and that of C. diandra of 35%. C. acuti/ormis remained vegetative throughout their two year study. Shaver et al. (1979) found a population of Carex aquatilis in Alaska that persisted only by vegetative reproduction. van der Valk and Davis

(1979) observed that annual seed production in Carex atherodes was nearly zero in a prairie glacial marsh. Bernard (1975) states that he had never seen a seedling of Carex iacustris or

Carex rostrata in more than three years of working with the species. Costello (1936) wrote that he had not seen a seedling of Carex stricta in six years of studying the species in

Wisconsin. He described the number of fruiting specimens seen during that time as 78

"comparatively few" for the thousands of tussocks observed. He was also unsuccessful at all attempts to germinate Carex stricta seeds in the lab.

Explanations ranging from environmental to genetic have been offered for low rates of flower and seed production in sedge species. As that subject was not pursued in the present study, it is unknown which, if any may apply to the species and locations studied. Grootjans and van Tooren (1984) found that reduced groundwater discharge had a negative influence on the development of inflorescences in Carex aqllatilis in The Netherlands. They also observed that a dry period in the spring preceded withering of flowers before maturity. They noted that perigynia of Carex aquatilis collected in The Netherlands "have only been found to contain undeveloped fruits." They attributed this to Carex aqllatilis being at the southern edge of its range in The Netherlands, where climatic conditions may prevent regular fruit production.

Bernard (1975) described the life history of Carex lacllstris in New York. He suggested that water depth the previous season had a role in flower production. That species initiates flower primordia in October. Water depth influences shoot length, and flowers do not form on shoots that are too short. If shoots are too long, they die during the winter. Therefore, water depth during the growing season one year influences flowering the next spring. Smith (1973) found that high water levels in the spring promoted flowering of Scolochloajestllcacea. Apparently flower primordia may be damaged by late freezes in the spring if they are not submersed.

A possible reason for poor seed quality and quantity, suggested by Naylor (1984), is that more seeds are initiated than can be supplied with reserves. Similarly, seed abortion in ryegrass was attributed to high nitrogen inputs, which stimulated vegetative tiller growth at the expense of seed development. Carex aqllatilis, Carex facustris and Carex rostrata produce the majority of their new shoots after seed production is complete. In this way they avoid competition for resources between vegetative and sexual reproduction (Bernard and

Gorham, 1978 ; Bernard, 1975). 79

Poor seed set in the Carex species studied may involve self-incompatibility or self­ infertility. Aspinwall and Christian (1992) investigated the causes of poor seed set (in spite of high flower production) in the perennial, rhizomatous, herb Filipendula rllbra. They found that most of the individuals at the site belonged to the same clone. The species is self­ incompatible, so most pollination was between incompatible clones, resulting in very little seed set. They believed this supported Handel's (1985) hypothesis that "seed set in clonal, self­ incompatible species should decrease as clone size increases."

It is unknown if Carex stricta, Carex aquatilis and Carex lacus/ris are self-incompatible.

Handel (1978) reports two woodland sedges, Carex plantaginea and C. platyphyUa, are self­ compatible and will set high percentages of seeds when selfed. Only a few other species of

Carex are reported to be self-compatible: Carex pedunclliata, the bog species Carex pillriflora, C. obnllpta, C. callescens and C. pallciflora, the salt marsh species C. lyngbyei, and subalpine meadow species, Carex rossii, C. spectabilis (Pojar, 1974). Schmid (1984) reported that sedges of the Carexflava group are self-compatible (and presumebly self-fertile) and those of the sections Acutae and Vesicariae (including Carex rostrata ) are self­ incompatible and self-infertile. This accounted for the higher seed set observed in the C. flam group. The C. flava group has the cespitose growth form. Self-incompatible sedges tend to be rhizomatous rather than cespitose and to have unisexual spikelets rather than androgynous spikelets (Britton and Brown, 1970; Ford et al., 1991; Pojar, 1974). Ford et al. (1991) suggest that self-incompatibility drives selection for rhizomatous rather than cespitose growth form. Based on these criteria, Carex stricta and C. aqllatilis, which have a cespitose growth form and androgynous spikes, are likely to be self-compatible. C. lacllstris, having a more rhizomatous growth form and unisexual spikes would be more likely to be self-incompatible.

Kik et at. (1990) hypothesized that the low sexual reproductive capacities of non­ colonizing, long-lived (> 100 years old) clonal popUlations of Agrostis slolonifera were the 80 result of natural selection for competitive ability in relatively stable conditions. In stable environments, they found fewer, more widespread clones than in unstable environments.

