Literature Review and Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Montana

Prepared in cooperation with the U.S. Fish and Wildlife Service

Scientific Investigations Report 2009–5098

U.S. Department of the Interior U.S. Geological Survey

Literature Review and Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Montana

By Robert A. Gleason, Brian A. Tangen, Murray K. Laubhan, Raymond G. Finocchiaro, and John F. Stamm

Prepared in cooperation with the U.S. Fish and Wildlife Service

Scientific Investigations Report 2009–5098

U.S. Department of the Interior U.S. Geological Survey U.S. Department of the Interior KEN SALAZAR, Secretary U.S. Geological Survey Suzette M. Kimball, Acting Director

U.S. Geological Survey, Reston, Virginia: 2009

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Suggested citation: Gleason, R.A., Tangen, B.A., Laubhan, M.K., Finocchiaro, R.G., and Stamm, J.F., 2009, Literature review and database of relations between salinity and aquatic biota—applications to Bowdoin National Wildlife Refuge, Montana: U.S. Geological Survey Scientific Investigations Report 2009–5098, 76 p. iii

Contents

Abstract...... 1 Introduction...... 1 Purpose and Scope...... 3 Description of Bowdoin National Wildlife Refuge...... 3 Methods...... 5 Primer on Salinity Concepts...... 5 Salinity Definitions and Measurements...... 5 Sodicity...... 5 Factors Influencing Salinity...... 6 Tolerances and Adaptations of Biota to Salinity...... 13 Plants ...... 13 Invertebrates...... 14 Database Summary Statistics...... 16 Plants ...... 16 Invertebrates...... 16 Considerations in Using Information from the Plant and Invertebrate Databases...... 16 Application to Bowdoin National Wildlife Refuge...... 19 Plants ...... 19 Invertebrates...... 22 Summary...... 22 Acknowledgments...... 22 References...... 27 Appendix 1. Plant Taxa Identified at Bowdoin National Wildlife Refuge...... 30 References...... 31 Appendix 2. Invertebrate Taxa Identified at Bowdoin National Wildlife Refuge...... 32 References...... 32 Appendix 3. Description of Variables Found in the Plant and Invertebrate Databases...... 33 References...... 34 Appendix 4. Description of Each Data Source Used to Develop the Plant and Invertebrate Databases...... 35 References...... 40 Appendix 5. Summary of Plant Database...... 42 References...... 64 Appendix 6. Summary of Invertebrate Database...... 65 References...... 76 iv

Compact Disc

[In pocket]

Plant and Invertebrate Database (http://pubs.usgs.gov/sir/2009/5098/downloads/databases_21april2009.xls) Format: American Standard Code for Information Interchange (ASCII), Microsoft Office Excel 97–2003 Worksheet, Windows XP Professional operating system

Figures

1. Map showing the location of Bowdoin National Wildlife Refuge in relation to Beaver Creek, Milk River, Dodson South Canal, and Nelson Reservoir...... 2 2–16. Graphs showing: 2. Concentration of major ions in Lake Bowdoin during April–August 2006...... 6 3. Comparison of common reporting units for salinity in relation to plant community salinity categories and the salinity of seawater...... 7 4. Mean annual (± standard error) specific conductance for Lake Bowdoin from 1975 to 2007...... 9 5. Palmer Hydrological Drought Index (PHDI) from 1895 to 2007 and yearly mean specific conductance for Lake Bowdoin from 1975 to 2007...... 10 6. Palmer Hydrological Drought Index (PHDI) from 1895 to 2007 and deliveries to Bowdoin Wildlife Refuge from the Milk River via the Dodson Canal from 1938 to 2007...... 10 7. Palmer Hydrological Drought Index (PHDI) from 1895 to 2007 and Beaver Creek flows near Bowdoin National Wildlife Refuge from 1950 to 2007 (monthly) and from 1982 to 2007 (peak)...... 11 8. Data on reservoirs in the Beaver Creek watershed created between 1898 and 2004 for watering stock, providing wildlife habitat, and irrigation supply...... 12 9. Number of taxa from Prairie Pothole Region wetlands in each plant community salinity category...... 13 10. Number of taxa from the plant database by various categories...... 17 11. Number of taxonomic orders from the invertebrate database for each category...... 18 12. Specific conductance of water and soil for each plant physiognomy/life-span category and each wetland indicator category...... 20 13. Specific conductance of water for common plant species at Bowdoin National Wildlife Refuge that are associated with wetlands...... 23 14. Specific conductance of soil for common plant species at Bowdoin National Wildlife Refuge that are associated with wetlands...... 24 15. Specific conductance of water for representative taxa in the invertebrate database that could occur at Bowdoin National Wildlife Refuge...... 25 16. Specific conductance of water for invertebrate families from representative taxonomic orders in the invertebrate database...... 26 v

Tables

1. Slope and area of primary soils on Bowdoin National Wildlife Refuge...... 4 2. Plant community salinity categories and the corresponding ranges of specific conductance values of water for three common reporting units...... 7 3. Classification of saline and sodic soils and related soil condition based on specific conductance, pH, and the sodium adsorption ratio...... 7 4. Reported ranges of specific conductance, pH, and sodium adsorption ratio of soils located at Bowdoin National Wildlife Refuge...... 8 5. Sodium chloride concentration at which seed germination of halophytic plant species was reduced from 75–100 percent to approximately 10 percent...... 15 6. Summary information from the plant database (210 total species) for 110 species documented on Bowdoin National Wildlife Refuge ...... 19 vi

Conversion Factors

Inch/Pound to SI

Multiply By To obtain Length mile (mi) 1.609 kilometer (km) Area acre 0.4047 hectare (ha) Volume acre-foot (acre-ft) 0.123 hectare meter (ha-m) Mass ton, short (2,000 lb) 0.9072 megagram (Mg) Flow rate cubic foot per second (ft3/s) 0.02832 cubic meter per second (m3/s)

SI to Inch/Pound

Multiply By To obtain Length kilometer (km) 0.6214 mile (mi) Area hectare (ha) 2.471 acre Volume hectare meter (ha-m) 8.107 acre-foot (acre-ft) Mass megagram (Mg) 1.102 ton, short (2,000 lb) Flow rate cubic meter per second (m3/s) 35.31 cubic foot per second (ft3/s)

Specific conductance is given in microsiemens per centimeter at 25 degrees Celsius (µS cm-1 at 25°C). Concentrations of chemical constituents in water are given in milligrams per liter (mg L-1). Throughout this report (including the plant and invertebrate databases) mg L-1 is related to µS cm-1 by using the following formula: mg L-1 = µS cm-1 × 0.64 (Tchobanoglous and Burton, 1991). Literature Review and Database of Relations Between Salinity and Aquatic Biota: Application to Bowdoin National Wildlife Refuge, Montana

By Robert A. Gleason, Brian A. Tangen, Murray K. Laubhan, Raymond G. Finocchiaro, and John F. Stamm

Abstract Introduction

Long-term accumulation of salts in wetlands at Bowdoin Bowdoin National Wildlife Refuge (NWR) was estab- National Wildlife Refuge (NWR), Mont., has raised concern lished in 1936 to provide habitat for breeding and migrating among wetland managers that increasing salinity may threaten waterfowl. The refuge is located about 11 km to the east of plant and invertebrate communities that provide important Malta, in the Milk River Valley of Phillips County, Mont. habitat and food resources for migratory waterfowl. Currently, (fig. 1). Characteristic of refuges located in the arid North- the U.S. Fish and Wildlife Service (USFWS) is evaluating Central United States, Bowdoin NWR is at risk to increased various water management strategies to help maintain suitable salinization and accumulation of trace elements that can ranges of salinity to sustain plant and invertebrate resources of threaten wetland biota (DuBois and others, 1992; Nimick, importance to wildlife. To support this evaluation, the USFWS 1997; Laubhan and others, 2006). Bowdoin NWR has been requested that the U.S. Geological Survey (USGS) provide the subject of various investigations and model simulations information on salinity ranges of water and soil for common (Lambing and others, 1988; Hamilton and others, 1989; plants and invertebrates on Bowdoin NWR lands. To address Kendy, 1999; Bauder and others, 2007) that suggest that salt this need, we conducted a search of the literature on occur- concentrations in lakes and marshes are increasing and that the rences of plants and invertebrates in relation to salinity and long-term accumulation of salts could pose a significant threat pH of the water and soil. The compiled literature was used to to flora and fauna. The increasing salt concentrations on the (1) provide a general overview of salinity concepts, (2) docu- refuge have been attributed to changes in water availability ment published tolerances and adaptations of biota to salinity, and quality that, in conjunction with water management strate- (3) develop databases that the USFWS can use to summarize gies that stabilize water levels and enhance retention time, the range of reported salinity values associated with plant and have amplified the evapoconcentration of salts and reduced the invertebrate taxa, and (4) perform database summaries that effectiveness of natural salt removal processes such as flushing describe reported salinity ranges associated with plants and and deflation (removal by wind) (Hamilton and others, 1989). invertebrates at Bowdoin NWR. The purpose of this report is Further, this increase in salinity has been linked to alteration to synthesize information to facilitate a better understanding of vegetation communities that provide important wildlife of the ecological relations between salinity and flora and fauna habitat. For example, Hamilton and others (1989) observed when developing wetland management strategies. A primary that salt-tolerant plants had replaced cat-tails (Typha sp.) and focus of this report is to provide information to help evaluate bulrushes (Scirpus sp.), and submerged aquatic species such as and address salinity issues at Bowdoin NWR; however, the pondweeds (Potamogeton sp.) and widgeongrass (Ruppia sp.) accompanying databases, as well as concepts and informa- had largely disappeared. Additionally, declines in waterfowl tion discussed, are applicable to other areas or refuges. The use and productivity have been attributed to increased salinity accompanying databases include salinity values reported for and subsequent changes in vegetative composition. 411 plant taxa and 330 invertebrate taxa. The databases are To ensure the long-term success of Bowdoin NWR in available in Microsoft Excel version 2007 (http://pubs.usgs. providing healthy, sustainable habitat for waterfowl and other gov/sir/2009/5098/downloads/databases_21april2009.xls) and wildlife, the U.S. Fish and Wildlife Service (USFWS) is devel- contain 27 data fields that include variables such as taxonomic oping management strategies to maintain salt concentrations identification, values for salinity and pH, wetland classifica- within ranges suitable for the establishment of historical plant tion, location of study, and source of data. The databases are and invertebrate communities. Currently, the USFWS is evalu- not exhaustive of the literature and are biased toward wetland ating various water management scenarios that will enhance habitats located in the glaciated North-Central United States; their ability to export salts and manage salinity levels in lakes however, the databases do encompass a diversity of biota and marshes on the refuge. Water management scenarios commonly found in brackish and freshwater inland wetland considered by the USFWS range from no action to controlled habitats. and uncontrolled (for example, flushing caused by floods) 2 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

107°44′ 107°36′ 48°32′

ir o v er es R

River n lso Ne Milk

Lakeside Unit Lakeside Extension

Bowdoin National Dry Lake Wildlife Refuge Unit Dry Lake Lake Bowdoin Pond 48°24′

Creek

Dodson South Canal er Drumbo Unit av e B 0 2.5 5 10 KILOMETERS

0 1.25 2.5 5 MILES

Base from Environmental Systems Research Institute, Inc., 2006, various scales, and Montana Natural Resource Information System, various dates and scales Universal Transverse Mercator projection, Zone 13 North American Datum of 1983 (NAD 83)

R O Milk Study area C K Y River

River

M Fort Peck

O Missouri Lake U

N T MONTANA A

I N

48°16′

Figure 1. Location of Bowdoin National Wildlife Refuge in relation to Beaver Creek, Milk River, Dodson South Canal, and Nelson Reservoir. Description of Bowdoin National Wildlife Refuge 3 exports of saline water to creeks and rivers (oral commun., wetlands. Terrestrial soils of the refuge consist primarily of Comprehensive Conservation Plan planning committee). Bigsag clays and the Phillips-Elloam complex. The portion Each of these scenarios will result in varying levels of salinity (36 percent) of the refuge inundated by lakes is bordered, and control in lakes and marshes that will ultimately affect wildlife likely underlain, by Bigsag and Bowdoin soils (Kendy, 1999), habitat in terms of plant and invertebrate communities. which are relatively fine grained with low permeability and a To support the evaluation of water management scenar- high salt content. An inventory of primary soils and complexes ios, the USFWS requested that the U.S. Geological Survey on the refuge is presented in table 1. (USGS) provide information on suitable ranges of salinity Refuge habitats consist primarily of short- and mixed- for plants and invertebrates of importance to the refuge. To grass prairie, planted dense nesting cover, shrublands, and address this need, we present summarized data obtained from both brackish and freshwater wetlands. Wetlands make up various sources (for example, books, peer-reviewed publica- about 46 percent (2,924 ha) of the refuge, with the major- tions, reports, subject reviews, unpublished data) that relate ity of this area consisting of five large, shallow lakes: Lake the occurrences of common aquatic plant and invertebrate Bowdoin, Drumbo Unit, Dry Lake Unit, Dry Lake Pond, and species to various water and soil quality variables such as Lakeside Unit (fig. 1). Lake Bowdoin, the largest (2,209 ha) specific conductance and pH. As part of the review, we also and most prominent lake, is an oxbow of the preglacial discuss general physiological adaptations that allow flora and Missouri River. The modern Missouri River flows nearly fauna to survive in salt-affected environments. 113 km to the south of the refuge. Prior to the establishment of Bowdoin NWR, Lake Bowdoin was managed by the Bureau of Reclamation to store water from spring floods and irrigation return flows. Purpose and Scope Direct precipitation and diverted water from the Milk River via the Dodson South Canal are the main sources of The goal of this report is to provide the USFWS with water entering the refuge, while irrigation drainage returns scientifically based information to facilitate a better under- from nearby agricultural lands, overland runoff, floodwaters standing of the ecological relations between salinity and from adjacent drainages (for example, Beaver Creek), and flora and fauna when developing management strategies. groundwater discharges from saline seeps are secondary Although the primary focus is to provide information to help sources. An annual water allocation of 431 ha-m from the address salinity issues at Bowdoin NWR, the databases, as Milk River Irrigation Project that flows through Dodson South well as concepts and information discussed, are intended to Canal provides a relatively reliable source of water to the be applicable to other areas or refuges. To address our goal, refuge; however, this allocation can fluctuate annually and is we conducted an extensive search to locate existing scientific contingent on the amount of water available for distribution. literature that reported occurrences of plants and invertebrates Measured water deliveries from the years 1938 to 2006 ranged in relation to salinity and pH of the water and soil. Information from 70 to 1,419 ha-m and averaged 597 ha-m. Water losses from this literature search was incorporated into databases and are attributed primarily to evaporation and infrequent surface used to address the following objectives: outflows to neighboring Beaver Creek. The average annual 1. Provide a general overview of salinity concepts and the evaporation (92.71 cm) (Kendy, 1999) greatly exceeds the tolerances and adaptations of biota to salinity. mean annual precipitation (27.94–35.56 cm) (U.S. Department 2. Develop databases that the USFWS can use to summarize of Agriculture, 2004) in this semiarid region; thus, the refuge the range of reported salinity values associated with plant experiences a water deficit for much of the year. Currently, and invertebrate taxa. USFWS personnel manage refuge wetlands for waterfowl and 3. Perform database summaries that describe reported salin- other wildlife by using a system of channels and dikes that ity ranges associated with plants and invertebrates at allows for the capture, storage, and movement of water within Bowdoin NWR. the refuge. A total of 210 plant species have been documented on the refuge (app. 1). Dominant emergent vegetation includes Description of Bowdoin National hardstem bulrush (Scirpus acutus), alkali bulrush (S. paludosus), common threesquare (S. pungens), and cat-tail (Typha Wildlife Refuge latifolia). Other notable species that occur along the shores of lakes and marshes include pickleweed (Salicornia rubra) The 6,294-ha refuge is located in the Glaciated Northern and saltgrass (Distichlis spicata). Common aquatic vegetation Grasslands (level IV ecoregion) (Woods and others, 2002) of includes sago pondweed (Potamogeton pectinatus), widgeon- northeastern Mont., which is characterized by rolling hills and grass (Ruppia maritima), and watermilfoil (Myriophyllum plains that are the result of glaciation. In general, glacial and exalbescens) (Weydemeyer and Marsh, 1936; Hamilton and alluvial deposits overlay bedrock, and the region is character- others, 1989; Johnson, 1990). Information on invertebrate ized by saline-sodic soils (sodium and sulfate are the domi- communities is relatively limited; however, 32 invertebrate nant ions) and grasslands that contain numerous depressional taxa have been documented within lakes and marshes on the 4 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. refuge (app. 2). Common aquatic macroinvertebrates documented on the refuge include midges (Chironomidae), scuds (Talitridae), water boatman (Corixidae), snails (Planorbidae and Physidae), damselflies (Coenagrionidae), mayflies (Caeni- dae), and water fleas (Cladocera) (Johnson, 1990).

Table 1. Slope and area of primary soils on Bowdoin National Wildlife Refuge.

[Data provided by U.S. Fish and Wildlife Service. Areas are based on 2005 color-infrared imagery and the Natural Resources Conservation Service’s Soil Survey Geographic Database. --, no data]

Slope range, Area, Cumulative percent Soil Percent of refuge in percent in hectares of refuge Water -- 2,269.13 35.68 35.68 Bigsag clay 0–2 889.50 13.99 49.67 Phillips-Elloam complex 0–4 463.07 7.28 56.95 Bowdoin clay 0–2 387.58 6.09 63.04 Creed-Absher complex 0–4 372.11 5.85 68.89 Scobey-Kevin clay loams 2–8 349.75 5.50 74.39 Scobey-Kevin-Elloam clay loams 2–8 279.08 4.39 78.78 Ferd-Gerdrum complex 0–4 237.77 3.74 82.52 Telstad loam 0–4 219.79 3.46 85.98 Creed-Gerdrum complex 0–4 167.26 2.63 88.61 Lardell clay loam 0–2 154.30 2.43 91.04 Telstad-Joplin loams 2–8 131.13 2.06 93.10 Vanda clay 0–2 113.52 1.79 94.89 Scobey-Phillips complex 0–4 104.53 1.64 96.53 Scobey-Elloam-Absher gravelly clay loams 2–8 53.78 0.85 97.38 Phillips-Absher complex 0–4 44.50 0.70 98.08 Joplin-Hillon loams 2–8 35.15 0.55 98.63 Sunburst-Kevin gravelly clay loams 8–15 28.67 0.45 99.08 Scobey clay loam 0–4 19.03 0.30 99.38 Scobey-Elloam clay loams 2–8 11.70 0.18 99.56 Harlake-Lostriver clays 0–2 10.93 0.17 99.73 Nishon clay loam 0–2 8.18 0.13 99.86 Degrand loam 0–4 4.09 0.06 99.92 Scobey-Kevin complex 2–8 2.91 0.05 99.97 Kobase silty clay 0–2 0.81 0.01 99.98 McKenzie clay 0–2 0.64 0.01 99.99 Hillon-Kevin complex 8–15 0.08 <0.01 99.99 Primer on Salinity Concepts 5

Methods solution to conduct an electrical current and is highly corre- lated with the concentration of ions in solution. Conductance measurements also are highly dependent on temperature We developed plant and invertebrate databases by because an increase in the temperature of a solution will conducting an extensive search to locate existing scientific decrease viscosity and increase the mobility of ions in solu- literature that reported occurrences of plants and invertebrates tion; a rise in temperature could also increase the number in relation to salinity and pH of the water and soil. Literature and keyword (for example, wetlands, salinity, salt, inver- of ions in solution because of dissociation of molecules. tebrates, plants) searches were conducted by using various Because of the need to compare conductivity values collected electronic databases such as FirstSearch, Cambridge Scientific at various temperatures by using differing equipment (that is, Abstracts, Water Resources Abstracts, Web of Science, Google probes that differ with respect to electrode distance and area), Scholar, and others. Additionally, we used unpublished data measurement devices are typically temperature-compensated gathered as part of various investigations conducted primarily and provide standardized readings at 25°C with an electrode -1 by the USGS Northern Prairie Wildlife Research Center. distance of 1 cm (for example, 1,000 µS cm at 25°C) (Lind, Information obtained from this search was entered 1985). Specific conductance is commonly reported as millisie- into spreadsheet databases to facilitate data summaries. The mens (mS), microsiemens (µS), millimhos (mmhos), or detailed databases contain 27 fields that include variables micromhos (µmhos). Siemens and mhos are equivalent units such as taxonomic identification, values for salinity and pH, that are reciprocal of the ohm, the International System (SI) wetland classification, location of study, and source of data. unit of electrical resistance. Appendix 3 provides a detailed description of each variable Total dissolved solids is a measure of all inorganic and found in the databases, and appendix 4 includes an annotated organic substances dissolved in water and is usually presented description of the data obtained from each source to provide as parts per thousand (ppt, ‰), percent (%), milligrams per -1 users information on the utility or applicability of the data to liter (mg L ), or total mass (in grams [g]). The concentration their region of interest. of TDS is typically determined by using gravimetry, which involves evaporating a known volume of liquid solvent to leave a residue which is then weighed to determine mass of dissolved solids. Total dissolved solids also may be approxi- Primer on Salinity Concepts mated from specific conductance measurements. For example, specific conductance is typically multiplied by an empirical Salinity Definitions and Measurements factor that generally varies from 0.5 to 1.0 to estimate TDS (Lind, 1985). Throughout this report (including the plant and Salinity refers to the concentration of dissolved or invertebrate databases) mg L-1 is related to µS cm-1 by using soluble salts in water and soil. Salts are generally defined as the following formula: mg L-1 = µS cm-1 × 0.64 (Tchobano- ionic compounds which in water solution yield a cation other glous and Burton, 1991). As a reference, in figure 3, three + - than hydrogen (H ) and an anion other than hydroxyl (OH ) common salinity reporting units (µS cm-1, mg L-1, and ppt) are - or dioxygen (O2 ). As an example, the following equation compared in relation to plant community salinity categories displays the chemical formula for common table salt, sodium (table 2) described by Stewart and Kantrud (1972). chloride:

