University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln
USGS Northern Prairie Wildlife Research Center US Geological Survey
9-1989
PRAlRlE BASIN WETLANDS OF THE DAKOTAS: A COMMUNlTY PROFILE
Harold L. Kantrud USGS Northern Prairie Wildlife Research Center
Gary L. Krapu USGS Northern Prairie Wildlife Research Center, [email protected]
George A. Swanson USGS Northern Prairie Wildlife Research Center
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Kantrud, Harold L.; Krapu, Gary L.; and Swanson, George A., "PRAlRlE BASIN WETLANDS OF THE DAKOTAS: A COMMUNlTY PROFILE" (1989). USGS Northern Prairie Wildlife Research Center. 249. https://digitalcommons.unl.edu/usgsnpwrc/249
This Article is brought to you for free and open access by the US Geological Survey at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in USGS Northern Prairie Wildlife Research Center by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Biafog ica! Report 85t7.28) September 1989
PRAIRIE BASIN WETLANDS OF THE DAKOTAS: A COMMUNITY PROFILE Copies of this publication may be obtained !(om the Publications Unii,, U.S. Fish and Wildlife Sewice, !U!b and C Streets, N.W., Mail Stop 711?* Arlington Square Building, Washi~gton,DC 20240, or may be purchased from the National Teci7nicai Intorms?iol? Service (NTIS), 5285 Port Royai Road, Springfield, VA
Cover :
Upper ?eft: Palustrine emergent vegetation (Scirpns gcutus) in an oligosaiine, semipermanently f l ooded wetland, upper middle: Falustrine aquatic bed vegetation (samoqeton qramineus) in a fresh, seasonally flooded prairie wetlard. 7 - vpper right: Snow geese (cnen caerui escensj t.isi~~t-ja .---'&:-v-*-d+.UiLlVULLul temporarily flooded vetland during spring migration through the Prairie Pothole region. Boxton: Basin wetlands of various sizes and with different hydrokogieaY regines are a major feature of the landscape In croplands, haylands, and pasture types 2: land-use patterns in the Prairie Pothole reqbsn, ~iologicalReport 85 (7.28) September 1989
PRAliRliE BASIN WE"FANDS OF THE DAKOTAS: A COMMUNlTY PROFILE
Harold A. Kantrud Gary L. Krapu George A. Swanson U.S. Fish and Wildlife Service Northern Prairie Wildlife Research Center P.O. Box 2096 Jarnestown, ND 58402
Project Officer James A. Allen U.S. Fish and Wildlife Service National Wetlands Research Center 1010 Gause Boulevard j lid ell, LA 70458
U.S. Department of the Interior Fish and Wildlife Service Research and Development Washington, DC 20240 Suggested citation:
Kantrud, H.A., G.L. Krapu, and G,A, Swanson, 1989, prairie basin wetlands of the Dakotas: a cemmunity prafila- U.S, Fish Wildl. Sew. Biol. Rep. 85(7.28). 116 pp, PREFACE
This description of prairie basin basins in the Prairie Pothole Region wetlands of the Dakotas is part of of the Dakotas. This profile a series of community profiles on outlines the wetland subsystems, ecologically important wetlands of classes and subclasses that occur in national significance. The shallow these basins, and provides a useful wetlands of the Dakotas form the reference to their geologic, bulk of the portion of the Prairie climatic, hydrologic, and pedologic Pothole Region lying within the setting. United States. This region is famous as the producer of at least half of Detailed information on the biotic North America's waterfowl and an environment of the wetlands dealt unknown, but large, proportion of with in this profile will be useful other prairie-dwelling marsh and to scientists and resource managers. aquatic birds. Special recognition is paid to the macrophyte and invertebrate communities, which have dynamic The wetlands described here lie in qualities found in few other of the relatively small, shallaw basins world's wetland ecosystems. that vary greatly in their ability to maintain surface water, and in The most noteworthy animal their water chemistry, which varies inhabitants of these basins are from fresh to hypersaline. These waterfowl, which are a resource of wetlands occur in a wide variety of international concern. Because hydrological settings, in an area af where annual and seasonal precipi- the importance of this resource, tation varies greatly in form and much research on the habitat use and amount. Thus the presence of surface feeding ecology of breeding water- water in these wetlands is largely fowl has been conducted in the unpredictable. Superimposed on these region. These topics receive special ~henomena are the effects of a attention in this profile, ;ariety of land uses, including The Prairie Pothole Region is a pasture, cultivation, mechanical major world supplier of cereal forage removal, idle conditions and grains. Consequently, wetlands in burning. All those factors greatly the region are often drained for affect the plant and animal crop production or otherwise cropped communities found in these basins. when water conditions permit. These practices degrade the value of This prof ile covers lacustrine and wetlands for nost species of wild- palustrine basins with temporarily life and conflict with the aims of flooded, seasonally flooded, and conservationists. The subject of semipermanently flooded water human uses and impacts to prairie rec;irr,ss. Basins with these water we5Lands is thrrs an impcrstant: part regimes compose about 90% of the of this profile. CONVERSION TAI3LE:
Metric to U. S.--.- Cii~,t.om:iry .. .
-MII I ti.!Y mi 1 l imeters (iilm) ce~ltin~et,ers(cm) met.er.5 (m) meters (m) ki lomet,ers (km) kilornr~ter:, (km)
square metet.5 (ni2) 10. 76 5clilare feet square ki lometers (kd) 0 . 3236, I iqii~3r.emi les ht?ctares (!?a) 2.411
1 i tfsrs ( 1 ) iJ
ki lu~,iloric:s (kcal) Ce l s i u5 tlr~q:-ccs("C)
I1. . .-. :> i nciies illcilei teet (f t) f atiioms st,at.iit.e ini les (irii) iiaut,ical mi let, (rinii)
s(4iidr.e Sect (fi.;' ) sqc1ar.e iii i 1 es (ni i :!) aLr'eS
gallons (yai) cubic feet. (f t:') acre-feet
o1irici.i { OL) ounces (or) poiii~dsj l b) ;;;j5 (!!>) short ton5 (too) British ihvlmal ui>it,s (Htii) Fahrenheit degrees (OF) CONTENTS
PREFACE ...... iii CONVERSION FACTORS ...... iv FIGURES ...... vi TABLES ...... vii CHAPTER 1 . INTRODUCTION ...... 1 1.1 Geographic Area Covered ...... 1 1.2 Definition of Prairie Basin Wetlands ...... 1 1.3 Classification of Prairie Basin Wetlands ...... 3 1.4 Distribution and Abundance of Wetlands ...... 8 CHAPTER 2 . ABIOTIC ENVIRONMENT ...... 10 2.1 Geology ...... 10 2.2 Climate and Weather ...... 13 2.3 Hydrology ...... 15 2.4 Water Quality ...... 18 CHAPTER 3 . BIOTIC ENVIRONMENT ...... 23 3.1 Phytoplankton, Periphyton. and Metaphyton ...... 23 3.2 Macrophytes ...... 24 3. 3 Invertebrates ...... 35 3.4 Fish ...... 44 3.5 Amphibians and Reptiles ...... 45 3.6 Birds ...... 46 3.7 Mammals ...... 55 CHAPTER 4 . ECOLOGICAL PROCESSES ...... 65 4.1 Physical Functions ...... 65 4.2 Biological Functions ...... 67 CHAPTER 5 . HUMAN USES AND IMPACTS ...... 70 5.1 Economic Functions ...... 78 5.2 Wetland Ownership ...... 71 5.3 Wetland Degradation and Drainage ...... 72 5.4 Wetland Management and Restoration ...... 73 5.5 Prospects forthe Future ...... 74
REFERENCES ...... es*~~...... 77 APPENDIXES A . Planktivorous Algae of prairie wetlands ...... 91 B . Qcminance Types cf Emergent vegetation ...... ,,,7 nQ C . Emergent Wydrophytes of prairie Wetlands Arranged According to Salinity Tolerance ...... 105 Number Paqe I Prairie Pothole Region of the Dakotas...... 2 2 Arrangement of water regimes around basin wetlands ...... 6
3 Physiographic regions in the Prairie Pothole Region ...... 9 4 Extent of Pleistocene glaciation in North America ...... 11 5 Water level fluctuations in prairie wetland^...... ^^ 16 6 Factors influencing wetland ecology ...... 27 7 Sources of water in prairie wetlands ...... 19 8 Flow pattern in a flow-through type prairie wetland ...... 20 9 Flow pattern in a closed-system type prairie wetland ..... 20 10 Flow pattern in a sump type prairie wetland ...... 21 11 Spatial relation of vegetational zones ...... 25 12 A generalized vegetational cycle in an Iowa wetland ...... 27
13 A sequence of wetland phases as related to climate ...... 30 14 Cultivated basins in an agricultural landscape ...... 32 15 Wetland PI of the Cottonwood Lake study area ...... 36 16 Photograph of wetlands PI and P8 of the Cottonwood Lake study area ...... 37 17 Wetland P8 of the Cottonwood Lake study area ...... 38 TABLES
Number Paqe
1 Wetlands that occur in the Prairie Pothole ~egion of the Dakotas...... ,...... +..... 2 Climatic data for Bismarck. ND ...... 3 Stages of a habitat cycle in a semipermanent Iowa wetland ...... 4 Invertebrate zooplankton of prairie wetlands ......
5 Aquatic coleoptera of prairie wetlands ...... 6 Distribution of breeding marsh and aquatic birds in prairie wetlands ...... 7 Density of breeding marsh and aquatic birds in prairie wetlands ...... 8 Frequency of breeding marsh and aquatic birds in prairie wetlands ...... 9 General use of prairie wetlands by mammals ...... 10 Primary production in prairie wetlands ...... CHAPTER 1. INTRODUCTION
nThe entire face of the country is 1.2 DEFINITION OF PRAIRIE BASIN WETLANDS covered with these shallow lakes, ponds and puddles, many of which are, however, dry or undergoing a There is no single scientifically process of gradual drying out." So acceptable definition of wetlands stated Charles Froebel (1870) as he because of their tremendous diver- described the lands along the sity and because they lie along a Sheyenne and James River Valleys, continuum or gradient between deep- Dakota Territory, during General water habitats and uplands or Alfred Sully's 1865 expedition. between purely aquatic and terres- Thus the uniqueness of North trial ecosystems (Cowardin et al. America's Prairie Pothole Region has 1979). The term "basinw as used in been recognized for well over 100 this report refers to a depression years. In this Community Profile, capable of holding surface water, we briefly describe the biotic and but not to the entire watershed or abiotic settings and features of the "drainage basin" that contributes most common kinds of wetlands found surface water runoff ta that in the portion of this region that depression. lies in the Dakotas, and outline the natural and manmade ecological According to Cowardin et al. processes that affect these (1979), wetlands are lands wetlands. transitional between terrestrial and aquatic systems where the water table is usually at or near the l .I GEOGRAPHIC AREA COVERED surface or the land is covered by shallow water. Additionally, a site The Prairie Pothole Region of must have one or more of the three North America stretches from central following attributes to be defined Alberta to central Iowa and encom- as a wetland: passes well over 700,000 km2. This community profile covers the 168,000-km2 portion of the region (1) The site must, at least lying in North and South Dakota periodically, support hydrophytes. (Figure 1). This area is bounded on the north by the International (2) The land at the site must be Boundary, east by the Red and predominantly undrained hydric soil. Minnesota River Valleys and portions Hydric soils are defined by the U.S. of the western boundaries of Soil Conservation Senrice (1985) as Minnesota and Iowa, and on the south those that in their undrained and west by headwaters of numerous condition are saturated, EPooded, or small streams and rivers along the ponded long enough during the north and east sides of the ~issouri growing seasoc tc deveLop a~aerobic River. A few small areas of pot- conditions f avaring the growth and holed land not shown in Figure 1 lie regeneration of kydrophytkc west of the Missouri River. vegetation, L I miles SCALE - Boundary of Prairie Pothole Region -. - Extent of glaciation
-- . Missouri River
Figure 1. Prairie P0th0le Region of the Dakotas showing relation to maximum extent of glaciation and Missouri River. The Missouri Escarpment separates two major biogeographic regions of North America, the Great Plains, and Central Lowlands (after Biuernlo 1977). (3) Tile site substrate is nonsoil encounter some with relatively and is saturated with water or cov- convoluted shorelines composed of ered by shallow water at Zorne time several bays or peninsulas, or, more during the growing season each year. rare1y , with one or several is1 ands . Nonsoils arc defined by the U,S. Soil Conservation Service (1975) as barren areas such as ice, rock, or 1.ll CLASSiFlCATiON OF PRAlRlE BASIN WETLANDS substrates underlying deep water.
Nearly all natural basins in the -Systems and Subsystems Prairie Pothole Region of the Dakotas have both the first and Under the Cowardin et aX. (1979) second of these attributes, and thus wetland classification, the prairie are wetlands according to the wetlands referred to in this report Cowardin et al. (1979) classifi- are palustrine and lacustrine cation. The basins are underlain by systems. There are no subsystems in hydric soils, and when surface water the palustrine system. For the is present, can support hydrophytes. lacustrine system, only the littoral Many of the basins are intensively subsystem is dealt with in this cultivated; the soil may be bare or report. This subsystem extends from commercial craps may be present the shoreward boundary to a depth of during dry conditions, but when 2 m below low water or to the water is replenished durinq the maximum extent of nonpersistent growing season, communities of emergents, if these grow at depths hydrophytes quickly dcvclop. >2 m. The latter conditions are rare or absent in the pothole region Pow basins in the rcgion contain of the Dakotas. An outline of the deep-water habitat. This habitat cowardin et al. (1979) elassifi- will not be dcalt with in this cdtion system, as it applies to the report. Deep-water habitats arc Prairie Pothole Region of the permanently flooded lands lyinq Dakotas, is shown in Table 1. Only below the deep-water boundary of subclasses dominated by plants are wetlands. For the wetlands des- includcd in Table 3. because these cribed in this report, the boundary subclasses are by far the most between them and dcep-water habitats common in the region. Little is lies at a depth of 2 m below low known about the animals that water; however, if ernergents, dominate unvegetated substrates that shrubs, or trccs grow beyond this are known to exist in several depth dt any time, their deep-water wetland classes found in the region, edge is the boundary. Substrates of deep-water habitat are nonsoil whose -Cl . asses water depths are in excess of that required to support emergent vege- Classes of the palustrine system tation (Cowardin et al. 1373; U.S. found in the Prairie Pothole Region Soil Conservation Service, Soil of the Dakotas are emergent, aquatic Survey Staff 1975). In addition, a bed, unconsolidated share, small propartion of the wetldnds in unconsolidated bottom, forested, and the region are uf the water regimes shrub-scrub, Classes af lacustrine "intermittently exposedf1 and system represented are aquatic bed, P'saturated,53 These will also be emergent wetland, unconsolidated omitted from this rcport. shore, unconsolidated bsttsm, and rocky shore, Prairie wetlands occur in glacially or postglacially derived --Emerqent wetla&, This class of basins in the northern grassland wellan3, containing erect rcete2 bicme (see section 2.1). The basins herbaceous hydrophytes, is by far are roughly round or oval in shape, the mest common wetland class in the although it is fairly comon to region. Emergent wetland supparts Table 4. Wetland systems, subsystems, classes, and subclasses ofCowardin eta!.(1979) that occur in the Prairie Pathole Region 011 the Dakotas, and for which dominance types are planis. ------System Subsystem Class SubcPass
Riverine Lower Perennial Aquatic Bed Algal Aquatic Moss Rooted Vascular Floating Unconsolidated Vegetated Shore Emergent Wetland Non-persistent Upper Perennial Aquatic Bed Algal Aquatic Moss Rooted Vascil! ar Floating Unconsolidated Vegetated Shore Intermittent S treambed Vegetated l,acust~ine Limnet ic Aquatic Bed Alyal Aquatic Moss Rooted Vascular Ff oat inq Littoral Aquatic Hcd Algal Aquatic Moss Rooted Vascular Float.inq Emcrgent Wetland Non-persistent PsXustsine (no subsystenas) Aquatic Bed Algal* Aquatic Moss* Rooted Vascular* Floating* Unconsol Ldated Vegetated Shore Moss-Lichen Moss Wetland Lichen Emergent Wetland Persistent* Non-persistent* Scr-ub-Shrub Broad-leaved Wetland Deciduous Forested Broad-leaved Wetland IJec iduous
'" "" ,- . -. *See Appendix B for common dominance types.
veqetaticsn that is largely peren- under disturbances such as heavy rtial, and present during most of the grazing or cultivation, nonpersis- growing season most years, Erner- tent emergents can dominate. gent wetland is often called marsh, meadow, fen, slough, or swamp in ----Aauatic bed. The class "aquatic the regian. Persistent emergent~ bede?nciud@s wetland dominated by are by far the most common, although piants that grow principally on or below the water surface for most of seasonally flooded, and semiper- the growing season during most man@ntly flooded. Basins where the years. This is the second most central or deepest portion is common wetland class in the region, subject to these water regimes compose nearly 90% of all basins in UnconsoPidated shore, This class the Prairie Pothole Region. The is mostly associated with the concentric pattern of these regimes erosional and depositional shoreline in lacustrine and palustrine wetland zones of wetlands not dealt with in systems is shown in Figure 2. this report, but the class comonly occurs in many shallow prairie wet- In the basins considered in this lands as bottoms are exposed during report, "emergent wetlandw occurs drought. with water regimes temporarily flooded, seasonally flooded, and Unconsolidated bottom. This class semipermanently flooded. Water is mostly associated with the un- regimes of scrub-shrub and forested stable bottoms of wetlands with more wetland are restricted to tem- permanent water regimes than are porarily flooded areas. Aquatic bed dealt with in this report, but can is seasonally or semipermanently occur in some deeper prairie wet- flooded. Unconsolidated shore is lands as bottoms are exposed during seasonally or temporarily flooded, extreme drought. but the water regime of uncon- solidated bottom is restricted to Scrub-shrub. The class "scrub- semipermanently flooded. shrubw includes wetlands dominated by woody vegetation <6 m tall. In Subclasses prairie wetlands, this class is mostly limited to temporarily Emergent wetland contains vege- flooded sites long protected from tation of the subclasses persistent fire and grazing by domestic or nonpersistent. wPersistentqt livestock. vegetation in prairie wetlands is normally perennial in habit and Forested. The class IfforestedM remains standing at least until the includes wetlands dominated by woody beginning of the next growing vegetation 6 m or more tall. In season. Persistent vegetation is prairie wetland, this class is the predominant subclass in nearly mostly limited to temporarily all prairie wetlands, often referred flooded sites long protected from to as "wet grasslands. " Nearly Eire and grazing by domestic live- always annual in habit, "nonpersis- stock. The class can sometimes tentw vegetation increases with develop where artificial disturbance falling water levels and disturbance creates sites favorable to germina- in prairie wetlands. tion and growth of seeds from trees with wind-disseminated propagules. Aquatic bed can include the subclasses '@algal,""aquatic moss," Rocky shore. This class is found eYooted vascular," or floating. only on a few large wetlands in the "Rooted vasculart"is the predominant Prairie Pothole Region where glacial subclass because the highly slumping and ice action have strewn mineralized and alkaline waters of boulders along high-energy shore- the region are not conducive to the lines. This class is seldom found growth of most moss@s, and the in the wetlands dealt with in this relatively windy conditions that report. prevail in the region usualPy move floating algae and vascular floating Water Reaimes plants into emergent wetland, where they can appear below the canopy, The water regimes dealt with in The subclasses aquatic moss and this report are temporarily flooded, floating vascular are found mast iACUSTRINE WETLAND SYSTEM
(non -wetland)
lnterrnitter~tlyExposed Sern~perrnanentlyFlooded Seasonally Flooded 1 Teniporarily Flooded
PALUSTRINE WETLAND SYSTEM
FRESH OR MlXOSALlNE EUSALINE OR HYPERSALINE
Intermittently \ \ \ Flooded
(zones of short persistent and errnaricntly Flooded Satur-atedl L.... non-persistent ernerqents (rone of tall persistent dependent on caoillary 11-ernporariiy Flooded ernergents clt?pendenl on groundwater and dtrect groundwate: seepage) precipitatron)
Figure 2. Arrangeinent of Cowardin et al. (1979) water regimes around lacustrine and palustrine wetlands in the Prairie Pothole Region of the Dakotas (from Kantrud et a!. 1989).