Clones in the unstable environments were smaller and more numerous, and had more sexual reproduction. The extent of the spread of individual clones of Carex species in the natural sedge meadows studied is unknown. It may be quite extensive, based on the stability of the environments in the sedge meadows. The monodominance of the vegetation suggests these sites are not subject to much disturbance, a condition that reduces species diversity (van der

Valk and Davis, 1980).

Poor seed production may also be due, simply, to poor pollination due to rainy weather during pollen dispersal, as these sedges are all wind pollinated. Fungal or insect infestation may also have a role. Rates of viable seed production seem to vary from year to year and from site to site (and possibly from place to place at any given site), suggesting an environmental cause. Exact causes of poor seed production by some Carex species are unknown at this time.

Carex seed's relatively low cost compared with the cost of planting seedlings makes seeds a potentially economical alternative for establishing sedge meadow Carex. However, the low germination rates and uncertainty associated with using seeds makes seedlings the recommended alternative. Until seed suppliers are willing and able to provide accurate estimates of germination rates of batches of seeds, and adjust prices accordingly, Carex seed should be avoided or evaluated carefully before planning sedge meadow establishment based on the use of seeds. A compromise solution would involve purchasing seed, germinating it in controlled conditions in a greenhouse, then transplanting the seedlings to the field. This would take advantage of the lower cost of seed than seedlings, better germination in a controlled environment and better survival in the field of seedlings grown for some time in the greenhouse.

Seed suppliers should be required to provide a standardized expression of pure live seed 81 and germination rates of seeds sold, as is done for agricultural seed. A seed certification program regulating the expression of seed quality would encourage research and development and provide a standard for pricing wetland seed. Certification will be necessary for progress to continue in sedge meadow restoration techniques.

Seed storage Germination tests of fresh seed suggest that seeds of Carex aquatilis, C. iaclIstris and C. stricta are not dormant when mature. They may require a few weeks of afterripening to permit germination. It is unknown whether they re-enter dormancy at some later date in response to seasonal influences or storage conditions (Baskin and Baskin, 1989).

Testing of germinability in seeds of different ages suggested that, for the species tested, germination rates declined with time. The role of storage conditions in this decline is unknown. There wass some suggestion that some storage conditions increased germination rates beyond what they were prior to storage in some species. This may have been an artifact of sampling, or it reflects something of the biology of the seeds, either by breaking dormancy or by fulfilling germination requirements, by providing a dampening or chilling effect.

Seed viability was reduced in all storage treatments. In general, germination rates were highest after wet, cold storage. The effect on germination was more dependent on species.

Carex iacllstris seemed to do well stored at room temperature, but was intolerant of freezing of any kind. Carex aquatilis germinated poorly after dry storage in the first study, but in the second, dry storage was optimal. It too was intolerant of freezing wet, particularly the alternating freezing/cold. There may be an advantage to storing the Carex seed in water, as long as it never freezes. Freezing of wet seed seriously reduced germination in almost all cases. Seed could tolerate freezing if stored dry. Personal observations of storage facilities at wetland nurseries are that they are not temperature controlled. Wet storage is not recommended unless a cold storage facility is available that will prevent freezing of the seed.

It is likely that effects of storage conditions on germination are not related to seed 82 survival, which declined in all cases, but rather to a cold-damp conditioning which improved germination after storage. Schramm (1978; 1990) reports that seeds of many prairie forb and grass species germinated better after cold, damp conditioning.

Larson and Stearns (1990) found that seeds of Care x scoparia were dormant when mature. They recommended cool-dry storage, and found that germination was reduced in seed stored in water. They suggested that this decline in germination in moist storage was a sign that Carex scoparia was adapted to sites with little standing water. Amen and Bonde (1964) report that two alpine Carex species, C. ebenea and C. albonigra, were dormant when mature, and stratification did not break dormancy. Johnson et af. (1965) reported that cool, moist storage reduced germination in several Carex species in Wyoming. Comes et af. (1978) reported that seeds of Carex lanllginosa germinated better after wet than dry storage. The trend evident in these studies is toward a more positive germination response to wet storage by Carex species favoring wetter locations, and a negative response to wet storage by species growing naturally in drier habitats.