Na+ + Cl- ↔ NaCl Sodicity

Common ions (cations and anions) found in northern Sodicity refers to the sodium ion concentration in a soil wetlands, including those within Bowdoin NWR (fig. 2), are or solution. Substrates classified as sodic have high sodium calcium (Ca2+), magnesium (Mg2+), potassium (K+), sodium ion concentrations relative to other cations such as calcium or + - 2- - magnesium. High sodium concentrations cause soil aggre- (Na ), bicarbonate (HCO3 ), carbonate (CO3 ), chloride (Cl ), 2- gates to swell and disperse, resulting in destruction of soil and sulfate (SO4 ) (Swanson and others, 1988; Nimick, 1997; Bauder and others, 2007). When removed from solution (for structure. In contrast, calcium and magnesium ions cause soil example, through evaporation), principal salts in water and particles to flocculate, forming aggregates that enhance and stabilize soil structure. The extent of soil structure destruction soils at Bowdoin NWR include sodium sulfate (Na2SO4), or enhancement caused by cations is not only a function of sodium bicarbonate (NaHCO3), calcium carbonate (CaCO3), concentration but also is related to the organic matter content, calcium sulfate (CaSO4), and magnesium sulfate (MgSO4); minor amounts of chlorine (Cl-) and fluoride (F-) salts are also carbonate and sulfate concentrations (for example, calcite and found (Bauder and others, 2007). gypsum), leaching potential, and texture of the soil. The loss Measures of salinity for both soil and water are usually of soil structure can affect plant growth by impairing hydrau- expressed as conductance (that is, specific conductance, elec- lic conductivity (water movement through soil), drainage, trical conductivity) or total dissolved solids (TDS). Specific gas exchange, and rooting depth. Additionally, sodicity—and conductance is a measure of the capacity of an aqueous salinity to a certain extent—is often accompanied by high 6 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. pH, which may impact plant growth by causing nutrient a relatively wet and dry year. LaBaugh (1989) reviewed data imbalance. from northern prairie wetlands and lakes and reported ranges Soils are commonly classified as saline, sodic, or saline- of specific conductance and TDS of 42–472,000 µS cm-1 and sodic on the basis of a combination of three factors: the 31–342,000 mg L-1 (approximately 48–534,375 µS cm-1), sodium adsorption ratio (SAR), specific conductance, and respectively. Additionally, LaBaugh (1989) described studies pH (table 3). The SAR is the ratio of sodium to calcium and in which specific conductance increased twofold to sixfold and magnesium cations in solution, defined as salinity increased by 80 ppt (approximately 125,000 µS cm-1) within a season. Variation in salinity is attributable to many factors, including climate, connectivity to regional and local groundwater flow systems, and intra-annual and interannual In general, when the SAR is above 12, physical properties of fluctuations in water levels that result in the dilution, evapo- soil may be altered, and plants can become stressed because concentration, or deflation (blowing of salts from mudflats) of ion toxicity and reduced water absorption (osmotic stress). of salts (Lieffers and Shay, 1983; LaBaugh, 1989; LaBaugh We do not have specific field data (see table 3) necessary to and others, 1996). For example, between 1975 and 2007 the classify soils found at Bowdoin NWR (table 1); however, on mean annual salinity in Lake Bowdoin varied from 4,113 (in -1 the basis of reported ranges of SAR, specific conductance, and 1990) to 28,908 (in 1984) µS cm (fig. 4). During this 33-year pH values (table 4) for soils on the refuge, all three soil classes period, Lake Bowdoin was classified as moderately brack- likely occur, especially if the classifications were based on the ish 9 percent, brackish 79 percent, and subsaline 12 percent upper end of the reported values (table 4). of the time. Such variation is characteristic of saline lakes, which often experience dynamic shifts in salinity in response to extreme climatic fluctuations that operate at decadal time scales. The influence of climatic variation on Lake Bowdoin Factors Influencing Salinity salinity flux is illustrated when salinity is examined in relation to the Palmer Hydrological Drought Index (PHDI), which When evaluating the suitability of habitats for aquatic shows long-term cumulative drought and wet conditions biota, it is important to recognize that water and soil salin- (fig. 5). Variation in the PHDI over the past 30 years, or since ity can vary considerably both temporally and spatially. salinity has been monitored in Lake Bowdoin, indicates two Swanson and others (1988) reported that specific conductance drought periods (1980–85, 1999–2007) and two relatively wet measurements for 178 prairie lakes in North Dakota ranged periods (1975–80, 1986–98) (fig. 5). During these alternating from 365 to 70,000 µS cm-1 and that measurements from wet and dry periods, salinity levels in Lake Bowdoin tended to 1 pothole wetland varied from 522 to 7,700 µS cm-1 between decrease during wet periods and increase during drought.

14,000

12,000 EXPLANATION Sulfate (SO 2- ) 10,000 4 Sodium (Na+) - 8,000 Bicarbonate (HCO3 ) Magnesium (Mg2+) 6,000 Chloride (Cl-) 2- Carbonate (CO3 ) 4,000 Potassium (K+) Calcium (Ca2+ ) 2,000

0 CONCENTRATION, IN MILLIGRAMS PER LITER APRIL MAY JUNE JULY AUGUST

Figure 2. Concentration of major ions in Lake Bowdoin during April–August 2006 (data provided by the U.S. Fish and Wildlife Service). Primer on Salinity Concepts 7

100,000 60,000 60

80,000 50,000 50 Saline

Seawater 40,000 40 60,000

30,000 30 40,000

Subsaline 20,000 20 PARTS PER THOUSAND MILLIGRAMS PER LITER 20,000 10,000 10 MICROSIEMENS PER CENTIMETER Brackish

0 0 0 Moderately brackish Slightly brackish Fresh

Figure 3. Comparison of common reporting units for salinity in relation to plant community salinity categories (see table 2) and the salinity of seawater (approximately 35 parts per thousand).

Table 2. Plant community salinity categories and the corresponding ranges of specific conductance values of water for three common reporting units.

[Categories from Stewart and Kantrud, 1972. µS cm-1, microsiemens per centimeter; mg L-1, milligrams per liter; ppt, parts per thousand]

Salinity category µS cm-1 mg L-1 ppt Fresh 0–500 0–320 0–0.3 Slightly brackish 500–2,000 320–1,280 0.3–1.3 Moderately brackish 2,000–5,000 1,280–3,200 1.3–3.2 Brackish 5,000–15,000 3,200–9,600 3.2–9.6 Subsaline 15,000–45,000 9,600–28,800 9.6–28.8 Saline >45,000 >28,800 >28.8

Table 3. Classification of saline and sodic soils and related soil condition based on specific conductance, pH, and the sodium adsorption ratio.

[All three criteria—specific conductance, pH, and sodium adsorption ratio (SAR)—are considered for classifying soils as saline, sodic, and saline-sodic (modified from Havlin and others, 1999; North Dakota State UniversityAgriculture and University Extension Service Web site, accessed October 1, 2008, at http://www.ag.ndsu.edu/pubs/plantsci/soilfert/eb57-2.htm). µS cm-1, microsiemens per centimeter]

Specific conductance, Classification Soil condition pH SAR µS cm-1 Saline Normal1 >4,000 <8.5 <13 Sodic Poor <4,000 >8.5 >13 Saline-sodic Normal1 >4,000 <8.5 >13 1Soil retains “normal” physical conditions as long as sodium does not dominate the exchange complex of the soil. 8 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

The increased salinity during drought is attributed to Creek not only dilute salts but also flush salts from the system. evapoconcentration of solutes; however, drought also provides The most notable Beaver Creek flood event that resulted in mechanisms for removal of salt by deflation and seepage to significant salt flushing during the past 30 years occurred groundwater (Weydemeyer and Marsh, 1936; Hamilton and in 1986 when peak flows exceeded 20,000 ft3/s (fig. 7B); others, 1989; LaBaugh and others, 1996). Superimposed over this event coincided with a sixfold salinity decrease in Lake these natural processes of deflation and seepage is the effect Bowdoin (fig. 5). There are insufficient data to calculate the of water deliveries to the refuge via the Dodson South Canal frequency of Beaver Creek flood events (fig. 7), but refuge (fig. 6), which can be used to dilute salts or maintain water personnel estimate that the historical average of floodwater levels during dry periods that would normally result in defla- occurrence on the refuge was once every 3–5 years (Rodney tion events. Consequently, manipulating water levels provides and Mohrman, 2006; 2007). More recent observations by an opportunity to both trigger and halt deflation episodes. refuge staff suggest, however, that the frequency of flood Decreases in salinity during wet periods are attributed events has decreased to once every 7–10 years. The reduced to increased freshwater inputs that dilute salt concentrations. flood frequency is believed to have been caused by the estab- Further, periodic floodwaters that enter the refuge from Beaver lishment of small impoundments and irrigation diversions in

Table 4. Reported ranges of specific conductance, pH, and sodium adsorption ratio of soils located at Bowdoin National Wildlife Refuge (see table 1).

[Soil series information obtained from the Natural Resources Conservation Service’s Official Soil Series Descriptions, accessed July 2008, athttp://soils.usda. gov/technical/classification/osd/index.html. µS cm-1, microsiemens per centimeter; SAR, sodium adsorption ratio; --, no data]

Total area, Specific conductance, Soil series Taxonomic family pH SAR in hectares1 µS cm-1 Bigsag 889.5 Fine, smectitic, calcareous, frigid Typic Halaquepts >16,000 7.9–9.0 13–20 Scobey 820.8 Fine, smectitic, frigid Aridic Argiustolls -- 6.6–8.4 1–8 Elloam 807.7 Fine, smectitic, frigid Aridic Natrustalfs 2,000– 8,000 6.6–9.0 8–25 Kevin 660.5 Fine-loamy, mixed, superactive, frigid Aridic Argius- -- 6.6–7.8 -- tolls Phillips 612.1 Fine, smectitic, frigid Aridic Haplustalfs 0–2,000 6.1–7.3 <13 Creed 539.4 Fine, smectitic, frigid Aridic Natrustalfs 0–4,000 6.1–8.4 8–20 Absher 470.4 Fine, smectitic, frigid Leptic Torrertic Natrustalfs 8,000–16,000 6.6–9.6 18–70 Gerdrum 405.0 Fine, smectitic, frigid Torrertic Natrustalfs 1,000–8,000 7.4–9.0 10–20 Bowdoin 387.6 Very-fine, smectitic, frigid Sodic Haplusterts 8,000–16,000 7.4–9.0 5–13 Telstad 350.9 Fine-loamy, mixed, superactive, frigid Aridic Argius- 2,000–4,000 6.6–7.8 -- tolls Ferd 237.8 Fine, smectitic, frigid Aridic Haplustalfs 0–2,000 6.6–7.8 <13 Joplin 166.3 Fine-loamy, mixed, superactive, frigid Aridic Argius- 0–4,000 6.6–8.4 -- tolls Lardell 154.3 Fine-loamy, mixed, superactive, frigid Typic Aquisalids >16,000 7.9–10 8–50 Vanda 113.5 Fine, smectitic, calcareous, frigid Torrertic Ustorthents 2,000–8,000 7.8–9.6 1–30 Hillon 35.2 Fine-loamy, mixed, superactive, calcareous, frigid -- 7.9–9.0 -- Aridic Ustorthents Sunburst 28.7 Fine, smectitic, calcareous, frigid Torrertic Ustorthents -- 7.9–8.4 -- Harlake 10.9 Fine, smectitic, calcareous, frigid Aridic Ustifluvents 0–2,000 6.6–8.4 0–8 Lostriver 10.9 Fine, smectitic, calcareous, frigid Aridic Ustifluvents 8,000–16,000 7.4–9.6 13–30 Nishon 8.2 Fine, smectitic, frigid Typic Albaqualfs -- 6.1–8.4 -- Degrand 4.1 Fine-loamy over sandy or sandy-skeletal, mixed, super- -- 6.6–7.8 -- active, frigid Aridic Argiustolls Kobase 0.8 Fine, smectitic, frigid Torrertic Haplustepts 0–4,000 6.6–8.4 0–4 McKenzie 0.6 Fine, smectitic, frigid Chromic Endoaquerts ------1Total area of all soil complexes that include the soil series (see table 1). Primer on Salinity Concepts 9 the Beaver Creek watershed (fig. 8). For example, Rodney and Mohrman (2006) estimated that irrigation diversion and reservoir retention have reduced total mean annual runoff in the Beaver Creek watershed upstream of Bowdoin NWR by 45 percent. The observed salinity fluctuations in Lake Bowdoin that occur in relation to changes in climate and associated export mechanisms provide a basis for understanding how management actions and human alterations in the watershed may alter the sustainability of salt-affected lakes. Hence, understanding and defining the key temporal scales of variability in relation to ecological processes are fundamental to saline lake management.

50,000 Saline

40,000

30,000 Subsaline

20,000

SPECIFIC CONDUCTANCE, 10,000 Brackish

IN MICROSIEMENS PER CENTIMETER Moderately brackish Slightly brackish 0 Fresh 1975 1980 1985 1990 1995 2000 2005

YEAR

Figure 4. Mean annual (± standard error) specific conductance for Lake Bowdoin from 1975 to 2007. Data were provided by the U.S. Fish and Wildlife Service. Horizontal dashed lines define plant community salinity categories (see table 2) and allow for a general characterization of vegetative composition in Lake Bowdoin from year to year. 10 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1980 1998 1975 1985 10 30,000

20,000 5 10,000

0 0

-5 SPECIFIC CONDUCTANCE,

PHDI IN MICROSIEMENS PER CENTIMETER Specific conductance -10 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 PALMER HYDROLOGICAL DROUGHT INDEX (PHDI) YEAR

Figure 5. Palmer Hydrological Drought Index (PHDI) from 1895 to 2007 and yearly mean specific conductance for Lake Bowdoin from 1975 to 2007. Positive PHDI values indicate wetter periods, and negative PHDI values indicate dryer periods. Vertical dashed lines represent transitions between recent wet and dry periods. Specific conductance data were provided by the U.S. Fish and Wildlife Service.

1980 1998 1975 1985 10 1,500

1,000 5

500

0 0

-5

PHDI Deliveries SOUTH CANAL, IN HECTARE METERS

-10 DELIVERIES FROM MILK RIVER VIA DODSON 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 PALMER HYDROLOGICAL DROUGHT INDEX (PHDI) YEAR

Figure 6. Palmer Hydrological Drought Index (PHDI) from 1895 to 2007 and deliveries to Bowdoin Wildlife Refuge from the Milk River via the Dodson South Canal from 1938 to 2007. Positive PHDI values indicate wetter periods, and negative PHDI values indicate dryer periods. Vertical dashed lines represent transitions between recent wet and dry periods. Delivery data were provided by the U.S. Fish and Wildlife Service. Primer on Salinity Concepts 11

1980 1998 1975 1985 10 30,000 A

20,000 5 10,000

0 0

-5 IN HECTARE METERS

PHDI MONTHLY BEAVER CREEK FLOW, Flow -10 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 PALMER HYDROLOGICAL DROUGHT INDEX (PHDI) YEAR 10 B 20,000

5 10,000

0 0

-5 IN CUBIC FEET PER SECOND

PHDI ANNUAL BEAVER CREEK PEAK FLOW, Peak flow -10 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 PALMER HYDROLOGICAL DROUGHT INDEX (PHDI) YEAR

Figure 7. Palmer Hydrological Drought Index (PHDI) from 1895 to 2007 and Beaver Creek flows near Bowdoin National Wildlife Refuge from 1950 to 2007 (monthly) and from 1982 to 2007 (peak). A, Monthly flow in hectare meters (data obtained from Parrett, 2005; the streamflow data represent recorded and estimated values). B, Annual peak flow in cubic feet per second (data obtained from the U.S. Geological Survey stream gage 06166000). Vertical dashed lines represent transitions between recent wet and dry periods. 12 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1,400 A All reservoirs 1,200 Stock Wildlife 1,000 Irrigation

800

600

400

200 CUMULATIVE NUMBER OF RESERVOIRS 0 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000

YEAR 4,000 B All reservoirs Stock Irrigation 3,000 Wildlife

2,000

1,000

0 CUMULATIVE CAPACITY, IN HECTARE METERS 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000

YEAR

Figure 8. Data on reservoirs in the Beaver Creek watershed created between 1898 and 2004 for watering stock, providing wildlife habitat, and irrigation supply. A, Cumulative number of reservoirs. B, Water-storage capacity, in hectare meters. Data were provided by Jana Mohrman, Hydrologist, U.S. Fish and Wildlife Service. Tolerances and Adaptations of Biota to Salinity 13

Tolerances and Adaptations of Biota to The ability of plants to survive saline environments depends on their capacity to tolerate salts throughout various Salinity life cycle stages. Plants capable of growth, flowering, and seed production when rooted in soils with sodium chloride Plants concentrations greater than 0.5 percent are commonly referred to as halophytes (Baskin and Baskin, 1998). In contrast, plants Differences in salinity are often reflected in the species incapable of survival and reproduction in these environments composition of vegetation; thus, salinity has been incorpo- are referred to as glycophytes, or salt-sensitive plants. rated as a modifier in many lake and wetland classification The ability of halophytes to grow and survive in saline systems (Stewart and Kantrud, 1971; Cowardin and others, environments is possible because of various physiological and 1979). Stewart and Kantrud (1972) related the occurrence structural adaptations. Seed germination is affected by many of 135 plant species in prairie pothole wetlands to six salin- abiotic factors such as temperature, photoperiod, soil mois- ity categories (table 2). Of these species, nearly 70 percent ture, and pH; however, salinity is one of the primary factors did not occur in wetlands classified as brackish, subsaline, that limits seed germination in both glycophytes and halo- and saline (salinity levels greater than 5,000 µS cm-1), and phytes. Baskin and Baskin (1998) reported that concentrations about 35 percent did not occur when salinities exceeded of sodium chloride ranging from about 3,000 (0.3 percent, 2,000 µS cm-1 (moderately brackish, fig. 9). Hence, rela- approximately 4,688 µS cm-1) to 100,000 mg L-1 (10 percent, tively few inland wetland plant species can persist in saline approximately 156,250 µS cm-1) were sufficient to reduce environments. germination rates of many halophytes to about 10 percent

NUMBER OF PLANT TAXA 10

Fresh Saline Brackish Subsalin

Slightly brackish Moderately brackish

Figure 9. Number of taxa from Prairie Pothole Region wetlands (data from Stewart and Kantrud, 1972) in each plant community salinity category (see table 2); plant taxa were included only in the highest category in which they were found. For example, if a species was found in fresh, slightly brackish, and moderately brackish sites, it was included only in the moderately brackish category for this graph. 14 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