often in small basins, small bays of "vegetated." Unconsolidated shore larger basins, or openings in can quickly appear in seasonally emergent wetland, whereas the flooded wetlands, n~rmally of the subclass algal is most often seen in class aquatic bed, if drought small, highly eutrophic wetlands. conditions prevail early in the growing season. Unconsolidated share in the wetlands dealt with in this report Uncansolidated bottom in the is usually of the subclass prairie region is usually of the subclasses "sand" or "mud. a' Areal wetland system, subsystem, class, cover of vegetation in this subclass and water regime. These combi- usually does not exceed 30% nations are shown as legends (Cowardin et al. 1979) . However, in (alphanumeric codes) on the National semipermanently flooded wetlands of Wetland Inventory maps that are the Prairie Pothole Region, annuals currently being prepared for the can germinate and grow to cover United States. Copies of the values exceeding 30% during years of pamphlet Photointerpretive Conven- extreme drought (L.M, Cowardin, tsfoE. National Wetlands Northern Prairie Wildlife Research &ventor%' (19871, which contains an Center, pers . eomm. ) . explanation of these legends, and the status sf the wetland mapping Only the subclasses "broad-leaved effort for the region, is available deciduous" and "dead" are found in from the U.S. Fish and Wildlife forested and scrub-shrub wetland in Service, Regional Wetlands the Prairie Pothole Region of the Coordinator, National Wetlands Dakotas. Inventory, Federal Center, Denver, CQ 80225, The legends can be used to locate temporary, seasonal, and Relation to Other wetland semipermanent wetland basins by -.Classification noting instances where these codes are shown for the deepest or central This report includes only informa- area of a basin. The maps are tion on basins with the temporarily, intended to show the potential seasonally, and semipermanently natural vegetation.. Thus, culti- flooded water regimes of Cowardin et vated wetlands are designated as al. (1979). These water regimes emergent even khough emergent correspond to those found in the central zones of ponds and lakes vegetation may be absent. classified as temporary (Class 2), seasonal (Class 3), and semiper- Wetland Soils manent (Class 4) by Stewart and Kantrud (1971). s his system was The soils underlying wetland developed specifically fox the basins in North Dakota have been glaciated prairie region, and unlike classified, and classification is in the former system, was designed to progress for similar soils in South classify entire basins, rather than Dakota (Jimmie Richardson, North the central or concentric peripheral Dakota State University, pers. zones or bands of vegetation around comm.). The development of wetland the basins that are wetland of soils depends strongly on length of differing water regime in the flooding, water quality, and the cowardin et al. (1979) system. The terms "pondw and "lake" usually relative position of the basins with refer only to the water contained in respect to the ground-water table, the basins. Hereafter in this re- according to Bigler and Richardson port the terms temporary, seasonal, (L984), Richardson and BigPer (1984), and Fulton et al. (1986). and semipermanent wetlands will These sources and other material refer to entire basins, and the referenced by then show that corresponding zones will be termed temporary and seasonal wetlands are wet meadow, shallow marsh, and deep usually recharge areas that receive marsh, following Stewart and Kantrud water from the surxoundi ng uplands (1971). by runoEf or direct precipitation. Clays are dispersed and carried Wetland Inventorv Maps downward by an abundance of fresh water. This results in highly Use of the Cowardin et al. (1979) developed soil profiles with argil- classification system results in lic horizons. Soil series observed large numbers of combinations of in these situatians include Lindaas and Parnell (Typic Agriaqolls) and Dakota, averaged 6.2 (4.5-9.3), 0.46 Tonka, an Argiacquic Agialboll. (0.42-0,50), 12 (7-4-171, 0.04 (Q,Q2-0.87), and 0-99 (0.43-1.6) Most semipermanent wetlands in mg/kg dry weight of these metals, North Dakota are flow-through respectively. Sediments f rom wetlands that receive ground-water palustrine wetlands have discharge and also recharge the significantly higher concentrations ground water. Evapotranspiration, of arsenic, cadmium, lead, and frost action, and a near-surface selenium than those from riverine water table allow water to move from wetlands, but, with one exception, the wetland body to the soils in the concentrations were within normal or wet meadow. Reducing conditions in background ranges. the water column aid in mobilizing salts into the wet meadow. Several other little-understood factors result in higher soil salinity in 1.4 DlSTRIBUTlOM AND ABUNDANCE OF WETLANDS the wet meadow than in the adjacent uplands or the interior zones of the wetland basin. These flow-through During the 1960's and 197Q1s,2.3 wetlands have a large range of million temporary, seasonal, and salinity, and salinization of the semipermanent wetland basins, with whole wetland occurs in some cases. a total area of 1.04 million ha, Soils in these wetlands range from were estimated to be present in the Typic Haplaquolls and Galciaquolls Prairie Pothole Region of the in the wet meadow to dominantly Dakotas (Ruwaldt 1975; Stewart and Fluvaquentic Naplaquolls such as the Kantrud 1973; H.A. Kantrud, unpubl. Southham series in the shallow marsh data). The estimates were based on and deep marsh zones (Bigler and relatively small numbers of randomly Richardson 1984). It is in these sampled plats. Approximate basin basins that drainage followed by numbers and areas by water regime tillage often results in crop were 698,000 temporary (113,000 ha) , Eailure because land salinization 1,474,000 seasonal (583,000 ha) , and occurs as the ground-water system 127,000 semipermanent (345,000 ha). delivers low-quality water to the These basins were estimated to freshly broken soils (Richardson compose 84.8% of the area and 89.3% of the number of natural basins in 1986), the region; subsequent drainage and A small number of semipermanent filling has further reduced the wetlands in North Dakota can be number of wetlands. considered discharge wetlands. These are underlain by saline soils Distribution of these wetlands such as the poorly developed correlates with the distribution of Minnewaukon and Lallie series (Typic various glacial landforms (for des- FLuvaquents) . If drainage is criptions see Section 2.1) . In both attempted on these soils, saliniza- Dakotas, the greatest proportional tion prevents the growth of crops area of semipermanent wetlands 0ther than hay. occurs in areas of dead-ice and terminal moraine, whereas the Little is known about heavy metals greatest proportional area of in soils sf prairie wetlands. temporary and seasonal wetlands Plartin and Hartman (1984) examined occurs in ground moraine and lake arsenic, cadmium, lead, mercury, and plain. The largest proportional area selenium concentrations in sediments of semipermanent wetlands is found sf riverine and palustrine wetlands in the Kissouri and prairie Coteaus, of the North Central United States. whereas the less gemanent wetlands Samples from three palustrine wet- are best represented in the lands in the Cottonwood Lake area Glaciated Plains and Lake Plains northwest of Samestown, North (Figure 3). Density of basins can be r40/km2 much less for the entire region, in some areas of dead-ice moraine or because some glacial and postglacial ground moraine, but the average is landforms contain few basins.
I 7 Prairie Cotcad
Figure 3. Major physiographic regions in! the Prairie Pothoie Region of the Dakotas. CHAPTER 2. ABlOTlC ENVIRONMENT
2.1 GEOLOGY The spruce-aspen forests of what are now the northern plains were suc- ceeded by grasslands, and since that Glaciation time, periods of warm, dry condi- tions have alternated with periods About 7 million kears ago, the of cool, wet conditions. Some addi- subtropical climate of what is now tional basins were formed during the Dakotas began to change to a this period from wind-worked sand continental climate of cool winters dunes, but nearly all of the basin and warm summers (Bluemle 1977). wetlands in the Dakotas were formed During the Pleistocene Epoch that as a direct result of glaciation or followed, a succession of great ice the melting of glacial ice. The sheets inched southward from Canada area of depressional topography and covered most of the Dakotas formed by a variety of geological (Figure 4). These huge glaciers processes comprises the famed Prai- transported vast quantities of rock rie Pothole Region, which, until the and soil. Large amounts of local advent of European map, was an silty and clayey bedrock outcrops approximately715,000-km grassland- were also pulverized and added to wetland complex that stretched from the mixture, forming glacial drift north-central Iowa to central or wtillN that was deposited as Alberta. This region was unique sediment across most of the area because of its abundance of shallow- glaciated. These deposits may be >250 m thick in places and are basin wetlands, which attracted most dotted with shallow basins left by of North America's waterfowl during the scourinq or shearing action of the breeding season. the glaciers, or from the collapse of ice blocks left to melt in the Stewart and Kantrud (1973) esti- deposits after the glaciers mated that 93% of the area and 94% retreated. According to Flint of the number of wetlands in the (1955), many basins in South Dakota Prairie Pothole Region of North were formed when glacial drift Dakota were composed of natural materials blocked ancient bedrock basin wetlands, whereas the valleys, Additional basins were remainder were mostly streams and formed by materials deposited by oxbows, stockponds and dugouts, water flowing from melting glaciers. reservoirs, road ditches and Thus, all the basin wetlands drainage channels, and minor types discussed in this report occur in a such as sewage lagoons. These qeologically young landscape. figures show the importance of alaciation on the composition of Getlands in the region. The retreat of the glaciers is marked by the beginning of the Fhvsiuqra~hie~ivisians Holocene Epoch about 10,000 years ago, as winters became cold and ~eobotanists have traditionally summers became hot (Bluemle 1977), divided the northern grasslands into Figure4. Extent of Pleistoceneglaciationin North America in relation to Nonh and South Dakota (modified from BIuemle 1977). two large areas called the Great Missouri Coteau drains to the Gulf Plains and Central Lowland (Fenneman of Mexico via the Missouri River. 1931) (Figure 1). The more arid Great Plains supports native grass- By far the largest number and area land that is shorter than that in of basin wetlands in the Dakotas the moister Central Lowland to the occurs in the Central Lowland, east. The Prairie Pothole Region of Most of this land mass drains either the Dakotas encompasses portions of to Hudson's Bay (North Dakota) or both these areas. the Gulf of Mexico (South Dakota). Within the Central Lowland lie six major physiographic regions (Figure The Great Plains portion of the 3) . These are, in decreasing arder Prairie Pothole Region of the of area, the Glaciated Plains Dakotas contains a single physio- (Q?. 000 h2),Prairie Coteau (15.000 graphic division, the Missouri km j , Dakata Lake Plain (5700 km2), Coteau (Figure 3). This division is Souris Lake Plain (3600 km2), approximately 52,000 km2 in area. Devil's Lake Plain (14002km2), and The entire west slope of the Turtle Mountains (1200 kna f. All seven physiographic divisions deposited at the edge of a glacier contain a variety of glacial or while the ice margin was melting postglacial landforms. Those back at about the same rate as the landforms that contain relatively ice was moving forward (Bluemle numerous wetland basins will be 1977). Till is a general term for discussed in the following section. the mixture of materials ranging in size from clay particles to boulders of many tons that were pushed Glacial Morainic Landfoms forward by and carried on top of advancing glaciers. Terminal Most natural basin wetlands in the moraines are most common in the Prairie Pothole Region of the Glaciated Plains, but also occur in Dakotas are found in five glacial the Missouri and Prairie Coteaus morainic landforms (Bluemle 19771, (Figure 3). These moraines are Moraine means any materials commonly 2-15 km wide and 5-90 km deposited directly by glaciers. long. Basins in terminal moraine are-highly variable in size, depth, Ground moraine. This the and density. predominant glacial landform of the Glaciated plains (Figure 3), and can Dead-ice moraine. This form is be recognized by a gently rolling responsible for some of the most landscape with numerous shallow rugged glacial topography in the saucer-shaped depressions, but few Dakotas; it formed when glaciers hills or deep cup-shaped depressions advanced over steep escarpments. (Bluemle 1977). This landform Shearing action carried material occurs where moderate amounts of into and on top of the glacier glacial till were deposited at the (Bluemle 1977). This insulated the base of a moving glacier and by underlying ice, which took several collapse from within the glacier thousand years to melt and collapse. when it finally melted. When the overlyinq materials slumped and slid, thousands of basins of all Washboard moraine. This form shapes and sizes were formed. Dead- appears as small areas of irregu- ice moraine is the most common larly spaced ridges of material landform in the Missouri Coteau, thought to have been carried upward Turtle Mountains, and Prairie through the ice along shear planes Coteau. Smaller amounts of dead-ice parallel to the edge of the glacier moraine occur in the Glaciated (Bluemle 1977). Small basins are Plains. numerous in washboard moraine. This landform is mostly found in associa- Glacial Meltwater Landforms tion with ground moraine in the Glaciated Plains. Three glacial meltwater landforms contain significant numbers of basin Thrust moraine. This is perhaps wetlands. These landforms were the mast spectacular glacial land- created by water from melted glacial fsm, as it is the result of large- ice, including precipitation on scale glacial shearing tpt moved glaciers as they were melting blocks of land up to 20 km in area (Bluemle 1977). for short distances (Bluemle 1977). The "hole1@ left by these blocks Glacial outwash plain. This form commanly resulted in a large lake, consists of sand and gravel that was whereas the hilly blocks often con- deposited by water flowing from tain numerous small but relatively melting glaciers. In some of these deep basins. Most thrust moraine is broad plains are numerous large found in the Glaciated Plains, lakes, whereas in others are large numbers of very shallow depressions. Terminal moraine, This form This landform is mastly found in the resulted when glacial till was Glaciated Blains. Collapsed slacial outwash, This middle latitudes. Temperature and form resulted when glacial outwash precipitation data for Bismarck, ND, was deposited on stagnant ice. When have been summarized for a 30-year the ice melted, the sand and gravel period by Court (1974) and are shown slumped to form an irregular hilly in Table 2. This city lies at the surface with numerous wetlands, many west-central edge of the Prairie of them large, shallow lakes Pothole Region of the Dakotas. The (Bluemle 1977) . Most collapsed continental nature of the climate is glacial outwash occurs in the shown by the wide (85 "C) tempera- Missouri Coteau, but several large ture extremes and relatively low examples can also be found in the (385 mm/yr) precipitation. Glaciated Plains. Many of the largest basins in North Dakota are in collapsed glacial outwash, and The following data on climate and many of the lakes formed in these weather have been summarized from basins are highly saline. Visher (1966), Black (1971) and Jensen (n.d. ) . Glacial lake lai ins. These areas occur today where lakes of glacial Temperature meltwater stood for hundreds or thousands of years (Bluemle 1977). Temperatures form roughly south- Glacial Lake Agassiz was the largest to-north gradients in the Prairie of such lakes in what now are the Pothole Region of the Dakotas. Dakotas. This giant water body once Normal annual temperature ranges occupied a vast area encompassing from about 3 'C in northern North most of southern Manitoba and Dakota to about 8 "C in southern Ontario, as well as huge areas of South Dakota. In January, the Saskatchewan and northern Minnesota. coldest month, temperatures average Nearly all the glacial lakes -19 "C in northeastern North Dakota vanished when the glaciers receded and -8 "C in southeastern South and outlets were established. Topo- Dakota. During July, the warmest graphy is very flat in these areas, month, the range is 16 "C to 23 "C and wetland basins are very shallow. for the two areas, respectively. However, near the center of some Rapid plant growth begins when mean glaciated lake plains, a few large daily temperatures rise to 6.1 "C. remnant basins are still present. The date for this event varies from Most glacial lake plain in the about 26 April in northeastern North pothole region occurs in the Dakota, Dakota to about 3 April in south- Souris, and Devil's Lake Plains eastern South Dakota, a difference (Figure 31, but small areas also can of over three weeks. Conversely, be found in the Glaciated Plains and vegetative growth largely ceases Missouri Coteau. In the latter area when mean daily temperatures fall to and the Turtle Mountains, there are 1.7 OC. This event occurs about 20 also some small areas where glacial October in northern North Dakota and lakes flooded stagnant ice for about 10 November in southeastern shorter periods, resulting in hilly South Dakota. The normal length of topography with numerous basins when the frost-free season varies from the stagnant ice finally melted. about 110 days in northern North Dakota to about 150 days in south- eastern South Dakota. The frost- 2.2 CLIMATE AND WEATHER free season may be slightly shorter in basin wetlands, as cold air flows The Dakotas have a climate downhill and can accumulate in topo- characterized by relatively short, graphic depressions, Crops planted moderately hot summers and in and around wetlands often are the relatively l~ng, cold winters first to be injured by freezing because these states lie in the temperatures, giving rise to what middle of a large continent at some f amers cal h "frost pockets. *t Table 2. Cfirnalic data lor Bismarck, ND,1931-60 (from Coiirt 1974). -I--_. .--.__I- _ """ I-.--^II
.-- re~c.an_~~xcsj&i%at-ion Eean sn_owfaI$- ("C) (Elm 1 tmm) Flonth mean max. min.
Jan. -12. R Feb, -10.8 Mar. - 3.8 Apr. 6. a. May 13.0 June 18.1 July 22.3 Bug. 21.0 Sept . 14.8 Oct. 7.8 Nav . - 2.0 UQC. - 8.4
The higlraczst summer telnperatures ranges from 25 per year in northern otficially recorded in North and North Dakota to 35 per year in South Dakota to 1'345 were 49.4 "C southeastern South Dakota. This and 48.9 "C, rc~pectively, whereas combination of temperature and wind the lowcst official winter tempcra- differences cause the n~rmalannual turcs prinr to 1952 (the last year evaporation from pans to vary from for wftictl summarized data arc 1.02 m in northeastern North Dakota <%v&ilable) were -51.1 "C and -50 "C, to 1.65 m in southeastern South respectively. Soils usually freeze Dakota. to dcpths af 0.9-1.8 m in northern FJorth Dakota and 0.5-0.9 m in southc~ssternSouth Dakota. The relatively small amount of precipitation in the Canadian prairies has been attributed by Hare Surface winds average 21 km/hz in and Hay (1974) to the weakness of nostl~eastcrnNorth Dakota and 17,7 atmospheric disturbances and their kmPhr in svuthedstcrn South Dakota. associated uplift* Air masses move Mctroruloyists plot witlcis as net ex- eastward from the Rocky Mountains cesses of wind in a given direction and fall steadily toward lower during vimxiaus time periods, Far elevat ions in the northern prairies. the I"~~/QPI,there is a relatively The rate of fall is sufficient to large excess of wind from the nnrth- reduce cyclonic action appreciably, west in January, and a relatively thus seducing the effectiveness of small excess of wind from the north- the mechanism that causes precipi- east in July. Many low-pressure tation. This phenomenon also cells track fram west-narthxest to reduces precipitation in the Prairie east-southeast across the region Pathole Region of the Dakotas. The during the year, The nu&@s. crf southern part of the region has more spring and summer thunderstornns precipitation because it receives more moisture-laden aimasses from originating as Alberta lows. South the Gulf oE Mexico, Dakota has 2.1 blizzards per year, with 52% of them originating as Annual precipitation in the region Colorado lows. Snow from February ranges Erom 33 cm in northwestern and March blizzards and snowstorms North Dakota to 58 cm in south- is often an important source of eastern South Dakota. Urger spring runoff for prairie wetlands. amounts of spring and summer pre- cipitation in the southeastern part Nail falls in moderate amounts of the region account for most of compared to the rest of the nation, this difference. About 70% of the on an average of 2-4 days/yr. Brief annual precipitation falls as rain heavy rains occur more frequently during spring and summer, with June than in the arid west, but are only the wettest month. Distinctly dry rare or occasional compared to the years, having <75% of normal pre- southeastern United States and the cipitation, occur with 10% frequency gulf coast. Normal annual water in northwestern North Dakota, but loss by runoff and evaporation is only 4% in southeastern South Dakota. There is a large gradient 0.36 m in northwestern North Dakota across the Prairie Pothole Region of and 0.56 m in southeastern South the Dakotas in the length of the Dakota; the rest enters the ground. relatively dry season, that is, when weekly normals of <1.27 cm of pre- cipitation can be expected. In 2.3 HYDROLOGY northwestern North Dakota, this season averages 8-10 months, whereas Wetlands exist because specific in southeastern South Dakota, this geologic settings and hydrologic season lasts only 2-5 months. processes favor ponding of water or soil saturation. A unique combination of glaciation and Snowfall in the region averages <1 climatic conditions in the Prairie m/yr, much less than areas directly Pothole Region has produced a large to the east or west. In the dry number of dynamic aquatic ecosystems northwestern corner of the region, that have a tendency to be non- 25% of normal annual precipitation integrated (not receiving or falls as snow, whereas in south- contributing to channelized surface eastern South Dakota only 12% falls flow). Nonintegrated basins have in this form. Appreciable amounts the potential to impound large of snow fall on >60 days/yr in volumes of water and undergo long- northeastern North Dakota, whereas term, rather extreme changes in significant amounts fall on only water depth and biotic conditions in about 28 days/yr in southeastern response to climatic trends. Water- South Dakota. Dry winters, when level fluctuations in typical <7.6 cm of precipitation falls, seasonal and semipermanent North occur one year in five. Dakota wetlands are shown in Figrare 5. Note the greater rate of water Blizzards, as defined by the loss and greater frequency of National Weather Service, are storms dryness in the seasonal wetland. The with wind speeds ~51.5 km/hr, low-grade shorelines of prairie temperatures <-6.67 "C, and wetlands combine with semiarid visibility not >152 m. Reduced climate to produce dynaxaic wetlands; visibility is caused by snow, e. g. , small increases in water level blowing snow, or mixtures of snow cause great increases in the and soil. North and South Dakota proportion of a basin inundated, and have a far greater frequency of conversely, hot, dry conditions blizzards than any 05 the adjacent often remove surface water Erom states. North Dakota has 2,2 large areas of a basin in a blizzards per year, with 54% of them relatively short time. SEMIPERMANENT WETLAND
SPILLS
MONTH
SEASONAL WETLAND
MONTH
Figure 5. Water level changes during the ice-free season over a 6-year period in a seasonai(iower) and semipermanent (upper) basin wetland inNorth Dakota (from Kantrud et al. f 989). Hydrologic regimes are dictated transpiration are controlled largely by climate and geology (Figure 6) by wind and vapor fluxes above the that establish the environment for wetland. Areas like the Prairie hydrologic processes (Winter 1989). Pothole Region with persistent Atmospheric, surface, and ground winds, high summer temperatures, and water interact with basin topo- dry overlying air masses can be graphic setting and the hydraulic expected to have high evaporation characteristics of glacial tills to and transpiration rates during the establish wetland hydrologic func- summer months. tions. Precipitation and a combination of evaporation and Lissey (1971) found that most of transpiration are, respectively, the the water available for ground water major components of water gain and recharge is derived from spring loss in prairie wetlands (Winter and snowmelt. Snow collects in topo- Woo, in press). Evaporation and graphic depressions and 50% to 90%
CLIMATE BASIN PHYSIOGRAPHY BASIN MORPHOLOGY TEMPERATURE WATERSHED CHARACTERISTICS BASIN CONTOUR PRECIPITATION SUBSTRATE GEOLOGY (Zone Ratios) EVAPORATION SUBSTRATE PEDOLOGY SIZE MOISTURE ELEVATION STORAGE CAPACITY A. SURPLUS B. DEFICIT.
RECHARGE FLOWTHROUGH A SURFACE B GROUND DISCHARGE (Dead End) HYDROPERIOD CHEMISTRY TOTAL DISSOLVED SOLIDS NUTRIENTS BIOTA INTERACTIONS PLANT COMMUNITIES INVERTEBRATE COMMUNITIES VERTEBRATE COMMUNITIES LAND USE (Long Term) IDLE GRAZED CULTIVATED MOWED BURNED
Figure 6. Environmentai factors influencing prairie wetland ecology. af the meltwater from topoyrapfilc seasonally flooded; they hold water highs collects in depre~sisns for only a few months each year, and because snow melts before the yround the water is generally low in thaws. Iissey concluded thnt except dissolved solids. Some basins are for occasi onnl intense storms, through-f low systems with respect to summer rdinfall is not a major ground water; that is, ground water contributor to rec:harqc in the flows in throuyh parts of their bed up! antls. while other parts recharge ground water, Through-f low basins hold G l clcint, tnpocgr~aphy and qeoloqi- water over longer periods and the r.3 l i y yoverncd pcrmcability water tends to have higher concen- v,irintionn hnvc a sign1f icant ef tcct trations of dissolved solids. Some on surf at-e and yround-water hydro- basins serve only as discharge areas 1uqy . "rills in maraines are for ground water. Lakes that qcnerally silty and claycy materials rcceive discharge from both regional thnt &re not very permeG\ble, Out- and local ground-water flow systems w,l:;ii deposits, on the other hand, and do not lase water to seepage or qcnerdl l y consist of strat i f ied sand surface outflow are highly saline, and gravel, materials thdt dre very having specific canductance as high pcrmc(3bla. The ti11s ixl tht castern as 70 mS/cm. i7akotar; tend to be mur-h tliqhrr in chcrIe?-dcrivecl material. These tills Wetland hydrologic functions <%PC more clayey and less permcable control the chemical characteristics to wntor than those to thc west and of prairie lakes, and as a result, south, which have a larger propor- plant and invertebrate communities. C iorr of 1 imestone, r;'tndct-.o~~c.,anti ci ltstonc dcrzvatives (Winter 1983). ilocnuli~ pra i r io wc?t 1 ancis are 2.4 WATER QUALITY t-.f~~~r-rrct-erist~caX 1 y nun intoysdtcd, ground-w<~ter f-law systems play a The shallow, eutrophic character- doxninnrlt r-cr le in their hytlrnl oq ips. istics of mast wetlands in the I,] n:;ay (1971) d~*scribrd the Prairie Pothole ~egion,coupled with pr i ncip lcs of dtlprcss io:1-f.caci1r;cetl warm summers and cold winters, cjrc~und-watox recharge <$rid dlscharqe routinely produce a chemical en- in lhc.2 ylt~c:l~stcdprEfirie c)t south- vironmcnt that is hostile to many wr8:;tcrn M~~raitob~~.l,~tI\nucjh et dl+ aquatic vertebrates. Wetlands that jZ'a87) studicd the k~grdroloqy of a expcriencc anaerobic conditions wd-t l,rrrcr: ccsmplex in tirc Cot t otlwood during summer or winter are referred Ipakeb &rc$il ak Sttltrzman County, NU, to as summcrkill (Kling 1975) and thot c~nte%i ned wet l .r~ncfss i t U
EVAPORATION AND PRECIPITATION TRANSPIRATION
Deep marsh Deep marsh Permanent Permanent Low pralrje Wet meadow Shallow marsh Erner~enl Open woter Open woter Open water Pilore Phase jL1ttot01) (Profundal) SURFACE SURFACE RUNOFF OUTFLOW
GROUND-WATER
DISCHARGE Ep~l>mnioo
Thermocline
GPOUWO-W 4TER RECHARGE
Figure 7. Dominant sources of water in prairie wetlands. CVAPORATION AND PRECIPITATION+ TRANSPlRATIOhl+
SURFACE OUTFLOW
Figure 8. Dominant ffow pattern of a flow-through type of pralrle wetland contarnlng pooled water low in dlssoivsd salts.