The decline in viability in storage observed may be as variable as was initial viability. It could be due to local environmental stresses during seed formation. Naylor (1984) suggests that seed that is less vigorous due to fungal attack or due to less stored reserves devoted to them by vegetatively spreading adults is more likely to perish in stressful environments, such as anoxia. Nearly all Carex stricta stands were observed to have the some sort ofleaf spot fungus. It is also possible that the vigorously vegetative Carex species, simply do not produce long-lived seed. The regularity of seasonal drawdowns means seeds never have to wait long for suitable conditions for germination. As in the shallow water emergents observed by van der Valk and Davis (1979), there is no selective pressure to produce long-lived seed in the

Carex species studied.

Interpreting results obtained in studies of the effect of storage conditions on germination 83 was complicated by several factors, which may limit the ability to make any useful conclusions or recommendations. Results were inconsistent between the first and second studies. In the first seed storage study, seeds were obtained anywhere from a few months to more than 1 Y2 years after they were harvested. During that time they were subjected to uncontrolled conditions that had unknown effects on their viability and germinability. The second complicating factor is the difficulty in obtaining representative samples due to the small size of

Carex seeds, the high frequency of empty perigynia and the extreme variability in quality from year to year and site to site.

A third complicating factor is the use and interpretation of tetrazolium dye testing, for reasons just mentioned, and because Carex seeds would probably be classified as "hard seed," meaning impermeable to water, making tetrazolium testing more difficult. Much of the information generated in the present study regarding viability may be inaccurate due to misinterpretation of tetrazolium reactions or misapplication of methods.

Growth studies Although greenhouse studies suggest that growth of Carex stricta In soils taken from the experimental wetlands was improved by all soil amendments tested

(Tables 30-31), field studies indicate that recruitment and growth of volunteer species was also improved by the soil amendments (Table 36-37).Topsoil probably carried seeds of ruderals, which may have inhibited growth of Carex stricta in plots amended with topsoil

(Table 34). Only compost had a consistently significant positive effect on Carex stricta growth in the field (Table 35). Compost and topsoil significantly increased recruitment and growth of seed mix species (Table 37). However, compost and topsoil also improved recruitment and growth ofruderal species (Table 37). Fertilizer improved recruitment of ruderal species, but was ineffective on seed mix species and on Carex stricta growth.

Greenhouse studies suggest that growth of Carex stricta increased in direct proportion to percent compost in the growing medium, up to 100 % compost, although differences were not 84 significant above about 33 to 50 % compost. Costello (1936) reported that Carex stricta was only found on organic substrates. He believed soils must be somewhat peaty before it will even invade.

Conclusions and recommendations

In conclusion, these findings suggest caution should be used when attempting to utilize seed for propagating Carex species, due to their low germination rates and short-lived seed.

Transplanting seedlings is more expensive, but may be worth the money in the greater likelihood of successful establishment gained by their use. Amending the soil in the experimental wetlands with compost would be beneficial to both recruitment of seed mix species and to the growth of Carex stricta seedlings, if invasion by volunteer species could be managed. Managing the sedge meadow vegetation would be aided by annual early spring burning and late-spring water level increases or herbicide application to control ruderal speCIes.

Adding compost would also improve soil physical properties and, therefore, soil moisture and hydrologic characteristics, making them more like those found in natural sedge meadows.

Planning for the hydrologic regime in created sedge meadow must account for the fact that in natural sedge meadows located on muck soils, saturated soil conditions extend far above the level of the water table, often to the soil surface. 85

LITERATURE CITED

Amen, R.D. and E.K. Bonde. 1964. Dormancy and germination in alpine Carex from the Colorado front range. Ecology. 45(4):881-883.

Aspinwall, N. and T. Christian. 1992. Pollination biology, seed production, and population structure in Queen-of-the-Prairie, Filipenduia ntbra (Rosaceae) at Botkin Fen, Missouri. American Journal of Botany. 79(5):488-494.

Auclair, AN.D. 1982. Seasonal dynamics of nutrients in a Carex meadow. Canadian Journal of Botany. 60:1671-1678.

Barendregt, A, S.M.E. Starn and MJ. Wassen. 1992. Restoration offen ecosystems in the Vecht River plain: cost-benefit analysis of hydrological alternatives. Hydrobiologia. 233:247-258.

Baskin, J.M. and C.c. Baskin. 1989. Physiology of dormancy and germination in relation to seed bank ecology. P. 53-66. In: Leck, M.E., Y.T. Parker and R.L. Simpson (eds.) Ecology of Soil Seed Banks. Academic Press, New York.

Bernard, J.M. 1973. Production ecology of wetland sedges: The genus Carex. Polskie Archiwum Hydrobiologii. 20(1):207-214.