(table 5). Reduced germination in halophytes is generally saline soils sequester water, making it unavailable to plants. In attributed to salinity-induced osmotic stress that inhibits water addition, as water evaporates from shallow, saline lakes (such uptake and production of compounds (for example, enzymes) as Lake Bowdoin), salts often accumulate as surficial crusts. necessary for germination and embryo development (Baskin This crusting can reduce water infiltration, thereby decreas- and Baskin, 1998). The inhibitory effects also vary by species ing the availability of water to plants growing on the edges of of salt; for example, potassium chloride (KCl) or magnesium lakes. chloride (MgCl2) may affect the germination of a certain plant Effective management of salt concentrations to promote species more than do other salts, such as sodium chloride or the development of vegetation for wildlife habitat will require calcium chloride (CaCl2). Additionally, natural ecosystems an understanding of plant adaptations to salinity, which vary typically contain a mixture of salts (Swanson and others, by life stage. The general overview provided above is intended 1988), and the presence of one salt may enhance or reduce the only to highlight some of the more common adaptations and adverse effects of another (Baskin and Baskin, 1998; Tester life-cycle strategies that should be considered in concert with and Davenport, 2003). other important determinants of plant community composi- Many halophytes exhibit physiological adaptations that tion. For example, the simple plant/salinity relations discussed allow seeds to remain dormant and not germinate during above should be considered in relation to many other factors hypersaline conditions. This trait allows seeds to persist in the that significantly influence establishment, growth, and repro- soil seed bank until environmental conditions are favorable duction. Soil pH, texture, temperature, redox potential, mois- for germination; for example, seeds will not germinate until ture, and water depth all influence plant establishment and climatic events lower salinity through dilution or leaching. growth in wetland environments (for example, Stewart and Ultimately, dormancy release mechanisms help improve Kantrud, 1972; van der Valk and Davis, 1978; van der Valk, survival during seedling development, which is one of the 1981; Fredrickson and Laubhan, 1994; Winter, 2003). most salt-sensitive periods of the halophyte life cycle. After germination and early establishment, salinity can affect halophyte metabolism (for example, decreased rates of Invertebrates photosynthesis and respiration), as well as stimulate anatomical and morphological changes (for example, increased As with that of plants, the composition of aquatic succulence, stomata number and size, cuticle thickness, tylose invertebrate communities can vary greatly in response to (bladder-like growth) development, early onset of lignifica- changes in salinity. In general, relatively fresh aquatic systems tion, and changes in diameter and number of xylem vessels) contain diverse invertebrate communities, whereas highly (Poljakoff-Mayber, 1975). These anatomical adaptations allow saline systems tend to be dominated by relatively few taxa. halophytes to tolerate osmotic gradients associated with saline It is important to note, however, that even though species environments. richness declines in highly saline systems, biomass often Mature plants absorb salts from the soil solution through does not (for example, Euliss and others, 1991). Common their root systems and lose water to the atmosphere via invertebrates of highly saline systems include copepods and transpiration. These processes result in the accumulation of rotifers (particularly some species of the genera Brachionus salts in plant tissues and often lead to high salt concentra- and Hexarthra), as well as a small number of insects from tions and death in nonhalophytes. In contrast, halophytes have the orders Diptera (typically from the families Chironomi- evolved various mechanisms to survive saline environments. dae and Culicidae [genus Aedes]) and Coleoptera (Pennak, Generally, two principal types of halophytes are recognized: 1989). Shifts in invertebrate community structure that result those that tolerate excess salts and those that resist excess from increased salinities in prairie wetlands typically include salts (Ungar, 1974; Baskin and Baskin, 1998). Salt-tolerant displacement of amphipods (for example, Hyalella) and halophytes either withstand high intracellular salt concentra- anostracans (for example, Branchinecta) by more salt-tolerant tions (salt-enduring) or secrete excess salts (salt-excluding) by brine shrimp (Artemia). In addition, a narrow range of insect means of salt glands, by accumulating salts in special hairs, taxa such as brine flies (Ephydridae) and salt-tolerant water or by transporting salts accumulated in plant structures back boatmen (Corrixidae) have been found to dominate systems to the roots. In contrast, salt-resistant halophytes either do not with high salt concentrations (Swanson and others, 1988; absorb salts from the soil solution or do not transport salts to Euliss and others, 1999). For example, the insect community sensitive organs or leaves. of hypersaline (up to 300 mS cm-1) ponds in California was High salt concentrations in the soil can also affect plant dominated by 1 corixid (Trichocorixa reticulate) and 1 midge growth by reducing the water potential gradient. For water to (Tanypus grodhausi) (Euliss and others, 1991), whereas move from the soil to plant tissues, the water potential of the 1 midge species dominated the benthic community of an plant must be lower (that is, more negative) than that of the alkaline lake in central Washington where salinity levels were soil. The water potential of the soil is related to the concen- about 17,000 mg L-1 (approximately 26,563 µS cm-1). Species tration of dissolved solids in the soil-water solution; a high richness in this lake increased two to eight times as salinity concentration of salts in the soil reduces the soil water poten- levels were reduced to about 1,500 mg L-1 (approximately tial and decreases the water potential gradient. Essentially, 2,344 µS cm-1), with the highest number of species occurring Table 5. Sodium chloride concentration at which seed germination of halophytic plant species was reduced from 75–100 percent to approximately 10 percent.

[Modified from Baskin and Baskin, 1998. NaCl, sodium chloride; mg -1L , milligrams per liter]

NaCl concentration NaCl concentration Genus Species Genus Species Molarity mg L-1 Molarity mg L-1 Melilotus indica 0.05 2,922.0 Sporobolus virginicus 0.26–>0.60 15,194.4–>35,064.0 Salicornia bigelovii 0.05–0.06 2,922.0–3,506.4 Suaeda nudiflora 0.26 15,194.4 Sarcobatus vermiculatus 0.06–0.15 3,506.4–8,766.0 Aster tripolium 0.34–0.60 19,869.6–35,064.0 Zygophyllum qatarense 0.07 4,090.8 Atriplex halimus 0.34 19,869.6 Distichlis spicata 0.09 5,259.6 Atriplex patula 0.34 19,869.6 Glaux maritima 0.09 5,259.6 Cotula coronopifolia 0.34 19,869.6 Plagianthus divaricatus 0.09 5,259.6 Mesembryanthemum australe 0.34 19,869.6 Scirpus americanus 0.09 5,259.6 Phragmites communis 0.34 19,869.6 Scirpus olneyi 0.09 5,259.6 Polypogon monspeliensis 0.34 19,869.6 Selliera radicans 0.09 5,259.6 Prosopis farcta 0.34–0.6 19,869.6–35,064.0 Spergularia salina 0.09 5,259.6 Salicornia patula 0.34 19,869.6 Triglochin striatum 0.09–0.17 5,259.6–9,934.8 Atriplex griffithii 0.35 20,454.0 Iva annua 0.13–0.17 7,597.2–9,934.8 Cochlearia danica 0.43 25,129.2 Zannichellia pedunculata 0.13 7,597.2 Rumex crispus 0.43 25,129.2 Ceratoides lanata 0.17–0.34 9,934.8–19,869.6 Puccinellia distans 0.45 26,298.0 Hordeum jubatum 0.17–0.31 9,934.8–18,116.4 Puccinellia lemmoni 0.45 26,298.0 Juncus maritimus 0.17 9,934.8 Halopeplis amplexicaulis 0.50 29,220.0 Tolerances and Adaptations of Biota to Salinity Melaleuca ericifolia 0.17 9,934.8 Atriplex triangularis 0.51 29,804.4 Myrica cerifera 0.17 9,934.8 Limonium bellidifolium 0.60 35,064.0 Schoenus nitens 0.17 9,934.8 Limonium vulgare 0.60 35,064.0 Scirpus robustus 10.17–0.20 19,934.8–11,688.0 Salsola kali 0.60 35,064.0 Spergularia marina 0.17–0.34 9,934.8–19,869.6 Salicornia pacifica 0.68 39,739.2 Suaeda linearis 0.17 9,934.8 Spartina alterniflora 0.68 39,739.2 Melilotus segetalis 0.20 11,688.0 Puccinellia festucaeformis 0.75 43,830.0 Melilotus messanensis >0.20 >11,688.0 Cressa cretica 0.85 49,674.0 Sesuvium portulacastrum 0.2–>0.60 11,688.0–>35,064.0 Salicornia europaea 0.85 49,674.0 Atriplex canescens 0.21 12,272.4 Suaeda depressa 0.85 49,674.0 Salicornia brachystachya 10.24 114,025.6 Tamarix pentandra 10.85 149,674.0 Salicornia dolichostachya 10.24 114,025.6–14,025.6 Suaeda japonica 0.90 52,596.0 Atriplex prostrata 0.26 15,194.4 Kochia americana 1.02 59,608.8 Atriplex polycarpa 0.26 15,194.4 Zygophyllum dumosa 1.51 88,244.4

Plantago coronopus 0.26 15,194.4 Salicornia herbacea 1.70 99,348.0

Salicornia emerici 0.26 15,194.4 1 5 1Highest concentration of NaCl tested; seeds could germinate at higher concentrations. 16 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. at levels ranging from 2,000 to 3,000 mg L-1 (approximately Database Summary Statistics 3,125 to 4,688 µS cm-1) (Wiederholm, 1980). A majority of aquatic invertebrates are able to tolerate a relatively large range of salt concentrations, but few Plants species can survive in highly saline waters. Most invertebrate The plant database (http://pubs.usgs.gov/sir/2009/5098/ species are unable to maintain their internal salt and water downloads/databases_21april2009.xls) contains information balance when placed in a solution that contains greater than on 411 taxa obtained from 17 studies conducted in 9 States 0.9 percent (9,000 mg L-1 or approximately 14,063 µS cm-1) and 3 Canadian Provinces, as well as literature that did not salts, or roughly 25 percent the salt content of seawater specify a location (apps. 4 and 5). Figure 10 is a graphic (Pennak, 1989; Hart and others, 1991). Only a narrow assem- blage of euryhaline organisms (for example, inhabitants of portrayal of the compiled data by the number of plant taxa estuaries and tide pools) can thrive in both freshwater and in relation to source (fig. 10A), location (fig. 10B), wetland marine environments. In general, habitats that support a large classification (fig. C10 ), wetland indicator category (fig. 10D), number of common freshwater fauna have concentrations of plant physiognomy/life-span (fig. 10E), and salinity categories TDS that range from about 10 to 1,000 mg L-1 (approximately (fig. 10F). We used this database to compute the minimum and 16 to 1,563 µS cm-1) (Pennak, 1989; Hart and others, 1991). maximum reported specific conductance and pH by substrate Salinity levels in inland lakes and wetlands, however, can vary (soil, water) for each plant taxon (app. 5). considerably, and the few species that can tolerate extremely high salt concentrations require highly specialized and effi- Invertebrates cient osmoregulatory systems to maintain a proper internal balance of salt and water. Since water will tend to move into The invertebrate database (http://pubs.usgs.gov/ the hypertonic (high salt) tissues of freshwater invertebrates sir/2009/5098/downloads/databases_21april2009.xls) contains through permeable surfaces such as epithelia, cuticle, chitin, 330 taxa obtained from 7 sources (apps. 4 and 6) that represent and gills (Pennak, 1989), these species have developed various studies from 3 States, 2 Canadian Provinces, and Australia. adaptations such as contractile vacuoles, flame bulb systems, Figure 11 provides a summary of the number of invertebrate nephridia, and various glandular structures that are capable taxa by source (fig. 11A), location (fig. 11B), wetland clas- of forming highly dilute urine (hypotonic to body fluids) that sification (fig. 11C), taxonomic classification (fig. 11D), and is excreted in great quantities. As a result, concentrations of salinity maxima (fig. 11E). Appendix 6 contains the minimum internal salts and water remain somewhat constant. and maximum specific conductance and pH for invertebrate Similar to that of plants, the salinity tolerance of aquatic taxa included in the database. invertebrates often varies by life stage, with immature organisms often exhibiting less tolerance to salts than do adults (Euliss and others, 1999; Brock and others, 2005; Pinder and Considerations in Using Information from the others, 2005). Many species of Coleoptera and Hemiptera Plant and Invertebrate Databases are able to exist in prairie wetlands in spite of highly variable salinities because their adult stages are capable of flying to The plant and invertebrate databases represent infor- suitable habitats. Conversely, many nonflying aquatic inver- mation obtained from an extensive search of the scientific tebrates and immature stages of flying insects must rely on literature and unpublished data; however, it is not consid- various adaptations that allow them to endure highly saline ered comprehensive. Data are derived primarily from field habitats, including burrowing into substrates or produc- studies conducted in natural wetland ecosystems rather than ing ephippia (resting eggs), cysts, or waterproof secretions from experimentally derived tolerance values. Users should (Euliss and others, 1999). For example, cladocerans such consider the following aspects and limitations when creating as Daphnia sp. produce ephippia that remain viable in the summaries or conducting analyses: substrate until conditions are favorable for hatching. Because 1. The data represent information from numerous geo- of the high mobility of some adult invertebrates and the inabil- graphic locations (for example, Iowa, North Dakota, ity of many immature aquatic invertebrates to cope with high Prairie Pothole Region, Manitoba) and wetland types (for salinities, the presence of adults in highly saline wetlands does example, palustrine, lacustrine, salt-marsh); users may not necessarily indicate the presence of a persistent population want to focus on, or exclude data from, a certain region or (Pinder and others, 2005). wetland type. Database Summary Statistics 17

350 300 A B 300 250

250 200

200 150 150

100 100 NUMBER OF PLANT TAXA NUMBER OF PLANT TAXA 50 50

0 0 a Ohio Iowa Alberta Kansas VariousMontana Manitoba Nebraska Minnesota Ungar, 1970 Ungar, 1967Ungar, 1966Ungar, 1974 South DakotNorth Dakota Kantrud,Kantrud, 1990Kantrud, 1991 1996 Johnson, 1990 Saskatchewan Kansas; Oklahoma Ungar and Riehl, 1980 USGS, unpublished a Prairie Pothole Region Ungar and others,Ungar 1979 and others, 1969 Kantrud and others, 1989 Bowdoin NWR, Montana Rawson and Moore, 1944 Stewart and Kantrud, 1972 LOCATION OF STUDY Grosshans and Kenkel, 1997 Hammer and Heseltine, 1988 SOURCE OF DATA 350 140 C D 300 120

250 100

200 80

150 60

100 40 NUMBER OF PLANT TAXA NUMBER OF PLANT TAXA 50 20

0 0

N/A UPL CELL PEM FAC- FAC OBL PEMB PEMA PEMC PEMF FACU- FACU FACW FACW- FACW+ Unknown Palustrine Lacustrine Salt-marsh WETLAND INDICATOR CATEGORY WETLAND CLASSIFICATION 200 160 E F Water 180 140 Soil 160 120 140 100 120

100 80

80 60 60 40 NUMBER OF PLANT TAXA NUMBER OF PLANT TAXA 40 20 20

0 0 e

Tree Fresh H-Vine B-Forb Shrub A-Forb P-Forb Saline A-Grass P-Sedge P-Grass A-Sedge Brackish Subsalin Cryptogam PLANT PHYSIOGNOMY/LIFE-SPAN CATEGORY Slightly brackish ModeratelySALINITY brackish CATEGORY

Figure 10. Number of taxa from the plant database by various categories. A, Data source. B, Location. C, Classification. D, Wetland indicator category. E, Plant physiognomy/life-span category. See appendix 3 for definitions of variables within each category. F, The number of taxa from the plant database in relation to plant community salinity categories presented in table 2. Plant taxa were included in each category on the basis of the maximum reported salinity of soil and water. 18 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

25 25 A B

20 20

15 15

10 10 TAXONOMIC ORDERS TAXONOMIC ORDERS 5 5 NUMBER OF INVERTEBRATE NUMBER OF INVERTEBRATE

0 0

Australia Nebraska North Dakota Johnson, 1990 Saskatchewan British Columbia McCarraher, 1970 Kay and others, 2001 Tangen, unpublished USGS, unpublished b Bowdoin NWR, Montana Rawson and Moore, 1944 Lancaster and Scudder, 1987 SOURCE OF DATA LOCATION OF STUDY 25 180 C D 160 20 140

120 15 100

80 10 60 TAXONOMIC ORDERS 5 40 NUMBER OF INVERTEBRATE

NUMBER OF INVERTEBRATE TAXA 20

0 0

PEMF PEMC Class Order Riverine Phylum Family Genus Lacustrine Species TAXONOMIC CLASSIFICATION WETLAND CLASSIFICATION 20 E

Figure 11. Number of taxonomic orders from the invertebrate 5 database for each category. A, Data source. B, Location. C, Classification. See appendix 3 for definitions of variables within each category. D, Depiction of the number of taxa that NUMBER OF INVERTEBRATE FAMILIES 0 were reported at each level of taxonomic classification. E, The e number of taxa from the invertebrate database in relation to plant Fresh Saline Brackish community salinity categories presented in table 2. Invertebrate Subsalin taxa were included in each category on the basis of the maximum Slightly brackish Moderately brackish reported salinity of water. SALINITY MAXIMA Application to Bowdoin National Wildlife Refuge 19

2. To avoid misinterpretation, users should always consider fluctuations. Therefore, salinity or other measurements the database variables DATA_TYPE and CONDUCTIV- recorded on a given day or year may not be representative ITY_CODE. DATA_TYPE defines whether the water- of the site during other time periods. Consequently, the quality value (for example, specific conductance) was occurrence of plant and invertebrate taxa recorded at a site reported as a single measurement or as a summarized may be influenced by factors, or values of factors, other value such as a mean, median, minimum, or maximum. than those occurring at the time of sampling. For consistency, salinity units were converted to micro- 7. For both plants and invertebrates, tolerance to salinity can siemens in the databases. The variable CONDUCTIV- vary significantly depending on life stage. Mature plants ITY_CODE identifies whether the original study reported may be able to survive in saline conditions as long as the the value as microsiemens, or if it was converted (see proper conditions existed during the period of germina- Methods, above). tion, which is often earlier in the year when snowmelt and 3. For consistency, taxonomic and common names for plants spring rains dilute waters and result in lower salinities. were based on The Great Plains Flora Association (1986) Similarly, presence of mobile adult invertebrates does not when possible. Therefore, the variables GENUS and SPE- necessarily indicate the presence of a persistent popula- CIES do not necessarily represent the scientific names tion. reported by the original source; the variables GENUS_R and SPECIES_R present the scientific name reported by the original study. Application to Bowdoin National 4. Users of the invertebrate database must first determine which taxonomic classification is most appropriate since Wildlife Refuge individual studies report invertebrate occurrences at varying degrees of resolution (for example, Order, Family, Plants Genus, Species). Resolution will be lost when using a higher taxonomic classification such as Order, whereas There are about 210 plant species documented on using a lower taxonomic classification such as Genus may Bowdoin NWR according to a vegetation survey conducted result in the loss of some information on occurrence. in the late 1980s and early 1990s (app. 1). Of these plants, 5. Certain taxa in both databases may have only been the database compiled for this report includes information reported in a single or small number of studies. There- for 110 species that was obtained from 13 known locations fore, salinity values for specific taxa may be limited by distributed among 9 types of aquatic systems, 8 wetland a paucity of data. Additionally, many studies that report indicator categories, and 11 plant physiognomy/life-span salinity values are conducted in highly saline systems and categories (table 6). On the basis of this information, at least focus only on salt-tolerant taxa; therefore, many common some taxa in each plant physiognomy and wetland indicator freshwater taxa may not be well represented. category can tolerate fresh to moderately brackish habitats 6. It is important to consider that many abiotic factors that (fig. 12; based on 10th and 90th percentile ranges); however, control wetland plant and invertebrate community compo- most taxa within each category (based on mean and median as sition are highly dynamic and exhibit seasonal and annual measures of central tendency) tolerate only slightly brackish

Table 6. Summary information from the plant database (210 total species) for 110 species documented on Bowdoin National Wildlife Refuge (app. 1).