CVAPORAT ION AND PFII CIf'II A I ION TRANSPIRATION
Prrs,>rir;t.,rs O;it.il u0,l.r ,P.ot,no.:
liHO1JMI.>-WATf R i 1,. 0 r, . ,, r. ------ihnrrnot:iino
GROUND-WATER RECHARGE
Figure 9, lfiomlraant flow pHtern af a cfased-systemtype of prairie wetland containing pooled water lnaemecllate rn dissolved salts.
20 EVAPORATION AND PREClPfTATiON TRANSPIRATION
€pilimnlo- GROUND-WATER DISCHARGE ...... Thermocline
inirolittoiol
Figure 10. Dominant flow pattern of a hydrologic-sump type of prairie wetland containing pooled water that accumulates dissolved salts.
removed from dry basins or the dry Topography and the geologic shorelines of wet basins through characteristics of glacial tills wind erosion. Wetlands that lose provide the framework within which water to surface outflow remove ground water flow systems develop. salts at rates that are a function of annual turnover rates (volume Till in south-central North Dakota lost versus wetland storage has a high percentage of silt and capacity) . As turnover rates clay, is poorly sorted, and has low increase, the salt content of the permeability; outwash is largely wetland approaches that of the sand and gravel, is well sorted, and dominant water source. Water loss has high permeability (Swanson et to ground water also removes salts al. 1988) . Significantly higher from the wetland. The rate of loss average values of specific is influenced by the permeability of conductance and concentrations of the till that controls the rate of sodium, potassium, sulfate, and export. Wetland basins that do not alkalinity were found in wetlands lose water to surface outflow, but located in outwash, whereas signifi- are ground-water through-flow cantly higher average concentrations systems, tend, because of low perme- of calcium occurred in wetlands ability, to be higher in dissolved located in till (Swanson et al. salts. Wetlands that lose water 1988). Most wetlands that had primarily to the atmosphere by values of specific conductance evapotranspiration tend to concen- greater than 20 mS/cm in south- trate salts to high levels (Swanson central North Dakota were located in et al. 1988). outwash, and nearly all wetlands located in till had values less than Wetlands differ from one another 10 mS/cm. Saline wetlands located in their chemical characteristics in outwash are topographically low, based on their location in ground- nonintegrated ground water discharge water flow systems (LaBaugh 1989) . areas that function as hydrologic
CHAPTER 3. BlOTlG ENVIRONMENT
PHYTOPLANKTON, PERIPHYTON, AND a ground-water recharge area and METAPHYTON dries up by midsummer (LaBaugh et al. 1987) . Following snowmelt and flooding of dead shallow-marsh vege- All algae, including wetland tation, dominated by Carex atherodes algae, are grouped according to the and Pol~qonumcoccineum, decomposi- habitats they occupy. Floating tion begins. Shallow-marsh vegeta- algae are referred to as metaphyton. tion decomposes rapidly when flooded Algae suspended in the open water and releases nitrogen and phosphorus column or pelagic zone are referred that are available for use by algae to as planktonic algae (phytoplank- because this wetland has few sub- ton). Algae that grow attached to merged vascular plants competing substrata and are a component of the with the algae for nutrients. When periphyton (aufwuchs) are termed the water recedes, filamentous algae periphytic algae (Crumpton 1989). dominate the wetland surface until Periphytic algae can be further complete drawdown. New growth of divided into those that grow within shallow marsh vegetation and leach- bottom sediments (epipelic algae), ing of nutrients from the decom- those that grow on plants (epiphytic posing algae complete the cycle. alaae) , and those that arow on rocks (epilithic algae) (~ru<~ton1989) . Semipermanently flooded wetlands also establish seasonal patterns of Aquatic macrophytes serve as a algal growth that tend to alternate substrate for epiphytic algae. Thus with the growth of submerged vascu- seasonal changes in macrophyte lar plants and periodic drawdowns. abundance can influence the abun- Wetland P11 in the cottonwood Lake dance and distribution of epiphytic study area functions as a ground- algae as well as epipelic and water discharge area and is domi- epilithic algae that are often nated by open water and beds of the shaded by the macrophytes. submersed macrophyte Potamoaeton pectinatus. During spring the shallow water in this wetland is The role of algae in the primary turbid (Secchi disc depth=l0 em) and production of prairie wetlands is dominated by planktonic algae. As not well defined (see Chapter 4). the summer progresses, epipelic and epiphytic algae and P. pectinatus Patterns of algal growth in stabilize the flocculent substrate seasonally flooded wetlands are and the water clears. During fall, influenced by hydroperiods and the when the P. peetinatus and benthic apparent uptake and leaching of algae break down, the water again nitrogen and phosphorus associated becomes turbid and planktonic algae with shallow marsh vegetation dominate, Waterfowl contribute to (LaBaugh et al. 1987) . Wetland T8 the onset of turbid conditions by in the Cottonwood Lake study area excavating the substrate in search (Winter and Carr 1980) functions as of P. pectinatus turians. Aquatic invertebrates, which arc composition of hydrophyte com- important toods ot laying female munities in the region. Fire is dabbl i ng ducks and their broads, considered a human-related dis- tend to graze on pcrlphyton or turbance and is included in this filter phytoplankton, bacteria, and section, even though lightni ng-set organic detritus from the wdter (abiotic) fires commonly occurred on column. the prairies in pristine times (Higgins 1986). Lists of all hydro- Phytoplankton species occupying a phytes found in North and South wetland complex in the Cottonwood Dakota are found in Reed (1988a, b). 1,iike area of North Dakota are presented in Appendix A (1,aBaugh The early studies of Shunk (1917) and Swansun 1988). Wetland T8 is and Metcalf (1931) identified the seasondl l y f loaded and Eunct ions as major features of prairie wetlands d ground-water recharge area. Wet- (central or concentric peripheral land TJ is seclsonallly flooded and zones of vegetation) and the factors functions as a through-flow system. that seem to control plant species Wctland Pf is a semipcrmdnent wet- composition and distribution of land that does not have a surface these zones (water depth, length of outlet, but Ptjnction:;
The vegetation of nearly all prairie basin wetlands is composed of a central zone bounded peri- I nf ormcat 1 on on the c:nn;position, pherally by concentric zones (Figure r.tru~:ture, and olnssif ic Pond Class Ill Seasonal Pond or Lake MARSH 7ONE Class IV Semi-permanen t Pond or Lake Class V Permanent Pond or Lake Class Vii Fen Pond Class VI Alkal~ Pond or Lake Figure 11. Spatial relation of vegetational zones in Stewart and Kanlrud ('1971) classes of natural basin wetlands in the glaciated prairies. of the zones even though individual shallow-marsh zones in central North species have discrete distributi.ons. Dakota probably averages 40 mm/day The number of zones present in a from early .4pril until the end of basin increases with the degree of June (Kantrud and Stewart 1977). water permanency; that is, the length of time the zones can be Deep marsh zon~.These zones are expected to contain surface water. either dominated by tall, coarse The best accepted names for these cattails or bulrushes with an under- zones in the basins dealt with in story of submersed or floating this report are, in order of in- plants, or by beds of submersed creasing degree of water permanency, vascular plants, unconsolidated wet meadow, shallow marsh, and deep bottom devoid of vegetation is marsh. Wet-meadow zones can be sometimes found. Water regime is dominated by woody vegetation and usually semipermanent, with surface not appear as meadow. The appear- water present from spring through ance of large amounts of woody fall and frequently overwinter. vegetation around prairie wetlands Water loss in centrally located is often attributed to fire suppses- deep-marsh zones in North Dakota sion by European man (Bird 1961). wetlands likely averages 25 mm/day Shallow-marsh and deep-marsh zones (Kantrud and Stewart 1977). can be dominated by submersed plants. Wet-meadow and shallaw- Some characteristics of these marsh zones are commonly cultivated zones are evident. In a given and can be devoid of hydrophytes basin, water depth increases with when dry or for short periods after zonal water pemanency, but this reflooding. often is not true among or between basins of different hydrological Wet meadow zones. These zones of setting. For example, shallow-marsh prairie wetlands are mostly domi- zones in many freshwater wetlands nated by low to medium-height are often deeper than deep-marsh grasses, rushes, sedges, and forbs zones in more saline wetlands. In or woody vegetation. Submersed or all but the most saline wetlands, floating plants are absent. Inun- the stature of emergent vegetation dation is usually temporary far a increases along the gradient from few weeks or months in spring or temporarily to semipermanently briefly after heavy summer rains. flooded. Species richness of the Water loss in centrally located wet- emergent component of the vegetation meadow zones in North Dakota can in these zones tends to decrease average >50 mm/day from early April with water permanence. Deviations till the end of May (Kantrud and from the '*normalw zonation pattern Stewart 1977). occur regularly in the form of zonal inversions or deletions brought Shallow marsh zones. These zones about by fluctuating water levels are usually dominated by coarse (Smeins 1967; Millar 1976). Zonal emergent grasses, sedges, and deviations can be used to interpret burreed of midheight or a few, often the recent hydrologic history of a nonpersistent, forbs. Submersed or basin wetland. floating vascular plants or aquatic mosses or liverworts are fairly Veqetation Dynamics common, and can occur in the under- story of emergents or in shallow Prairie wetland vegetation changes open water, Except for the spring more dramatically than most other snow-melt period, the presence of natural vegetation in North America open water in shallow marsh zones is because of the effects of regional largely a function of land use. climatic instability on hydrcfogic Inundation is normally seasonal from regimes. Late winter snowfall, spring to mid- or late summer. spring runoff, summer precipitation, Water loss in centrally located and evapotranspiration are all highly variable between years and Stewart and Kantrud (1971) con- within seasons. ~xtended droughts sider the degree of interspersion of as well as short series of years cover and open water mostly a func- with much higher than average tion of water depth rather than precipitation are common. water permanency. Studies by Weller and Spatcher (1965), Weller and High water levels can kill up to redr ricks on (1974) and van der Valk 25% of the emergent vegetation in and Davis (1976, 1978a) on semiper- prairie wetlands, but when water manent Iowa wetlands outline the levels fall, new bands of vegetation cyclic effects of physical and quickly develop on newly exposed biological forces on the species shorelines (Walker 1959, 1965). composition and distribution of Many emergent species such as Ty~ha wetland vegetation. spp. reproduce vegetatively except during low-water conditions, when The stages in this vegetation reproduction by seed is common cycle are shown in Figure 12 and its (Bedish 1964). Extremely high water effects on the biota are outlined in levels can also kill trees and other Table 3. In the dry-marsh stage, woody vegetation that have developed water is absent or low in the in wet meadow zones, leaving a ring central zone and muskrats can be of dead trees that can persist for absent. Vegetation on the marsh years. floor consists of mudflat annuals DRY MARSH ,I , ,%: I SENESCENCE MARSH SEASE SECTS BANK GH WkTE9 LAKE MARSH ~EEZ: CE3hxa!tar.TIr.*, DEGENERATfNG MARSH Figure 12. A generalized vegetation cycle in an lowa prairie wetland (van der Valk and Davis 1978a). Table 3. Stages of the typicat habitat cycte in semipemanent towa wetfands (Wefler and Spalcher 19651. Staqc karcr ;n relac:?- )J4s,.r3% Bird nanc :c pasic capsc-tj :'e.~et-sz:an popu:atlons pcpdlat ions -- Dry mar%h Aase?t cr lo*: 3ense re;.egatat lor:: Low to absent ; Red-uinqed Red-wlnqed b:ackbrrds; e-argents dry rjr nost scec:es fina populatrons blackblrds few muskrat lodges: nearly dry at base a s..-table seedcec centrally sparse; sone low water Locared .,-,,e by upland t ;rds Dense rn~lrsh: mere :?crc3sing vater '.'ex-) dense; rate of :ncreasl;lq Numbers and Hed-winged blackbirds vegetat ron than 1eve:s: ezergents openkng dependent varlety Increase; f lrst ye1 1 ow- open water t Loaded upon mzskrat poptila- increasing headed blackbirds appear tions and rnfidence near sparse open pools: of f:oodina cn cer- few coots and ta.n specles grebes tjernl-mapsh: open water Ee~ranto near Yaskrat eat-out; Tncreaslnq Maxlmum specles Many red-winged and and vegetatron LT! r~xlnupi flotation and death; rapldly; well drverslty and yellow-headed black- about equal amounrs decllne in sha1;ch- distributed productron for birds uniformly dlstrl- water specles; veg- most specles buted; coot and pied- etatlve propaqatlon billed grebes abundant Open marsh; more Submergents and deep- Maximum or nost specles in sparse bird populaticns open wa'er than water emergents per- declining decline; a few and emergent vegetatron vegetation slst; others absent swrxnrng specles or dlsappearlnq tolerate as long as long as some vegetation persrst Ope? wazer marsh; Maxlmun or as 1o.x Sclrp,~sacutus may Sparse: bank Red-wlnged Red-winged blackbirds vlrrually a a- nedaan persist ln sparse dens corrmon blackbrrds use use shoreline shrubs eutrophic laiie populations shcrelrne vege- and trees tatlon; other specles vrr- tually absent and seedlings of emergent species. hydrological conditions of prairie The annuals can complete their life wetlands. cycle and add seeds to the seed bank during this stage. With the resum- ption of normal precipitation, the Effects of Water Oualitv wetland enters the regenerating Much information has been gathered marsh stage. Muskrat populations on the effects of salinity (concen- begin to recover. Mudflat annuals tration of total dissolved solids in decompose and are eliminated, but the soil or water column) on prairie propagules of submersed and floating wetland vegetation since the early species from the seed bank germi- obsewations of Bailey (1888) and nate, and within a few years the Visher (1912). Such information has emergent species spread vegetatively been summarized recently by Kantrud and normal patterns of zonation are et al. (1989). Areas containing established. Major inputs of seeds of the emergent species to the seed large numbers of wetlands of rela- bank occur at this time. If water tively high salinity are mostly levels continue to rise, emergent restricted to glacial outwash plains vegetation begins to die from excess lying in the western and north- water depth, senescence, disease, western portion of the Prairie and insects. This is the degenerat- Pothole Region of the Dakotas. ing marsh stage. Continued high water results in the. lake-marsh The gradient from fresh to hyper- stage dominated by submersed vege- saline water is a continuum, and any tation only, because of excess divisions are arbitrary. In this muskrat damage and the inability of report we refer only to the salinity seeds of emergents to germinate subclasses of Stewart and Kantrud under deep water. Finally, drought (1971) and the water-chemistry modi- or artificial drawdown return the f iers of Cowardin et al. (1979). wetland to the dry-marsh stage. Surface water in temporary and seasonal wetland basins in the The cycle described above for Dakotas is usually fresh or slightly semipermanent wetlands in Iowa is brackish, although a few moderately idealized, and the role of the brackish seasonal basins have been muskrat is more important there than observed (Stewart and Kantrud 1971). in most of the pothole region of the The fresh and slightly brackish sub- Dakotas (see Section 3.7). Much more complex patterns of vegetation classes of Stewart and Kantrud change occur in nature because of (1971) correspond to the fresh (a0.8 partial drawdowns or multiple mS/cm) and lower 01 igosal ine (0.8- drawdowns over short time periods. 2.0 mS/cm) water-chemistryrnodifiers The less permanent temporary and of the Cowardin et al. (1979) seasonal wetlands do not underso classification system. such complex cycles, but alternafe on a wet-dry basis nearly every Semipermanently flooded wetland year. In addition, in the more arid basins, and their associated wet- Dakotas, water levels can fall so meadow, shallow-marsh and deep-marsh rapidly during drought that germina- zones, can be fresh, slightly tion of mudflat annuals cannot keep brackish, moderately brackish, up with the receding waters. This brackish, or subsaline. Most sf results in the drawdown bare-soil these basins are slightly brackish. phase described by Stewart and The moderately brackish and brackish Kantrud (1971) (Figure 13). Never- subclasses of Stewart and Kantrvd theless, the rapid recruitment of (1971) correspond to the upper plants a few species at a time from oligosaline (2 .. 6-8.0 mS/cm) and a much more species-rich seed bank lower mesosaline (8.0-15 mS/cm) seems to be a universal phenomenon modifiers of Cowardin et al. (1979), related to the rapidly changing whereas the subsaline subclass NOKMAI WATER CONDITIONS includes the range sf the to estimate the salinity tolerance mesasaline (15-30 mS/cm) and poll'- of many emergent and aquatic bed saline (30-45 mS/cm) modifiers. species in Appendix C. These are crude estimates because specific The general effect of increased conductivities of surface waters and salinity in any zone of wetland soil extracts from the root zone vegetation is to decrease the number often differ greatly. of species. In polysaline wetlands, only the single submersed angiosperm Plany species have a broad range of Ruppia maritha occurs. A similar salinity tolerance. This is a com- reduction in species is noted during mon trait among hydrophytes of the drawdown conditions (Stewart and Prairie Pothole Region. Species Mantrud 1971, 19721, when mono- that are tolerant of high salinities dominant stands of Kochia scaparia can survive low salinities (Ungar are often found in dry polysaline 1966). This suggests that compe- basins. There also is a tendency tition or other factors play an towards succulence in plants found important role in determining in more saline basins. species composition in less saline sites (Ungar 1974) Salinity levels can fluctuate . widely within and among seasons, Disturbance particularly in smaller wetlands and those of intermediate and high salinity (Rozkowska and Rozkowski Human-related disturbance is one 1969, Stewart and Kantrud 1972) . of the most important factors af- These changes can be accompanied by fecting species composition of wet- changes in vegetative species corn- land vegetation in the northern position (Ungar et al. 19791. For prairies (Smeins 1967, Walker and example, aquatic bed dominated by Coupland 1968, 1970). The intensity Chara sp., Zannichellia palustris, of disturbance is often more impor- and Potamo~eton pectinat2 often tant than the type of disturbance, alternates with stands of Ruppia and the native hydrophytes in the area are adapted to respond quickly maritima in certain prairie wetlands to changing conditions and form as surface waters are respectively dense stands. Common dominance diluted and cancentrated. types of wetland vegetation in various water regimes are related to Common dominance types of emergent disturbance and idle conditions in vegetation under various land uses Appendix B. and water chemistry conditions are shown for wetlands with temporary, Drainage and cultivation are the seasonal, and semipermanent water rnost extreme disturbances seen in regimes in ~ppendix B, along with most prairie wetlands in the similar information for the class Dakotas, although in some instances aquatic bed. the basins themselves have also been destroyed by filling with earth or It is digficult to establish rocks, or used for solid-waste meaningful salinity tolerances for disposal (see Section 5.31, individual species in their natural habitats, because of the complex of Basins that are cultivated but not factors associated with salinity drained are extremely cormon in the fLuctuations and ecotypic variations Dakotas (Figure 14). Stewart and among species (Kantrud et a1 . 1989) . Kantrud (1973% estimated that 29% of Neverthelessr we present the com- the area and 52% of the individual bined data of Smeins {196Sj, ~isrud wetlarids in the Prairie Pcthole (1.968) , sletten and Larson (1984), Region of Msrth Dakota had been and H.W. Kantrud (unpubl. ) , as cultivated far enrap production in published in Kantrud et aP. (1989) 3,967, a year of excellent water FIgsrrB 14 @ultiv@t& lemp~rarilyend seasenalty t1-M baslns dat the landscape In the lntosnslvely famed rrgrons 06 thc Dakotris (near takota, PIorlh Dakol;~.Jiinc 1978; photograph by Alan Sargeant). c-ox-itl-r t 1 ow. These lnrld~ were cultivated shallow-marsh zones as p~.~rn~tri1y tt*m~xrary drld sease3naX. canopies of emergents are opened and 'I'l~ea w~f.".me-,.xdow and stra? law-marsh much easif y cslonizable eanconsol i- rnrj.cbt;(1 ,I matry semipcrrnan~ntwet 1 nnds dated shore is created. ,ax r2 elso reyularl y ruX t ivated. X~ux'sl'lq P lip pralorlycd cxt rcmc dt'auqht I>sawdown species invade cwltivated ~)r't 1 1 0,t kac? b)ot lcmn of -an~%l~y basins very rapidly as water l3.s:. I~P;with f;blmipik~-m~incntly f toodcd recedes, farming plant associations .srr~l i t19: tar m i t t c*~~ti y cxpuscd water dif fererlt f rola those found in r tZrj>me!: wtlrc USCC~lor crop pr~duc- similar basins in grassland. This t.irrn irx 9 hr* I>,%koC,tu. .ii~dimkbrltnt inn is the cropland drawdown phase of 16: t~xtP f*:n~~ly ca~rin10~3 t n bas ins la- Stewart and Kanttud f 197 2) , c ,rtc.d krt c.rnpla.tnd in this IZXC~, as tll~-rcx i:; no hrz i er to runof l water Basins cultivated intensively t u asm t ha^. ~lpl seasonally and semipermanently several wetland zones (Stewart and flooded prairie wetlands, Twenty- Kantrud 1971) that are detemined by one species of snails have been interactions between basin hydro- identified in the Hudson Bay logic functions and basin moqfao- drainage of North Dakota and logy. Large shallow-marsh zones sf eighteen species in the Mississippi semipemanently flooded wetlands River drainage. Mussels and most contain invertebrate species that pill clams are confined to flowing are similar to those found in water and permanent lakes. isolated seasonally floded basins. Climatic conditions cycle wetland The distribution of snails in basins between extremes of flooding prairie wetlands is influenced by and drawdown that alter zmes and hydroperiod and salinity. Molluskan their assseiated invertebrate fauna species associations described by {Swanson 198Pb). Wancara (1983) for North Dakota are controlled by wetland hydrologic e ircum- functians. Each wetland contains striatus associations are found in vata train Helisorna anceas, COTTONWOOD LAKE $8 1980 Physa qyrina, Gyraulus panus, and SE1f4, SEG.24, T142N, R67W Staqnicola elodes (Cvancara 1983). The first five species are found STUTSMAN COUNTY, NORTH DAKOTA primarily in permanent lakes, How- ever, permanent lakes that contain large semipennanently and seasonally flooded zones will contain molluskan species that are associated with these zones. Wetland basins dominated by deep- marsh and shallow-marsh zones sup- port pulmonate snails that produce relatively thin shells and do not have gills. Members of this sub- class can occupy habitats that routinely exhibit anaerobic condi- tions. They respond to low dissolved oxygen by moving to the surface where they can use atmo- SCALE spheric oxygen. Factors influencing *-"------.