Bernard, J. M. 1975. The life history of shoots of Carex lacus/ris. Canadian Journal of Botany. 53 :256-260.

Bernard, J.M. 1990. Life history and vegetative reproduction in Carex. Canadian Journal of Botany. 68: 1441-1448.

Bernard, J.M. and E. Gorham. 1978. Life history aspects of primary production in sedge wetlands. P.39-51. In: Good, R. E., D.F. Whigham and R.L. Simpson. (eds.) Freshwater Wetlands: Ecological Processes and Management Potential. Academic Press. New York.

Bernard, J.M. and J.G. Macdonald. 1974. Primary production and life history of Care x iacllstris. Canadian Journal of Botany. 52:117-123.

Bernard, J.M., D. Solander and J.Kvet. 1988. Production and nutrient dynamics in Carex wetlands. Aquatic Botany. 30: 125-147.

Bernard, J.M. and B.A Solsky. 1977. Nutrient cycling in a Carex laclIslris wetland. Canadian Journal of Botany. 55:630-638.

Boelter, D.H. 1966. Hydrologic characteristics of organic soils in Lakes states watersheds. Journal of Soil and Water Conservation. 21:50-53.

Brady, N.C. 1984. The Nature and Properties of Soils. Ninth Edition. Macmillan. New York. 750p. 86

Britton, N. and A Brown. 1970. An Illustrated Flora of the Northern United States and Canada. VoU. Dover Publications, Inc. New York. 680 p.

Broome, S.W., E.D. Seneca and W.W. Woodhouse, Jr. 1988. Tidal salt marsh restoration. Aquatic Botany. 32: 1-22.

Brown, M.T. 1991. Evaluating constructed wetlands through comparisons with natural wetlands. EPN600/3-911058. US Environmental Protection Agency, Corvallis, Oregon.

Carpenter, Q.l 1990. Hydrology and vegetation on a calcareous peat mound in southeast Wisconsin. Master's Thesis. University of Wisconsin, Madison.

Carter, V., M.S. Bedinger, RP. Novitzki and W.O. Wilen. 1978. Water resources in wetlands. In: Greeson, P.E., lR. Clark and lE. Clark, (eds.) Wetland functions and values: the state of our understanding. Proceedings of national symposium on wetlands. Lake Buena Vista, Florida. American Water Resources Association. Technical Publication TPS 79-2. Minneapolis, Minnesota.

Causton, D.R 1988. An Introduction to Vegetation Analysis. Unwin Hyman Ltd. London. 342 p.

Chapin, F.S. and M.C. Chapin. 1980. Revegetation of an arctic disturbed site by native tundra species. Journal of Applied Ecology. 17:449-456.

Comes, RD., V.F. Bruns and AD. Kelley. 1978. Longevity of certain weed and crop seeds in fresh water. Weed Science. 26(4):336-344.

Confer, S.R. and W.A Niering. 1992. Comparison of created and natural freshwater emergent wetlands in Connecticut (USA). Wetlands Ecology and Management. 2(3): 143-156.

Costello, D.F. 1936. Tussock meadows in southeastern Wisconsin. Botanical Gazette. 97:610-648.

D'Avanzo, C. 1989. Long-term evaluation of wetland creation projects. P. 75-84. In: Kusler, J.A and M.E. Kentula (eds.). Wetland Creation and Restoration: The Status of the Science Volume II: Perspectives. EPN600/3-89/038. US Environmental Protection Agency, Corvallis, Oregon.

Ebinger, lE. 1977. Checklist of vascular plants of the Chain O'Lakes region, Lake County, Illinois. P. 35-43. In: Ebinger, lE., RE. Andrews and He. Nilsen (eds.). Papers on the Ecology of the Chain O'Lakes Region. Eastern Illinois University, Charleston.

Environmental Laboratory. 1987. Corps of Engineers Wetlands Delineation Manual. Department of the Army, Waterways Experiment Station Technical Report y-89-1. Vicksburg, Mississippi.

Faulkner, S.P., W.H. Patrick and RP. Gambrell. 1989. Field techniques for measuring wetland soil parameters. Soil Science Society of America Journal. 53:883-890. 87

Ford, B.A, P. W. Ball and K. Ritland. 1991. AIlozyme diversity and genetic relationships among North American members of the short-beaked taxa of Care x sect. Vesicariae (Cyperaceae). Systematic Botany. 16(1): 116-131.

Fredrickson, L.H. and T.S. Taylor. 1982. Management of seasonally flooded impoundments for wildlife. Resource Publication 148. United States Department of the Interior. Fish and Wildlife Service. Washington, D.C.