[Data were obtained from 13 known locations and represent 9 types of aquatic systems (classification), 8 wetland indicator categories, and 1 plant physiognomy/life-span categories]

Location Alberta, Iowa, Kansas, Manitoba, Minnesota, Montana, Nebraska, North Dakota, Ohio, Oklahoma, Prairie Pothole Region, Saskatchewan, South Dakota, “unknown” Classification1 CELL, LACUSTRINE, PALUSTRINE, PEM, PEMA, PEMB, PEMC, PEMF, SALT-MARSH, “unknown” Wetland indicator category2 FAC (10), FAC- (1), FACU (26), FACU- (2), FACW (9), FACW+ (1), OBL (30), (number of taxa) UPL (24), n/a (7) Plant physiognomy/life-span3 category A-forb (19), A-grass (6), A/P-forb (3), B-forb (1), cryptogam (2), H-vine (1), (number of taxa) P-forb (45), P-grass (17), P-sedge (7), shrub (5), tree (4) 1See variable “CLASSIFICATION” in app. 3 for definitions. 2See variable “WIC” in app. 3 for definitions. 3See variable “PHYSIOGNOMY” in app. 3 for definitions. 20 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

Slightly Moderately Subsaline Fresh brackish brackish Brackish Saline

A Tree

Shrub

P-sedge

P-grass

A-grass

P-forb

A/P-forb

A-forb

Cryptogam

PLANT PHYSIOGNOMY/LIFE-SPAN CATEGORY 10 100 1,000 10,000 100,000 SPECIFIC CONDUCTANCE OF WATER, IN MICROSIEMENS PER CENTIMETER Slightly Moderately Subsaline Fresh brackish brackish Brackish Saline

H-vine B Tree Shrub P-sedge P-grass A-grass P-forb B-forb A/P-forb A-forb Cryptogam

PLANT PHYSIOGNOMY/LIFE-SPAN CATEGORY 10 100 1,000 10,000 100,000 SPECIFIC CONDUCTANCE OF SOIL, IN MICROSIEMENS PER CENTIMETER

Slightly Moderately Subsaline Fresh brackish brackish Brackish Saline

C UPL

FACU-

FACU

FAC

FACW

FACW+

WETLAND INDICATOR CATEGORY OBL

10 100 1,000 10,000 100,000 SPECIFIC CONDUCTANCE OF WATER, IN MICROSIEMENS PER CENTIMETER

Slightly Moderately Subsaline Fresh brackish brackish Brackish Saline

D UPL

FACU-

FACU

FAC

FACW

FACW+

WETLAND INDICATOR CATEGORY OBL

10 100 1,000 10,000 100,000 SPECIFIC CONDUCTANCE OF SOIL, IN MICROSIEMENS PER CENTIMETER

Figure 12. Specific conductance of water and soil for each plant physiognomy/life-span category and each wetland indicator category (see variables “PHYSIOGNOMY” and “WIC” in app. 3). A, Specific conductance of water for each plant physiognomy/life-span category. B, Specific conductance of soil for each plant physiognomy/life-span category. C, Specific conductance of water for each wetland indicator category. D, Specific conductance of soil for each wetland indicator category. The data represent taxa identified at Bowdoin National Wildlife Refuge (app. 1) that are found in the plant database. The box plots show the median (solid line); mean (dotted line); 10th, 25th, 75th, and 90th percentiles (horizontal boxes with error bars); and outliers (dots). Vertical dashed lines delineate boundaries for plant community salinity categories (table 2).—Continued 22 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. conditions (figs. 13, 14). Most of the dominant plant species have the potential to occur in wetlands with varying ranges of found on the refuge (see Description of Bowdoin National salinity that should facilitate evaluation of various manage- Wildlife Refuge) exhibited the greatest tolerance to salinity. ment practices and strategies aimed at enhancing wildlife Examples include hardstem bulrush (Scirpus acutus), alkali habitat. As an example, wetlands on Bowdoin NWR are bulrush (S. paludosus), common threesquare (S. americanus), characterized by elevated salinity levels that have been saltgrass (Distichlis spicata), pickleweed (Salicornia rubra), exacerbated by changes in land use and past management sago pondweed (Potomogeton pectinatus), and widgeon- activities that enhance water inputs into a closed basin. The grass (Ruppia maritima). Few of these species, however, high salinity levels have had a negative impact on the ability appear to tolerate salinities above brackish (approximately of wetlands to support waterfowl, which is one of the primary 15,000 µS cm-1). objectives of the refuge. Refuge staff can use the information presented in this report to evaluate various management strategies designed to remove salts from wetlands and create Invertebrates environmental conditions that favor establishment of plant and invertebrate communities that are highly attractive to Most of the taxonomic orders and families of aquatic waterfowl. In addition, refuge staff can use this informa- invertebrates known to occur on the refuge (app. 2) are repre- tion to help establish specific (for example, obtain a specific sented in the database (figs. 15, 16). Examination of inverte- range of salinity levels) rather than general (for example, brates at the taxonomic level of family (fig. 16) indicates that reduce salinity levels) management goals. A primary focus of most families can occur in fresh to brackish habitats; however, this report was to provide information to address the salinity the greatest diversity tends to occur within slightly to moder- problem at Bowdoin NWR, but information in the database, ately brackish habitats. Similar to that of plants, information as well as concepts and information discussed, is intended to on invertebrates compiled in the database suggests that few be applicable to other areas or refuges. Ideally, information in taxa appear capable of tolerating salinities above brackish the database will be augmented with additional field studies by (approximately 15,000 µS cm-1). resource managers (for example, salinity monitoring, vegetation inventories) to facilitate a better understanding of the ecological relations between salinity and flora and fauna when Summary developing management strategies.

We reviewed and summarized relevant literature to assess the relation between salinity and composition of biotic Acknowledgments communities associated with freshwater ecosystems. This is not a comprehensive literature review and should not be Funding for this project was provided by the U.S. Fish considered exhaustive. Further, we do not report experimen- and Wildlife Service and the U.S. Geological Survey. We tally derived threshold values (for example, minimum or thank Charles Dahl, Betty Euliss, Rachel Laubhan, Dave maximum salt tolerance) for individual plant or invertebrate Mushet, and Kathy Tribby for assisting in the literature search species; instead, we present data obtained from field surveys and providing data and technical assistance, and Suzanne that describe species occurrences across naturally fluctuat- Roberts for the cover art on this report. Additionally, we thank ing ranges of salinity and other environmental factors (for the following people for comments on previous versions of example, salt composition, temperature, pH). This approach this report: Mike Artmann, Wayne King, Jana Mohrman, Rick allows for the prediction of broad species assemblages that Sojda, Amy Symstad, and Kathy Tribby. Acknowledgments 23

Slightly Moderately Subsaline Fresh brackish brackish Brackish Saline

Agrostis stolonifera Sagittaria latifolia Elodea canadensis Ammannia coccinea Ranunculus macounii Callitriche hermaphroditica Carex rostrata Lysimachia ciliata Phalaris arundinacea Potamogeton friesii Potamogeton richardsonii Polygonum lapathifolium Mentha arvensis Echinochloa crusgalli Ceratophyllum demersum Eleocharis acicularis Sagittaria cuneata Potamogeton pusillus Potamogeton filiformis Myriophyllum exalbescens Utricularia vulgaris Beckmannia syzigachne Alisma subcordatum Typha latifolia Rumex maritimus Heliotropium curassavicum Scirpus acutus Eleocharis macrostachya Salicornia rubra Potentilla anserina Juncus balticus Hordeum jubatum Triglochin maritima Scirpus americanus Distichlis spicata var. stricta Potamogeton pectinatus Scirpus paludosus Ruppia maritima

10 100 1,000 10,000 100,000 SPECIFIC CONDUCTANCE OF WATER, IN MICROSIEMENS PER CENTIMETER

Figure 13. Specific conductance of water for common plant species at Bowdoin National Wildlife Refuge that are associated with wetlands (FACW, FACW+, OBL; see variable “WIC” in app. 3). The box plots show the median (solid line); mean (dotted line); 10th, 25th, 75th, and 90th percentiles (horizontal boxes with error bars); and outliers (dots). Vertical dashed lines delineate boundaries for plant community salinity categories (table 2). 24 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

Slightly Moderately Subsaline Fresh brackish brackish Brackish Saline

Potamogeton richardsonii Sagittaria latifolia Sagittaria cuneata Ranunculus macounii Eleocharis acicularis Mentha arvensis Polygonum lapathifolium Solidago gigantea Lysimachia ciliata Potamogeton pusillus Phalaris arundinacea Ceratophyllum demersum Utricularia vulgaris Rumex maritimus Beckmannia syzigachne Alisma subcordatum Potentilla anserina Myriophyllum exalbescens Typha latifolia Scirpus acutus Echinochloa crusgalli Agrostis stolonifera Scirpus americanus Potamogeton pectinatus Hordeum jubatum Juncus balticus Triglochin maritima Distichlis spicata var. stricta Scirpus paludosus Salicornia rubra

100 1,000 10,000 100,000 SPECIFIC CONDUCTANCE OF SOIL, IN MICROSIEMENS PER CENTIMETER

Figure 14. Specific conductance of soil for common plant species at Bowdoin National Wildlife Refuge that are associated with wetlands (FACW, FACW+, OBL; see variable “WIC” in app. 3). The box plots show the median (solid line); mean (dotted line); 10th, 25th, 75th, and 90th percentiles (horizontal boxes with error bars); and outliers (dots). Vertical dashed lines delineate boundaries for plant community salinity categories (table 2). Acknowledgments 25

Slightly Moderately Fresh brackish brackish Brackish Subsaline Saline

Gnathobdellida Conchostraca Pelecypoda Pharyngobdellida Limnophila Rhynchobdellida Collembola Ephemeroptera Decapoda Hemiptera Odonata Amphipoda Tricoptera Cladocera Anostraca Eucopepoda Coleoptera Diptera

100 1,000 10,000 100,000

SPECIFIC CONDUCTANCE OF WATER, IN MICROSIEMENS PER CENTIMETER

Figure 15. Specific conductance of water for representative taxa in the invertebrate database that could occur at Bowdoin National Wildlife Refuge. The box plots show the median (solid line); mean (dotted line); 10th, 25th, 75th, and 90th percentiles as horizontal boxes with error bars; and outliers (dots). Vertical dashed lines delineate boundaries for plant community salinity categories (table 2). 26 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

Slightly Moderately Fresh brackish brackish Brackish Subsaline Saline

Gammaridae Amphipoda Talitridae Artemiidae Branchinectidae Chirocephalidae Anostraca Bosminidae Cladocera Chydoridae Daphnidae Leptodoridae Moinidae Polyphemidae Sididae Curculionidae Coleoptera Dytiscidae Gyrinidae Haliplidae Helodidae Hydrophilidae Staphylinidae Ceratopogonidae Diptera Chaoboridae Chironomidae Corethrinae Culicidae Dixidae Dolichopodidae Ephydridae Psychodidae Sciomyzidae Stratiomyidae Syrphidae Tabanidae Tipulidae Belostomatidae Hemiptera Corixidae Gerridae Hebridae Mesovelidae Notonectidae Pleidae Velidae Aeshnidae Odonata Coenagrionidae Lestidae Libellulidae

100 1,000 10,000 100,000 SPECIFIC CONDUCTANCE OF WATER, IN MICROSIEMENS PER CENTIMETER Figure 16. Specific conductance of water for invertebrate families from representative taxonomic orders in the invertebrate database. The box plots show the median (solid line); mean (dotted line); 10th, 25th, 75th, and 90th percentiles (horizontal boxes with error bars); and outliers (dots). Vertical dashed lines delineate boundaries for plant community salinity categories (table 2). References 27

References Hamilton, D.B., Roelle, J.E., and Schafer, W.M., 1989, A simulation model of water and salt balance at Bowdoin National Wildlife Refuge: Fort Collins, Colo., U.S. Fish Baskin, C.C., and Baskin, J.M., 1998, Seeds—ecology, bioge- and Wildlife Service, National Ecology Research Center, ography, and evolution of dormancy and germination: San NERC-89/08, 60 p. Diego, Calif., Academic Press, 666 p. Hammer, U.T., and Heseltine, J.M., 1988, Aquatic macro- Bauder, J.W., Hershberger, K., and Sessom, H., 2007, Bow- phytes in saline lakes of the Canadian prairies: Hydrobiolo- doin National Wildlife Refuge salt mitigation impact gia, v. 158, p. 101–116. review: Bozeman, Montana State University, Department of Land Resources and Environmental Sciences, Open-file Hart, B.T., Bailey, P., Edwards, R., Hortle, K., James, K., report to Montana Reserve Water Rights Compact Commis- McMahon, A., Meredith, C., and Swadling, K., 1991, A sion, Technical Advisory Team, April 2007, 81 p. review of the salt sensitivity of the Australian freshwater biota: Hydrobiologia, v. 210, p. 105–144. Brock, M.A., Nielsen, D.L., and Crossle, K., 2005, Changes in biotic communities developing from freshwater wetland Havlin, J.L., Beaton, J.D., Tisdale, S.L., and Nelson, W.L., sediments under experimental salinity and water regimes: eds., 1999, Soil fertility and fertilizers (6th ed.): Upper Freshwater Biology, v. 50, p. 1376–1390. Saddle River, N.J., Prentice Hall, 515 p.

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Euliss, N.H., Jr., Jarvis, R.L., and Gilmer, D.S., 1991, Stand- Kantrud, H.A., 1991, Widgeongrass (Ruppia maritima L.)—a ing crops and ecology of aquatic invertebrates in agri- literature review: Washington, D.C., U.S. Fish and Wildlife cultural drainwater ponds in California: Wetlands, v. 11, Service, Fish and Wildlife Research 10, 58 p. p. 179–190. Kantrud, H.A., 1996, The alkali (Scirpus maritimus L.) and Euliss, N.H., Jr., Wrubleski, D.A., and Mushet, D.M., 1999, saltmarsh (S. robustus Pursh) bulrushes—a literature Wetlands of the Prairie Pothole Region—invertebrate spe- review: Washington, D.C., National Biological Service cies composition, ecology, and management, in Batzer, Information and Technology Report 6, 77 p. D.P., Rader, R.B., and Wissinger, S.A., eds., Invertebrates Kantrud, H.A., Millar, J.B., and van der Valk, A.G., 1989, in freshwater wetlands of North America—ecology and Vegetation of wetlands of the prairie pothole region, in van management: New York, N.Y., John Wiley & Sons, Inc., p. 471–514. der Valk, A., ed., Northern prairie wetlands: Ames, Iowa State University Press, p. 132–187. Fredrickson, L.H., and Laubhan, M.K., 1994, Managing wetlands for wildlife, in Bookhout, T.A., ed., Research and Kay, W.R., Halse, S.A., Scanlon, M.D., and Smith, M.J., 2001, management techniques for wildlife and habitats: Bethesda, Distribution and environmental tolerances of aquatic macro- Md., The Wildlife Society, p. 623–647. invertebrate families in the agricultural zone of southwestern Australia: Journal of the North American Benthological Grosshans, R.E., and Kenkel, N.C., 1997, Dynamics of Society, v. 20, p. 182–199. emergent vegetation along natural gradients of water depth and salinity in a prairie marsh—delayed influences of Kendy, E., 1999, Simulation of water and salt budgets and competition, in Goldsborough, L.G., and Hann, B.J., eds., effects of proposed management strategies for Bowdoin University of Manitoba Field Station (Delta Marsh) thirty- National Wildlife Refuge, northeastern Montana: Helena, second annual report: Portage la Prairie, Manitoba, Canada, Mont., U.S. Geological Survey Water-Resources Investiga- p. 83–93. tions Report 98–4260, 86 p. 28 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

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Ungar, I.A., Benner, D.K., and McGraw, D.C., 1979, The distribution and growth of Salicornia Europaea on an inland salt pan: Ecology, v. 60, p. 329–336. Ungar, I.A., Hogan, W., and McClelland, M., 1969, Plant communities of saline soils at Lincoln, Nebraska: The American Midland Naturalist, v. 82, p. 564–577. Ungar, I.A., and Riehl, T.E., 1980, The effect of seed reserves on species composition in zonal halophyte communities: Botanical Gazette, v. 141, p. 447–452. U.S. Department of Agriculture, 2004, Soil survey of Phil- lips County area, Montana, part I: Washington, D.C., U.S. Department of Agriculture, Natural Resources Conservation Service, 226 p. van der Valk, A.G., 1981, Succession in wetlands—a Gleaso- nian approach: Ecology, v. 62, p. 688–696. van der Valk, A.G., and Davis, C.B., 1978, The role of seed banks in the vegetation dynamics of prairie glacial marshes: Ecology, v. 59, p. 322–335. Weydemeyer, W., and Marsh, V.L., 1936, The bird life of Lake Bowdoin, Montana: The Condor, v. 38, p. 185–198. Wiederholm, T., 1980, Effects of dilution on the benthos of an alkaline lake: Hydrobiologia, v. 68, p. 199–207. Winter, T.C., ed., 2003, Hydrological, chemical, and biological characteristics of a prairie pothole wetland complex under highly variable climate conditions—the Cottonwood Lake area, east-central North Dakota: Washington, D.C., U.S. Geological Survey Professional Paper 2003–1675, 109 p. Woods, A.J., Omernik, J., Nesser, M., Shelden, J.A., Com- stock, J., Azevedo, J.A., and Sandra, H., 2002, Ecoregions of Montana (2d ed.), (color poster with map, descriptive text, summary tables, and photographs): Corvallis, Oreg., U.S. Environmental Protection Agency, Western Ecology Division, map scale 1:1,500,000. 30 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

Appendix 1. Plant Taxa Identified at Bowdoin National Wildlife Refuge

Plant taxa (genus and species; common name) identified at Bowdoin NationalWildlife Refuge during a preliminary vegetation survey conducted from 1987 to 1993 by researchers from the U.S. Fish and Wildlife Service, U.S. Army Corps of Engi- neers, and the Southeastern Wisconsin Regional Planning Commission. Taxa followed by an asterisk (*) are in the plant/salinity database. Nomenclature follows The Great Plains Flora Association (1986).

Acer negundo; box elder Chrysothamnus nauseosus; rabbit brush Achillea millefolium; yarrow* Cirsium arvense; Canada thistle, field thistle* Agropyron cristatum; crested wheatgrass* Cirsium undulatum; wavy-leaf thistle Agropyron repens; quackgrass* Cirsium vulgare; bull thistle* Agropyron smithii; western wheatgrass* Clematis pseudoalpina; virgin’s bower Agropyron spicatum; bluebunch wheatgrass Clematis terniflora; virgin’s bower Agrostis stolonifera; redtop* Cleome serrulata; rocky mountain bee plant Alisma subcordatum; common water plantain* Collomia linearis; collomia Allium textile; onion Convolvulus arvensis; field bindweed* Amaranthus retroflexus; rough pigweed* Dactylis glomerata; orchard grass* Ambrosia artemisiifolia; common ragweed, short ragweed* Distichlis spicata var. stricta; inland saltgrass* Ammannia coccinea; toothcup* Echinacea angustifolia; purple coneflower Anemone patens; pasque flower Echinochloa crusgalli; barnyard grass* Apocynum androsaemifolium; spreading dogbane Elaeagnus angustifolia; Russian olive* Arnica fulgens; arnica Eleocharis acicularis; needle spikerush* Artemisia cana; dwarf sagebrush Eleocharis macrostachya; common spikerush* Artemisia dracunculus; silky wormwood* Elodea canadensis; Canadian waterweed* Artemisia frigida; prairie sagewort* Elodea nuttallii; waterweed Artemisia ludoviciana; white sage* Elymus canadensis; Canada wild rye* Aruncus pubescens; bride’s feathers Elymus glaucus; blue wild rye Asclepias speciosa; showy milkweed* Epilobium angustifolium; willow-herb Asparagus officinalis; asparagus Equisetum arvense; field horsetail* Aster ericoides; white aster* Equisetum fluviatile; water horsetail* Aster floralvinda; wild aster Eragrostis cilianensis; stinkgrass Aster puniceus; swamp aster Erigeron glabellus; fleabane Aster sp.; wild aster* Erigeron sp.; fleabane Astragalus pectinatus; tine-leaved milk-vetch, narrow-leave Eriogonum flavum; yellow wild buckwheat Astragalus racemosus; alkali milk-vetch, creamy poison-vetch Escobaria vivipara; spinystar Avena fatua; wild oats* Euphorbia esula; leafy spurge* Beckmannia syzigachne; sloughgrass* Euphorbia glyptosperma; ridge-seeded spurge Betula papyrifera; paper birch, canoe birch Fraxinus pennsylvanica; green ash* Bouteloua gracilis; blue grama Galium boreale; northern bedstraw* Brassica kaber; charlock* Gaura coccinea; scarlet gaura Bromus inermis; smooth brome* Glycyrrhiza lepidota; wild licorice* Bromus japonicus; Japanese brome* Grindelia squarrosa; curly-top gumweed* Bromus porteri; nodding brome Gutierrizia sarothrae; snakeweed Bromus tectorum; downy brome* Helianthus laetiflorus; sunflower Buchloe dactyloides; buffalo grass Helianthus maximiliani; maximilian sunflower* Callitriche hermaphroditica; northern water starwort* Heliotropium curassavicum; seaside heliotrope* Cardaria draba; hoary cress Hordeum jubatum; foxtail barley* Carex brevior; fescue sedge* Hydrochloa carolinensis; water grass Carex deweyana; sedge Hystrix patula; bottlebrush grass Carex hookerana; sedge Iva axillaris; poverty weed Carex rostrata; beaked sedge* Juncus balticus; baltic rush* Castilleja sulphurea; Indian paintbrush Juncus sp.; rush* Centaurea maculosa; spotted knapweed Juniperus communis; common or dwarf juniper Cerastium arvense; prairie chickweed Kochia scoparia; kochia* Cerastium brachypodum; nodding chickweed* Koeleria cristata; junegrass Ceratophyllum demersum; hornwort, coontail* Lactuca oblongifolia; blue lettuce Cheilanthes feei; lip fern Lactuca serriola; prickly lettuce* Chenopodium sp.; goosefoot, lamb’s quarter* Lappula echinata; blue stickseed Chrysanthemum leucanthemum; ox-eye daisy, marguerite Lappula occidentalis; flatspine stickseed Chrysopsis villosa; golden aster Lappula redowskii; stickseed Appendix 1. Plant Taxa Identified at Bowdoin National Wildlife Refuge 31