-- - gastropod distribution include water 0 100 M permanency, total dissolved salts, dissolved oxygen, and substrate characteristics. Waters with >20 rnS/cm conductivity do not support. 1 DFEP-MARSH ZONE (OPEN-WATER PHASE) mollusks. Some species, however, 2 DEEP-MARSH ZONE (EMERGENT PHASE) such as Staqnicola elodes, Physa jennessi., and Armiqer crista, tend 3 SHALLOW-MARSH ZONE to tolerate elevated concentrations 4 WET-MEADOW ZONE of dissolved salts (Cvancara 1983). Mollusks occupy spring seeps asso- ciated with saline lakes (Swanson et al. 1984). Mollusk scientific Figure 17, Wetland P8 of the Cottonwood Lake, ND, names used in this report follow study area showing vegetative zones. those used by Clarke (1981). Aqua- tic mollusks play an important role in the food web of prairie wetlands (Cvancara 1983; Swanson et al. 1974; seasonally flooded basins or sea- Pennak 1978) . sorlcill y f loodcd zones of semiper- nianerrtly f looded basins. Other Crustaceans are a major component molluskan species found in this of the invertebrate fauna of prairie association inci ude -EPrgr;~cngt~~s wetland complexes. Species struc- -urnbiii~:atcl?gs_, -- - Armiqeg c~i-%L@~ture within a wetland complex is S&=j- GP 2.-a ~-i~-~\__r3~il.,Phvsa influenced by hydroperiod and jcnnessi , p?pmene"c. cxacuous , and salinity. Two species of amphipods, pisidium casertanurn (Cvancara 2983) . Gammarus lacustris and HyaleLla azteca, dominate the macrocrustacea Welisoma trivolvA-Lvmnaea stagna- of semipermanently flooded wetlands. --lis assaciations are found in perma- The former species occupies wetlands nent and semipermanent1y f looded that have an extended wet phase, are lakns. Xaflzskm species that are Icw in dissolved salts, anti have part of this association include relatively deep, cool water. Cinci~natia cincinnatie~isis~kmni- Hya11ela azteca occupies most semi- cola limosa, Pisidium nitidurn, Val- permanently flooded wetlands. Both species routineYy invade seasonally Insects, particularly Coleoptera, flooded wetlands by attaching them- Odonata, and Wemiptera, play an selves to aquatic birds and mammals important role in the ecology of during the summer months, but are prairie wetlands by assuming the eliminated during drawdown (Swanson role of top predators within the L984), Psnphipods are replaced by water column. Wetland insects salt-tolerant Anostraca as salt con- differ from the gastropods and centrations increase (Swanson et al. crustacea of prairie wetlands in 1984) ., that most are capable of flight and can quickly invade shallow wetlands soon -after-they receive water. This Oecapods are not found in noninte- applies especially to adult grated wetland basins in south- Coleoptera; members of this group central North Dakota, but are found are found in seasonally flooded in oxbow lakes and integrated wetlands soon after spring snowmelt wetlands in the headwater areas of is complete (Ranson and Swanson tributaries that feed perennial 1989). river systems. Descriptions of insects often omit Eubranchiopods, such as Anostraca information on habitat. Light traps (fairy shrimp), Conchostraca (clam that attract adults from a rela- shrimp), and Notostraca (tadpole tively large area are rautinely used shrimp), are the dominant macrocru- for collecting. This procedure, staceans occupying temporarily and while efficient for taxonomic seasonally flooded basins in the studies, cannot be used to associate Prairie Pothole Region. Fairy adults with the types of aquatic shrimp are also found in large wet- habitat that support larvae, meadow and shallow-marsh zones of semipermanently flooded wetlands. Insects are similar to gastropods Species composition of fairy shrimp and crustacea in that their species populations change with increasing structure is influenced by hydro- salt concentrations of the water period and salinity. A study of the column. Amphipods and fairy shrimp Coleoptera of seasonally and semi- are occasional 1y present in the same permanently flooded wetlands in a wetland, ~rine shrimp (~rtemia Cottonwood Lake wetland complex salina) predominate where salt demonstrated differential use of cancentrations are >35 mS/cm. these wetlands bv srsecies (Table 5). Of the 57 specie; izentified, 47 and Invertebrates occupying the water 38 were collected on seasonally and column of a prairie wetland complex semipemanently flooded wetlands, in the Cottonwood Lake, ND, study respectively (Hanson and Swansan area are listed in Table 4. Wetland 1987); 18 and 11 species were found T8 is seasonally flooded and func- only on seasonally and semipema- tions as a ground-water recharge nently flooded wetlands, respec- area. Wetland T3 is a seasonally tively. Individual species showed flooded through-flow system. Wet- preferences for one of the wetland land PI is a nonintegrated, semi- classes even though they occupied permanent wetland that has no both. For example, only one Asabus surface outlet and functions as a antennatus was found on seasonally ground-water discharge area. Wet- flooded wetlands, compared ta 172 land P8 is a through-flow, semiper- specimens collected sn semiperma- manent wetland that contains a nently flooded wetlands. On the surface outlet. Wetland PI1 is a other hand, 88 Wvdroporous semipermanent wetland that has no fusci~enniswere found on seasonally surface outlet, functions as a flooded wetlands, but only 2 on ground-water discharge area, and semipemanently flooded wetlands. concentrates salts by loss of water Qf the CoPeoptera reported for the to the atmosphere. State of Narth Dakota 39% were found ------in- io 0' Lnm L?P 13- WPiD ,, ,, ,, ,, -13%------m --..-moor- mr-ma r winmm - -4------ . .- -- - m- r-m-- --wr- -m ..?-r- --mm I Ii] I m m ma - oomo t- md0 h ---*.- -- -*nslrc; P, \DPQPP or- 0dUU.c ------nm I 034J E-~mdCIm do- N 0 lll rn rumx au3 "" m u ------mr-w -w - w -In nmn -? p. N d 2'0 E "0 sn~~aznqnssn~ueyyj[ 8'0 6'0 L'O 4'9 PVP 2'8 srpquo~3snq.uey;y 2-0 P'O sr~-pursuoasnquem T*O> T*O> 5'0 2'0 L"0 s~u~7~esnssapor? X"O> T'O Z'O Z-0 Z '0 snap~ouoaSTU~O~~P? 9 '0 0'1 snsoxnaeu snx~ydo33e? 8 ' 0 P'T: 2'0 8"0 5.T snJeq3nbrq SnT?ydO33P? T'O> x-0 snTn3lnbTq SnrqATI 2'0 2.0 T"0 sn-tnaxa3eq snTqnrI Z"O 1'0 2.0 O'E 5.0 0.s snpTqxn3 sn3ozbnH 1.0 -r -0 Snq€?~~aSSnqOXbAH L"0 E 'T T -0 T '0 1 "0 f-nes snqoxbn~ T"O> T'O> T 'O> snqe~~asnqoxbng 2'0 E'O TmO> S'T fl'Z 8'0 stfanzqea sn~oxbng Z'O T'O Z"0 E*T Z'O T'Z sn~eq3unaossa~am~snqo~bn~ ST E'Z 8 ' 0 JPClUoD SnTOlbAH 8'0 1.0 Ei " T srsuap~u~asnqoxbxjq T'O> T'O> sapyoxezm snqox~n~ Tea> T'O> T'O> ST E'Z 8'0 snsoxqaua? snoxoaozpnH L'O ZsX C'O snu~a~~aasnolodoxpn~ T'O> T'O> 1 'o> T 'o> sjx~qeqouSnoxouoxpnR T'O> T'O> z-z 9'2 6.1 s~uuaa~3sn3snoxouo~pn~ T 'O> 1'0> snoa~yaS~DT~P~AH 1'0 1.0 T'O> X'O T.0 snqsapom snarqepn~ 2'0 E-0 T"0 snxaxaxaa snzapoqaez3 2'0 T '0 C'O 6-0 9'1 SrTequapyaDa snxapoqdex3 C"O T'O s.0 snprxqny snDs?qna T'O T 'O> 1.0 1.0 2'0 ?xa?pxo5 sn~srqna Te0> Too> T'O> T'O> sn3~u?~utnaz?3sn3s~qna T'O> T'O> snueysexe snas~qna T'O 2 ' 0 T'O X'O 1.0 SnpbUOT snuoqoquo3 6 ' 0 E 'T 4'0 9'2 9'Z 9'Z s~x?qaxn3ssaqaquhTo3 2'2 2'2 Z'Z snqeTnqauna snqeby xeo> 1-0 TTT~Jsnqew 6'0 E'O T "T srsuapeue~snqeby TOO> T'Q> T'O> ST O'T 8"T snyzej~qsnqebv P '1: 9 ex T 'T 1 "O> T 'O> sn3euuaque snqPbp 'I '0 Z'O T'O> snqebv E '9 8'8 6°C 8"6Z G"ZZ L'GE aep$3s?3lia L"L 8.01 L ' t. 6"OP 0'82 57s ejcaqdoaxog Table 5. (Concluded). --" Seasonal Semipermanent wetlands wetlands - -- T3 T8 Total PI P8 Total Hal iplidae Haliplus ho~~inqi Bit ipl.us immaculic011 is paliplus strisatus 1 in1us mbgutt-s --Peltodytes gdentulps -Peltodvtes tortulosus Hydrophilidae Berosus fraterngs Berosu~hatch i g~~-osu,sstriat!.a* Mel~pmob3,on~us &&oahorus linearis ~~cwhorusl ineatua Bx@mb&~LLL.~&B HJ&~&K&~b&2&,8 HYdrosk&w ~Q&~.LLEK ~JX~I~~QiAgs @AX~"L~ r.i 5s '!' ~~~l2"%~~rrn~s &&%? xi3 1i ,5 PI iL&3-% US on the 97-ha Cottonwood Lake study 3A area (Hanson and Swanson 19139). The subject of fish use af prairie wetlands has recently been reviewed No1 son and Butler ( 1087 ) examined by Peterka He states that :;c.alaanal. abundance of larval chiro- (1989). nomids and emergence patterns of fathead minnaws (Pime~hales for two years on two wctland promelas) and brook sticklebacks rici~lt~ (Culaea in-cpnstans) are the only two classcs an the Cottonwood L,akc; study fotlnd that seasonally native fishes that can tolerate rc They shallow water depths low L loodud wetlands had lowcr larval (<5rn), avnsities than semipermanently dissolved-oxygenconcentrations,and occasional high concentrations of f Xooded wet1 ands, and that dens it les of emerging adults wcrci lower in the sulfates and bicarbonates found in these wetlands. Law dissolved oxy- s~asonafy flooded s itcs. t sen concentrations cause both ;interkill and summerkill. We also Fresh and oligosaline wetlands noted that most wetlands in the (Cowardin et ax. 1979) in the region lie in closed drainage Prairie Pothole Region support a basins, thereby limiting fish typical species structure of aq'clatic dispersal. Fish are limited to an invertebrates as described for the even greater extent in the wetlands United States by Pennak (1378), but dealt with in this report, as only as salt concentrdtiuns increase, palustrine systems and lacustrine only the most sal t-to1 erant groups littoral systems, which extend to a such as the share flies (Ephydridaef depth of 2 rn below low water, are persist. included. Fish cannot survive year round in limit for survival of native fish in temporary or seasonal wetland basins magnesium sulfate and sodium in the Prairie Pothole Region, chloride type waters in the prairie because these wetlands dry out or region (Rawson and Moore 1944; freeze completely or nearly to the Burnham and Peterka 1975). For bottom each winter. Interpolated sodium sulfate and sodium bicar- data of Schoenecker (1970) and bonate type waters, 12,000 mg/l and Barica (1979) for Nebraska and 2,000 mg/l seems to be the upper Manitoba, respectively, indicate limits (Held and Peterka 1974; that wetlands in the Dakotas remain McCarraher 1971) . frozen for about 4-5 months each year with maximum ice depths of 3.5 AMPHIBIANS AND REPTILES about 0-7-0.8 m. It is likely that a few temporary and seasonal basins According to Wheeler and Wheeler with riparian connections to more (1966), the herpetofauna of South permanent wetlands receive an influx Dakota includes 40 species (12 of fish in the spring during years amphibians and 28 reptiles), and of of high water levels, but unless North Dakota, only 25 species (11 water levels are maintained, these amphibians and 14 reptiles). Of fish will soon perish. It is also these, only the tiger salamander possible that some temporary or (Ambvstoma tisrinum), American toad seasonal wetlands are temporarily (Bufo americanus) , Great Plains toad stocked with fish, fry, or eggs by (B. coanatus) , Dakota toad (B. waterbirds, but this method of hemio~hrvs), Rocky Mountain toad (B. dispersal is undocumented (Peterka woodhousei) , chorus frog (Pseudacris 1989). niarita), leopard frog (Rana pipiens), wood frog (R. sylvatica), only in the few semipermanent painted turtle (Chrvsemvs picta), wetland basins with reliable plains garter snake (Thamnophis connections to deepwater habitat or radix), and red-sided garter snake other refugia can one expect to find (T. sirtalis) use prairie basin fish, unless they have been intro- wetlands. The only species duced. Lawler et al. (1974) found intimately associated with these fathead minnows and brook stickle- wetlands are the tiger salamander, backs in 10%-20% of the wetlands in leopard frog, and chorus frog. the prairie region of Manitoba, but Tiger salamanders are biologically if the wetlands that contained water well adapted to the dynamic physical >2 m deep had been eliminated from and chemical characteristics of this sample, the percentages would prairie wetlands, while major undoubtedly have been much lower. aquatic predators and competitors such as fish are poorly adapted to Wetlands where fish survive in their harsh chemical environment and high-water years can lose all fish recurring drought (Peterka 1989). when low water levels prevail. In Larval tiger salamanders have been some of these wetlands, refugia from collected in West Stump Lake, NR, in winterkill can be provided by water with specific conductivities ground-water seepage areas where of 12.5 mS/cm (Deutschman and some oxygen is present (Peterka Peterka 1988). Larval densities in 1989). North Dakota wetlands can exceed 5,00O/ha and maximum annual produc- Fish survival in a few semiper- tion of 565 kg/ha has been recorded manent wetlands can be limited by (Deutschman and Peterka 1988). Sal- excess dissolved solids, as well as amander larvae at high densities are shallowness and low supplies of dis- an important component in the energy solved oxygen at various times of flow of the wetlands they inhabit. the year. Total dissolved solids concentrations of 17,000-18,000 mg/l Amphipods (Gamarus lacustris) seem to be the approximate upper accounted for 78% of the volume of food items in stomach contents of when gad water canditians prevail, salamanders in North Dakota (Myers %out 23% of the Prairie Pothole 1973) . Gamarus spp, tend to occupy Region lies in the Dakotas. Haray wetlands that function as thraugh- wetlands in the region are also flow systems; these wetlands are importan$ fall staging and mignratioa relatively stable and low in salts. areas (Kantrud 1986ks1, The opera, Large salanrander lama@ (neotenics) shallaw wetlands found in cultivated occasionally overwinter in wetlands fields are also heavily used by and at this time can impact the spring migrant waterfowl. For invertebrate comunity, example, arctic-nesting geese stop in the Prairie Pothale Region to Aquatic birds that normally feed fatten while enroute to their on fish also feed on lama1 breeding grounds (Alisauskus 19881, salamanders. Wiedenheft (1983) Availability of eshallaw wetlands in reparted finding wounds on the open, agricultural landscapes also dorsal surface of the tails of 10% allows the birds to disperse widely, of a salamander population. thereby reducing or eliminating Potential predators include the potential hazards from disease, grebes, herring gu11s (Xfasus particularly avian cholera (G. arsent&us) , American white pelicans Krapu, pers, comm. ) . Faanes and Stewart (1982) list 23 species of Anatidae that have nested at least once in Morth Dakota. The trumpeter swan (Cvgnus buccinator), Leopard frogs have an immense white-winged scoter (pelanitta range in North America. On the fusca) , and coanmon merganser (- aakata prairies, these frogs seem to merclanser) have been extirpated as prefer temporary and seasonal breeding species in the state. The wetlands for courtship activities, snow goose (Chen caerulescens) is but lay most aE their eggs in listed as an occasional breeder, but semipemanent wetlands where the has nested only once in the state, tadpole% require about 3 months to and that under unusual circum- motamarphose inta adults. stances. Rare breeders include the American black duck (Enas rubri~es), The diminutive chorus frog is cinnamon teal (Anas cvano~tera), faun& aver most of North America. cornwon goldeneye ( clanau- The frogs e~ngregateto breed in the &) , and buff lehead (E3. albeola) . early sprinq in seasonal and semipemgnent- wetlands. After breeding, they wander away from The remaining 15 species are well- water bodies, but still require damp established breeders in the Prairie places to avaid desiccation. Pothole Region of Morth Dakota. These include the Canada goose (Branta canadensis) , woad duck (& 9,green-winged teal (Anas crecca) , mallard (A. platyxhvnchos) , northern pintail (A, acuta) , blare- winged teal (A. discors) , northern shoveler (A. clvpeata), gadwall (A. The North Aneri~anPrairie Pothole &e~era), American wigeon (W, Region produces the bulk oE North --americana), canvasback (AvtbVa &America % cluck population, Smith et valisineria), redhead (A, ameri- al. (2364) estimated that the canaf, ring-necked duck (11, csl- region, camprising only 10% oE the laris), lesser scatap af f inis) , waterfowl breeding area sf the con- hasded merganser (Lopsdvtes cucul- tinent, produces 50% af the ducks in +atus) and ruddy duck (Oxvura an average year, and more than that ~amaicensis), 46 Ruf fiehead a.nd common galdeneye do breed in the Dakotas and throughout not breed in South Dakota (Whitney the Prairie Pothole Region. et al. 2978). The trumpeter swan has been restored as a breeding Wetland use by waterfowl can be species in the state, but nut in the expressed in various ways. Use Prairie Pothole Region. The expressed on an area basis results American black duck, hooded in high relative values for small merganser, and common merganser are basins, whereas use expressed on a rare, but probably breed in the wetland unit basis results in state. Cinnamon teal breed relatively high values for large regularly west of the Missouri River basins, even though many large in South Dakota, but are rare in the basins have large central areas of Prairie Pothole Region, leaving 14 open water that may receive little species of waterfowl as well- use. Use of basins by breeding established breeders in the Prairie waterfowl has also been expressed in Pothole Region of South Dakota. pairs per unit length of shoreline. This also tends to result in high values for small basins. Waterfowl In 1973-74, about 69%-79% of an use of basins has also sometimes estimated 439,600 to 1,067,500 pairs been expressed in terms of basin of breeding waterfowl in South area without regard to the presence Dakota were located in the pothole or absence of surface water (Kantrud region (Brewster et al. 1976;. The proportion of birds in the region and Stewart 1977). This is done in was undoubtedly much higher before attempts to evaluate an entire the construction of thousands of wetland class, not only those basins dams and dugouts for livestock within a class that happen to con- watering facilities in the western tain ponded water at any particular part of the state. time. Tem~orarv wetland basins, The Stewart and Kantrud (1973) esti- primary function of these wetlands mated that about 84% of 2,281,000 is to provide isolation for breeding pairs of breeding waterfowl were pairs and supply invertebrate foods located in the pothole region of for waterfowl early in the nesting North Dakota in 1967. This region period. The rapid warming of these comprises 51.5% of the state area. shallow wetlands in the spring They estimated that the breeding results in early development af waterfowl population in the pothole invertebrate populations (Swanson et region of the state averaged about al. 1974). Temporary basins, 1,600,000 pairs (995,000-1,955,000 especially those containing flooded pairs) during 1967-69. crop residues, receive considerable use by spring migrant waterf owl, but Because prairie wetlands are of are usually dry during the fall extreme importance to an inter- migration period. PJeverthePess, national resource, the numbers of these basins are occasionally filled breeding, molting, staging, and by fall rains, in which case they migrating waterfowl observed in may be used to a considerable extent these basins, and the foods consumed by fall migrant dabbling ducks, by these birds in various stages of their life cycle, have received Up to 6,8% of the South Dakota considerable attention by wildlife breeding population can be found an biologists. A current review of these basins when water conditions these topics is available (Swanson are relatively good, and the water and Duebbert 1989). The temporary, ponded there makes up about 4% af seasonal, and semipemanentwetlands the surface water available in all dealt with in this report are by far wetlands (Rrawaldt et al. 1979). the most important types of wetlands About 2%-4% of the breeding water- for most species of waterfowl that fowl population was observed on temporary wetlands in North Dakota grea$es$ pair densities sn these during the 1967-69 seasons (Stewa~ wetlands, on the basis af Baath basin and Kantrud 1973) . For both of area and basin mi%, were, in these studies, the proportional use decreasing ordear sf magnitude, found of the basins was underestimated B b because many intensively tilled wet- rd lands, many undoubtedly of the sn temporarily flooded water regime, green-wemged could not be classified as to water ct gadwall (&& regime, Cowardin et al. (1988 and unpubl. Very silnilar results were rqearted data) used wetland data from the by Kantrud and Stewart (1977) for late 1970's to early 1980's and temporary basins in Morth Dakota. waterfowl pair counts from the mid- They also observed use of these 1980% to estimate the density of basins by small numbers of &erican five common species of breeding wigeon (& gmericana) . They found dabbling ducks in various classes of that, on the basis of area of basins wetland basins during a wet year in with surface wader, temporary wet- the Dakotas. This analysis showed lands showed greater use by dabbling that 9,9%-13.4% of the mallards, ducks (364.2 pairs/)om2) than any 6.7%-10.6% of the gadwall, 4.3%-6.3% other wetland class. But when all of the bluewinged teal, 7.7%-11.2% basins were considered without of the northern shoveler, and 13.3%- regard to the presence or absence of 17.1% of the northern pintail surface water, temporary basins papulations could be expected to be supported 126.6 pairs/km . This was found on temporary wetlands. The only about 47% and 79% of the dab- relatively greater use of temporary bling duck density found on seasonal wetlands by early-nesting species, and semipermanent wetlands, respec- mallard and northern pintail, is in tively. agreement with the food-habit studies of Swanson et al. (1985) for Seasonal wetland basins. These the mallard and Krapu (1974a, b) for basins are a major source of inver- the northern pintail. These studies tebrate protein for laying female showed that temporary wetlands are ducks early in the breeding season, a major source of the animal protein and also provide isolation for pairs needed by nesting hens to produce and sites for averwater nests. clutches of viable eggs. Poor- During wet years, these basins are quality diets lead to reduced clutch also highly attractive as brood and egg size, laying rate, and habitat (Talent et al. 1982) and nurslber of nesting attempts (Eldxridge molting areas. Seasonal wetlands and Mrapu 1988) . That mallard usually receive considerable use by breeding populations correlate spring migrant waterfowl, but better with acreages of water 15 em normally are dry by fall and little OP more deep than with acreages sf used by fall migrants unless water 30 as 46 cm or more deep unusually wet conditions prevail. provides further evidence of the Under these circumstances, seasonal importance of shallow water to wetlands can be heavily used by early-nesting species (Stewart and dabbling ducks, especially those Kantrud unpubl,). species that regularly feed in