Galinato, M.I. and AG. van der Valk. 1986. Seed germination traits of annuals and emergents recruited during drawdowns in the Delta Marsh, Manitoba, Canada. Aquatic Botany. 26:89-102.

Garbisch, E.W. 1989. Information needs in the planning process for wetland creation and restoration. P. 9-13.1n: Kusler, J.A and M.E. Kentula (eds.). Wetland Creation and Restoration: The Status of the Science Volume II: Perspectives. EPN600/3-891038. US Environmental Protection Agency, Corvallis, Oregon.

Gartner, B.L., F. S. Chapin III and G.R. Shaver. 1983. Demographic patterns of seedling establishment and growth of native graminoids in an Alaskan tundra disturbance. Journal of Applied Ecology. 20:965-980.

Gillespie, J. 1987. The germination and propagation of Care x stricta. MEPD Thesis. University of Wisconsin, Whitewater.

Gillham, RW. 1984. The capillary fringe and its effect on water table response. Journal of Hydrology. 67:307-324.

Godwin, H. and F.R Bharucha. 1932. Studies in the ecology of Wick en Fen. II. The fen water table and its control of plant communities. Journal of Ecology. 20: 157-191.

Gorham, E. 1974. The relationship between standing crop in sedge meadows and summer temperature. Journal of Ecology. 62:487-492.

Gosselink, J.R and RE. Turner. 1978. The role of hydrology in fresh water wetland ecosystems. p63-78. In: Good, R. E., D.F. Whigham and R.L. Simpson. (eds.) Freshwater Wetlands: Ecological Processes and Management Potential. Academic Press. New York.

Grabe, D.F. (ed.) 1970. Tetrazolium testing handbook for agricultural seeds. Contribution number 29 to the handbook on seed testing. Tetrazolium testing committee of the Association of Seed Analysts.

Grootjans, AP. and B.F. van Tooren. 1984. Ecological notes on Carex aqua/ilis communities. Vegetatio. 57:79-89.

Hammer, D.A(Ed.). 1989. Constructed Wetlands For Wastewater Treatment: Municipal, Industrial, and Agricultural. Lewis Publishers, Inc. Chelsea, Michigan.

Hammer, D.A 1992. Creating Freshwater Wetlands. Lewis Publishers, Inc. Chelsea, MI. 88

Haggas, L., RW. Brown and RS. Johnston. 1987. Light requirement for germination of Payson sedge. Journal of Range Management. 40(2): 180-184.

Handel, S.N. 1978. Self-compatibility in Carex pial1taginea and C. piatyphylla (Cyperaceae). Bu\1etin of the Torrey Botanical Club. 105(3):233-234.

Handel, S.N. 1985. The intrusion of clonal growth patterns on plant breeding systems. American Naturalist. 125:367-384.

Heikurainen, L., I Paivanen, and I Sarasto. 1964. Groundwater table and water content in peat soil. Acta Forestalia Fennica. 77.1: 1-18.

Heinselman, M.L. 1970. Landscape evolution, peatland types, and the environment in the Lake Agassiz Peatlands Natural Area, Minnesota. Ecological Monographs. 40(2):235-261.

Heliotis, F.D. and e.B. DeWitt. 1987. Rapid water table responses to rainfall in a northern peatland ecosystem. Water Resources Bulletin. 23(6): 1011-1016.

Hughes, D.A 1977. Wet ground plant communities from the Chain O'Lakes region, Lake County, Illinois. P. 8-14.111: Ebinger, lE., R.E. Andrews and H.C. Nilsen (eds.). Papers on the Ecology of the Chain O'Lakes Region. Eastern Illinois University, Charleston.

Johnson, W.M., 10. Blankenship and G.R. Brown. 1965. Explorations in the germination of sedges. Rocky Mountain Forest and Range Experiment Station, U.S. Forest Service Research Note RM-51.

Kelsey, P.D. and R.G. Hootman. 1992. Relationships between water tables, plant communities and hydric soils: West Chicago Prairie, West Chicago, Illinois. Soil Survey Horizons. 33(3):53-58.

Kentula, M.E., RP. Brooks, S.E. Gwin, C.C. Holland, AD. Sherman and IC. Sifneos. 1992a. An approach to improving decision making in wetland restoration and creation. Al Hairston (ed.). U.S. Environmental Protection Agency. Environmental Research Laboratory, Corvallis, Oregon.