Lappula texana; cupseed stickseed Rhus aromatica; fragrant sumac, polecat bush Lepidium perfoliatum; clasping peppergrass Ribes odoratum; buffalo currant Lepidium sp.; peppergrass Rosa arkansana; prairie wild rose* Liatris punctata; dotted blazing star* Rosa woodsii; western wild rose* Linaria vulgaris; butter-and-eggs Rudbeckia sp.; coneflower Lupinius flexuosus; lupine Rumex maritimus; golden dock* Lygodesmia juncea; skeletonweed* Rumex sp.; dock, sorrel* Lysimachia ciliata; fringed loosestrife* Rumex venosus; wild begonia Machaeranthera spinulosus; aster Ruppia maritima; ditchgrass, widgeon grass* Malva moschata; musk mallow Sagittaria cuneata; arrowhead* Malva neglecta; common mallow* Sagittaria graminea; arrowhead Medicago lupulina; black medick* Sagittaria latifolia; common arrowhead* Medicago sativa; alfalfa* Salix exigua subs. interior; sandbar willow* Melilotus officinalis; yellow sweet clover* Salix sp.; willow* Mentha arvensis; field mint* Salsola iberica; russian thistle* Monarda spicata; horse mint, beebalm Sarcobatus vermiculatus; greasewood Myriophyllum exalbescens; American milfoil* Scirpus acutus; hardstem bulrush* Oenothera villosa; common evening primrose Scirpus americanus; chairmaker’s bulrush* Opuntia polyacantha; plains prickly pear Scirpus paludosus; cosmopolitan bulrush* Oxytropis lambertii; purple locoweed Senecio integerrimus; groundsel Oxytropis sericea; white locoweed Setaria sp.; foxtail* Panicum miliaceum; broom-corn millet Setaria viridis; green foxtail Penstemon albidus; white beardtongue Shepherdia argentea; buffaloberry Petalostemum purpurea; purple prairie clover Silene cserei; smooth catchfly Phalaris arundinacea; reed canarygrass* Sisymbrium altissimum; tumbling mustard Phleum pratense; timothy* Solidago gigantea; late goldenrod* Phlox douglasii; phlox Solidago juncea; early goldenrod Pinus ponderosa; ponderosa pine Solidago missouriensis; prairie goldenrod* Plantago lanceolata; English plantain, buckhorn Solidago rigida var. rigida; rigid goldenrod* Plantago major; common plantain* Sonchus arvensis; field sow thistle* Poa pratensis; Kentucky bluegrass* Sparganium emersum; bur-reed Poa sandbergii; Sandberg’s bluegrass* Sphaeralcea coccinea; red false mallow Polanisia dodecandra; clammy-weed Sporobolus airoides; alkali sacaton* Polemonium haydeni; Jacob’s-ladder Sporobolus asper; rough dropseed Polygonum aviculare; knotweed* Stipa comata; needle-and-thread* Polygonum buxiforme; knotweed Stipa spartea; porcupine-grass Polygonum convolvulus; climbing or wild buckwheat* Stipa viridula; green needlegrass* Polygonum lapathifolium; pale smartweed* Symphoricarpos occidentalis; western snowberry* Polygonum persicaria; lady’s thumb Symphoricarpos orbiculatus; coralberry, buckbrush Polygonum ramosissimum; bushy knotweed* Taraxacum officinale; common dandelion* Populus deltoides; cotton-wood* Thlaspi arvense; field pennycress* Potamogeton filiformis; slender pondweed* Tragopogan dubius; goat’s beard, western salsify Potamogeton friesii; Fries’ pondweed* Triglochin maritima; seaside arrowgrass* Potamogeton pectinatus; sago pondweed* Typha latifolia; broad-leaved cat-tail* Potamogeton pusillus; baby pondweed* Utricularia vulgaris; common bladderwort* Potamogeton richardsonii; claspingleaf pondweed* Verbena bracteata; prostrate vervain Potentilla anserina; silverweed* Vicia americana; American vetch* Potentilla argentea; silvery cinquefoil Yucca glauca; soapweed, yucca Potentilla arguta; tall cinquefoil* Zigadenus venenosus; death camass Potentilla rivalis; brook conquefoil* Prunus virginiana; choke cherry Psoralea argophylla; silver-leaf scurf-pea* References Ranunculus longirostric; white water crowfoot Ranunculus macounii; Macoun’s buttercup* The Great Plains Flora Association, 1986, Flora of the Great Ratibida columnifera; prairie coneflower* Plains: Lawrence, University Press of Kansas, 1,402 p. 32 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

Appendix 2. Invertebrate Taxa Identified at Bowdoin National Wildlife Refuge

Invertebrate taxa identified at Bowdoin National Wildlife Refuge by Johnson (1990) and DuBois and others (1992). All taxa are in the invertebrate/salinity database.

Phylum Class Order Family Annelida Hirudinea Annelida Oligochaeta Arthropoda Arachnoidea Hydracarina Arthropoda Branchiopoda Cladocera Arthropoda Copepoda Arthropoda Insecta Coleoptera Arthropoda Insecta Diptera Ceratopogonidae Arthropoda Insecta Diptera Chironomidae Arthropoda Insecta Diptera Culicidae Arthropoda Insecta Diptera Ephydridae Arthropoda Insecta Diptera Stratiomyidae Arthropoda Insecta Diptera Tabanidae Arthropoda Insecta Diptera Arthropoda Insecta Ephemeroptera Baetidae Arthropoda Insecta Ephemeroptera Caenidae Arthropoda Insecta Ephemeroptera Arthropoda Insecta Hemiptera Corixidae Arthropoda Insecta Hemiptera Notonectidae Arthropoda Insecta Hemiptera Arthropoda Insecta Odonata Aeshnidae Arthropoda Insecta Odonata Coenagrionidae Arthropoda Insecta Odonata Libellulidae Arthropoda Insecta Tricoptera Leptoceridae Arthropoda Insecta Tricoptera Phryganeidae Arthropoda Insecta Tricoptera Polycentropodidae Arthropoda Insecta Tricoptera Arthropoda Malacostraca Amphipoda Gammaridae Arthropoda Malacostraca Amphipoda Talitridae Mollusca Gastropoda Limnophila Lymnaeidae Mollusca Gastropoda Limnophila Physidae Mollusca Gastropoda Limnophila Planorbidae Nematoda

References

DuBois, K.L., Palawski, D.U., and Malloy, J.C., 1992, Bowdoin National Wildlife Refuge contaminants biomonitoring study: Helena, Mont., U.S. Fish and Wildlife Service, Contaminant Report R6/207H/92, 53 p. Johnson, K.M., 1990, Aquatic vegetation, salinity, aquatic invertebrates, and duck brood use at Bowdoin National Wildlife Ref- uge, Montana: Bozeman, Montana State University, master’s thesis. Appendix 3. Description of Variables Found in the Plant and Invertebrate Databases 33

Appendix 3. Description of Variables Found in the Plant and Invertebrate Databases

Variable Description and units CHLORIDE Concentration of chlorides, reported as % CLASS Taxonomic Class of invertebrate taxa CLASSIFICATION Wetland classification; in some cases this was provided, and in others the class was assigned on the basis of site descriptions provided by the authors of each data source. • LACUSTRINE = lake; classification of Cowardin and others, 1979 • PALUSTRINE = marsh or wetland; classification of Cowardin and others, 1979 • PEM = palustrine emergent; classification of Cowardin and others, 1979 • PEMA = palustrine emergent, temporarily flooded; classification of Cowardin and others, 1979 • PEMB = palustrine emergent, saturated; classification of Cowardin and others, 1979 • PEMC = palustrine emergent, seasonally flooded; classification of Cowardin and others, 1979 • PEMF = palustrine emergent, semipermanently flooded; classification of Cowardin and others, 1979 • SALT-MARSH = general term for saline wetlands commonly used in scientific literature • CELL = artificial wetland or pond • UNKNOWN = data were presented for multiple wetland classes, or the wetland class could not be determined COMMON_NAME Common name of plant CONDUCTIVITY_CODE This variable identifies conductivity measurements (µS cm-1) that were reported as µS cm-1 or that were converted to µS cm-1 from other units (for example, mg L-1, ppm, %). • CONVERSION = value was converted from other salinity unit. Total dissolved solids and specific conductance were related using the following forumula: mg -1L = µS cm-1 × 0.64 (Tchobanoglous and Burton, 1991). • REPORTED = value was reported as µS cm-1 DOM_SALT Dominant salt identified for the water body FAMILY Taxonomic Family of invertebrate taxa GENUS Genus of the plant or invertebrate • Plant genus terminology following The Great Plains Flora Association (1986) when possible GENUS_R Genus of the plant reported by the original source LOCATION Location of the study from which the data were obtained (for example, State, Province, country, region) MG_L Salinity, reported as milligrams per liter (mg L-1) OPTIMUM_MG_L Salinity reported as optimum for the growth of the plant species, reported as milligrams per liter (mg L-1) ORDER Taxonomic Order of invertebrate taxa PERCENT Salinity (for example, dissolved solids, total salts), reported as % PH pH of the water or soil where a plant species occurred PHYLUM Taxonomic Phylum of invertebrate taxa PHYSIOGNOMY Plant physiognomy (for example, forb/herb, graminoid, shrub, tree, vine) and life span (A, annual; B, biennial; P, perennial) according to The Great Plains Flora Association (1986) and the U.S. Department of Agriculture, National Conservation Service web page, accessed September 2007, at http://plants. nrcs.usda.gov/growth_habits_def.html PPM Salinity, reported as parts per million SODIUM Concentration of sodium, reported as % SOURCE Source of the data 34 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

Variable Description and units SPECIES Species of the plant or invertebrate • Terminology following The Great Plains Flora Association (1986) when possible SPECIES_R Species of the plant reported by the original source SUBSTRATE Substrate from which the variable (for example, salinity, pH) was obtained. • S = soil • W = water SULFATE Concentration of sulfate, reported as % DATA TYPE: Data type; these variables identify whether the measurement (for example, salinity, pH) was reported as • TYPE_CHL (chloride) a mean, median, range, etc. For example, if TYPE_US_CM = ‘MEAN,’ then the value was reported • TYPE_MG_L (mg L-1) as a mean value by the original source; if TYPE_PH = ‘MIN,’ then the value was reported as a mini- • TYPE_PCNT (%) mum value (for example, smallest value of a range) by the original source, etc. • TYPE_PH (pH) • MAXIMUM = reported as maximum value, typically as part of a range (for example, • TYPE_PPM (ppm) 1–10) • TYPE_SOD (sodium) • MEAN = reported as a calculated mean • TYPE_SUL (sulfate) • MEDIAN = reported as a calculated median • TYPE_US_CM • MINIMUM = reported as minimum value, typically as part of a range (for example, (µS cm-1) 1–10) • OCCURRENCE = a single value was reported, and it was not specified as a mean, median, etc. US_CM Salinity, reported as microsiemens per centimeter (µS cm-1); if salinity was reported in other units (for example, milligrams per liter [mg L-1], percent [%]), the data were converted to provide a consistency among studies. The original data/units are also provided, and the data that were converted are identified by the variable “CONDUCTIVITY_CODE” (see above). WIC Wetland indicator category according to the U.S. Department of Agriculture, Natural Resources Conser- vation Service Web page, accessed September 2007, at http://plants.usda.gov/wetinfo.html: • OBL = Obligate Wetland: Occurs almost always (estimated probability 99%) under natural conditions in wetlands. • FACW = Facultative Wetland: Usually occurs in wetlands (estimated probability 67%–99%), but occasionally found in nonwetlands. • FAC = Facultative: Equally likely to occur in wetlands or nonwetlands (estimated probability 34%–66%). • FACU = Facultative Upland: Usually occurs in nonwetlands (estimated probability 67%–99%), but occasionally found on wetlands (estimated probability 1%–33%). • UPL = Obligate Upland: Occurs in wetlands in another region, but occurs almost always (estimated probability 99%) under natural conditions in nonwetlands in the regions specified. If a species does not occur in wetlands in any region, it is not on the National List. • A positive (+) or negative (-) sign was used with the Facultative Indicator categories to more specifically define the regional frequency of occurrence in wetlands.The positive sign indicates a frequency toward the higher end of the category (more frequently found in wetlands), and a negative sign indicates a frequency toward the lower end of the category (less frequently found in wetlands).

References

Cowardin, L.M., Carter, V., Golet, F.C., and LaRoe, E.T., 1979, Classification of wetlands and deepwater habitats of the United States: Washington, D.C., U.S. Fish and Wildlife Service, FWS/OBS-79/31, 131 p. Tchobanoglous, G., and Burton, F.L., 1991, Wastewater engineering; treatment, disposal, and reuse (3d ed.): New York, N.Y., Metcalf & Eddy, Inc., McGraw-Hill Inc., 1,024 p. The Great Plains Flora Association, 1986, Flora of the Great Plains: Lawrence, University Press of Kansas, 1,402 p. Appendix 4. Description of Each Data Source Used to Develop the Plant and Invertebrate Databases 35

Appendix 4. Description of Each Data Source Used to Develop the Plant and Invertebrate Databases

Grosshans and Kenkel, 1997 General description: Species composition (7 dominant plant species) and specific conductance were reported for the Marsh Ecology Research Complex (MERC) located on the southern end of Lake Manitoba, Manitoba, Canada. The MERC consists of 10 sand-diked marshes (cells) that are approximately 5–7 ha. The data were obtained from table 2. Description of data: Mean soil conductivities were presented for each species for a range of water depths. Specific conductance reporting units: micromhos cm-1 (µS cm-1), no conversion Description of data “type”: ●● Mean: reported as mean value

Hammer and Heseltine, 1988 General description: Plant species composition (aquatic macrophytes), pH, specific conductance, and makeup of the dominant salt were reported for 35 lakes located in Alberta and Saskatchewan, Canada. The data were obtained from tables 1 and 2. Description of data: Ranges or single values of specific conductance, single values of pH, and makeup of the dominant salt were presented for all lakes. Additionally, aquatic macrophytes occurring in each lake were identified. Since specific conductance was presented as a range and as a single value, the low and high ranges are presented in this database as minimum and maximum values and the single values are presented as an occurrence (the authors do not state that the values are measures of central tendency). Plants occurring in each lake were assigned the specific conductance (single value or range), pH, and dominant salt of the lake. Specific conductance reporting units: mS cm-1 ●● converted to µS cm-1 (mS cm-1 × 1,000 = µS cm-1) Description of data “type”: ●● Occurrence: single value presented ●● Min: lowest value presented in range ●● Max: greatest value presented in range

Johnson, 1990 General description: Salinity gradients and their relation to vegetation and aquatic invertebrates were reported for habitats on Bowdoin National Wildlife Refuge, Mont. Description of data: Table 1 provides the specific conductance values for each transect. Table 2 describes aquatic macrophytes found along 8 transects categorized as slightly brackish; table 3 describes aquatic macrophytes found along 5 transects categorized as moderately brackish; table 4 describes aquatic macrophytes found along 9 transects categorized as brackish; table 6 describes aquatic invertebrate taxa and their abundance per salinity category. Aquatic macrophytes were assigned salinity values based on sample transect data presented in table 1. Invertebrate species were assigned a range of conductivity values based on the salinity categories from table 6. Specific conductance reporting units: µS cm-1, no conversion Description of data “type”: ●● Min: lowest conductivity value reported for each taxa ●● Max: greatest conductivity value reported for each taxa

Kantrud, 1990 General description: Data were obtained from a literature review of sago pondweed (Potamogeton pectinatus L.) and represent information from multiple sources. Description of data: Table 5 presents the salinity concentrations of locations where sago pondweed occurred. The data represent numerous sources and consist of ranges or single observations of salinity and optimum salinity. Table 6 presents a range of pH values that represent data from multiple sources. Specific conductance reporting units: mg L-1 ●● converted mg L-1 to µS cm-1 (mg L-1 / 0.64) Description of data “type”: ●● Min: ○○Salinity: lowest value presented in table 5, which contains ranges from multiple sources (that is, multiple minimum values) ○○pH: lowest value from single range presented in table 6, which represents data summarized from multiple sources 36 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

●● Max: ○○Salinity: greatest value presented in table 5, which contains ranges from multiple sources (that is, multiple maximum values) ○○pH: greatest value from single range presented in table 6, which represents data summarized from multiple sources

Kantrud, 1991 General description: Data were obtained from a literature review of wigeongrass (Ruppia maritima) and represent information from multiple sources. Description of data: Table 5 presents the salinity concentrations of locations where wigeongrass occurred. The data represent numerous sources and consist of ranges or single observations of salinity and optimum salinity. Tables 6 (water) and 7 (soil) present ranges of pH values that represent data from multiple sources. Specific conductance reporting units: mg L-1 ●● converted mg L-1 to µS cm-1 (mg L-1 / 0.64) Description of data “type”: ●● Min: ○○Salinity: lowest value presented in table 5, which contains ranges from multiple sources (that is, multiple minimum values) ○○pH: lowest value from single range presented in tables 6 or 7, which represents data summarized from multiple sources ●● Max: ○○Salinity: greatest value presented in table 5, which contains ranges from multiple sources (that is, multiple maximum values) ○○pH: greatest value from single range presented in tables 6 or 7, which represents data summarized from multiple sources

Kantrud, 1996 General description: Data were obtained from a literature review of the alkali (Scirpus maritimus L.) and saltmarsh (Scirpus robustus Pursh) bulrushes and represent information from multiple sources. Description of data: Tables 6 (water) and 10 (soil) present the salinity concentrations of locations where alkali and saltmarsh bulrush occurred. The data represent numerous sources and consist of ranges or single observations of salinity and optimum salinity. Tables 8 (water) and 10 (soil) present ranges of pH values that represent data from multiple sources. Specific conductance reporting units: g L-1 (water) and mS cm-1 (soil) ●● converted g L-1 to µS cm-1 (g L-1 × 1,000 = mg L-1 / 0.64) ●● converted mS cm-1 to µS cm-1 (mS cm-1 × 1,000) Description of data “type”: ●● Min: ○○Salinity: • g L-1: lowest value presented in table 6, which contains ranges from multiple sources (that is, multiple minimum values) • mS cm-1: lowest value from single range presented in table 10, which contains ranges from multiple sources • Total salts (%): lowest value from single range presented in table 10, which contains ranges from multiple sources ○○pH: lowest value from single range presented in tables 8 or 10, which contains ranges from multiple sources ●● Max: ○○Salinity: • g L-1: greatest value presented in table 6, which contains ranges from multiple sources (that is, multiple maximum values) • mS cm-1: greatest value from single range presented in table 10, which contains ranges from multiple sources • Total salts (%): greatest value from single range presented in table 10, which contains ranges from multiple sources ○○pH: greatest value from single range presented in tables 8 or 10, which contains ranges from multiple sources

Kantrud and others, 1989 General description: Vegetation occurrence (hydrophytes, submerged and floating aquatic plants) and specific conductance data were summarized for Prairie Pothole Region wetlands by combining data from relevant literature and unpublished data. The data were obtained from tables 5.10, 5.11, and 5.12. Description of data: Mean, minimum, and maximum values for specific conductivity were presented for each plant species (when available). Additional information included wetland class and/or water regime. Specific conductance reporting units: mS cm-1 Appendix 4. Description of Each Data Source Used to Develop the Plant and Invertebrate Databases 37

●● converted to µS cm-1 (mS cm-1 × 1,000 = µS cm-1) Description of data “type”: ●● Mean: reported as mean value ○○Values for submerged and floating aquatic plants (table 5.12) are presented as either a mean, or as a single measurement. In this report, all values are considered a mean since the authors do not specify which are mean values or single measurements. ●● Min: reported as minimum value ●● Max: reported as maximum value

Kay and others, 2001 General description: Aquatic macroinvertebrates were collected and conductivity and pH were measured at 176 river sites in Australia during the spring of 1997. Description of data: Table 3 presents ranges (min/max) of specific conductance and pH for each invertebrate family. Specific conductance reporting units: mS cm-1 ●● converted to µS cm-1 (mS cm-1 × 1,000 = µS cm-1) Description of data “type”: ●● Min: lowest value reported in range ●● Max: greatest value reported in range

Lancaster and Scudder, 1987 General description: Communities of aquatic Coleoptera and Hemiptera were examined in eight fishless lakes of varying salinities in central British Columbia, Canada. Description of data: Table 1 presents seasonal (monthly; May–October) measurements of specific conductance for each lake. Table 2 presents the invertebrate species composition for each lake. Invertebrate species were assigned the range of conductivity values for the lakes where they occurred. Specific conductance reporting units: µS cm-1, no conversion Description of data “type”: ●● Mean: mean conductivity value for each lake from the six sample periods ●● Min: lowest conductivity value for each lake from the six sample periods ●● Max: greatest conductivity value for each lake from the six sample periods

McCarraher, 1970 General description: Fairy shrimps (Anostraca) were collected and specific conductance and pH were measured in 246 sites in the sandhills region of Nebraska. Description of data: Table 1 presents specific conductance and pH values for 6 fairy shrimp species from 23 locations. Specific conductance reporting units: µS cm-1, no conversion Description of data “type”: ●● Occurrence: value represents single conductivity/pH measurement from each location (the author does not state whether the values are single measurements or measures of central tendency).