upland grainfields. Temporary basins are used only by breeding dabbling ducks (Anatini) in During May and June of relatively the Dakotas, Use of these basins wet years, 14.5%-16.2% of the decreases greatly during relatively breeding population can be found on dry springs, as ponded water can be seasonal basins in South Dakota. absent or maintained for only a few These basins contain 9,8%-11.0% of weeks. In South Dakota, eke the surface water available in all wetlands under these conditions pairs/b' of colllbined diving and (Ruwalc?t et al. 1979). In North stiff-tailed ducks. Unclassifi- Dakota, over 47% of the breeding able, intensively tilled wetland duek population was obsercred on basins, many of which are also these basins over a three-year likely of the seasonal water regime, period (Stewart and Xantrud 1973) . supported 193.2 pairs/km2 dabbling Again, the proportion of the water- ducks and 6,8 pairs/km2 of cowined fowl population that was obsemed on diving and stiff-tailed ducks. They these basins is ~Llnerrstimaked in also observed that, when the bath these studies, because sf the presence or absence of surface water difficulty ~f classifying inten- was not considered, seasonal wetland sively tilled wetlands early in the basins supported a greater density growing season, (287.2 pairs/)un2) of breeding water- fowl than any other elass of natural Because of their greater average basin wetland. depth and water-holding ability, seasonal basins are used by breeding Semi~ermanent wetland basins. diving ducks (Avthvini) and stiff- These basins supply most of the tailed ducks (Oxvurini), as well as needs of all common prairie-nesting dabbling ducks. In recent years, waterfowl and their broods. Use of use of seasonal wetland basins near semipermanent wetlands by breeding wooded areas by wood ducks (Aix waterfowl seems to be greatest when sponsa, tribe Cairinini) has also amounts of emergent cover and open increased in the Dakotas. water are approximately equal Schultz On the basis of both basin area (Weller and Spatcher 1965) . (1987) showed that livestock grazing and basin unit, seasonal wetlands in of Typha stands in semipermanent South Dakota received the greatest wetlands could result in a tenfold use by blue-winged teal, followed in increase in pairs of breeding ducks. decreasing order by mallard, north- Use by broods can increase greatly ern pintail, gadwall, northern during years when seasonal wetlands shoveler, redhead (Avthya ameri- are dry (Talent et al. 1982). A cana) , green-winged teal, ruddy duek notable exception to the relatively (Oxvura i amaicensis) , and American high use of these wetlands occurs in wigeon (Ruwaldt et al. 1979). early spring. Semipermanent wetlands are among the last to become ice- Seasonal basins are used even more free in the spring, and thus they do heavily in North Dakata , where not provide an early source of diving ducks are more abundant and invertebrate foods for nesting dab- widespread than in South Dakota. In bling duck hens, particularly those addition to the species recorded in that can begin nesting in late HarcPn seasonal basins in South Dakota by or April. Ruwaldt et al. (1979), similar basins in North ~akota-also sup- Semipermanent wetlands are also ported lesser scaup (Avthva af fin- the nain general habitat: for staging ) canvasback (A, valisineria) , and fall migrant waterfowl in the and ring-necked duck (& collaris) Dakotas and the rest sf the Prairie (Kantrud and Stewart 1977). As in Pothole Region, althaugh internit- South Dakota, the main users of tently exposed and pemanenkly seasonal wetlands were blue-winged flooded lacustrine wetlands are niost teal, northern pintail, mallard, heavily used by certain species, qadwall, and northern shoveler, ~antrud and Stewart (1977) also found that, on the basis of basins Use of semipemanent wetland with pondekl wares, classifiable basins by dabbling dau~ks increases seasonal wetlands in North Dakota during the breeding season. In supported 317.2 pairs sf dabbling South Dakota during 1973-74, Ruwaldt ducks per square kikonaeter and 16.6 et al, 41979) found 29.8%- 44.6% and 41.7%-47.2%, respectively, of the men water ~onditisnsare goad in May and June breeding duck South Dakata, semipemaneat wetXand population on these basins. During ba~liaag suppa& %he highest: densities these months semipermanent basins of blue-winged teal, followed in accounted for 35%-42% of the decreasing order by he mallard, surface-water area in the state. nosthern pintail, gadwall, remead, Semipemanent wetlands composed 18% northern shoveler, ruddy duck, of the area and 3% of the number of green-winged teal, and Arnericatn all wetlands, and 16% of the wigeon (Ruwaldt et af. 1979). In breeding duck population was found North Dakota, highest pair densities an these basins in the Prairie on ponded sexnipemanent basins an Pothole Region of North Dakota in the basis of area were also recorded 1967 (Stewart and Kantrud 1973), fox blue-winged teal, followed in The following year was relatively decreasing order by remead, gad- dry, and 41% of the breeding duck wall, mallard, ruddy duck, northern population was observed on pintail, northern shoveler, canvas- semipemanent wetlands, back, green-winged teal, lesser scaup, ring-necked duck, and Ameri- Although the proportion of the can wigeon (Kantrud and Stewart breeding duck population found on 1977) - Use by all dabbling ducks semipermanent basins can increase and combined diving and stiff-tailed greatly during relatively dry years, ducks was 189.1 pairs/kmz and 73.2 duck densities on these basins can pairs/km2, respectively. decrease, Ruwaldt et al. (1979) observed 6-73 duck pairs per wet Semipelrananent wetland basins are basin (2.57 pairs/ha) when water the principal breeding habitat for conditions were good and the water diving and stiff-tailed ducks in the in the basins nade up 42.3%- 44.6% Dakotas. During 2973-74, these of the surface water area. At this basins supported 72.2%-100% of the time, 24.8%-42.7% of the total duck populations of these birds in South population was observed on these Dakota (Ruwaldt et al. 1979). In basins, During the following North Dakota, 74.9% of these birds relatively dry year thase basins were found on semipermanent basins still held J5,6% of the available during 1967-69 (Kantrud and Stewart surface water, but duck densities in 1977). The redhead, canvasback, and them fell to 3,6 pairs per wet basin ruddy duck nearly always nest over (2.07 pairs/ha) . At this time, water in emergent vegetation in 26-68-47.2% of the duck population these wetlands, whereas nests of the was found sn these basins. This lesser scaup and ring-necked duck phenomenon is presumably caused by are usually found in adjacent wet the lack ax! enough water to praduce xneadaws. Redheads often lay their foods in nearby temporary and eggs in the nests of other seasonal baains (Krapu 1974a, overwater-nesting ducks. Swansan et aP, 1974) and the loss of surface water in the shallow wet- Breeding waterfowl obtain most of meadow and shallow-marsh zones of their foods from wetlands, although the semipemanent weelands them- field feeding in upland habitats is selves. Northern pilltail, blue- eoman during both spring and fall winged teal, srnd narthexn shoveler migration. Food consumption by seem particularly subject to such actult and juvenile Anatini (dabbling annual declines in density in ducks), Aythyinae (diving ducks), semipermanent basins {Ruwaldt e@ al and Qwyurini (ruddy duck) during the 1979). Annual population densities breeding season in the Prairie of these species have been shown ta Psthale Region was recently reviewed be highly correlated with the by Swanson and fllPebbert (1989). Ereequency and area of wetlands containing surface water in North Food consumption varies greatly Dakata (Stewart and Rantrend 1974), among species and between sexes of adults of a species; it also depends largely an relative availability of the food items as well as the Approximately 39% of tne 353 valid wetland erne, salinity, and vege- species on the North Dakota bird tative zone of the wetlands where fist (Faanes and Stewart 1982) use the birds were collected. For these wetlands. Of the 223 species with reasons, only a superficial account known or inferred breeding status in af this complex subject can be North Dakota, 57 (26%) are marsh or presented here, aquatic birds other than waterfowl. Consumption of invertebrates by Of these 57 species, the spectacu- the most camon species of laying lar whooping crane (Grus americana) female Anatini is very high, varying has been extirpated as a breeding from 72% by volume for the gadwall bird in the state. Distribution of and mallard to 99% by volume for the nests and records of adults during northern shoveler and blue-winged the breeding season indicate this teal. Foods of male Anatini during species once nested throughout the the breeding season vary from 70% Prairie Pothole Region of North seeds by volume for northern Dakota (Stewart 1975). pintail to 97% invertebrates by volume for northern shoveler. Of the 56 remaining species, 15 are generally not associated with Juvenile Anatini of all species, the shallow-basin wetlands dealt especially during the first few with in this report, although many weeks of growth, rely heavily on of them can be seen resting or animal foods. All species inves- feeding in or around these wetlands, tigated to date consume large both during the breeding season and amounts of either Insecta, Crustacea in migration. This group is corn- or Nollusca. Exceptions are older posed of the common loon (Gavia juvenile gadwall and American immer), red-necked grebe (Pociiereps wigeon, which consume mostly 9rrseaena) osprey (Pandion vegetable material. heliaetus), bald eagle (Hafiaeetus leucocewhalus), yellow rail Adult female Aythyini and Oxyurini (Coturnico~snoveboracensisj, cornon also rely heavily on invertebrate snfpe (GalJinaaa aallinaso) , foods during the laying period. American woodcock (golo~axminor), Consumption varies from a low of ring-billed gull (Lams deb- warensis) , California gull {Lams 70% by volume for the redhead to a californisus), Caspian tern (Sterna high of 99% by volume for the ruddy casrria) common tern hirundo), duck (Woodin and Swanson 1989). , (S, Chironomidae (Diptera) are important least tern (S. antillarum), belted for several species. Invertebrate kingfisher flycatcher ( use by most breeding male ~natiniis swamp sparro similar to the females. An excep- tion is the canvasback, which con- sumes about 98% vegetable foods by There are eight rare to uncommon volume during the breeding period. species that likely nest, have nested, or are inferred to nest in Nearly all juvenile Aytbyini and vegetative zones found in wetlands Qxyurini studied to date consume 80% dealt with in this report, but ar more invertebrates, by either little is known af their habitat weight or volume, mostly mphipada, use, This group sf birds includes Trichoptera, and Diptera, The only the least bittern (Ixobmchus exception is the redhead, which exilisj, little blue heron (Earetta Bartonek and Hickey (1969) showed caemhea) , tricolored heron (Earetta to consume 43% invertebrates by tricoLsr), cattle egret (BubuPcus volume. ibis), green-backed heron (Butorides mtus) king rail ( density because they are based on elesans), sandhill crane the entire wetland, rather khan on ) and white-faced ibis specific zones or plant cornunities 1 - The sandhill used by the birds. On this basis, crane likely was a fairly common me highest densities of birds breeding species in North Dakota occurred on semipemanent wetlands, before the turn of the centuq These wetlands are especially (Stewart 1975). attractive to Werican coot and black tern. Temporary wetlands An additional 11 fairly common to supported high densities of red- abundant species regularly breed or winged blackbirds and savannah nest in various vegetative zones sparrows. Killdeer and marbled found in wetlands dealt with in this godwit were also found in greatest report, but little quantitative densities on these wetlands, habitat use information is available although most of the use was of for these birds. These species shoreline areas. Horned grebe, include the American white pelican eared grebe, and willet were ( ervthrorhvnchos), double- especially dense on seasanal crested cormorant ( wetlands. Agricultural use of wetlands obviously decreased their value to most species except for the American avocet and Wilsonqs Forster's tern phalarope, both of which reached highest densities on wetlands with frequently tilled soils. Faanes (1982) censused breeding birds of all species in seasonal and semipermanent basin wetlands and other habitat types in a study area in central North Dakota during For the remaining 22 species, a April;Sune 1980. Me recorded 13Ei7.8 fair amount of infomation is pr/h in semipermanent wetlands. available on use af wetland habitat This density was exceeded only in in North Dakota. These 22 species shelterbelts (2658.7 pr/h2] . Sea- are most characteristic of the sonal wetlands supported 574 pr/km2. wetlands dealt with in this report; This density was exceeded only by the distribution of 21 of them among shelterbelts, semipermanent wet- the subject wetlands is shown in lands* and prairie thickets (878.5 Table 6, Nearly two-thirds of the Pr/b 1 total population were found on semipermanent wetlands, and a13 Faanes (1982) also found the species used this wetland class, highest nesting or breeding densi- Far 11 species, over half the ties of Forster's tern, red-winged population was found sn semiperma- blackbird, and brown-headed cowbird nent: wetlax3ds, Seasonal wetlands (Eo1othrus w)in semipermanent were used by 20 species and wetlands, and noted that breeding supparted about one-third of the song sparrows also used the peri- total population, From 53% to 64% phery of this wetland class. He of the popuhation of seven of the also observed breeding pairs sf species occupied seasanal wetlands. bobolink ( 1 I Temporam and undifferentiated- savannah sparrow, and shanp-tailed tillage wetlands were of little sparrow in seasonal wetlands. importance ts the birds listed. Infomatian on frewency sf use of Densities are expressed in pairs temporaq, seasonal, and semiperma- per square kilometer in Table "7'. nent wetlands was garnered for 13 These are crude measurements of selected n~nwaterfowl species in 52 - 3- m- 8: Ch 6-3 'dG916X nn'n r-n 0 QJC 52D P-Y 3 Dl 3t-J IU Y r. r+ mn'm 13 m m r rca Y P m=rrert rt 5 r. F-' P- Q -1; C 3 C P m3QJr.3 8-e V1 P-m @. TY3t? mom P, PC mmm Q t~ r-Y r (1 Y rm&I.;l r.Y m 3c+ rplpl t GZrsP, m PI 8 o rr ri.P n rm mvmrur: Y n' 0 r-Y r 00 QJ V - a2E.e 13P- $2 0 tf V vl V C1 ro N i-t w WIP 0 IP ra0WmululNawh)W h) ul Cn WQF~W@MOW4CnWPulPWQCnW & & Table 7. Density (pairs/km2) of breeding marsh and aquatic birds in Nafih Dakota wet!ands in 4965 and 1968-69(from Kantrud and Stewart 1984). Wetland class" Species 2 3 4 T Horned grebe Eared grebe Western grebe Pied-billed grebe Black-crowned night heron American bittern Northern harrier Virginia rail Sora American coot Piping plover (Charadrius melodus ) Killdeer Willet Marbled godwit American avocet Wilson's phalarope Black tern Marsh wren Common yellowthroat Yellow-headed blackbird Red-winged blackbird Savannah sparrow Total 633.1 431.8 723.8 89.3 'Temporary (Class 21, seasonal (Class 3), semipermanent (Class 4) and undifferentiated tillage (Class T) wetlands. Souf;ln Dakota in 1975 and 1976 (Weber migrants and summer visitors. Cul- et ale 1982). Among the wetland tivated and non-cultivated seasonal classes listed (Table 81, the and temporary wetlands provide habi- highest frequencies for eight tat far millions of arctic- and species occurred in semipermanent subarctic-nesting shorebirds that wetlands that were used by 92% af pass through the Prairie Pothole the species. Seasonal wetlands were Region each spring. Ten species of used by 85% af the species, and most shorebirds studied in North Dakota fremently by the black-crowned foraged primarily an crustaceans and night heran and Wilson9s phalarope. insects in these shallow basins Temporary wetlands were used by 54% during spring migration (Eldridge sf the species and most frequently and Krapu in prep.), In one study by the American avocet, lesser area in central North Dakota, Faanes yellowlegs a~dmarbled godwit> f1982) found use of semipermanent wetlands by white pelican, great Very Little infomatian is avz%i2- blue heron, tundra swan (m able on use of prairie wetlands by colurribianus),bufflehead,Califarnia 54 Table 8. Frequency of occurrence of "1 selecad species of marsh and aquatic birds in South Dak~ta wetlands during May and June suwreys In 6975-76(from Weber et at. 4982). Wetland classa Species 2 3 4 merican bittern 0.8 6.3 Great blue heron 1.2 Green-backed heron 0.4 Black-crowned night heron 3.3 1.2 Sora 3.4 5.5 American avocet 1.4 0.4 Lesser yellowlegs 2.7 1.9 1.5 Willet 4.1 3.4 4.7 Marbled godwit 4.1 1.5 2.3 Wilsonqs phalarope 8.2 9.8 7.4 Black tern 1.1 6.3 Yellow-headed blackbird 2.7 6.8 37.5 Red-winged blackbird 37.0 51.9 74.2 '~emporary (Class 21, seasonal (Class 3) and semipermanent (Class 4) wetlands. gull, ring-billed gull, and rusty quantitative data on this subject blackbird (Euphaaus carolinus) to be for the Dakotas, but it is obvious equal Lo or greater than other to anyone familiar with the habits wetland types. Similar data for of this species that it readily seasonal wetlands indicated high use seeks the thermal cover and protec- of these basins by greater yellow- tion from predators and hunters legs (Trinqa melanoleucus), solitary afforded by the dense stands of sandpiper (zsolitaria), short- emergent hydrophytes. There are billed dowitcher (Limnodromus large areas in both states where ariseus), Bairdts sandpiper (Cali- wetlands provide pheasants nearly dris bairdii), pectoral sandpiper the only protection available from (C. melanotos) , stilt sandpiper the harsh winter winds common to the (Micro~almahimantopus), and water northern prairies. This species is pipit (Anthus ~pinoletta). The an important game bird in both highly variable water conditions Dakotas. In 1986, 42,000 hunters that prevail across the region harvested an estimated 122,008 birds emphasize the need to maintain a in North Dakota (Grandah1 1987) . Rn widely dispersed and intact base of estimated 800,000 birds were har- shallow basins to serve the function vested in South Dakota in 1985 QRsn of spring migration habitat, as deep Fowler, South Dakota Game, Fish and water does not provide an availabPe Parks Dept . , pers . comm. ) . invertebrate faod source for birds that are moqhologically adapted to 3.7 MAMMALS feeding in shallow water. Historic Use sf Prairie WetPands W discussion of the use of prairie wetlands by birds would not be com- There can be little doubt that: the plete without mention of the impor- activities sf the wild bison (Bison tance of these basins t~ wintering bison), which was extirpated fro% populations of ring-necked pheasant the Prairie Pothale Region sf the (Phasianus colchicus). There are no Dakotas in the 19th centuq, had a major biotic influence on prairie species to his list and added infor- wetlands in pristine times. Unfor- mation on range and habitat use. tunately, there is no documentation The latter was presented in Jones et of how wetlands were impacted by the al. (1983) in an attempt to cat@- feeding, drinking, dusting, or other gsrize mammals into groups, often activities of millions of these arbitrary, that would indicate the huge, shaggy beasts as they roamed degree of dependence of current. wild the prairies. other grassland mammals upon basin wetlands in the mammals extirpated from the region region (Table 9). Only wild maimals are the grizzly bear (Ursus arctos), in Groups 1 and 2 will be discussed kit fox (Vul~esvelox) and plains in this report. wolf (Canis lupus). These carnivores probably made only minor In the following discussions note use of prairie wetlands. that all data on the sale of North and South Dakota pelts by hunters Uncounted numbers of wapiti and trappers are for in-state sales (Cervus ele~hus) and pronghorn only, resulting in a probable under- (Antiloca~raamericana) and smaller statement of their total worth. numbers of mule deer (Odocoileus hemionus), the only other large Pelts of the beaver have been herbivores of open grasslands, once sought after since the opening of inhabited the region and undoubtedly the region to the fur trade in the used the wetlands, at least for 18th century. Beaver occasionally drinking. These three species are occupy semipermanent wetlands in the still found in small numbers in the prairie Pothole Region of the region. Also nearly extirpated from Dakotas, especially basins in the the prairie region are the river northwestern portion of the region otter QLutra canadem), mountain where aspen (Populus tremuloides) is lion (Felis concolor), lynx (F. sometimes common in the wet-meadow lynx), and bobcat (F. rufus) . zone and adjacent uplands (A.B. Although once distributed throughout Sargeant, Northern Prairie Wildlife the region, it is unlikely that any Research Center, pers . corn,) , but of these species were strongly even there drought can cause mass associated with the wetlands dealt emigration or decimate populations. with in this report. Fritzell states that beaver are occasionally reported, but do not Modern Use of Prairie Wetlands establish permanent breeding popu- lations in semipermanent wetlands in A review of modern use of prairie the southern portion of the region. wetlands by mammals has recently been completed by Fritzell (1989). This species occupies primarily He states ehat no mammals other than riverine wetlands in the region. muskrats (Ondatra zibethicus) are Populations have increased in recent typically considered major elements years, but, compared to its former of prairie wetland ecosystems, but abundance, beaver nunLbers are still cites strong evidence to show that relatively low. Beaver often cause wetlands are vital to many prairie extensive damage to shade and mamals. We lists 17 species of ornamental trees where rivers flow terrestrial or semiaquatic mammals through residential areas; their whose geographic range encompasses dam-building activities can also most of the Prairie Pothole Region cause nuisances along watercourses, and which comonly use wetlands for and offending animals usually are cover or obtain a substantial por- quickly destroyed. tion of their food from wetland- dependent organisms. He also The beaver is a moderately i~por- mentions several other species that tant furbearer in the Dakotas. have been trapped in small numbers Dealer surveys indicate an average in prairie wetlands, He have added of 9,025 animals were sold annually Tabie 9. General currerrl use a8 basin wetlands in the Prairie Pothole Region eB the Dakotas by wild FnamFnaIS. Group and Definition cornon name Scientific name 1. Depends on wetlands with Beaver Casto4: canadensis surface water to complete Muskrat Ondatra Life cycle zibethicus Mink Bustela vison 2, Range encompasses 9kQ% of Arctic shrew Sorex qrcticus the region; nakes extensive Masked shrew Sorex cinereus seasonal use of wetlands or Pygmy shrew Microsorex hovi Likely can complete life- Eastern Svlvilams cycle in wet-meadow zones cottontail floridanus Snowshoe hare ~ewus am- Western harvest mouse mesalot& Meadow vole Mierotus gennsvlvanicu~ Meadow jumping Zawus budsonius mouse Western jumping mouse Coyote Canis latrans Red fox Vul~esvul~es Raccoon Procvoq lotox LQng-tailed Mustela