Kentula, M.E., Ie. Sifneos, IW. Good, M. Rylko and K Kunz. 1992b. Trends and patterns in Section 404 permitting requiring compensatory mitigation in Oregon and Washington, USA Environmental Management. 16(1): 109-119.

Kerans, K 1990. The restoration business: Country Wetlands Nursery Ltd. Restoration and Management Notes. 8(1 ):29-31.

Khaleel, R, KR. Reddy and M.R Overcash. 1981. Changes in soil physical properties due to organic waste applications: A review. Journal of Environmental Quality. 10(2):133-141.

Kik, C., J. van Andel, W. van Delden, W. Joenje and R. Bijlsma. 1990. Colonization and differentiation in the clonal perennial Agrostis stolonifera. Journal of Ecology. 78:949-961. 89

Koerselman, W. 1989. Groundwater and surface water hydrology ofa small groundwater­ fed fen. Wetlands Ecology and Management. 1(1):31-43.

Konings, H., E. Koot and A Tijman-de Wolf. 1989. Growth characteristics, nutrient allocation and photosynthesis of Carex species from floating fens. Oecologia. 80: 111-121.

Kusler, lA and M.E. Kentula, (eds.) 1989. Wetland Creation and Restoration: The Status of the Science. Volume II. Perspectives. U.S.Environmental Protection Agency 600/3-98/038. Environmental Resources Laboratory, Corvallis, Oregon.

Larson, lL. and F.W. Stearns. 1990. Factors influencing seed germination in Carex scoparia Schk. Wetlands. 10(2): 277-283.

Lyle, E.S., Jr. 1987. Surface Mine Reclamation Manual. Elsevier Science Publishing Co., Inc. New York. 268 p.

MacRae, R.I. and G.R. Mehuys. 1985. The effect of green manuring on the physical properties of temperate-area soils. Advances in Soil Science. 3:71-94.

Millar, lB. 1971. Shoreline-area ratio as a factor in rate of water loss from small sloughs. Journal of Hydrology. 14:259-84.

Moran, R. C. 1981. Prairie fens in northeast Illinois: Floristic composition and disturbance. P. 164-168. In: Stuckey, R.L. and K.l Reese. (eds.) Proceedings of the Sixth North American Prairie Conference. Ohio Biological Survey Biological Notes, 15.

Naylor, R.E.L. 1984. Seed Ecology. Advances in Research and Technology of Seeds. 9:61- 93.

NOAA. 1991. Climatological Data, Illinois. 96(1-12). National Climatic Data Center. Asheville, North Carolina.

NOAA. 1992. Climatological Data, Illinois. 97(1-9). National Climatic Data Center. Asheville, North Carolina.

Novitzki, R.P. 1982. Hydrology of Wisconsin wetlands. U.S. Geological Survey Information Circular 40. Wisconsin Geological and Natural History Survey. Madison, Wisconsin.

O'Brien, AL. 1977. Hydrology of two small wetland basins in eastern Massachusetts. Water Resources Bulletin. 13(2):325-340 .

. O'Brien, AL. and W.S. Motts. 1980. Hydrogeological evaluation of wetland basins for land use planning. Water Resources Bulletin. 16(5):785-789.

O'Neal, AM. 1949. Soil characteristics significant in evaluating permeability. Soil Science. 67:403-409. 90

Owen, C.R., Q.l Carpenter and C.B. DeWitt. 1989a. Comparative hydrology, stratigraphy, microtopography and vegetation of natural and restored wetlands at two Wisconsin mitigation sites. P.119-132. In: Webb, F.l, Jr. (ed.). Proceedings of sixteenth annual conference on Wetlands restoration and creation. Hillsborough Community College, Tampa, Florida.

Owen, C.R., Q.J. Carpenter and C.B. DeWitt. 1989b. Evaluation of three wetland restorations associated with highway projects. Report to the Wisconsin Department of Transportation. Institute for Environmental Studies. University of Wisconsin, Madison. 89 p.

Owen, C.R and H.M. Jacobs. 1992. Wetland protection as land-use planning: the impact of Section 404 in Wisconsin, USA. Environmental Management. 16(3):345-353.

Paivanen, l 1973. Hydraulic conductivity and water retention in peat soils. Acta Forestalia Fennica. 129:4-70.

Pearson, C.J. and S.G. Shah. 1981. Effect of temperature on seed production, seed quality and growth of PaSpalllnl dilatatllnl. Journal of Applied Ecology. 18:897-905.