Rawson and Moore, 1944 (plants) General description: Plant species (common rooted aquatic plants) composition and salinity concentration were reported for lakes of varying salinities in Saskatchewan, Canada. Description of data: Table VIII presents plant species occurrences and salinities (total solids) for 9 lakes. Plants occurring in each lake were assigned the salinity reported for the lake. Specific conductance reporting units: ppm (total solids) ●● ppm converted to µS cm-1 (ppm = mg L-1 / 0.64 = µS cm-1) Description of data “type”: ●● Occurrence: single value presented

Rawson and Moore, 1944 (invertebrates) General description: Salinity ranges were reported for invertebrate taxa inhabiting lakes of varying salinities in Saskatchewan, Canada. Description of data: Table X reported salinity ranges for numerous species of Cladocera and Copepoda and one species of Anostraca. Data for additional invertebrates (for example, insects, snails, leeches) were obtained by estimating salinity ranges presented graphically in figure 7. 38 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

Specific conductance reporting units: ppm (total solids) ●● ppm converted to µS cm-1 (ppm = mg L-1 / 0.64 = µS cm-1) Description of data “type”: ●● Occurrence: single value presented ●● Min: lowest value reported (table X) or estimated (figure 7) ●● Max: greatest value reported (table X) or estimated (figure 7)

Stewart and Kantrud, 1972 General description: Plant species (primary and secondary) characteristic of Prairie Pothole Region wetlands in North Dakota were reported as (1) frequently common or abundant, (2) frequently fairly common/occasionally common or abundant, or (3) occasionally fairly common for various wetland zones (for example, wet-meadow, shallow-marsh, deep-marsh). Each plant was reported to occur in wetlands characterized as fresh, slightly brackish, moderately brackish, brackish, subsaline, or saline. Each of these salinity categories was associated with a normal range of specific conductance values.The data were obtained from figures 4, 9, 20, 27, 35, 37. Description of data: Ranges (minimum/maximum) of salinity were assigned to each plant based on the salinity categories of the type of wetland where each species occurred. Specific conductance reporting units: micromhos cm-1 (µS cm-1), no conversion Description of data “type”: ●● Min: lowest salinity value associated with the “most fresh” salinity category reported for the species ●● Max: greatest salinity value associated with the “most saline” salinity category reported for the species

Tangen, unpublished General description: Aquatic macroinvertebrates were collected and conductivity and pH were measured from 24 wetlands (PEMF) within the Prairie Pothole Region of North Dakota during 2000. Description of data: Data from individual wetlands were summarized across the three sample periods (June, July, August) and include invertebrate taxa occurrence in relation to specific conductance and pH of the water. Invertebrates occurring in each wetland were assigned the specific conductance and pH of the wetland. Specific conductance reporting units: µS cm-1, no conversion Description of data “type”: ●● Mean: mean value (specific conductance, pH) for each wetland (n=24) calculated from measurements collected during each sample period (n=3). ●● Min: minimum value (specific conductance, pH) for each wetland (n=24) calculated from measurements collected during each sample period (n=3). ●● Max: maximum value (specific conductance, pH) for each wetland (n=24) calculated from measurements collected during each sample period (n=3).

Ungar, 1966 General description: Salt tolerance of plants growing in saline areas of Kansas and Oklahoma was examined. Description of data: Table I presents ranges of soil salt content in which plant species were found in saline areas of Kansas and Oklahoma. Values reported are for the upper 10 cm of the soil profile and are expressed as percentage total salts on a dry soil weight basis. Mean, minimum, and maximum salinity values were reported for each species. Specific conductance reporting units: percent ●● converted percent to µS cm-1 (% × 10,000 = ppm = mg L-1 / 0.64) Description of data “type”: ●● Mean: reported as mean value ●● Min: reported as minimum value ●● Max: reported as maximum value

Ungar, 1967 General description: Vegetation-soil relationships on saline soils in northern Kansas were examined. Description of data: Table 3 presents ranges of soil salinity and pH, as well as the concentration of chloride (%) for vegetation communities found in salt marshes of northern Kansas. Salinity values reported are for the upper 10 cm of the soil profile and are expressed as conductivity and total solids. Range (minimum and maximum) of conductivity, pH, chloride, and total solids were reported for the dominant species for each vegetation community. Specific conductance reporting units: mmhos cm-1 and total solids (%) ●● converted mmhos cm-1 to µS cm-1 (mmhos cm-1 × 1,000) Appendix 4. Description of Each Data Source Used to Develop the Plant and Invertebrate Databases 39

Description of data “type”: ●● Min: lowest value reported in range ●● Max: greatest value reported in range

Ungar, 1970 General description: Species-soil relationships on sulfate-dominated soils of South Dakota were examined. Description of data: Plant species-soil (upper 10 cm) relationships were presented for Stink and Bitter lakes in South Dakota. Table 2 presents single and median values for pH, conductivity, total salts (%), sulfate (%), chloride (%), and sodium (%) for each plant species. Specific conductance reporting units: mmhos cm-1 and total salts (%) ●● converted mmhos cm-1 to µS cm-1 (mmhos cm-1 × 1,000) Description of data “type”: ●● Occurrence: single value reported ●● Median: value reported as median

Ungar, 1974 General description: The data were acquired from a book chapter containing an extensive literature review of halophytes. Salin- ity and pH values were reported in the text as well as in tables. Description of data: Salinity and pH data from numerous sources were reported for various plant species. These data were summarized for this report. Specific conductance reporting units: mmhos cm-1 and/or total salts (%) ●● converted mmhos cm-1 to µS cm-1 (mmhos cm-1 × 1,000) ●● converted percent to µS cm-1 (% × 10,000 = ppm = mg L-1 / 0.64) Description of data “type”: ●● Min: lowest value reported (data summarized from entire chapter) ●● Max: greatest value reported (data summarized from entire chapter)

Ungar and others, 1969 General description: Plant species-soil (upper 10 cm) relationships at salt marshes near Lincoln, Nebraska, were examined. Description of data: Tables 1 and 2 present single values of conductivity, total salts (%), sodium (%), chloride (%), sulfate (%), and pH. Specific conductance reporting units: mmhos cm-1 and total salts (%) ●● converted mmhos cm-1 to µS cm-1 (mmhos cm-1 × 1,000) Description of data “type”: ●● Occurrence: single value reported ●● Median: median value reported ●● Min: lowest value reported ●● Max: greatest value reported

Ungar and others, 1979 General description: The distribution and growth of Salicornia europaea and other halophytes along a soil salinity gradient were described for a salt marsh in Ohio. Description of data: Ranges of specific conductance for vegetation zones (represented by dominant species) were presented. Specific conductance reporting units: mmhos cm-1 ●● converted mmhos cm-1 to µS cm-1 (mmhos cm-1 × 1,000) Description of data “type”: ●● Min: lowest value reported ●● Max: greatest value reported

Ungar and Riehl, 1980 General description: Soil salinities were presented for 5 vegetation zones (represented by dominant species) from an inland saline pan in Ohio. Description of data: Soils from the 5 vegetation zones were collected and ranges of soil salinities for the growing season were presented. Each plant species was assigned the range of soil salinities for the vegetation zone where it was present. Specific conductance reporting units: percent ●● converted percent to µS cm-1 (% × 10,000 = ppm = mg L-1 / 0.64) 40 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

Description of data “type”: ●● Min: lowest value reported ●● Max: greatest value reported

USGS, unpublished a General description: Vegetation surveys were conducted and specific conductance and pH (water and soil) measurements were collected from 204 depressional wetlands (PEMC, PEMF) located in 5 States (Iowa, Minnesota, Montana, North Dakota, and South Dakota) across the Prairie Pothole Region during 1997 (USGS, Northern Prairie Wildlife Research Center, Study Plan 168.01). Description of data: Data from individual wetlands were summarized by state and wetland class (that is, PEMC, PEMF) and include plant species occurrences in relation to specific conductance and pH of the soil and/or water. Plants occurring in each wetland were assigned the specific conductance and pH of the wetland and/or soil sample corresponding to each vegetation zone. Specific conductance reporting units: µS cm-1, no conversion Description of data “type”: ●● Occurrence: value represents occurrence of plant species in a single wetland basin ●● Mean: mean value for the species calculated from all wetlands located in each state/wetland class combination ●● Min: lowest value for the species based on data from all wetlands located in each state/wetland class combination ●● Max: greatest value for the species based on data from all wetlands located in each state/wetland class combination

USGS, unpublished b General description: Aquatic invertebrates were collected from 17 wetlands (PEMC, PEMF) located at the Cottonwood Lake long-term study area in Stutsman County, N. Dak. This database represents samples collected monthly (May–October) from 1992–2006 (USGS, Northern Prairie Wildlife Research Center, Study Plan 140.01). Description of data: 1. Data from individual wetlands (3 transects, 3 sample methods [funnel traps, benthic core sampler, sweep net], 2–4 sampling zones [wet-meadow, shallow-marsh, deep-marsh, open-water]) were summarized by month and year. 2. Specific conductance data were summarized for individual wetlands by month and year. 3. Data for all wetlands were summarized by wetland class (PEMC, PEMF) and year. Specific conductance reporting units: µS cm-1, no conversion Description of data “type”: ●● Mean: mean value for the taxon calculated for each year/wetland class combination. ●● Min: lowest value for the taxon for each year/wetland class combination. ●● Max: lowest value for the taxon for each year/wetland class combination.

References

Kantrud, H.A., Millar, J.B., and van der Valk, A.G., 1989, Vegetation of wetlands of the prairie pothole region, in van der Valk, A., ed., Northern prairie wetlands: Ames, Iowa State University Press, p. 132–187. Kay, W.R., Halse, S.A., Scanlon, M.D., and Smith, M.J., 2001, Distribution and environmental tolerances of aquatic macroin- vertebrate families in the agricultural zone of southwestern Australia: Journal of the North American Benthological Society, v. 20, p. 182–199. Lancaster, J., and Scudder, G.G.E., 1987, Aquatic Coleoptera and Hemiptera in some Canadian saline lakes—patterns in community structure: Canadian Journal of Zoology, v. 65, p. 1383–1389. McCarraher, D.B., 1970, Some ecological relations of fairy shrimps in alkaline habitats of Nebraska: American Midland Natu- ralist, v. 84, p. 59–68. Rawson, D.S., and Moore, J.E., 1944, The saline lakes of Saskatchewan: Canadian Journal of Research, v. 22, p. 141–201. Stewart, R.E., and Kantrud, H.A., 1972, Vegetation of prairie potholes, North Dakota, in relation to quality of water and other environmental factors: Washington, D.C., U.S. Geological Survey Professional Paper 585-D, 36 p. Ungar, I.A., 1966, Salt tolerance of plants growing in saline areas of Kansas and Oklahoma: Ecology, v. 47, p. 154–155. Ungar, I.A., 1967, Vegetation-soil relationships on saline soils in northern Kansas: American Midland Naturalist, v. 78, p. 98–120. Ungar, I.A., 1970, Species-soil relationships on sulfate dominated soils of South Dakota: American Midland Naturalist, v. 83, p. 343–357. Ungar, I.A., 1974, Inland halophytes of the United States, in Reimold, R.J., and Queen, W.H., eds., Ecology of halophytes: New York, N.Y., Academic Press, Inc., p. 235–305. Ungar, I.A., Benner, D.K., and McGraw, D.C., 1979, The distribution and growth of Salicornia Europaea on an inland salt pan: Ecology, v. 60, p. 329–336. Ungar, I.A., Hogan, W., and McClelland, M., 1969, Plant communities of saline soils at Lincoln, Nebraska: The American Mid- land Naturalist, v. 82, p. 564–577. Ungar, I.A., and Riehl, T.E., 1980, The effect of seed reserves on species composition in zonal halophyte communities: Botani- cal Gazette, v. 141, p. 447–452. 42 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 9 10 10 10 10 10 10 10 10 10 10 10 17 10 10 10 10 10 10 10 9,10 4,10 4,10 10,17 10,17 10,13 4,9,10 Source 3,4,9,10 10,14,15,16,17 7.9 8.0 8.1 9.7 8.8 7.7 9.8 8.6 8.3 Max pH 7.7 7.7 7.0 7.2 6.7 7.4 7.2 Min 490 620 630 600 1,301 4,688 2,790 2,450 1,623 7,000 5,000 1,850 5,000 6,460 2,200 1,623 5,500 5,000 5,500 9,500 4,550 1,050 1,850 3,800 5,500 3,330 Max ) 20,313 20,313 15,000 -1 (µS cm Conductivity 40 40 40 113 345 300 350 730 307 400 300 140 650 400 345 340 300 400 300 400 640 200 290 400 Min S S S S S S S S S S S S S S S S S S S S W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name ticklegrass ticklegrass ticklegrass common ragweed, short ragweed wheatgrass western wheatgrass rough pigweed short-awn foxtail western wheatgrass quackgrass short-awn foxtail quackgrass pink wild onion intermediate wheatgrass wild onion intermediate wheatgrass water plantain common water plantain tall wheatgrass thickspike wheatgrass common water plantain narrowleaf water plantain crested wheatgrass narrowleaf water plantain slender wheatgrass redtop sweet flag redtop yarrow —Continued

Species scabra scabra hyemalis artemisiifolia sp. smithii retroflexus aequalis smithii repens aequalis repens stellatum intermedium canadense intermedium triviale subcordatum elongatum dasystachyum subcordatum gramineum cristatum gramineum caninum stolonifera calamus stolonifera millefolium Summary of plant database.

Genus Agrostis Agrostis Agrostis Ambrosia Agropyron Agropyron Amaranthus Alopecurus Agropyron Agropyron Alopecurus Agropyron Allium Agropyron Allium Agropyron Alisma Alisma Agropyron Agropyron Alisma Alisma Agropyron Alisma Agropyron Agrostis Acorus Agrostis Achillea Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 43

1 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 9,10 4,10 9,10 4,9,10 Source 7.7 9.5 8.6 8.6 8.4 9.6 8.6 9.7 8.6 Max pH 7.2 7.2 8.5 7.6 8.1 8.0 7.3 Min 774 429 550 900 850 850 590 729 1,623 5,500 2,830 5,000 2,000 1,975 1,730 2,940 1,340 1,500 2,620 1,725 2,200 3,330 2,000 2,380 4,000 3,970 1,582 Max ) 15,000 -1 (µS cm Conductivity 40 40 40 40 326 350 140 310 300 355 140 300 274 350 295 525 290 400 900 290 560 140 290 326 Min S S S S S S S S S S S S S S S S S S W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name dogbane dogbane wormwood common milkweed wormwood common milkweed indian hemp dogbane, prairie showy milkweed indian hemp dogbane, prairie showy milkweed meadow anemone milkweed meadow anemone ovalleaf milkweed little bluestem swamp milkweed big bluestem swamp milkweed false indigo white sage lead plant prairie sagewort toothcup ragweed silky wormwood western ragweed biennial wormwood western ragweed biennial wormwood common ragweed, short ragweed —Continued

Species absinthium syriaca absinthium syriaca cannabinum speciosa cannabinum speciosa canadensis sp. canadensis ovalifolia scoparius incarnata gerardii incarnata fruticosa ludoviciana canescens frigida coccinea sp. dracunculus psilostachya biennis psilostachya biennis artemisiifolia Summary of plant database.

Genus Artemisia Artemisia Asclepias Artemisia Asclepias Apocynum Asclepias Apocynum Asclepias Anemone Asclepias Anemone Asclepias Andropogon Asclepias Andropogon Asclepias Amorpha Artemisia Amorpha Artemisia Ammannia Ambrosia Artemisia Ambrosia Artemisia Ambrosia Ambrosia Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 44 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 4 4 10 10 10 10 10 10 14 10 10 10 10 10 10 10 10 10 10 9,10 9,10 14,17 10,17 14,17 4,9,10 4,9,10 4,9,10 4,9,10 Source 10,12,13,14,15,17 8.4 8.3 9.7 9.7 8.2 7.3 8.1 8.1 8.4 8.7 8.4 Max pH 8.0 7.4 6.7 6.7 7.8 7.0 8.2 Min 590 850 410 590 471 630 330 630 540 700 9,844 2,500 5,500 9,500 4,550 5,500 2,000 1,000 9,844 9,800 3,970 1,470 4,700 2,930 Max ) 16,100 76,400 55,000 15,000 15,625 -1 (µS cm Conductivity 40 40 40 40 290 391 300 300 295 313 400 400 820 492 500 380 500 620 530 620 Min 1,875 1,250 3,594 S S S S S S S S S S S S S S S S S W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name Canada milk-vetch nodding beggar-ticks saltmarsh aster wild aster nodding beggar-ticks sloughgrass wild aster panicled aster sloughgrass water hyssop panicled aster water hyssop arrow-leaved aster new england aster willow baccharis panicled aster water fern, mosquito fern white prairie aster water fern, mosquito fern white prairie aster wild oats white aster spearscale white aster spearscale rayless aster silver-scale saltbush silver-scale rayless aster whorled milkweed —Continued

Species canadensis cernua subulatus sp. cernua syzigachne sp. simplex syzigachne rotundifolia simplex rotundifolia sagittifolius novae-angliae salicina hesperius mexicana falcatus mexicana falcatus fatua ericoides subspicata ericoides subspicata brachyactis argentea brachyactis verticillata Summary of plant database.

Genus Astragalus Bidens Aster Aster Bidens Beckmannia Aster Aster Beckmannia Bacopa Aster Bacopa Aster Aster Baccharis Aster Azolla Aster Azolla Aster Avena Aster Atriplex Aster Atriplex Aster Atriplex Aster Asclepias Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 45

1 9 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 4,9 4,10 4,10 9,10 10,13 4,9,10 4,9,10 4,9,10 Source 7.8 7.7 9.9 9.5 9.6 8.2 9.8 9.5 8.4 Max pH 6.6 7.1 6.9 7.4 7.3 6.9 6.7 Min 326 650 530 900 400 500 820 1,623 8,500 3,075 5,387 3,800 4,000 1,475 1,430 1,150 5,156 2,000 2,000 6,800 2,500 3,480 4,700 2,000 3,800 6,673 3,480 3,465 Max ) 17,600 -1 (µS cm Conductivity 40 40 40 40 40 40 40 40 400 340 400 295 256 170 140 330 770 430 100 290 140 350 290 Min 1,320 S S S S S S S S S S S S S S S S W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name downy brome downy brome bebb’s sedge bebb’s japanese brome slough sedge japanese brome slough sedge smooth brome water sedge smooth brome water sedge foxtail sedge charlock musk thistle, nodding thistle sideoats grama hedge bindweed violet boltonia vernal water starwort boltonia northern water starwort slimstem reedgrass boltonia big devil’s beggar-ticks big devil’s slimstem reedgrass beggar-ticks bluejoint devil’s beggar-ticks devil’s bluejoint devil’s beggar-ticks devil’s —Continued

Species canadensis tectorum tectorum bebbii japonicus atherodes japonicus atherodes inermis aquatilis inermis aquatilis alopecoidea kaber nutans curtipendula sepium latisquama var. asteroides verna asteroides hermaphroditica stricta asteroides vulgata stricta sp. canadensis frondosa frondosa Summary of plant database.

Genus Calamagrostis Bromus Bromus Carex Bromus Carex Bromus Carex Bromus Carex Bromus Carex Carex Brassica Carduus Bouteloua Calystegia Boltonia Callitriche Boltonia Callitriche Calamagrostis Boltonia Bidens Calamagrostis Bidens Calamagrostis Bidens Bidens Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 46 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 4 4 4 4 4 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 4,9 4,10 10,17 4,9,10 4,9,10 4,9,10 Source 9.5 8.6 7.9 6.9 9.9 8.0 8.0 Max pH 7.2 7.5 6.9 Min 113 305 450 350 850 480 340 412 630 400 720 450 900 700 2,600 5,000 3,800 2,615 2,000 3,120 3,200 1,700 5,500 9,400 1,487 4,700 1,400 5,000 Max ) 32,600 -1 (µS cm Conductivity 40 40 40 40 40 200 620 290 300 900 900 100 192 410 290 391 450 400 Min S S S S S S S S S S S S S S S W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name sartwell’s sedge sartwell’s beaked sedge beaked sedge clustered-field sedge nodding chickweed clustered-field sedge branched centaury woolly sedge caraway fox sedge woolly sedge smoothcone sedge fox sedge hairy sedge rigid sedge interior sedge manyhead sedge deer sedge tussock sedge deer sedge sawbeak sedge meadow sedge sedge meadow sedge sedge brown bog sedge sartwell’s sedge sartwell’s brown bog sedge fescue sedge —Continued

Species sartwellii rostrata rostrata praegracilis brachypodum praegracilis pulchellum lanuginosa carvi vulpinoidea lanuginosa laeviconica vulpinoidea lacustris tetanica interior sychnocephala hallii stricta hallii stipata granularis sp. granularis sp. buxbaumii sartwellii buxbaumii brevior Summary of plant database.