frenata weasel Least weasel Mustela nivalis Striped skunk pe~hitisahitis White-tailed deer Qdocoileus virsinianus 3. Upland species whose Virginia opossum pidel~his range encompasses >lo% of virsiniana region; likely cannot Northern short- Blarina complete life-cycle in tailed shrew brevicauda wetlands, but occasionally Vespertilionid (Chiroptera: seen in or over them bats Ves~ertilianidae) White-tailed Lewus townsendii jackrabbit FranklinDsground squirrel Thirteen-lined ground squirrel Plains pocket gopher Deer mouse Northern grasshopper IllOuSe (Continued] Table 9. (Concluded). Group and Definition cornon name. Scientific name 3. (Continued) Southern red- backed vole Prairie vole House mouse Norway rat Gray fox Eastern spotted skunk 4. Makes extensive use of Water shrew Sorex ~alustris wetlands, but range Long-tailed vole Microtus encompasses ~10%of region lonqieaudus Southern bog Svnaptomvs cosweri lemming Ermine Mustela erminea Moose Alces alces in North Dakota during the 1983-84 unusual. Errington (1948) recorded to 1985-86 seasons, for an average maximum winter populations of 86 total of $106,766 (Steve Allen, muskrats/ha on a fertile Iowa marsh. North Dakota Game and Fish Dept., pers . comm .) . Buyers in South In the extreme southeastern por- Dakota paid a total of $57,669 for tion of the Dakota pothole region, 3,238 pelts during the 1986-87 sea- the importance of muskrat activity son (Larry Fredrickson, South Dakota on marsh ecolagy is great. The Game, Fish and Parks, pers. comm. ) . milder winter climate and more stable water levels there are simi- The muskrat is the only native lar to conditions in Iowa, where mama1 that still has a great impact high muskrat populations can deci- on wetland ecology in the region. raate nearly all emergent hydrophytes Semipermanent wetlands are the most and result in mass migration and important habitat for muskrat in the high mortality, as well as greatly region because these basins are affecting the use of the wetland by abundant, deep enough t~ sustain other animal species. Here, as under-ice activity in winter, and semipermanent wetlands cycle between suppart growth of emergent plants, extreme dryness and high water especially TYD~spp. and Sckr~us levels, muskrat activity greatly spp*, which provide both food and influences nearly all aspects of house-building materials. Muskrats marsh ecology (Weller and Spatcher also dig burrows in the banks of 1965; Weller and Fredrickson 1974; wetlands, van der Valk and Davis 1978a). Van der Valk and ~avis(1978a) recognize Consumption oE hydrophytes by high four stages in this cycle: dry muskrat populations often causes marsh, regenerating marsh, degen- great changes in the interspersion erating marsh, and lake anarsh of cover and open water in prairie (Appendix A, Figure hZ 1 , The food wetlands, FritzelP states that and cover requirements of expanding muskrat densities >50/ha are not muskrat populations is thought to be an important factor contributing or crepuscular species (A.B. to the decline in emergent hydro- Sargeant, Northern Prairie Wildlife phytes that marks the end of the Research Center, pers . corn,) . Data regenerating-massh phase. Xf high from Erringtsn (1943), however, water levels continue, muskrat suggest that when surface water is populations increase until emergent present about 50 ha of semipemanent plants are almost totally destroyed, weeland will support a female with This beads to the lake-marsh phase, young. Mink populations probably where submersed macrspbytes dominate fluctuate greatly in the Prairie and muskrats are forced to emigrate Pothole Region in response to or suffer high mortality (Errington drought (Adams 1962; Eberhardt 1974; et ale 1963). The lake-marsh stage Fkitzell 1989). will persist until water Bevels again decline. Dens are usually- located in The effects of muskrats on the abandoned muskrat burrows along ecology of semipemanent wetlands wetland shorelines (Schadweiler and become much less pronounced in North Storm 1969; Eberhardt 1973; Sargeant Dakota, where colder winter climates et al. 1973; Eberhardt and Sargeant prevail and wetlands often freeze to 1977). Females with young in dens the bottom, especially when fall can concentrate their activity in water levels are below average. relatively small (<20 ha) semiper- Under such conditions, foraging is manent wetlands (Sargeant et al. 197 3 ; Eberhardt and Sargeant 1977) curtailed and populations can be whereas the home ranges of males decimated (Seabloom and Beer 1963). average 646 ha from May through July Because of increased frequency of and include as many as 30 temporary, drought, muskrat populations in the more arid western portion of the seasonal, semipermanent, and region also do not usually build to permanent wetlands, at least in the levels high enough to greatly Prairie Pothole Region of Manitoba diminish emergent vegetation. (Arnold 1986) . Muskrats are an economically The concentration of nesting birds important furbearer in the region. in prairie wetlands and the relative Buyer surveys show an average of scarcity of fish, semiaquatic 64,181 pelts sold in North Dakota mammals, and large invertebrates, during the 1983-84 to 1985-86 such as crayfish, in these habitats trapping seasons, for an average result in levels of mink predation annual return of $183,331 to the that can cause significant mortality takers (Steve Allen, North Dakota of marsh birds. For example, Game and Fish Dept . , pers. corn. ) . Eberhardt and Sargeant (1977) found The harvest in South Dakota averaged that on a semipermanent North Dakota about 55,000 during the 1982-83 and wetland, a single mink fanrrily can 1983-84 seasons (Linscornbe and kill 8% of the adult and 52% of the Satterthwaite in press), but better juvenile coots and 20% of the adult water conditions during the 1986-87 and 6% of the juvenile pied-billed season resulted in sales of 248,126 grebes. They also Pound an average pelts, for which in-state buyers of 6.3 adult and 5-3 juvenile ducks paid a total of $617,834 (Larry taken per mink family, and noted Fredrickson, South Dakota Game, Fish that the preponderance of female and Parks, pers. corn. ) . diving ducks taken could account for the disparate proportion af males The mink is common throughout the commonly observed among this group Prairie Pothole Region and is of waterfowl, Amold and Fkitzell closely associated with basin (1987a, $1 estimated that avian prey wetlands. No reliable census comprised 55%-75% of the volme of techniqes have been developed for the Hay-July food of male mink in this solitaq, primarily nocturnal prairie wetlands in Manitoba. Economically , mink are a soma prairie wetieands is knndo~tedly moderately important furbearer in a prime attractant to mustelid and the Dakotas. An average of 4,571 canid carnivores. animals were sold to North Dakota buyers during the 1983-84 to 1985-86 The range of the snowshoe hare seasons, for an average return of encompasses roughly the northern $64,497 (Steve Allen, North Dakota half off Norm Dakota. This hare can Game and Fish Dept. , pers. corn. ) . be found fn wetland wet-meadow zones About 6,000-10,000 mink are dominated by willow or aspen estimated to be harvested annually . Jones eQ a%. in South Dakota (Linscombe and (1983) stated that alder (Ahus Satterthwaite in press) . During the spp. ) swamps and burned areas with 1986-87 season, 8,557 pelts were woody regrowth are especially sold to South Dakota buyers, for a favored. Snowshoe hares are herbi- return to takers of $203,828 (Larry vorous, but accasionally eat Fredrickson, South Dakota Game, Fish carrion. and Parks, pers. corn.). The western harvest mouse ranges The remaining mammals discussed in throughout %he southern two-thirds this report (Group 2 of Table 9) of the region. Jones et al. (1983) range across at least PO% of the list cattail as a typical habitat pothole region of the Dakotas, and and state that the species often either regularly make extensive use associates with meadow voles and of prairie wetlands, often during uses their runways. Seeds and small the winter months, or likely can amounts of insects are the main food complete their life-cycle in moist, of this small rodent, which is often shrubby, meadows. active year around. Up to 18% of the small mammals trapped in South Little is known about the life Dakota wetlands by Pendleton (1984) histories of the arctic, masked, and were western harvest mice, pyg~lyshrews in the grassland biome, except that marshes or moist Meadow voles are common herbivores habitats, often dominated by throughout the region, where they cattails or hydrophytic grasses, are inhabit surface runways and tussocks used by these tiny carnivores (Jones in stands of grasses, sedges, and et al. 1983) . Of the three, the rushes found in wet-meadow zones arctic shrew seems ta be the most (Jones et al. 1983). Cycles of wetland oriented, as aquatic abundance are very pronounced for invertebrates are a known food item. this species, and population However, a recent study in the densities as low as 2.5/ha and as Prairie Pothole Region of north- high as 617/ha have been reported central South Dakota indicated that (Jones et al. 1983). Up to 75% of pygmy shrews are more abundant than the small mammals trapped in North arctic: shrews in shallow-marsh zones Dakota wetlands by Eberhardt (1974) (GruebeEe and SLeuter 1988). were meadow voles. The eastern cottontail is This vale is prominent in the diet primarily a dweller in thickets and of the coyote, red fox, mink, long- brushlands, and the dense stands of tailed and least weasels, great vilPaw ( sgp.) that are horned owl Bubo virsinianus), sometimes found in wet-meadow zones shart-eared owl (Asio flameus) , and sf prairie wetlands provide northern harrier. The cyclical permanent habitat. This cottontail abundance of voles can greatly is mostly herbfvorsus, but in times affect the seasonal abundance of of food shortage may eat aullusks same of their avian predators. and even carcasses sf other cottontails. (Jones et ale 1983). The meadow jumping mouse occurs The presence sf cattontails around tktrourghout the pothole region of the Dakotas, but the western jumping hunts, travels through, and rests in mouse Is mostly restricted to the wetlands when they are dry or ice North Dakota portion (Jones et al. covered. Territories of red fox 1983). Both species are mostly families sympatric with coyotes in found in tall, lush herbaceous or soukh-central North Dakota averaged graminoid cover typical of low 11-9 km2 (Sargeant et al, 1987) . prairies or wet meadows in the northern plains. Adequate ground Red fox eat a huge variety of coves is essential for nests. Both foods, mostly animal matter. Foods species are active about 5 months of eaten depend on season and habitat. the year in this area. Material Common foods found in or at the cited in Fritzell (1989) shows that edges of prairie wetlands include up to 17% of the small mammals cottontails, meadow voles, harvest trapped in North Dakota wetlands mice, jumping mice, young raccoons, were meadow jumping mice. and birds and their eggs. The coyote is now the largest common carnivore in the Prairie Perhaps the most significant role Pothole Region of the Dakotas and of the red fox in the ecology of occurs throughout the area, but is prairie wetlands is as a predator of upland-nesting waterfowl and other most numerous in the western part. al. Territories for coyote families in semiaquatic birds. Sargeant et south-cezntrai North Dakota averaged (1984) estimated the average annual take of adult ducks (both killed and 61.2 km (Sargeant et al. 1987). Although primarily terrestrial, the scavenged) by the red fox in coyote hunts in peripheral wetland midcontinental North America at zones when surface water is present about 900,000. This impact was in interior zones, and uses dry and greatest in the high-density wetland ice-covered wetlands for hunting, area of the Missouri Coteau in North Dakota, where 5-year average of traveling, and resting. It also a feeds on prey associated with wet- 17.0% of the total prey biomass lands at various times of the year, consumed by foxes during the annual includins insects, birds and their denning season was dabbling ducks. eggs, c&ttontails, small rodents, muskrats, beaver, and deer (A. B. The red fox has recently been an Sargeant, Northern Prairie Wildlife economically important furbearer in Research Center, unpubl. data). the Prairie Pothole Region, although current (1987) pelt prices are Coyotes are an important furbearer relatively low. During a recent 10- in the region. During the 1983-84 year period (1970-L980), annual to 1985-86 seasons, North Dakota harvest in the pothole region sf buyers purchased an average of 7,913 North Dakota alone was fQ,OQO-62,000 pelts annually, for an average pelts that returned $53,000- annual combined return to takers of $2,389,000 ta takers (Sargeant et $255,458 (Steve Allen, North Dakota al. 3.984) . Buyer surveys conducted Game and Fish Dept . , pers. corn. ) . by Larry Fredrickson (South Dakota In 1986-87, South Dakota buyers Game, Fish and Parks Dept,, pers. purchased 8,149 pelts for a total of corn. ) indicate that $he 17,512 $349,674 to takers (Larry animals purchased there during 1986- Fredrickson, South Dakota Game, Fish 87 returned a total of $368,102 to and Parks Dept., pers. corn.). takers. Befare settlement of the Prairie The red fox is common in both Pothole Region by European man, the Dakotas, and, like the coyote, is omnivorous raecasn inhabited only primarily a terrestrial species that wooded river valleys in the pothole uses only the shorelines of wetlands region of the Dakotas, with greatest when surface water is present, but densities found in drainages in the eastern part of the region, How- total negative impace of this pre- ever, populations expanded greatly dation on waterfowl populakians after settlement, and raccoons can remains to be detemined, it will currently be found throughout the likely increase as the weeland region. Cowan (1973) and Fritzell habitat base continues ta decline (1978) found spring raccoon depsi- (Fritzell 1989). ties of 0.5-3.2 individuals/h in various parts of the region. These The raccoon has recently became an studies also showed that wetlands, important fur resource in the particularly the seasonal (Class 3) pothole region. During the 1983-84 and semipermanent (Class 4) basins to 1985-86 North Dakota trapping of Stewart and Kantrud (1971) were seasons, takers sold an average af commonly used habitat. P5,881 pelts annually in the state, for which buyers paid a total of Fritzell (1978) found that raccoon $202,006 (Steve Allen, North Dakota use of wetlands increased greatly Game and Fish Dept. pers. corn.). from spring to summer as foods Recent (1986-87) sales of 34,038 become available in these habitats. pelts to in-state buyers in South Greenwood (1882) found that 94% of Dakota resulted in a total return of the nocturnal raccoon activity in $715,138 to takers (Larry eastern North Dakota wetlands was Fredrickson, South Dakota Game, Fish related to foraging, and that the and Parks Dept., pers. comm.). most important food items gathered in these habitats were aquatic Few ecological data are available insects, birds, bird eggs, snails, for least and long-tailed weasels in and crustaceans. the Prairie Pothole Reqion. Both During the day, raccoons in the species range throughoutythe region. pothole region rest, and are seldom Weasels prey mostly on voles in this found in the same locations on area, and the wet-meadow zones of consecutive days from April to July. wetlands are important vole habitat. Use of wetlands for resting sites The long-tailed weasel is a known increases greatly during this predator of waterfowl eggs (Keith period, and by July, 70% of the 1961; Teer 19641, but the likely sites can be located there impact of this mammal on waterfowl (Schneider et al, 1971; Cowan 1973; is small. Fritzell 1978). Weasels are unimportant as a fur resource in both ~akotas, Reported Raccoons in the northern portions annual sales to buyers in both of the Prairie Pothole Region states in the mid-1980's was €100 probably do not use wetlands as animals, with prices paid to takers sites for winter dens, but wintering averaging <$0.90/animal (Steve animals have used dense cattail Allen, North Dakota Game and Fish qfteepeeswin South Dakota (Fritzell QeP , pers. comm. ; Larry 1989). Fredrickson, South Dakota Game, Fish The expansion of raccoon popula- and Parks D&&. , pers . cam.1 . tions in the prairie region has been assaciated with decreased waterfowl The Prairie Pothole Regisn lies nesting success aver the past 40 near the center of the range of the years (Kid et aX, 19722; IPuebbert striped skunk. This primarily and Kantrud 1974 ; Trauger and Stoudt nocturnal maxnmal is a aipitous 1974; Duebbert and Lokernoen 1976; predator often associated with Ssrgeant and Rrraald 1984) , Raccaatls wetlands. Densities in the prairie feed extensively on waterfowl eggs, segisn have varied from Q. 4 to 3. 1 especially those of the ovemater- animals/h2 (Scott and Salko 1939; nesting diving ducks (Cowan 1973; Uphagl 1967; Bjorge et al. 1981; Greenwood 15381, 1982'). Although the Greenwood et al, 1985), Literature reviewed by Fritzell functions of these wetlands are to indicates thak skunks are active in provide refuge and thermal cover, the Prairie Pothole Region from although some browse (Po~ulusspp. January to NoveMer. Eight of and Salix spp.) and forage (Sonchus twelve winter dens in a large spp. and Cirsium spp.) are also Manitoba marsh were in stands of obtained (Sparrowe and Springer Phrasmites australis (Mutch 1977) . 1970; Kucera 1976; Fritzell 1989) . Daytime rest areas can be under- In North Dakota, dry wetlands are ground or aboveground in dense also used as fawning sites grasses in uplands or aboveground in (Wamoning 1976). stands of emergent hydrophytes in portions of wetlands where surface Dense stands of Tmha spp., water is not present. Scirpus spp. , Phramites australxs, and Salix spp. seem to provide the Striped skunks in the Prairie most attractive cover for white- Pothole Region feed largely on tailed deer in wetlands in this area insects during the warmer months, (W .A. Kantrud pers. obs. ) . These although mice, frogs , birds, shrews, plants are mostly found in semiper- bird and turtle eggs, berries, manent wetlands. fruit, worms, and spiders are eaten White-tailed deer populations have (Jones et al. 1983) . Skunks are major predators of clutches of eggs expanded greatly in the Prairie of upland nesting dabbling ducks Pothole Region in the last 20 years, (Stoudt 1971; Duebbert and Kantrud likely because of a combination of better management and law enforce- 1974 ; Duebbert and Lokemoen 1980) , including many that are likely ment and the great increase in area encountered incidentally as skunks planted to row crops (corn, sorghum, search grasslands for insect prey and sunflower) that provide cover and high-energy winter foods. (A.B. Sarseant. Northern Prairie wild1 ife Research Center, pers. The white-tailed deer is an comm.) . Skunks can also destroy important game species in both overwater nests of diving ducks if states. Latest available estimates drought conditions increase nest place the 1986 harvest in the accessibility (Stoudt 1982). pothole region of North Dakota at 42,085 (Jim McKenzie, North Dakota The striped skunk is unimportant Game and Fish Dept ., pers . conun. ) . economically as a furbearer in both An estimated 30,424 animals were Dakotas. North Dakota dealers harvested in 1986 from the area of reported a total purchase of <200 South Dakota east of the Missouri animals/yr during the 1983-84 to River (Les Rice, South Dakota Game, 1985-86 seasons. Takers received an Fish and Parks, pers. corn,). average of only $2.4Q each for these Estimates of annual cash flaw pelts (Steve Allen, North Dakota associated with white-tailed dees Game and Fish Dept., pers. comm.). hunting in the United States Total purchase by South Dakota indicate that a single animal is dealers was 1,878 animals during the worth $1,657 (Williamson and Doster 1986-87 season; these pelts returned 1981). Although this figure 5s an average of $2.22 each to takers undoubtedly high for the Dakotas, it (Larry Fredrickson, South Dakota suggests that the income generated Game, Fish and Parks, pers. comm. ) . to state and local economies by sport hunting sf this species is Few quantitative data are avail- significant. able, but anyone who has observed or hunted white-tailed dees in the Prairie Pothole Region is aware of the importance of basin wetlands to Fritzell (2989) s this species, especially during the role of wetlands in the life cycle fall and winter. The primary sf mamals in the Prairie Pothole Region. He concluded that these the importance sf wetlands in habitats are of direct importance to predator-prey relationships that can. many species, and that some mammals signifisantly affect the production markedly affect other components of of waterfowl and other wetland wetland ecosystems and the values wildlife. He also noted that the humans extract from them. We cited harvest of furbearers and white- in particular the effects of the tailed deer in prairie wetlands is muskrat on nutrient exchange and an important contribution to local vegetative structure in wetlands and economies. CHAPTER 4. ECOLOGICAL PROCESSES 4.1 PHYSICAL FUNCTIONS Although the number of runoff- retaining basins in the Prairie The two main physical functions of Pothole Region of the Dakotas was prairie wetlands are their hydro- large and much drainage has been logic and water quality effects. completed, controversy as to the Although strongly interrelated, proven effects of wetland drainage these functions will be discussed as a cause of increased flooding separately in this section. Descrip- continues. Several completed tive material on hydrology and water studies have been criticized, quality is found in Sections 2.3 and largely because of problems of study 2.4, respectively. design and impacts of conditions antecedent to the studies (winter Hvdrolosic Functions 1988). Even so, the results of some of these studies will be reported The hydrology of prairie wetlands here. is known only superficially, even though it has been obvious for Ludden et al. (1983) in the Devils decades that this knowledge is Lake basin of North Dakota concluded crucial if the conflicts between that depressions can store as much agricultural water users and those as 72% of total runoff in a 2-year- concerned with flood protection, frequency runoff event and 41% of water quality, ground-water total runoff from a 100-year- recharge, and fish and wildlife frequency runoff event, with a production are to be resolved maximum storage capacity of nearly (Winter 1988)- The basin wetlands 8 x 10"' available. In an that are the subject of this undrained control block in North community profile form the core of Dakota" J. Clark Salyer National these conflicting interests. Wildlife Refuge, all local runoff plus 58% of the inflow was retained The value of prairie basin in wetlands (Malcolm 1979). wetlands as retainers of surface water flows has recently been Increased streamflows in the reviewed by Winter (1989). Me southern Red River Valley in North states that, before drainage, the Dakota were strongly correlated to numerous small depressions that the increase in area artificially characterize the topography made drained in each drainage basin (Brmm much of the region non-contributary et ale L98P), and wetlands in the to surface