Pietz, RI., C.R Carleson, Jr., lR. Peterson, D.R Zenz and C. Lue-Hing. 1989. Application of sewage sludge and other amendments to coal refuse material: II. Effects on revegetation. Journal of Environmental Quality. 18(2): 169-173.

Pojar, l 1974. Reproductive dynamics of four plant communities of southwestern British Columbia. Canadian Journal of Botany. 52: 1819-1834.

Roberts, J.A., W.L. Daniels, J.C. Bell and lA. Burger. Early stages of mine soil genesis as affected by topsoiling and organic amendments. Soil Science Society of America Journal. 52:730-738.

Richardson, C.l, D.L. Tilton, lA. Kadlec, lP.M. Chamie and W.A. Wentz. Nutrient dynamics of northern wetland ecosystems. P. 217-240. In: Good, R. E., D.F. Whigham and R.L. Simpson. (eds.) Freshwater Wetlands: Ecological Processes and Management Potential. Academic Press. New York.

Schennum, W.E. 1990. Wetlands: Reservoirs for prairie biota in "prairieless" landscapes. P. 95-100. In: Smith, D.D. and c.A. Jacobs (eds.). Proceedings of the Twelfth North American Prairie Conference. University of Northern Iowa, Cedar Falls.

Schmid, B. 1984. Life histories in clonal plants of the Carexflava group. Journal of Ecology. 72:93-114.

Schmid, B. 1986. Colonizing plants with persistent seeds and seedlings (Carexflava group). Botanica Helvetica. 96( 1): 19-26.

Schramm, P. 1976. The "do's and don'ts" of prairie restoration. P. 139-150. In: Glenn­ Lewin, D.C. and R.Q. Landers, Jr. (eds). Proceedings of the Fifth Midwest Prairie Conference. Iowa State University, Ames. 91

Schramm, P. 1990. Prairie restoration: A twenty-five year perspective on establishment and management. P. 169-177. In: Smith, D.D. and C.A Jacobs (eds.). Proceedings of the Twelfth North American Prairie Conference. University of Northern Iowa, Cedar Falls.

Shaver, G.R., F.S. Chapin and W.D. Billings. 1979. Ecotypic differentiation in Carex .aquatilis on ice-wedge polygons in the Alaskan coastal tundra. Journal of Ecology. 67:1025-1046.

Siegel, D.1. 1983. Ground water and the evolution of patterned mires, glacial Lake Agassiz peatlands, northern Minnesota. Journal of Ecology. 71:913-921.

Sjoberg, K. and K. Danell. 1983. Effects of permanent flooding on Carex-Eqllisetllnl wetlands in Northern Sweden. Aquatic Botany. 15:275-286.

Smith, AL. 1972. Factors influencing germination of Scolochloa jestllcacea caryopses. Canadian Journal of Botany. 50(11):2085-2092.

Smith, AL. 1973. Life cycle of the marsh grass Scolochloa jestllcacea. Canadian Journal of Botany. 51: 1661-1668.

Soil Conservation Service. 1970. Soil survey of Lake County, Illinois. Illinois Agricultural Experiment Station Soil Report nO.88.

Soil Conservation Service Plant Materials Centers. 1988. Plant Materials Handbook. Vol.l. Office of Surface Mining Reclamation and Enforcement.

Stout, AB. 1912. A biological and statistical analysis of the vegetation ofa typical wild hay meadow. Transactions of the Wisconsin Academy of Science, Arts and Letters. 17:404-469.

Tester, C.F. 1990. Organic amendment effects on physical and chemical properties ofa sandy soil. Soil Science Society of America Journal. 54:827-831. van der Valk, AG. and c.B. Davis. 1978. The role of seed banks in the vegetation dynamics of prairie glacial marshes. Ecology. 59:322-335. van der Valk, AG. and C.B. Davis. 1979. A reconstruction of the recent vegetational history of a prairie marsh, Eagle Lake, Iowa, from its seed bank. Aquatic Botany. 6:29-51. van.der Valk, AG. and C.B. Davis. 1980. The impact ofa natural drawdown on the growth of four emergent species in a prairie glacial marsh. Aquatic Botany. 9:301-322. van der Valk, AG. and R.L. Pederson. 1989. Seed banks and the management and restoration of natural vegetation. P. 329-346. In: Leck, M.E., V.T. Parker and R.L. Simpson (eds.) Ecology of Soil Seed Banks. Academic Press, New York. van der V Alk AG. and J.T.A. Verhoeven. 1988. Potential role of seed banks and understory species in restoring quaking fens from floating forests. Vegetatio. 76:3- 13. 92

Verhoeven, IT.A. and H.H.M. Arts. 1987. Nutrient dynamics in small mesotrophic fens surrounded by cultivated land. II. Nand P accumulation in plant biomass in relation to the release of inorganic Nand P in the peat soil. Oecologia. 72:557-56l.