Genus Carex Carex Carex Carex Cerastium Carex Centaurium Carex Carum Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 47

1 2 9 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 17 10 10 10 10 10 1,10 4,10 9,10 10,17 4,9,10 Source 3,4,9,10 9.6 8.6 8.7 8.6 9.1 8.5 8.6 9.7 8.6 7.7 9.9 Max pH 7.2 7.2 8.3 7.4 7.8 8.2 7.0 Min 311 850 450 550 537 575 615 729 500 5,387 1,263 5,500 1,260 7,500 2,200 1,720 1,870 4,000 2,830 1,623 4,000 1,700 5,100 3,330 3,300 Max ) 15,000 39,800 15,000 39,800 -1 (µS cm Conductivity 40 40 40 40 720 326 140 140 274 450 550 860 345 300 350 450 500 140 300 380 480 300 Min 3,500 S S S S S S S S S S S S S S S S W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name tapeleaf flatsedge Canada thistle, field thistle hawk’s-beard Canada thistle, field thistle hawk’s-beard common water hemlock northern hawthorn common water hemlock northern hawthorn redosier dogwood common water hemlock goosefoot, lamb’s quarter goosefoot, lamb’s redosier dogwood goosefoot, lamb’s quarter goosefoot, lamb’s plains coreopsis alkali blite horse-weed alkali blite field bindweed oak-leaved goosefoot oak-leaved goosefoot field bindweed lamb’s quarters lamb’s bull thistle lamb’s quarters lamb’s bull thistle hornwort, coontail flodman’s thistle flodman’s hornwort, coontail —Continued

Species acuminatus arvense runcinata arvense runcinata maculata angustifolia rotundifolia maculata rotundifolia sericea maculata sp. sericea sp. tinctoria rubrum canadensis rubrum arvensis glaucum glaucum arvensis album vulgare album vulgare demersum flodmanii demersum Summary of plant database.

Genus Cyperus Cirsium Crepis Cirsium Crepis Cicuta Crataegus Cicuta Crataegus Cornus Cicuta Chenopodium Cornus Chenopodium Coreopsis Chenopodium Conyza Chenopodium Convolvulus Chenopodium Chenopodium Convolvulus Chenopodium Cirsium Chenopodium Cirsium Ceratophyllum Cirsium Ceratophyllum Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 48 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 4 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 4,9 9,10 4,10 10,17 4,9,10 2,9,10 4,9,10 Source 2,4,9,10 10,11,14,15,16,17 9.1 9.5 9.1 9.8 9.1 8.2 9.8 10.0 Max pH 6.7 6.6 6.8 7.9 6.8 7.8 7.2 Min 620 530 900 850 770 900 1,150 2,000 5,000 5,500 2,597 3,160 5,000 2,380 2,000 5,000 2,400 1,730 1,470 5,800 2,000 1,975 6,460 2,000 Max ) 15,000 10,938 41,000 -1 120,000 104,000 (µS cm Conductivity 40 40 40 40 40 40 40 113 550 290 290 500 400 305 560 500 290 100 395 400 500 310 350 480 900 Min S S S S S S S S S S S S S S S S S S W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name russian olive canadian waterweed barnyard grass waterpod spikerush barnyard grass drepanocladus moss spikerush inland saltgrass common spikerush inland saltgrass dichanthelium common spikerush blunt spikerush flixweed blunt spikerush purple prairie clover common spikerush flatstem spikerush orchard grass lady’s-slipper flatstem spikerush needle spikerush umbrella sedge needle spikerush fragrant flatsedge redrooted cyperus silverberry tapeleaf flatsedge —Continued

Species commutata angustifolia canadensis crusgalli nyctelea sp. crusgalli sp. sp. stricta spicata var. palustris stricta spicata var. oligosanthes palustris ovata obtusa var. sophia ovata obtusa var. purpurea macrostachya compressa glomerata sp. compressa acicularis sp. acicularis odoratus erythrorhizos acuminatus Summary of plant database.

Genus Elaeagnus Elaeagnus Elodea Echinochloa Ellisia Eleocharis Echinochloa Drepanocladus Eleocharis Distichlis Eleocharis Distichlis Dichanthelium Eleocharis Eleocharis Descurainia Eleocharis Dalea Eleocharis Eleocharis Dactylis Cypripedium Eleocharis Eleocharis Cyperus Eleocharis Cyperus Cyperus Cyperus Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 49

1 4 4 10 10 10 10 10 10 10 10 10 10 10 10 10 14 10 10 10 10 10 10 10 10 4,9 4,10 4,10 4,10 10,17 Source 8.6 9.7 8.6 8.0 8.6 7.3 8.5 7.3 Max pH 8.0 7.9 Min 540 729 412 450 300 450 460 148 700 760 2,200 1,260 1,270 1,613 2,050 3,800 3,800 2,100 2,100 1,730 1,520 1,232 2,570 4,000 1,880 5,000 2,930 3,800 Max ) 15,156 -1 (µS cm Conductivity 40 500 300 401 400 340 345 192 140 140 470 100 450 400 140 938 570 290 600 500 148 400 400 Min S S S S S S S S S S S S S S S S S S W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name narrowleaf cottongrass daisy fleabane bedstraw, cleavers bedstraw, Philadelphia fleabane northern bedstraw Philadelphia fleabane smooth scouring rush green ash wild strawberry smooth scouring rush wild strawberry common scouring rush flat-top goldentop water horsetail field horsetail flat-top goldentop field horsetail showy prairie gentian willow-herb, fireweed spotted spurge narrow-leaved willow-herb spotted spurge narrow-leaved willow-herb willow-herb leafy spurge joe-pye weed, spotted weed willow-herb joe-pye weed, spotted weed Canada wild rye —Continued

graminif var. graminif var. Species polystachion strigosus sp. philadelphicus boreale philadelphicus laevigatum pennsylvanica virginiana laevigatum virginiana hyemale graminifolia fluviatile arvense graminifolia arvense grandiflorum sp. maculata leptophyllum maculata leptophyllum ciliatum esula maculatum ciliatum maculatum canadensis Summary of plant database.

Genus Eriophorum Erigeron Galium Erigeron Galium Erigeron Equisetum Fraxinus Fragaria Equisetum Fragaria Equisetum Euthamia Equisetum Equisetum Euthamia Equisetum Eustoma Epilobium Euphorbia Epilobium Euphorbia Epilobium Epilobium Euphorbia Eupatorium Epilobium Eupatorium Elymus Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 50 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 9 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 4,9 4,10 9,10 4,10 10,17 4,9,10 Source 7.6 8.6 9.8 9.9 8.1 8.6 7.6 7.8 7.6 7.6 Max pH 8.1 6.7 7.4 7.4 7.4 Min 355 630 450 800 760 856 590 326 590 630 355 2,615 1,730 2,000 1,487 4,000 2,200 2,200 4,000 1,290 3,800 1,000 5,000 3,800 4,000 3,480 2,500 Max ) 20,313 15,000 -1 (µS cm Conductivity 40 40 40 40 300 450 450 500 450 140 300 400 260 350 400 590 330 400 450 288 140 300 340 400 Min 2,000 1,290 S S S S S S S S S S S S S S S W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name dame’s rocket dame’s rough false pennyroyal dame’s rocket dame’s seaside heliotrope curly-top gumweed hedge hyssop false sunflower, ox-eye false sunflower, wild licorice sunflower wild licorice fowl mannagrass stiff sunflower stiff fowl mannagrass plains sunflower tall mannagrass nuttall’s sunflower nuttall’s tall mannagrass nuttall’s sunflower nuttall’s northern mannagrass maximilian sunflower sea milkwort maximilian sunflower sea milkwort common sunflower small bedstraw common sunflower small bedstraw sneezeweed bedstraw, cleavers bedstraw, —Continued

Species matronalis hispidum matronalis curassavicum squarrosa neglecta helianthoides lepidota sp. lepidota striata rigidus striata petiolaris grandis nuttallii grandis nuttallii borealis maximiliani maritima maximiliani maritima annuus trifidum annuus trifidum autumnale sp. Summary of plant database.

Genus Helenium Hesperis Hedeoma Hesperis Heliotropium Grindelia Gratiola Heliopsis Glycyrrhiza Helianthus Glycyrrhiza Glyceria Helianthus Glyceria Helianthus Glyceria Helianthus Glyceria Helianthus Glyceria Helianthus Glaux Helianthus Glaux Helianthus Galium Helianthus Galium Galium Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 51

1 9 4 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 4,9 10,17 4,9,10 4,9,10 4,9,10 4,9,10 Source 10,14,17 2,4,8,9,10 11,14,15,17 10,11,12,13,14,15,16,17 7.3 7.3 8.2 9.1 7.8 9.0 8.1 9.5 9.7 8.6 8.2 Max pH 6.9 8.6 7.2 7.8 6.4 7.3 8.1 7.3 Min 729 750 729 890 400 640 450 850 760 1,582 2,930 5,500 2,790 2,000 4,000 2,000 2,300 4,100 1,947 2,570 2,000 Max ) 45,000 20,313 10,000 24,000 45,000 48,600 39,800 43,750 -1 (µS cm Conductivity 40 40 40 40 40 40 40 40 300 340 490 302 600 274 380 270 530 290 290 267 964 850 Min 1,875 1,225 S S S S S S S S S S S S S S S W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name kochia marsh elder kochia torrey’s rush torrey’s marsh elder marsh elder torrey’s rush torrey’s blue flag rush blue flag inland rush iris, flag inland rush iris, flag dudley rush spotted touch-me-not toad rush toad rush yellow stargrass baltic rush foxtail barley foxtail barley baltic rush mare’s tail mare’s richardson’s rush richardson’s mare’s tail mare’s sweetgrass richardson’s rush richardson’s sweetgrass —Continued

Species scoparia xanthifolia scoparia torreyi xanthifolia annua torreyi versicolor sp. versicolor interior sp. interior sp. dudleyi capensis bufonius bufonius hirsuta balticus jubatum jubatum balticus vulgaris alpinus vulgaris odorata alpinus odorata Summary of plant database.

Genus Kochia Iva Kochia Juncus Iva Iva Juncus Iris Juncus Iris Juncus Iris Juncus Iris Juncus Impatiens Juncus Juncus Hypoxis Juncus Hordeum Hordeum Juncus Hippuris Juncus Hippuris Hierochloe Juncus Hierochloe Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 52 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 9 9 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10,14 4,9,10 2,4,10 Source 2,4,9,10 7.9 9.9 9.6 9.9 9.7 8.6 7.4 7.8 Max pH 7.0 7.2 6.6 7.8 Min 577 300 326 450 590 550 450 307 540 650 7,656 4,700 4,000 4,000 4,000 3,330 2,620 1,475 1,730 2,000 2,000 3,970 1,730 2,930 3,800 Max ) 13,900 24,000 15,000 45,000 -1 (µS cm Conductivity 40 40 40 40 469 330 350 274 300 350 670 192 770 380 340 290 340 Min 2,000 3,330 S S S S S S S S S S S S S S S S S S W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name rocky mountain blazing star bearded sprangletop bearded sprangletop peppergrass star duckweed rough bugleweed Amercian bugleweed star duckweed common duckweed Amercian bugleweed common duckweed Amercian bird’s-foot trefoil Amercian bird’s-foot rice cutgrass palespike lobelia rice cutgrass hoary puccoon marsh vetchling hoary puccoon blue flax marsh vetchling false pimpernel blue lettuce mudwort wild lily blue lettuce prairie blazing star prickly lettuce dotted blazing star prickly lettuce —Continued

Species ligulistylis fascicularis fascicularis densiflorum trisulca asper americanus trisulca minor americanus minor unifoliolatus oryzoides spicata oryzoides canescens palustris canescens perenne palustris dubia tatarica aquatica philadelphicum tatarica pycnostachya serriola punctata serriola Summary of plant database.

Genus Liatris Leptochloa Leptochloa Lepidium Lemna Lycopus Lycopus Lemna Lemna Lycopus Lemna Lotus Leersia Lobelia Leersia Lithospermum Lathyrus Lithospermum Linum Lathyrus Lindernia Lactuca Limosella Lilium Lactuca Liatris Lactuca Liatris Lactuca Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 53

1 9 9 4 2 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 4,10 10,17 4,9,10 4,9,10 4,9,10 Source 2,3,4,8,9,10 8.6 9.8 7.2 9.7 9.0 8.3 9.0 8.3 7.0 9.8 Max pH 7.3 7.2 7.4 7.7 6.7 6.7 6.8 Min 629 550 735 600 860 362 690 1,582 2,000 6,460 8,000 4,740 5,500 3,800 2,500 1,725 2,000 4,688 1,320 3,800 2,000 4,100 5,000 2,380 2,653 2,780 5,500 Max ) 45,000 32,600 -1 (µS cm Conductivity 40 40 40 40 40 40 326 140 450 300 400 300 700 880 192 450 370 330 290 166 500 375 Min 3,500 S S S S S S S S S S S S S S S W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name yellow sweet clover water milfoil American milfoil yellow sweet clover alfalfa American milfoil alfalfa mat muhly marsh muhly black medick scratchgrass western water clover common mallow scratchgrass tufted loosestrife wild bergamot tufted loosestrife alleghany monkey-flower loosestrife field mint field mint loosestrife fringed loosestrife field mint sweet clover fringed loosestrife skeletonweed sweet clover rough bugleweed —Continued

Species officinalis heterophyllum exalbescens officinalis sativa exalbescens sativa richardsonis racemosa lupulina asperifolia vestita neglecta asperifolia thyrsiflora fistulosa thyrsiflora ringens hybrida arvensis villosa arvensis hybrida ciliata arvensis sp. ciliata juncea sp. asper Summary of plant database.

Genus Melilotus Melilotus Myriophyllum Myriophyllum Melilotus Medicago Myriophyllum Medicago Muhlenbergia Muhlenbergia Medicago Muhlenbergia Marsilea Malva Muhlenbergia Lysimachia Monarda Lysimachia Mimulus Lysimachia Mentha Mentha Lysimachia Lysimachia Mentha Melilotus Lysimachia Lygodesmia Lycopus Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 54 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 4 9 9 4 4 4 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 3,4,9 14,17 4,9,10 4,9,10 1,10,16 Source 2,4,9,10 9.8 7.8 7.9 9.0 7.3 8.6 8.6 9.9 Max pH 7.8 7.6 8.3 6.9 Min 900 900 856 275 620 729 450 400 380 700 100 5,000 2,750 5,500 2,500 2,000 1,700 5,500 4,660 2,050 1,700 3,800 3,210 1,730 5,000 Max ) 20,313 15,000 20,100 32,600 -1 (µS cm Conductivity 40 40 40 40 260 300 400 500 290 350 290 100 580 450 600 400 140 300 300 500 400 Min 1,094 S S S S S S S S S S S S S S W W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name common lousewort, wood betony grass-of-parnassus Pennsylvania pellitory plains bluegrass switchgrass common plantain switchgrass common plantain common witchgrass alkali plantain popcorn-flower common witchgrass pearl millet Virginia ground cherry Virginia common reed pearl millet common reed violet wood sorrel timothy yellow wood sorrel timothy yellow wood sorrel reed canarygrass yellow water lily catnip reed canarygrass naiad whorl-leaf watermilfoil green parrot’s feather green parrot’s —Continued

Species variegatum var. canadensis glauca pensylvanica arida virgatum major virgatum major capillare eriopoda scouleri capillare americanum virginiana australis americanum australis violacea pratense stricta pratense stricta arundinacea lutea cataria arundinacea flexilis verticillatum pinnatum Summary of plant database.

Genus Pedicularis Parnassia Parietaria Poa Panicum Plantago Panicum Plantago Panicum Plantago Plagiobothrys Panicum Panicum Physalis Phragmites Panicum Phragmites Oxalis Phleum Oxalis Phleum Oxalis Phalaris Nuphar Nepeta Phalaris Najas Myriophyllum Myriophyllum Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 55

1 3 9 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 13 10 2,4 4,9 9,10 4,9,10 Source 3,4,9,10 10,14,17 8.3 7.6 9.7 9.9 7.9 7.4 9.7 7.2 9.7 8.6 Max pH 7.8 6.6 7.0 7.4 7.4 7.8 Min 660 355 630 500 429 632 550 400 629 1,560 2,380 4,100 8,000 5,000 5,000 2,620 3,020 1,385 1,850 1,623 5,500 5,000 2,450 5,500 5,000 5,156 2,450 Max ) 45,000 54,063 -1 (µS cm Conductivity 40 40 40 40 40 380 430 300 300 560 192 300 270 362 300 192 410 140 140 310 Min 2,000 1,145 1,250 S S S S S S S S S S S S S S S W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name pale smartweed erect knotweed variable pondweed climbing or wild buckwheat Fries’ pondweed Fries’ knotweed slender pondweed knotweed waterthread pondweed swamp smartweed quaking aspen water smartweed quaking aspen water smartweed cotton-wood bluegrass bluegrass cotton-wood Sandberg’s bluegrass Sandberg’s bushy knotweed Kentucky bluegrass bushy knotweed Kentucky bluegrass Pennsylvania smartweed fowl bluegrass Pennsylvania smartweed fowl bluegrass pale smartweed Canada bluegrass —Continued

Species lapathifolium erectum gramineus convolvulus friesii aviculare filiformis aviculare diversifolius emersum amphibium var. tremuloides amphibium tremuloides amphibium deltoides sp. sp. deltoides sandbergii ramosissimum pratensis ramosissimum pratensis pensylvanicum palustris pensylvanicum palustris lapathifolium compressa Summary of plant database.

Genus Polygonum Polygonum Potamogeton Polygonum Potamogeton Polygonum Potamogeton Polygonum Potamogeton Polygonum Populus Polygonum Populus Polygonum Populus Poa Poa Populus Poa Polygonum Poa Polygonum Poa Polygonum Poa Polygonum Poa Polygonum Poa Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 56 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 8 4 10 10 10 10 10 10 10 10 10 10 10 10 10 10 2,4,9 10,17 4,9,10 4,9,10 4,9,10 4,9,10 2,4,8,9 Source 2,4,9,10 10,16,17 3,4,8,9,10 2,3,4,8,9,10 1,5,10,16,17 2,3,4,5,8,9,10,11,17 9.6 9.6 8.6 8.9 7.3 9.5 9.9 8.6 8.9 8.2 7.2 7.7 9.9 8.3 10.8 Max pH 8.0 7.9 7.7 8.0 6.9 6.3 8.4 8.0 7.3 6.7 Min 890 267 148 300 520 496 535 777 500 630 4,688 1,870 5,000 1,990 6,700 2,000 3,000 2,940 1,520 4,700 5,000 2,000 Max ) 34,000 39,800 15,000 31,406 45,000 -1 120,000 162,500 (µS cm Conductivity 40 40 40 38 40 450 500 760 380 700 880 267 290 450 400 330 400 300 530 401 100 410 420 260 Min S S S S S S S S S S S S S S W W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name threadleaf buttercup shore buttercup shore buttercup nuttall’s alkaligrass nuttall’s flatstem pondweed sheathed pondweed nuttall’s alkaligrass nuttall’s narrowleaf pondweed pondweed silver-leaf scurf-pea silver-leaf claspingleaf pondweed brook conquefoil claspingleaf pondweed Pennsylvania cinquefoil baby pondweed Norwegian cinquefoil baby pondweed sago pondweed Norwegian cinquefoil sago pondweed tall cinquefoil longleaf pondweed silverweed longleaf pondweed silverweed floatingleaf pondweed flatstem pondweed floatingleaf pondweed variable pondweed —Continued

Species flabellaris cymbalaria cymbalaria nuttalliana zosteriformis vaginatus nuttalliana strictifolius sp. argophylla richardsonii rivalis richardsonii pensylvanica pusillus norvegica pusillus pectinatus norvegica pectinatus arguta nodosus anserina nodosus anserina natans zosteriformis natans gramineus Summary of plant database.

Genus Ranunculus Ranunculus Ranunculus Puccinellia Potamogeton Potamogeton Puccinellia Potamogeton Potamogeton Psoralea Potamogeton Potentilla Potamogeton Potentilla Potamogeton Potentilla Potamogeton Potamogeton Potentilla Potamogeton Potentilla Potamogeton Potentilla Potamogeton Potentilla Potamogeton Potamogeton Potamogeton Potamogeton Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 57

1 4 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 4,9,10 4,9,10 4,9,10 4,9,10 3,9,10 4,9,10 4,9,10 Source 8.7 9.0 8.6 7.9 8.4 7.8 7.9 9.7 8.0 9.9 9.6 Max pH 7.6 7.0 7.4 6.9 7.5 7.6 6.9 7.4 Min 900 740 729 550 740 391 630 550 2,270 4,500 3,800 2,790 3,200 8,500 2,930 1,263 3,200 1,440 1,120 6,673 2,100 4,000 1,530 4,700 5,000 1,520 2,867 2,100 2,500 Max ) -1 (µS cm Conductivity 40 40 40 40 40 40 40 113 140 400 500 200 300 290 307 375 450 400 530 355 535 410 Min 1,130 1,005 S S S S S S S S S S S S S S S S W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name western wild rose prairie coneflower white water crowfoot smooth wild rose buttercup, crowfoot prairie wild rose buttercup, crowfoot bog yellow cress cursed crowfoot cursed crowfoot bog yellow cress bristly crowfoot purple-fringed riccia purple-fringed riccia bristly crowfoot liverwort macoun’s buttercup macoun’s liverwort macoun’s buttercup macoun’s slender riccia white water crowfoot slender riccia white water crowfoot wild black currant small yellow buttercup wild black currant small yellow buttercup smooth sumac threadleaf buttercup —Continued

Species glabra woodsii columnifera subrigidus blanda sp. arkansana sp. palustris sceleratus sceleratus palustris pensylvanicus natans natans pensylvanicus sp. macounii sp. macounii fluitans longirostris fluitans longirostris americanum gmelinii americanum gmelinii flabellaris Summary of plant database.