runoff until depressions Red River system significantly were full of water and began to reduced flood levels downstream in overflow from one to the next. In major metropol itan areas (Jaha addition, when overflow did occur, 2.981)* the extremely low regional gradient of the land surface generally made 2% little-understood function of runoff velocities small, prairie we"slands is their rsfe in regional ground-water recharge and Most prairie wetlands thae have discharge (Winter 1989). Consider- escaped drainage now lie in water- ably more work needs to be done on sheds devoted primarily to agricul- this subject, especially on water- tural crop production. Hence, these flow rates through strata of various wetlands frequently experience large glacial deposits. Of eight hydro- inputs of sediment, fertilizer, and logical studies of palustrine and other agricultural chemicals trans- lacustrine wetland systems conducted ported in sediment, surface runoff in the glaciated prairies of North water, and subsurface drainage. America, seven showed that these These contaminants are often removed wetlands contributedto ground-water from the water in passing through recharge (Adamus and Stockwell wetland basins, Prairie wetlands 1983). The eighth study showed that can thus be important in preser- wetlands with permanently flooded vation of local water quality. water regimes contributed ground- water discharge year-round. Prelim- Contaminants transported mainly inary studies indicate that basin with sediment include chlorinated wetlands in North Dakota supply hydrocarboninsecticides;herbicides water to discharge sites up to 20 km such as trifluralin, profluralin, distant (Swanson et al. 1988). and paraquat; and total organic M and total P including phosphate. Water Quality Functions Ammonium nitrogen and a majority of the pesticides used in the mid- It was shown in Section 2.4 that western United States are carried to prairie wetlands naturally vary wetlands mostly with surface water greatly in the chemical composition runoff. Nitrate nitrogen is the of their surface waters. This is most important chemical carried by only one aspect of "quality. The subsurface drainage, and can be quality of water depends on what the transported mostly by surface water is to be used for. The human drainage in relatively impervious or users of water found in prairie wet soils. Grue et al. (1988) wetlands include cities, households, demonstratedthat aerial application ranches, and crop irrigators, who of methyl and ethyl parathion caused usually desire water low in dis- mortality on aquatic invertebrates salved minerals or organic com- for up to 18 days after spraying. pounds, as well as commercial chemical companies, who extract Only the role of prairie wetlands usable minerals most effectively as sinks or traps for N and P has from some of the most saline surface been studied to any great extent, waters found in the region. Other and most of the information comes users desire water of low tempera- from studies of a single Iowa wet- ture or low in suspended material. land (Davis and van der Valk 1978; For purposes of this section, water Davis et al. 1981; Neely 1982). of highest quality is considered These nutrients can be transferred that unchanged from its original from the water column of wetlands to cheaical cornpasition under pristine the atmosphere, to the interstitial conditions, water of sediments, or to biomass within the wetland. Each of these Of the 11 biological and nonbio- compartments can also return logical functions of: wetlands listed nutrients to the water column. by Adamus and Stockwell (19831, sed- iment trapping and nutrient removal The Iowa wetland mentioned above have the greatest positive impact on removed 86% of the NOIfnitrogen, 78% water quality, These two functions of the NH: nitrogen, and 20% of the have recently been reviewed for PO:+ phosphorus inputs from prairie wetlands by Neely and Baker agricultural runoff in a single (1989), and this section will draw year, but the wetland showed such heavily from their findings, high efficiency as a nutrient trap only when no outflow occurred, of herbivores and detritivores. Nevertheless, sequential processes Invertebrates, muskrats, and water- of nitrificatisn and denitrif lcation fowl are important herbivores in can result in large N losses to the prairie wetlands. A large group of atmosphere, to surface outflow, and invertebrates are known to ingest to the ground-water system in wet- living phytoplankton and attached lands experiencing frequent drying algae, but little is known of the and inundation. Thus it would seem role of algae in the support of that temporarily and seasonally these primary csnsmers in prairie flooded prairie wetlands would be wetlands. Some of the most impor- especially efficient in removal of tant of these herbivores in prairie excess N that entered their basins. wetlands include macro- and micro- crustaceans, dipteran and ephemero- Znvestigators of the Iowa wetland pteran insects, and freshwater referred to above also reported high mollusks. Domestic animals also N and P concentrations in tissues of consume much of the primary produc- the two most common emergent plants, tion from prairie wetlands, but are Ty~haglauca and Sparganium eurv- unimportant in many other of the carpum. If it is assumed that other world" wetland ecosystems. At emergents also accumulate these least one emergent hydrophyte, nutrients, then the removal of hay Scolochloa festucacea, is recognized and forage from prairie wetlands by as a highly valuable livestock humans and their livestock would be forage grass and is actively managed a means by which their function as for and harvested in prairie nutrient traps could be maintained wetlands (Neckles et al. 1985). for longer periods. Secondary consumers are extremely Many prairie wetlands likely re- important and provide foods for most tain sediment, nutrients, and other of the higher organisms found in anthropogenic substances simply prairie wetlands. Nurkin stated because the basins have no outlet. that knowledge of the basic ecolsgy No studies have addressed the and life-history of consumers in ability of these wetlands to with- prairie wetlands is so limited as to stand sustained inputs of these make work on secondary production contaminants. In addition, no study impossible at this time. has addressed the question of the fate of nutrients and pesticides A third, nontraphic user of pri- that enter highly saline prairie mary production is the large and wetlands. diverse group of organisms that depend on living and dead plants for thermal cover, nesting material, 4.2 BlOLC?GICPeL FUNCTIONS sites for egg attae-ent, et@. These organisms will not be dis- Primary production, which finks cussed here, but some infomation on inorganic resources (energy and nontraphic use is presented in the nutrients) to food-chain support and preceding chapters on various animal associated secondary production, is groups. certainly the most important bio- logical functi~nof all ecosystems, Primary Production and is the foundation for most human use of these ecosystems. The basis Het primary production, bstb above for faad-chain support in prairie and below greund level, is particu- wetlands has recently been reviewed larly high in prairie wetlands (van by Murkin (2989), and this section der Valk and Davis 1978b) , but shaws will draw heavily on hxs wark. great temporal variation, mainly because of the presence or absence The heterotrophic primary can- of surface water; water levels are sumers in prairie wetlands consist also important in this regard, In semipermanent wetlands, highest comon species in prairie wetlands primary productivity likely occurs are shown in Table 10. Bn many as emergent hydrophytes regenerate instances, algae do not conkribute when surface water is replenished greatly to standing-crop biomass, after prolonged drought, and lowest and thus are not usually considered productivity probably occurs after an important part of primary prsduc- several years of high water levels tion. However, recent iwvestiga- when emergent hydrophytes are tions by Hooper and Robinson (1976) replaced by submersed species (van and Shamess et al. (1985), and der Valk and Davis E978a). In unpublished data of van der Valk temporary and seasonal wetland (Murkin 1989), show that algal basins, productivity likely is production and biomass in prairie highest when water is present, as wetlands can be quite high (Table judged by increases in height, 10). Algal groups especially impor- biomass, and density of emergent tant as primary producers in prairie macrophytes (Neckles 1984; H.A. wetlands are epiphytes, which grow Kantrud, pers- obs.). on surfaces of submersed plant material; epipelics, which grow on Examples of productioc rates and bottom substrates; and metaphytes, above- and below-ground biomass for which usually form floating mats. various vegetation zones and some Prairie wetlands can produce high Table 10. Examples of biomass and production rates of some primary producers in prairie wetlands (modified from Murkin 1989). Primary producer Biomass or Reference production rate (g/m2 dry wt. ) or (g~/m2/yr) Vascular hydrophytes Above ground biomass Wet-meadow zones Barker and Fulton Shallow-marsh zones (1979) Deep-marsh zones emergent submergent Typha glauca van der Valk and Scimus validus Davis 1978b Scolochloa festucacea (dry zone) Neckles 1984 8t t; (wet zone) I1 Potamoaeton ~ectinatus Anderson 1978 Below ground biomass Tmha glauca 1442 van der Valk and Scir~usacutus 1870 Davis 197813 Phraamites australis 1565 1$ Algae Metaphytic (biomass) 268 van der Vahk (unpubl. ) Epiphytic (production rate) 48,5 Hooper and Robinson 1976 EpipePic (production rate) 300 Robinson pers, corn. cited in Plurkin (1989) midsummer populations of planktonic and leaves of emergents, submersed algae of sufficient densiky to cause macrophytes, and algae. Some living Eish kills (Kling 1975). emergent material can enter this compartment directly through muskrat Virtually nothing is known about activities (van der Valk and Davis the mast primitive producer group in 1978b) or those of man and his prairie wetlands, the ehemosymt'tretie livestock. The third component of bacteria. the detritus pool is the dissolved organic substances that beach into the water column from both standing and fallen litter. There is no doubt that huge herds of wild ungulates once removed De Pa Cruz (1979) recognized three considerable amounts of primary simultaneous processes whereby production from prairie wetlands. detritus is decomposed: (I) rapid This phenomenon has been continued leaching of soluble substances froln on many wetlands through mants newly dead plant tissue, (2) livestock and hay-cutting opera- weathering and mechanical fragmen- tions. Birds (primarily waterfowl) tation, and (3) biological decay are also known to consume up to 86.8 from bacteria and fungi. Particle gm/m2 vegetative biomass annually sizes rapidly decrease, and the dead from portions of certain prairie tissue and soluble organic sub- wetlands (Anderson and Low 1976). stances become food for a host of The emergence of aquatic flying invertebrates and other life forms insects may also remove significant whose excretory products can again primary production, although not be colonized by microbes. enough is known about the foods of larval insects to quantify this Detritivory relationship. However, as many as 19,713 adult chironomids/mz have There is no doubt that detritivory been trapped in a single month as is an important aspect of food chain they emerged from a semipermanent support in prairie wetlands, even North Dakota wetland (Nelson and though there are no estimates of Butler 1987). Mevertheless, the detrital consumption and utilization generally high productivity of most for any wetlands. Coarse particu- prairie wetlands ensures that a late matter likely enters prairie considerable amount of material wetlands as a pulse associated with enters the system as detritus. increased spring water temperatures. Bacteria, fungi, and other micro- An important part of the detritus organisms form the first level of pool in prairie wetlands is standing consumption of dead plant material dead emergent hydrophytes. Much of and are considered the base of the this litter enters the pool with the detrital food chain, with secondary onset of killing frosts at the end production assumed to occur at of the growing season, but in less higher trophic levels. Many of the permanent wetlands, many emergent invertebrate herbivores common to hydrophytes die from drought during prairie wetlands are also suspected late summer and early fall. Leaf to be detritivorres , but some arathars and shoot death of various emergent argue that such organisms Bay simply species can occur at any time during be assimilating micrsorganisms the growing season (van der Valk and associated with litter of small Davis 1978b). particle size, Thus Murkin (1989) concluded that the primary role of The fallen litter compartment is detritus in prairie wetlands may be usually considered to consist of ta provide habitak necessary for material in the water column, algae and invertebrate production in including fallen or bent-over stems these syste~ns, CHAPTER 5. HUMAN USES AND IIMPACTS 5.3 ECONOMIC FUNCTIONS Food Security Act of 1985 (Fam Bill) (W.R. 2100) by making agri- The natural wetlands of North and cultural producers ineligible for South Dakota supply important certain Federal assistance if they economic and recreational benefits alter or drain wetlands existing to the nation as a result of their after 23 December 1985, and produce prominence in the production af agricultural commodities on these waterfowl and other migratory birds. converted lands. Among the most important local, state, and regional benefits are No economic benefits derived from providing water and forage to prairie wetlands are more widely livestock, offering extensive disseminated than those resulting recreational opportunities through from waterfowl production. The hunting and fishing, flood control, North American waterfowl resource, ground-water recharge, and improving particularly the forms produced in water quality. The importance oE prairie wetlands, contribute signi- wetland habitat for wildlife, f icant economic benefits to most recreation, aesthetic values, and states and numerous local cornmuni- water quality is recognized by 9 out ties. At the national level, of 10 North Dakotans, based upon a waterfowl, including those produced recent poll (Fargo Forum, 31 in prairie wetlands, support December 1986; North Dakota Bureau recreation for about 2 million of Govermental Affairs, unpubl. waterfowl hunters (Novara et al. data). The same poll indicated 77% 1987). An estimated $638 million of the State's residents want was spent on migratory bird hunting tougher enforcement of laws con- in the United States in 1980 and an trolling wetland drainage, Yet estimated $6.6 billion is spent wetland protection is a politically annually for recreation related to contentious issue because grain nongarne wildlife (U.S. Department of growers, a dominant sector in the the Interior 1982). A significant economy of most comunities, cannot part of both expenditures results capture paylnent for many of the from migratory birds reared in values society derives from wet- prairie wetlands. lands. As a result, free-market incentives ta drain are often In North and South Dakota, water- stronger than those to maintain fowl hunting is a major recreational wetland habitat (=itch 1983) and activity; the former state ranks efforts to protect wetland habitat first in the nation in number of are often resisted. Drainage has waterfowl hunters per capita. Also, also been encouraged because Federal several thousand nonresidents travel price suppoes on agricultural to each of these states to hunt comodities are paid &-ally whether waterfowl each year and contribute crops are grown in the uplands or in several mill ion dollars to local drained wetland basins. The U,S. economies. In South Dakota, recrea- Congress addressed this issue in the tian associated with hunting the 70 ring-necked pheasant brings $35 easements protect c~ntracted wet- million annually &o the State (U.S. lands in from draining, Department of the Interior, Fish and burning, or filling by the landowner Wildlife Service 1987), and is a or operator, but the land can be significant source of income to many tilled, grazed, or hayed when rural communities. Pheasants rely natural conditions permit. on wetlands for a major part of their winter habitat requirements in State conservation agencies in the Prairie Pothole Region. North and South Dakota own approxi- Trapping and hunting of resident mately 9,300 and 21,870 ha of furbearers provides a supplemental natural wetlands, respectively income source to several thousand (Krapu and Duebbert 1989). A residents of the Dakotas, Highly significant but uninventoried area valued furbearers including the of wetlands is contained in the mink, muskrat, and raccoon occur unpatented State-owned lands lying primarily in association with below the meander line of natural prairie wetlands (See Section 3.7) . lakes and on State-owned school lands (Morgan 1971) . Meandered wetlands are mostly permanent or 5.2 WETLAND OWNERSHIP intermittent alkali lakes. An estimated 64,655 ha of meandered Wetlands in the Dakotas are pri- wetlands exist in South Dakota marily in private ownership, with (Wittmier 1982); similar statistics significant holdings by Federal and are not available for North Dakota. State agencies. The primary public Lands granted to the State for the landholders managing lands for wild- support of public schools include a life production are the U.S. Fish wide array of wetland types, includ- and Wildlife Service (FWS); the ing highly productive waterfowl South Dakota Department of Game, habitat, and are located primarily Fish and Parks; and the North Dakota within native grassland pastures Game and Fish Department. Private leased for grazing. A total of trusts and local and national con- 152,720 and 62,147 ha of State servation organizations own and school lands of all habitat types manage some wetland tracts for remain in public ownership in the wildlife production in both states. Prairie Pothole Region of North and Of an estimated 810,000 ha and South Dakota, respectively (State of 526,500 ha of natural wetland North Dakota, Commissioner of habitat existing in North and South University and School Lands 1987; Dakota, respectively, less than 10% South Dakota Office of School and is owned in fee title by public and Public Lands 1987). State school private wildlife interests. The lands are not protected from even- principal public agency owning tual sale and over three-quarters wetlands, FWS, has fee title to have been sold in North and South about 36,000 and 17,000 ha of Dakota. The current policy in North. wetlands in waterfowl production Dakota is ta maintain remiwing areas in the Prairie Pothole Region school lands in state ownership, of North and South Dakota, respee- whereas South Dakota is continuing tively (Krapu and Duebbert 1989). to sell its school lands (South There are also several thousand Dakota Office of School and Public hectares of wetland habitat on Lands 1987). A small amomt of National Wildlife Refuges ( wetland habitat is lo~atedon lands these states. In addition, as of administered by sther State September 30, 1988, perpetual ease- agencies, e.g. , in state parks and ments have been taken by FWS on state forests, Okher Federal 313,352 and 158,057 ha of privately agencies owning limited natural owned wetlands in North and South wetland habitat in the Prairie Dakota, respectively (U.S. Depart- Pothole Region af the Dakotas ment of Interior unpubl. ) . These include the Forest Service, Bureau of Reclamation, Amy Corps of Dakoka. Wittnnier (59882) estimated Engineers, Bureau of .Land Hanage- that 34,505 ha 0% temporary, ssa- ment, Bureau of Indian Affairs, and sonal , and semipemamrerat basins have Department of Defense. been drained in South Dakota since 2964. The latest private drainage sumey data for North Dakota (Memo- 5.3 WETLAND DEGRADATION AND DRAINAGE randum to the Area Hanager, Fish and Wildlife Sewice, Bismarck, Nastgh Dakota, 29 Hay 1980) stated that Artificial drainage and degra- 10,6% of the area of all privately dation of wetland habitat represent owned natural basin wetlands in the major threats to breeding popula- Prairie Pothole Region of the state tions of waterfowl and other marsh were drained from 1966 to 1980. The birds in the Prairie Pothole Region. survey did not include temporary It is estimated that approximately basins, so the loss rate is a 60% and 40% of the original wetland conservative estimate. area has been drained in North Dakota and South Dakota, respec- Wetland drainage has been most tively (Tiner 1984). Nearly all extensive in the Central Lowlands agricultural wetland drainage in the portion of the Dakotas (Figure 1) Dakotas has been by surface ditches, (Nelson et al. 1984). Drainage although underground tiles have been rates vary widely among lacalities, used to a small extent, mostly in in part depending on the proportion the southeastern part of the region. of the wetland base that is pro- Agricultural wetland drainage is tected by perpetual easements often closely linked to highway acquired by the U.S. Fish and construction projects that provide Wildlife Service under the Small the conduit for waters drained from Wetlands Acquisition Program. fields (U.S. Fish and Wildl. Sen, 1975). Some wetland basins no The primary cause of wetland longer hold water because of the degradation in the Prairie Pothole effects of drainage on shallow Region, includingthe part occurring aquifers that recharge them, while in the Dakotas, is agriculture the waters of a few basins are (Tiner 1984)- Intentional filling pumped or siphoned off during the of basins with soil from adjacent spring. Winter (1989) warned that uplands occurs, but is not common. the problems associated with More insidious is the gradual silta- modification of local hydrologic tion of basins caused by soil systems are self-perpetuating and erosion from adjacent cropland. that multiple modifications can This cause of wetland degradation ultimately lead to regional impacts. has become more severe in recent years with the advent' of larger, Systematic sampling of wetland more powerful farm, equipment that drainage in South Dakota by the U. S. allows cultivation of entire basins Fish and Wildlife Service for the and their steeply rolling water- period 2974-1980 indicated that 4.4% sheds. Increased siltation to of the area of seasonal and semi- basins can also be attributed partly pemanent wetland basins unprotected to expanded planting of row crops, at the end of 1974 in the survey particularly corn, sunflower, and areas were destroyed during the soybean. Row crops generally result periad (Memorandum to the Area in more soil erosion than small Wanager, Fish and Wildlife Service, grains, because of the additional Pierre, South Dakota, 17 December cultivation required during the 1980). Northeastern portions of the growing season. state had sralativePy low drainage rates fh,55 of area of wetlands Agricultural chemicals exert a destroyed), whereas rates were as subtle but pervasive influence on high as 7.5% in southeastern south prairie wetlands. Agricultural chemicals are widely used En temporaq and seasonal wetland association with growing intensi- component in many