Verhoeven, J.T.A., M.B. Schmitz and T.L. Pons. 1988. Comparative demograhic study of Carex rostrata Stokes, C. diandra Schrank, and C. aClftijormis Ehrh. in fens of different nutrient status. Aquatic Botany. 30:95-108.

Verhoeven, IT.A., S. van Beek and W. Storm. 1983. Nutrient dynamics in small mesotrophic fens surrounded by cultivated land. I. Productivity and nutrient uptake by the vegetation in relation to the flow of eutrophicated groundwater. Oecologia. 60:25-33.

Verry, E.S. and D.H. Boelter. 1978. PeatIand hydrology. P. 389-40l. In: Greeson, P.E., lR. Clark and lE. Clark, (eds.) Wetland functions and values: the state of our understanding. Proceedings of a national symposium on wetlands. Lake Buena Vista, Florida. American Water Resources Association. Technical Publication TPS 79-2. Minneapolis, Minnesota

Wein, R.W. and D.A. MacLean. 1973. Cotton grass (Eriophorum vaginatllm) germination requirements and colonizing potential in the Arctic. Canadian Journal of Botany. 51:2509-2513 .

Welling, C.H., R.L. Pederson and AG. van der Valk. 1988. Recruitment from the seed bank and the development of zonation of emergent vegetation during a drawdown in a prairie wetland. Journal of Ecology. 76:483-496.

Wiesner, L.E., A.E. Carleton and R.C. Bailey. 1967. Seed evaluation of sedge species. Proceedings of the Association of Official Seed Analysts. 57: 107-11l.

Wilcox, D.A., S.F. Apfelbaum and R.D. Hiebert. 1985. Cattail invasion of sedge meadows following hydrologic disturbance in the Cowles Bog wetland complex, Indiana Dunes National Lakeshore. Wetlands. 4:115-128.

Williams, R.E. 1968. Flow of groundwater adjacent to small, closed basins in glacial till. Water Resources Research. 4(4):777-783.

Zedler, lB. and M.W. Weller. 1989. Overview and future directions. P.465-473. In: lA. Kusler and M.E. Kentula (eds.). Wetland Creation and Restoration: The Status of the Science. Vol. 1 U.S. E.P.A/7600/3-98/038. U.S. Environmental Protection . Agency, Environmental Resources Laboratory, Corvallis, Oregon. 93

ACKNOWLEDGEMENTS

This project was funded by Wetlands Research, Inc. 94

APPENDIX

MAPS SHO\VING WELL PLACEMENTS A T THE

FIVE NATURAL SEDGE MEADOW SITES Lyons Prairie and Marsh Nature Preserve Lake Co., Illinois Barrington quad T43N RgE sec4

McHenry Co.

County Line Road Lake Co. North ..

10 well#4 VI well#3 well#2

19m well#1

.l ...... J...... l...... l...... 1 o 25m 50m 75m 100m Spring Grove Fen 100m'- • well#1 Nature Preserve 13m \ well#2 McHenry Co., Illinois 11 m well#3 Fox Lake quad 75m- T46N R9E sec 30

SOm - 62m 1.0 0\

25m:- Nippersink Creek well#4

~ 16m \ well#5 o - North ... Turner Lake Fen Nature Preserve Chain 0' Lakes State Park Lake Co., Illinois Fox Lake quad T49N R9E sec.21 Turner Lake

50m :--

7m 1 well#1 25m .-- \0 16m / well#2 -...J .- well#3 o ~

Turner Lake South North • Camping Area Wadsworth Prairie Nature Preserve Lake Co., Illinois Wadsworth quad.. T46N R11 E sec27 - Des Plaines River -

.v160m

North • 1.0 00

93m • ~ 31m well#4 ..------. well#3 well#1 well#2

1_._._._._._._._._. .1._._._._._._._._._._._.1.._._._._._._._._._._J_._._._._._._._._._._._J o 25m 50m 75m 100m Des Plaines River Trail Lake County Forest Preserve .--100m Lake Co .• Illinois Wadsworth quad \ well#1• T46N R11 E sec.27

o--75m

• \0 well#2 \0 ---50m 22m • 36m ~. well#3 well#5 --r well#4 '-25m

40m North ... <-0 well#6