localities, fication of land use in the Dakotas. thereby reducing productivity and Most cropland in the two states is making intensive management attrac- regularly treated with herbicides, tive as a means of enhancing water- but insecticide use is restricted fowl habitat, The potential for primarily to sunflowers, which are intensive management of water levels now a widely grown crop; 65% of the is limited by land topography to a sunflower hectarage in North Dakota relatively snall part of the wetland was treated with insecticides in base; e, g., a few State-owned water- 3.984 (Grue et al, 1888). Widespread fowl management areas and National aerial application of agricultural Wildlife Refuges. Intensive manage- ehemieals in the Dakotas exposes ment of marsh habitat in the Dakotas wetland animal life and plants lo is increasing, however, due pri- overspray and drift. Although more marily to habitat management efforts research is needed, preliminary by Ducks Unlimited on publicly owned studies in North Dakota indicate wetlands. that the ~otentialfor aaricultural chemicals-to enter prairie wetlands The need for wetland management in and influence survival and repro- the Prairie Pothole Region is not duction of wildlife is high, restricted to manipulation of water particularly for the most toxic and levels. Elimination of the natural widely used insecticides (Grue et biotic forces of fire and ungulate al, 1986) . The impact sf herbicides grazing on many nontilled wetlands on prairie wildlife is indirect and decreases insolation of surface comes primarily from elimination of waters and rates of nutrient turn- food and cover (Hudson et al. 1984; over, thereby decreasing produc- Hill and Camardese 1986). tivity of life forms adapted to If degradation and destruction of shallow, open water (see ~eccion2)" waterfowl breeding habitat continues Vegetation of shallow waters, if not at present rates into the next removed by herbivores or otherwise century, stocks of ducks can be cropped, often develops into rank expected to decline accordingly. and dense stands that discourage use Therefore, a wide array of measures by many aquatic species, including will be needed to partially cir- waterfowl. Some form of vegetation cumvent these losses, including manipulation on shallow wetlands Is increased production of waterfowl on desirable as part of the management publicly owned lands, restoration of of lands to maintain high cover lands that formerly were prime quality for waterfowl production. breeding habitat, - and slbwing further habitat loss. Wetland restoration has high potential for increasing produc- tivity of wetland complexes in Noeh and South Dakota that have lost 5.4 WETLAND MANAGEMENT AND RESTORATION critical wetland habitat through drainage. This management tool is Prairie wetlands are underlain by being used with increasing fre- fertile soils, and under natural quency, particularly on lands conditions, a high proportion retired from crop producti~n under undergo annual drawdowns and are 10-year contracts through the inherently productive of the foods Conservation Reserve Program (CRP), that support many species of migra- on lands deeded back to or on those tory birds, including waterfowl. lands where loans are aainistered kIowever, extensive drainage and by tke Farmers Home Administration degradation of wetlands in the 1, and to replace wetland Dakotas has reduced or eliminated a habitat lost during csnstkarction of major part sf the highly productive public works proj ects , With an estimated 1.5 million ha of drained prairie wetland consemation in the wetlands in the Dakotas, widespread Dakotas include the Emergency opportunities exist to restore Wetland Resources Act of 1986 (P.L. wetlands within the statesg cropplnd 99-645), which assures continued base. In conjunction with the CRP, funding of wetlands presewatkon FWS offers participating landowners through migratsry waterEowl hunting an additional payment in return for stamp sales and other means, and the the right to restore wetland habitat Swampbuster provision of the 1985 in the tract. In North Dakota, the Farm Bill (H.R. 2100) which sets reformulated wildlife mitigation substantial monetary penalties for plan for the Garrison Diversion Unit agricultural producers who drain includes planned restoration of wetlands and plant to annual crops 4,499 ha of former wetland as after December 23, 1985. The latter replacements for wetlands lost action, a marked departure fron past during project construction (U.S. Federal agricultural policies, will, Department of Interior 1987). The if strictly enforced, add a signi- scale of future wetland restoration ficant new tool for reducing drain- in North and South Dakota will age of prairie wetlands. This law depend primarily on the direction is potentially effective in the taken by Federal agricultural Dakotas because most agricultural programs. With sufficient incen- producers there rely upon Government tives to agricultural producers, price supports and other Federal large-scale wetland restoration subsidies for a significant part of programs are feasible with major their annual farm income (Heimlich potential benefits to North American and Langner 1986) . waterfowl and other wildlife populations. The greatest challenge to main- taining a productive wetland base in the Dakotas is presented in the 5.5 PROSPECTS FOR THE f UTURE Central Lowlands. There, in many locations, an extensive wetland base The importance of prairie potholes remains, in part because of per- to North American migratory bird petual easements obtained by FWS, populations is widely recognized but much habitat continues to be (U.S, Department of the Interior and degraded and/or drained by large Environment Canada 1986). Recogni- cereal-grain farming operations. tion of prairie wetland values and The Swampbuster provision may help the forces contributing to their to slaw wetland drainage, but degradation and destruction has additional measures are needed to resulted in several remedial actions more fully protect watersheds and being taken in recent years. Among provide nesting cover for waterfowl the most important has been the and other wildlife. Expansion of incorporation of stringent mitiga- Federal land retirement programs tion measures into Federal water such as Waterbank and the CRP are projects in the Dakotas to replace needed to reduce pressures to drain wetland habitat lost during project or degrade wetland habitats. There construction (Krapu and Duebbert is also a substantial land base 19854) . Replacement of wetland currently in private ownership in habitat lost through restoration on the Dakotas that offers limited a 121 basis with habitat of an ec~nomic returns to private ecologically equivalent type, as has Landowners, but makes or could make been done in the reformulated a major contribution to national and Garrison Diversion Unit, reflects internatisnal migratory bird habitat the growing concern of the Federal requirements, The gradual transfer Governxent cowarc2 prairie wetland of habitat management rights an a loss and the need far replacement substantial part of these lands when destruction occurs. Other through purchase in fee title andior recent measures of importance to the taking af consenration easements is essential for the long-term well- The plan is continental in scope, being of waterfowl and other with major elements proposed for the migratory birds that are dependent Prairie Pothole Region. One goal is on prairie wetlands during part sf to protect and improve 445,000 ha of their life cycles, additional waterfowl habitat in the midcontinent region. A substantial W coaperative agreement between part of that habitat could come from the United States and Canada, the the Dakotas through perpetual North merican Waterfowl Management easements on wetlands and grasslands Plan (U.S. Department of the Inter- and through acquisition in fee ior and Enviroment Canada 3986), title. Other important elements represents the foremost effort inelude financial incentives to currently undertaay to rebuild North landowners to manage lands to derican waterfowl populations. 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U.S. Fish Wildl. Serv. factors affecting waterfowl Tech. Rep. 28. 44 pp. preductian in the Saskatchewan parklands. U.S. Fish Wildl, Sen. Talent, L.G., G.L. Krapu, and R.E, Resour, Publ. 99. 49 pp, Jamis. 1982. Habitat use by mallard broods in south central stoudt, 3,H* 19.82, Habitat use and Morth Dakota. J. Wildl, Manage. productivity sf canvasback in 46: 629-635. southwestern Manitoba. U.S. Fish Wildl. Serar. Spec. Sci, Rep. Teer, J.G, 1964. Predation by lang- Wildh. 248, tailed weasels on eggs of blue- winged teal. J. WiLdl. Manaqe.- 28:404-486, Swanson, G.A* 1984, Dissemination sf amphipods by waterfowl. J. Wildl. Tiner, R.W. Jr, 1984. Wetlands of Planage. 48~988-991. the Urrited States: current status and recent treg'~ds,&is S . Goverment Restoring Uerica" Wildlife, Printing office, washington, BC. 1937-1987. U.S, Government 59 PP, Printing Office, Washington, DC. 20402, 394 pp. Trauger, D,L., and %.H. Stoudt. 1974. Lsoking out for the canvas- U.S. Department of the Interior, back, part XI, ~ucksUnlimited Fish and Wildlife Service/ 38: 30-31, 42, 44-45,48# 60s Enviroment Canada, eanadian Wildlife Service. 1986, North Ungar, I,A. 1966, Salt tolerance of American Waterfowl plianagement plants growing in saline areas of Plan. Washington, N/Ot%awa. 34 Kansas and Oklahoma. Ecolog PP 4 47:154-155. U.S. Fish and Wildlife Service. 1975. Task force report on effects Ungar, I .A. 1974. Inkand halophytes of road constnaction on wetland of the United States. Pages wildlife habitat. Twin Cities, 235-305 b R.J. ~eimoldand W.M. BIN, 14 pp. Queen, eds . Ecology sf halophytes. Academic Press, New York. U. S. Soil Conservation Service, Soil Survey Staff. 1975. Soil taxdnomy: Igar, I.A., D.K, Brenner, and D.C. a basic system of soil classifi- McGraw. 1979. The distribution cation for making and interpreting and growth of Salicornia eurwaea pan. soil surveys. U.S. Sail Cons. on an inland salt Ecology Serv. Agric. Handb. 436. 754 pp. 60: 329-336. U.S. Soil Conservation Service. Upham, L.L. 1967. Density, dis- 1985. Xydric soils of the United persal, and dispersion of the States 1985. 198 pp. striped skunk (Mewhitis me~hitis) M.S. in southeastern North Dakota. van der Valk, A.G., ed. 1989. Thesis. North Dakota State Northern prairie wetlands. Iowa Universi-ty, Fargo. 63 pp. State University Press, Ames. 400 U.S. Department of the Interior. 1982. The 1980 national survey of van der Valk, A.G., and C.B. Davis. fishing, hunting and wildlife- 1976. The seed banks of prairie associated recreation. Fish and glacial marshes. Can. 3. Bot. Wildlife Service, Washington, DC. 54:1832-1838, 156 pp. van der Valk, A.G., and C.B. Davis. 1978a. The role of seed banks in U.S. Department of the Interior. the vegetation dynamics of prairie 1987- Fish and wildlife resources glacial marshes. Ecology in relation to the Garrison 59:322-335. Diversion Reformulation Act. Fish and Wildlife Service, North Dakota van der Valk, A.G., and C.B, Davis, Field Office, Bismarck, ND. 30 pp. 1978b. Primary production of prairie glacial marshes, Pages 21-37 W.E. Good and D,F, U.S. Department of the Interior. Wigham, eds. Freshwater wetlands: [ 1988. ] wetlands status report, ecological processes and manage- FY88. Div. of Refuges and ment potential, Asadentic Press, Wildlife, Region 6, 6 PP. New 'Itork. krnpubl, MS. Visher, S.S, 5912. The biohsw sf U.S. Department of the Interior, south-central South Dakota, S.DI Fish and Wildlife Service, 1987. Geol. Biol, Sum, Bull* 5:61-930, Visher, S.S. 1966. Climatic atlas of Weller, M.W., and C.E, Spateher, the United States. Harvard Univer- 1965. Role of habitat in the sity Press, Cambridge, MA. 403 pp. dis.kributioxl and abundance of znarsh birds. Iowa State Univ. Vogl, R.J. 1973. Effects of fire on Agrie. Home Econ. Exp. Sta. Spec. the plants and animals of a Rep* 43. 31 pg. Florida wetland. Am. Midl. Nat. 89~334-347. Wheeler, G,C, and J. Weelere 5966. The amphibians and reptiles ~f Walker, B.H., and R.T, Coupland. North Dakota. University sf North 1968. An analysis of vegetation- Dakota Press, Grand Forks. 104 pp. environment relationships in Saskatchewan sloughs. Can. J. Bot. Whitman, W.R. 1976. Artificial 46:509-522. wetlands for waterfow8. Pages 336-344 M. Smart, ed. Inter- Walker, B.N., and R.T. Coupland. national conference an the 1970. Herbaceous wetland vegeta- conservation of wetlands and tion in the aspen grove and grass- waterfowl. International Water- land regions of Saskatchewan. Can. fowl Research Bureau, Slimridge, J. Bot. 48:1861-1878. England, Walker, J.M. 1959. Vegetation Whitney, N.R. Jr., (chairman), B.E. studies on the Delta Marsh, Delta, Harrell, B. K. Harris, M. Nolden, Manitoba. M.S. Thesis, University J.W. Johnson, B.J. Rose, and P.F. of Manitoba, Winnipeg. 203 pp. Springer. 1978. The birds of South Dakota: an annotated checklist. Walker, J.M. 1965. Vegetation The South Dakota Brnitholaglist's changes with falling water Levels Union, With the cooperation of the in the Delta Marsh, Manitoba. W.H. Over Museum. Vemillion, SD. PR.D. Dissertation. University of 142 pp. Manitoba, Winnipeg. 272 pp. Wiedenheft, W.D. 1983. Life history Ward, E. 1942, Phragmites manage- and secondary production of tiger ment. Trans. N. Am. Wildl. Conf. salamanders (Ambvstoma tisrinum) 7: 294-298. in prairie pothole lakes. M.S Thesis. North Dakota State Uni- Ward, P. 2968. Fire in relation to versity, Fargo. 47 pp. waterfowl habitat of the Delta marshes. Proc. Tall Timbers Fire Williamson, L.L., and G.E. Doster. Ecol. Gonf, 8:254-267. 1981. Socio-economic aspects of white-tailed deer disease. Pages Weber, H.J., P.A. Vnhs, r., and 434-439 in W.R. Davidson, ed. L,D, Flake. 1982. Use of prairie Diseases and parasites of white- wetlands by selected bird species tailed deer. Tall Timbers Research in Sauth Dakota. Wilson Bull, Station, Tallahassee, Florida. 94:550-554, Willson, M. F. 1966. Breeding ecology WeXler, M,W. 1978. Management of of the yellow-headed blackbird. freshwater marshes for wildlife. Ecol. Monogr. 36:51-77. Pages 267-284 in R.E. Good, D-F. Whigham, and R.L. Simpson, eds. Winter, T.C. 1988. A conceptual Freshwater wetlands. Academic framework for assessing cumulative Press, New Uork, impacts on the hydrology of nontidal wetlands. Environ. WePPer, M,W,, and L.W, Fredrickson, Manage* 12:605-620. 1974. Avian ecology of a managed glacial marsh. X,iving Bird Winter, T.C, i989. Hydrologic 12:269-291. studies of wetlands in the 88 northern prairie. Pages 16-54 in North America. Boulder, C0. In A,G. van der Valk, ed. Northern press. prairie wetlands. Iowa State University Press, Ues. Wittnier, Pf. 1982. Wetlands: their status and acquisition in South Winter, T.C., and M.R. Carr. 1980, Dakota. S.D. Bird Notes 34:33-38. Hydrologic setting of wetlands in the Cottonwood Lake area, Stutsman Woodin, M.C., and G.A. Swanson. County, North Dakota. U.S. Geol. 1989. Foods and dietary Sum. WRI 80-99. 42 pp. strategies of prairie-nesting ruddy ducks and redheads. Condor Winter, T.C., and M-K. Woo. 91:280-287. Hydrology of lakes and wetlands, -In N.E. Ross, M.G. Wolman, and Wright, W.A., and A.W. Bailey. 1982. H.C. Riggs, eds. Surface water Fire ecology, United States and hydrology. Geological Society of southern Canada. John Wiley & America, Vol. 0-1. The geology of Sons, Mew York. 501 pp. .-..A-awn --wln VVW -- cJ\MU?b co m COW -3' mmt-m m ~ll mm G N m W Fd P rl In d d -- -- tom mm V.4 -- WCn NW PC? NQ m mm CJ d \O t.mlnu)bula In'amIOeNCQ wewm d to -r C? w d m wmto Prod ul m m CO~ONO~COOICO mww WCrr-P-wdmm 4 N ONdmLnm&O 4 \a vcfl N m r( ct N d ma\ --- -- rtl 000 oon Ah.-.-- rr. ddd &dm WlOcO03P- w u-V YYY UVVVV w Q) cfl V1 a3m-F N 10 cn 10 EB d m d rn d rtl - -- .-.- - P..- \am Inw www VV r_rU --.., W t-. WP. COW m co w O\O mw mot-. t-. a Ln wmm r( 03 N rc cl 0\ r( m '3am h4-J +J C, a) (I) 4 O'G aka U (I) N -4, E E '3.4 Q) la C4-J k U C4-J m C d d iu -4 -4 u cn a-r( I- r-a Ln -4'- N 6- A/-. P r- 10 LC) m r- - i-. -- -- r-- LC) u'l 01 al al P mLn t--2'mLOm rl s-4 w 0 wi-iw i-i t-- CO N m i-i N W rn ow Cn CR t- LO N rl P) d d in I 01 aQ1 rn I Xi-' 1 G C [ Oi 0) r-i PC: 1 a L-r Q 3 (Li N -4E I $2 wr-4 aJ a I riik UC+, i 5 .?r i CEO, I .4 .-I 0 m E I (doc i o;r n -4 d.--. ,-.. OOO..- - 0- -- -0 drirtcnco 03 w rd r- ma Lord -vvvv V - -- -- ....- omcnwm rn ri cv m ~n~nP-N m mdoa m iO F-( QO CJW mNri C1 * ma) Pd m a C T2 -IZm a; riii r 3 8.~ .-!i .n !iig -"g z .- I1 QkF$ C :E P2 E Tim L 5 85 ,Xv, mSL 8 C k - -Vr $j2 P 22 c 03 o a E a 8% C: 42 trip g2K tT gg 4 a. - ffi V- Apwndix @. 1. GaCsmmon emergent hydrophfles of palusPrisle welfands with temporarily flcad@dm6bb~iQkl~e regimes (wet meaa%OwS)In the Prairie Pothole Region of the Dakotas, araangegd according to increasing maxlmum obsepve?cfitolerance of sfissolved salts (from SmeIns -8967;Disrud 1968; Kantrud et a!. 1989). - S~ecific conductivity fm~/cml Species Rean Min. Max. Vernonia fasciculata QLL Aarostis stolonifera -0.2 Lvcowus americanus 0.3 Botentilla rivalis &?A Carex sti~ata -0.4 Emisetum arvense -0.4 Juncus interior -0.4 Aster sasittifolius l.0 Plantaqo mai or l.0 Potentilla norvesiea -0.3 Juncus dudlevi -0.4 Carex buxbaumii 1.2 Lvsimachia hvbrida 0.1 Carex vulpinoidea 1.0 Ranunculus macounii 1.1 Rumex mexicanus 0.5 Juncus bufonius -2.3 Cirsium arvense -2.5 Bidens cernua 1.5 Helenium autumnale 1.5 Carex praesracilis -0.3 Echinochloa crussalli 1.3 Carex Paeviconica 1.5 Rorissa islandica 1.7 Poa palustris 1.4 Stachys walustris 1.8 Calamaqrostis canadensis 1.4 Carex sartwellii 1.5 Lvcosus aswer l.g Epilobium slandulosum 1.5 Mentha arvensis 1.6 Apoc~numsibiricum 1.8 Eleocharis comsressa 2.0 Carex tetanica 2.0 Potentilla anserina -1.6 Boltonia asteroides 1.4 Carex lanusinosa 2.0 Teucrium occidentale 3.1 Aster hesperius 2.4 Juncus forrevi 1.7 Aster simplex -1.8 Calamasrostis inexpansa 2.4 Juncus balticus 3.3 (Continued) fa7 S~ecific conduct~vitv~rn~/crnl@ Species Mean Min. Max. magracilis 9.0 Plantaqo eriopoda 9.8 Sonchus arvensis 5.2 S~artinapectinata 3. Cl Mublenber~ig asperifolia 11.0 -Hordeurn i ubatum 7.8 Trisloehin maritirna 12.5 Distichlis s~icata 17.0 Atri~lexpatula 23.0 Vnderlined means (Disrud 1968; Mantrud et al. 1989) indi- cate surface water measurements in wetlands where the species reached peak abundance; underlined ranges (ibi.d) are for instances where the species occurred in waters of greater or lesser salinity than that recorded by Smeins (1967). "ndicates measurements <0.05 mS/cm. Appendix C, 2. Cornman ernergent hydrophges of plustrine wetlands with seasonally flooded (SE) (shallow-marsh zones), wmlpmanenely floorfed (SP) (dwpmarsh sones), and saturated (SA) {fen zones) moisture regtmes in the Prairie Pothole Region of the Dakotas, arranged according to increipslng maxtrncmrss olsww& tolerance of dissolveel salts (from Srnelns 1967;Disrud 1968; Kantrud et a!. 49891. Species Water regime Specific ~onductivit~~mS/cm)' Mean Min. Max. Eauisetum f luviatile SE - - GaIium trif idus SA - - Scutellaria g.alericu2atq S A - - Impatiens biflora S A -- Mimulus rinsens SA - - -Eupatorium rnacuPaturn SA -- Sag_ittar&- cuneata SE - - Glyceria striata 5 A - - Ranunculus melini SA - - Asclepias incarnata SA M Parnassia glauca SA -- Glvceria borealis SE -- Salix interior SA -0.3 --Carex lacustris SA 0.9 Solidaao graminifolia SA F_;?y-gonum amuhibium SE 0.1 Scirpus atrovirens SA Cicuta maculata sa u Eriphorum anqustifol- SA -0.5 Carex rostrata S A e%l~ganum coccineum SE Phalaris arundinacea SE Pk;L Carex aquatilis S A 0.3 Lvsimachia thrmiflora SA O.ti GBvceria srandis SE Tr, Siurn suave - p- SE 8.1 Scirpus hetesochaetus SP 0.1 Alopecurus aeaualis SE LLJ' Sparqanium WcarDurn SE Tr. Eleocharis acicularis SE Q3.2. Scirpus validus SP, SA 0.2 Tvxrha X slauca SP Q3.J Sasittaria cuneata SE B.a Scirxrus fluviati1.i~. SP 0.3 Abisma qramineum SE 0.3 Carex atherodes SE 22.L" Ranunculus s~eleratus SE 0.1 Beckmannia svziaachne SE I6sL Alisma ~lantaso-aauatica SE Tr. panunculus cvmbalaria SE 0.6 Scolochloa festucacea SE -0.1 Phxaqmites australis SA OIli Typha latifolia SP, SA X'JJ~& X'JJ~& anaustifolia SP 9L4 (Continued) -- Species Water regime Specific conductivity ~rn~/cml! Mean Min. Max. EPeocharis pafustris SE 2.7 ---14.5 Scir~usnevadens is SE 15.7 BZ,O 2.QLi scir~us cutu us SP 4.3 0.2 24.0 Suaeda de~ressa SE 24.0 ELLQ 5.kL!2 SC~~DUSamericanus SE 4-9 ---0.5 70,0 SC~~DUSmaritimus SP 40.3 LL.!l 76.4 Puccinellia nuttalliana SE ZQ,O L4. 76.4 --- 'underlined means (Disrud 4966; Kantrud et al. 1989) indicate surface water measurements in wetlands where the species reached peak abundance; underlined ranges (ibid) are for instances where the species occurred in waters of greater or lesser salinity than that recorded by Smeins (1967). b~ndicatesmeasurements i0.05 mS/cm. AppesKaix C. 3. C~rnrnawsubmew and flmaing aquatic plants of the Prairie Pothole Region of the Dakotas arrl;sng& =cording ro Increasing maximum observed toferance of dissolved sails (from Srnei'ns 1967; Disrltd 3968; Kantrud et al. 1989). Specific conductivitv (rn~/crn)' Gean or single Species measurement Min. Max. -Q+1 Q.4 0.4 e4 0.6 u 0.8 Callitriche aalustris 0.3 zl;ip~urisvulqaris BeZ. Callitriehe hennaphraditica 1.2 Ranunculus flabellaris 2.0 -Potarnoaeton zosteriformis 1.0 S~irodela~olvrhiza 1.5 Ricciacar~usnatans 2.2 Dre~anocladusspp. 1.2 3.3 1.7 1.4 2.1 2.1 2.1 2.2 2.7 3.1 3.2 Rup~iamarltzma var-occidentalis Q,b 4.8 2.2 6.5 var. pastrata 36.1 Vnderlined means (Disrud 15368; Kantrud et aX. 1989) indicate surface water measurements in wetlands where the species reached peak abundance; underlined ranges (ibid) are for instances where the species occurred in waters of greater or lesser salinity than that recorded by Smeins (1967) . Prairie Basin Wetlands of the Dakotas: A comw~ityProfile - +-I ----- 7. hrhezr(d I k rr-8~ Or#anixsdion Re Harold A. Kantrud, Gary L. Krapu, and Geo *------" 9. PE&~cnnllv# %.nlz.tGen NOmm and Addnrr W.S. Fish and Wildlife Service , ------. Northern Prairie Wildlife Research Center i 11. CoRtrrcaic> w ~ormf~~HO P.Q. Box 2096 1 Jaaestown, ND 58402 ------1 fG)(C) 12. Swn+aitnl, br~nnisrtlonMame itnd 4ddrrrr - ^ -- U.S, Department of the Interior L3 TrW ~4 Rome d: Pmfiad C~rrrrd Fish and Wildlife Service National Wetlands Research Center Washington, DC 20240 I4 " IS. Sopp4rrnmtary Notrt The shallow basin wetlands of the Dakotas form the bulk of the portlon of the Prairie Pothole Region lying within the United States. This region produces a large proportion af North America's waterfowl and otrfles prairie dwelling marsh and aquatic birds. The Prairie Pothole Region is nlso a major world supplier of cereal grains. Consequently, wetlands in the region are often drained for crop production or are otherwise cropped when water conditions permit. Prairie basin wetlands vary greatly in their ability ta maintain surface water and in their water chemistry, which varies from fresh to polysaline. In addition, these wetlands are affected by a variety ot agricuPtura1 land USES and practises, including pasture, cultivation, rneckxanical farage removal, idle conditions, and burning. Xt is important to understand how these factors operate in prairie basin wetlands, since they qreatXy affect the plant and animal communities in these basins- Ecvl oqy nydrology waterfowl Gcoao~~y sails Prairie potholes Agric~xlturaf impacts Wetlands Ecusysteq processes Marshes 6 CDSAII fiaid(Grurto - - -- +- Ik AvrllmtM~tyStstamant Unlimited Distributian Unclassified OPTiO%AL SQRM I?? (4-7" (FM-rly MTIS-35)