Genus Rhus Rosa Ratibida Ranunculus Rosa Ranunculus Rosa Ranunculus Rorippa Ranunculus Ranunculus Rorippa Ranunculus Ricciocarpus Ricciocarpus Ranunculus Riccia Ranunculus Riccia Ranunculus Riccia Ranunculus Riccia Ranunculus Ribes Ranunculus Ribes Ranunculus Ranunculus Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 58 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 9 4 4 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 9,10 9,10 6,17 Source 10,14,17 2,4,8,9,10 11,12,13,16,17 2,3,6,8,9,11,17 8.5 9.5 9.0 8.6 8.8 9.8 9.9 9.5 9.7 7.7 8.8 6.9 10.8 Max pH 6.7 7.2 7.0 7.2 7.9 6.7 7.1 6.8 7.2 6.0 3.1 Min 113 885 782 650 900 700 1,582 2,700 2,620 6,673 5,500 5,000 2,930 1,444 5,000 4,550 6,700 1,400 1,623 1,623 3,800 Max ) 11,250 45,000 15,000 66,000 14,200 31,406 -1 142,813 609,375 (µS cm Conductivity 40 40 40 40 94 113 420 350 307 300 256 400 300 148 400 270 270 290 326 600 400 Min 2,344 5,000 5,500 S S S S S S S S S S S S S S W W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name dock, sorrel dock, sorrel willow dock peachleaf willow saltwort willow dock saltwort great water dock arrowhead western dock arrowhead western dock common arrowhead willow-leaved dock golden dock common arrowhead arrowhead golden dock curly dock arrowhead curly dock ditchgrass, widgeon grass black-eyed susan ditchgrass, widgeon grass ditchgrass, widgeon grass ditchgrass, widgeon grass black-eyed susan Eurasian dock western wild rose —Continued

Species sp. sp. salicifolius amygdaloides rubra salicifolius rubra orbiculatus sp. occidentalis sp. occidentalis latifolia mexicanus maritimus latifolia cuneata maritimus crispus cuneata crispus rostrata maritima var. hirta occidentalis maritima var. maritima maritima hirta stenophyllus woodsii Summary of plant database.

Genus Rumex Rumex Rumex Salix Salicornia Rumex Salicornia Rumex Sagittaria Rumex Sagittaria Rumex Sagittaria Rumex Rumex Sagittaria Sagittaria Rumex Rumex Sagittaria Rumex Ruppia Rudbeckia Ruppia Ruppia Ruppia Rudbeckia Rumex Rosa Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 59

1 7 7 4 4 10 10 10 10 10 17 10 10 10 10 10 10 10 10 2,9 4,9 4,10 4,10 10,17 4,9,10 4,9,10 Source 14,15,17 2,3,4,9,10 2,3,4,7,9,10,17 7,10,11,14,15,16,17 8.3 7.5 6.6 8.4 9.6 8.5 8.9 9.8 9.9 8.6 9.8 9.8 9.7 9.9 9.7 Max pH 4.0 3.1 7.2 8.3 7.9 5.2 6.4 6.6 7.2 7.7 6.9 8.3 6.6 7.4 Min 975 900 620 600 2,200 2,270 5,500 5,500 1,809 2,050 1,896 1,850 4,200 2,470 1,700 6,700 1,655 4,550 1,385 Max ) 11,200 60,938 70,000 37,000 39,800 45,000 -1 111,000 120,000 481,250 120,000 (µS cm Conductivity 40 40 40 113 450 313 500 580 100 500 465 410 270 565 340 201 410 300 300 300 192 300 330 192 Min 2,600 1,719 12,000 S S S S S S S S S S S S S S W W W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name bulrush river bulrush sturdy bulrush darkgreen bulrush sturdy bulrush darkgreen bulrush common threesquare chairmaker’s bulrush chairmaker’s common threesquare chairmaker’s bulrush chairmaker’s cosmopolitan bulrush hardstem bulrush cosmopolitan bulrush cloaked bulrush hardstem bulrush Russian thistle cloaked bulrush willow Nevada bulrush panicled bulrush willow slender bulrush black willow sandbar willow slender bulrush sandbar willow, coyote willow sandbar willow, river bulrush sandbar willow, coyote willow sandbar willow, peachleaf willow —Continued

Species subs. interior sp. fluviatilis robustus atrovirens robustus atrovirens pungens americanus pungens americanus paludosus acutus paludosus pallidus acutus iberica pallidus sp. nevadensis microcarpus sp. heterochaetus nigra exigua heterochaetus exigua fluviatilis exigua amygdaloides Summary of plant database.

Genus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Scirpus Salsola Scirpus Salix Scirpus Scirpus Salix Scirpus Salix Salix Scirpus Salix Scirpus Salix Salix Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 60 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 9 4 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 1,10 1,10 14,17 4,9,10 4,9,10 4,9,10 1,10,17 Source 2,4,9,10 9.7 9.6 9.7 7.2 9.1 8.6 9.8 9.9 9.7 9.7 Max pH 7.3 7.2 7.0 7.2 7.4 6.8 7.2 7.0 7.2 Min 800 964 629 661 760 725 300 340 530 500 629 2,500 4,700 1,260 6,460 5,000 1,220 2,450 2,620 4,700 5,000 4,660 3,800 6,673 4,550 Max ) 20,800 20,313 81,500 15,000 -1 (µS cm Conductivity 40 40 40 40 40 490 192 300 290 140 610 500 140 620 256 140 380 590 140 400 400 280 340 192 Min 3,594 S S S S S S S S S S S S S S S S S S W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name prickly sow thistle foxtail field sow thistle foxtail field sow thistle sea purslane rigid goldenrod falsegold groundsel soft goldenrod swamp ragwort prairie goldenrod swamp ragwort late goldenrod skullcap late goldenrod skullcap Canada goldenrod marsh skullcap figwort Canada goldenrod sprangletop water parsnip sprangletop water parsnip softstem bulrush white-eyed grass softstem bulrush catchfly, campion catchfly, bulrush —Continued

Species asper sp. arvensis sp. arvensis verrucosum rigida rigida var. pseudaureus mollis congestus missouriensis congestus gigantea sp. gigantea sp. canadensis galericulata lanceolata canadensis festucacea suave festucacea suave validus campestre validus sp. sp. Summary of plant database.

Genus Sonchus Setaria Sonchus Setaria Sonchus Sesuvium Solidago Senecio Solidago Senecio Solidago Senecio Solidago Scutellaria Solidago Scutellaria Solidago Scutellaria Scrophularia Solidago Scolochloa Sium Scolochloa Sium Scirpus Sisyrinchium Scirpus Silene Scirpus Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 61

1 9 10 10 10 10 10 10 10 17 10 10 10 10 17 10 10 4,9 4,9 10,17 14,15 13,17 10,17 14,15 14,17 4,9,10 4,9,10 4,9,10 4,9,10 Source 11,14,15,16,17 8.4 9.0 8.7 9.6 8.6 8.6 9.5 9.8 8.3 9.6 10.0 10.0 Max pH 7.8 6.9 7.4 8.9 8.1 6.8 6.6 6.9 7.4 6.6 6.5 Min 630 730 774 740 725 530 3,000 4,688 3,480 4,000 1,900 1,500 1,150 5,000 3,480 7,813 5,000 4,550 Max ) 96,000 20,313 66,000 15,000 39,800 33,500 20,313 45,000 72,000 42,656 -1 110,400 (µS cm Conductivity 40 40 40 40 326 300 330 588 525 725 350 400 620 290 700 290 270 156 Min 2,188 5,781 5,000 2,000 2,344 4,688 1,875 17,188 S S S S S S S S S S S S S S S S S S W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name alkali sacaton common dandelion duckmeat, greater duckweed common dandelion duckmeat, greater duckweed meadow-sweet salt cedar western snowberry meadow-sweet prairie wedgegrass western snowberry sea blite wedgegrass sea blite salt-marsh sand spurry green needlegrass salt-marsh sand spurry green needlegrass prairie cordgrass needle-and-thread prairie cordgrass hedge-nettle, marsh betony alkali cordgrass hedge-nettle, marsh betony alkali cordgrass giant burreed Texas dropseed Texas giant burreed Indian grass —Continued

Species airoides officinale polyrhiza officinale polyrhiza alba ramosissima occidentalis alba obtusata occidentalis depressa obtusa depressa marina viridula marina viridula pectinata comata pectinata palustris gracilis palustris gracilis eurycarpum texanus eurycarpum nutans Summary of plant database.

Genus Sporobolus Taraxacum Spirodela Taraxacum Spirodela Spiraea Tamarix Symphoricarpos Spiraea Sphenopholis Symphoricarpos Suaeda Sphenopholis Suaeda Spergularia Stipa Spergularia Stipa Spartina Stipa Spartina Stachys Spartina Stachys Spartina Sparganium Sporobolus Sparganium Sorghastrum Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 62 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

1 2 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 4,9 3,10 1,10 2,4,9 10,17 4,9,10 4,9,10 Source 2,4,8,9,10 2,4,9,10,11 8.2 7.8 9.9 9.0 9.9 8.5 7.4 9.9 8.6 9.9 9.5 9.6 9.6 Max pH 8.1 6.8 7.1 6.7 6.9 6.2 7.0 7.4 7.9 Min 640 326 930 520 740 590 450 860 856 8,100 3,370 5,387 2,000 3,800 6,600 4,550 2,380 5,387 4,660 1,730 5,500 4,550 9,100 1,720 Max ) 72,000 13,600 15,000 -1 120,000 (µS cm Conductivity 40 40 40 900 520 400 330 450 274 380 400 400 100 330 470 195 420 525 270 550 192 280 391 450 700 Min S S S S S S S S S S S S S S S S W W W W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name seaside arrowgrass seaside arrowgrass white clover red clover common bladderwort red clover common bladderwort alsike clover stinging nettle goat’s beard goat’s stinging nettle hybrid cat-tail spiderwort hybrid cat-tail field pennycress cat-tail early meadow rue cat-tail broad-leaved cat-tail early meadow rue purple meadow rue broad-leaved cat-tail narrow-leaved cat-tail purple meadow rue wood sage narrow-leaved cat-tail wood sage marsh arrowgrass wood sage —Continued

Species maritima maritima repens pratense vulgaris pratense vulgaris hybridum dioica dubius dioica x glauca bracteata x glauca arvense sp. venulosum sp. latifolia venulosum dasycarpum latifolia angustifolia dasycarpum boreale canadense var. angustifolia canadense palustris canadense Summary of plant database.

Genus Triglochin Triglochin Trifolium Trifolium Utricularia Trifolium Utricularia Trifolium Urtica Tragopogon Urtica Typha Tradescantia Typha Thlaspi Typha Thalictrum Typha Typha Thalictrum Thalictrum Typha Typha Thalictrum Teucrium Typha Teucrium Triglochin Teucrium Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; Appendix 5. Summary of Plant Database 63

1 9 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 9,10 3,4,9 4,9,10 Source 8.6 8.1 9.7 7.8 7.6 9.7 9.8 8.6 Max pH 7.4 8.6 7.2 7.4 7.7 7.6 7.4 Min 590 860 650 450 770 856 500 450 500 620 760 590 632 852 880 1,720 2,000 1,725 4,000 1,400 1,690 5,000 5,500 Max ) 45,000 -1 (µS cm Conductivity 40 40 40 391 535 580 140 400 590 400 192 400 310 530 355 300 415 300 192 410 Min S S S S S S S S S S S S S S W W W W W W W W W W Substrate , microsiemens per centimeter] , microsiemens -1 Common name downy blue violet downy blue violet yellow prairie violet vetch vetch American vetch golden alexanders purslane speedwell purslane speedwell golden alexanders water speedwell meadow parsnip ironweed meadow parsnip ironweed vervain white camass blue vervain horned pondweed cocklebur blue vervain tapegrass cocklebur tapegrass —Continued

Species sororia sororia nuttallii sp. sp. americana aurea peregrina peregrina aurea anagallis-aquatica aptera fasciculata aptera fasciculata sp. elegans hastata palustris strumarium hastata americana strumarium americana Summary of plant database.

1, Grosshans and Kenkel, 1997; 2, Hammer and Heseltine, 1988; 3, Johnson, 1990; 4, Kantrud and others, 1989; 5, Kantrud, 6, 1991; 7, 1996; 8, Rawson Moore, 1, Grosshans and Kenkel, 1997; 2, Hammer Heseltine, 1988; 3, Johnson,

Genus Source: 1 Viola Viola Viola Vicia Vicia Vicia Zizia Veronica Veronica Zizia Veronica Zizia Vernonia Zizia Vernonia Verbena Zigadenus Verbena Zannichellia Xanthium Verbena Vallisneria Xanthium Vallisneria Appendix 5. Association (1986), and the minimum maximum values represent all data types (that is, reported as minimum, The Great Plains Flora [Scientific and common names primarily follow water; µS cm W, maximum, mean, median, or single occurrence). S, soil; 1944; 9, Stewart and Kantrud, 1972; 10, USGS unpbublished a; 11, Ungar and others, 1969; 12, Ungar and others, 1979; 13, Ungar and Riehl, 1980; 14, Ungar, 1966; 15, Ungar, 1967; 16, Ungar, 1970; 1967; 16, Ungar, 1966; 15, Ungar, Ungar and others, 1969; 12, 1979; 13, Riehl, 1980; 14, Ungar, 1944; 9, Stewart and Kantrud, 1972; 10, USGS unpbublished a; 11, 1974. 17, Ungar, 64 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

References

Grosshans, R.E., and Kenkel, N.C., 1997, Dynamics of emergent vegetation along natural gradients of water depth and salinity in a prairie marsh—delayed influences of competition,in Goldsborough, L.G., and Hann, B.J., eds., University of Manitoba Field Station (Delta Marsh) thirty-second annual report: Portage la Prairie, Manitoba, Canada, p. 83–93. Hammer, U.T., and Heseltine, J.M., 1988, Aquatic macrophytes in saline lakes of the Canadian prairies: Hydrobiologia, v. 158, p. 101–116. Johnson, K.M., 1990, Aquatic vegetation, salinity, aquatic invertebrates, and duck brood use at Bowdoin National Wildlife Ref- uge: Bozeman, Montana State University, master’s thesis. Kantrud, H.A., 1990, Sago pondweed (Potamogeton pecinatus L.)—a literature review: Washington, D.C., U.S. Fish and Wild- life Service, Resource Publication 176, 89 p. Kantrud, H.A., 1991, Widgeongrass (Ruppia maritima L.)—a literature review: Washington, D.C., U.S. Fish and Wildlife Ser- vice, Fish and Wildlife Research 10, 58 p. Kantrud, H.A., 1996, The alkali (Scirpus maritimus L.) and saltmarsh (S. robustus Pursh) bulrushes—a literature review: Wash- ington, D.C., National Biological Service Information and Technology Report 6, 77 p. Kantrud, H.A., Millar, J.B., and van der Valk, A.G., 1989, Vegetation of wetlands of the prairie pothole region, in van der Valk, A., ed., Northern prairie wetlands: Ames, Iowa State University Press, p. 132–187. Rawson, D.S., and Moore, J.E., 1944, The saline lakes of Saskatchewan: Canadian Journal of Research, v. 22, p. 141–201. Stewart, R.E., and Kantrud, H.A., 1972, Vegetation of prairie potholes, North Dakota, in relation to quality of water and other environmental factors: Washington, D.C., U.S. Geological Survey Professional Paper 585-D, 36 p. Ungar, I.A., 1966, Salt tolerance of plants growing in saline areas of Kansas and Oklahoma: Ecology, v. 47, p. 154–155. Ungar, I.A., 1967, Vegetation-soil relationships on saline soils in northern Kansas: American Midland Naturalist, v. 78, p. 98–120. Ungar, I.A., 1970, Species-soil relationships on sulfate dominated soils of South Dakota: American Midland Naturalist, v. 83, p. 343–357. Ungar, I.A., 1974, Inland halophytes of the United States, in Reimold, R.J., and Queen, W.H., eds., Ecology of halophytes: New York, N.Y., Academic Press, Inc., p. 235–305. Ungar, I.A., Benner, D.K., and McGraw, D.C., 1979, The distribution and growth of Salicornia Europaea on an inland salt pan: Ecology, v. 60, p. 329–336. Ungar, I.A., Hogan, W., and McClelland, M., 1969, Plant communities of saline soils at Lincoln, Nebraska: The American Mid- land Naturalist, v. 82, p. 564–577. Ungar, I.A., and Riehl, T.E., 1980, The effect of seed reserves on species composition in zonal halophyte communities: Botani- cal Gazette, v. 141, p. 447–452. Appendix 6. Summary of Invertebrate Database 65 1 5 5 5 4 7 7 7 4 2 7 4 7 7 4 4 7 6 5 7 6 7 7 6 7 7 1,7 6,7 4,5 1,5,6,7 1,2,5,7 1,5,6,7 Source 7.8 9.0 9.3 9.9 9.7 9.7 9.4 9.3 10.0 12.9 12.9 10.3 10.0 10.6 Max pH 7.1 8.8 4.6 5.4 9.3 7.1 9.8 7.3 7.1 7.1 7.1 Min 10.0 ) -1 136 892 388 982 892 1,879 1,563 7,813 1,250 4,378 2,394 1,243 2,644 5,554 5,557 2,930 2,000 1,563 4,340 2,000 2,178 3,658 1,997 Max 31,250 15,000 15,625 33,600 69,100 36,342 15,625 , microsiemens per centimeter] -1 187,500 99 80 91 57 57 69 99 80 313 156 156 101 188 680 100 217 168 105 531 313 531 185 540 270 Min 1,312 5,130 26,622 Conductivity (µS cm Species rectangula costata obtusirostris seali ornatus bundyi texanus bundyi mackini stagnalis lindahli campestris salina gibbus Genus Alonella Alona Alona Bosmina Streptocephalus Eubranchipus Eubranchipus Streptucephalus Eubranchipus Chirocephalopsis Theromyzon Placobdella Branchinecta Helobdella Branchinecta Helobdella Glossiphonia Branchinecta Erpobdella Artemia Chydorus Chydorus Family Chydoridae Chydoridae Chydoridae Bosminidae Streptocephalidae Chirocephalidae Chirocephalidae Streptocephalidae Chirocephalidae Chirocephalidae Glossiphoniidae Glossiphoniidae Branchinectidae Glossiphoniidae Branchinectidae Glossiphoniidae Glossiphoniidae Branchinectidae Glossiphoniidae Erpobdellidae Artemiidae Chydoridae Erpobdellidae Chydoridae Hirudinidae —Continued

Order Cladocera Cladocera Cladocera Cladocera Cladocera Anostraca Anostraca Anostraca Anastraca Anostraca Hydracarina Acarina Anostraca Rhynchobdellida Rhynchobdellida Anostraca Rhynchobdellida Anostraca Rhynchobdellida Rhynchobdellida Anostraca Rhynchobdellida Pharyngobdellida Anostraca Cladocera Pharyngobdellida Anostraca Cladocera Gnathobdellida Class Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Arachnoidea Oligochaeta Arachnida Branchiopoda Hirudinea Hirudinea Branchiopoda Hirudinea Branchiopoda Hirudinea Hirudinea Branchiopoda Hirudinea Hirudinea Branchiopoda Branchiopoda Hirudinea Branchiopoda Branchiopoda Hirudinea Hirudinea Summary of invertebrate database.

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Annelida Arthropoda Arthropoda Annelida Annelida Arthropoda Annelida Arthropoda Annelida Annelida Arthropoda Annelida Annelida Arthropoda Arthropoda Annelida Arthropoda Arthropoda Annelida Annelida Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm 66 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. 1 5 5 5 5 7 5 5 5 7 5 7 7 5 7 7 7 5 5 5 7 5 7 5 5 7 7 5 5 5 1,7 5,7 Source Max pH Min ) -1 3,125 1,563 5,266 7,813 3,698 6,543 6,150 2,930 2,407 6,150 1,563 4,650 7,813 4,073 1,563 1,563 3,706 2,497 2,295 7,813 Max 15,625 15,625 31,250 31,250 15,000 15,625 31,250 31,250 31,250 , microsiemens per centimeter] -1 187,500 187,500 57 69 80 91 119 313 313 156 313 313 156 196 313 105 156 140 500 313 313 156 156 156 313 156 156 313 156 Min 1,104 31,250 Conductivity (µS cm Species leuckarti fimbriatus vetulus bicuspidatus albidus mucronata albuquerquensis pulex longispina crystallina quadrangula leuchtenbergianum denticulatus pediculus testudinaria macrocopa lamellatus sphaericus serrulatus kindtii Genus Cyclops Cyclops Simocephalus Cyclops Simocephalus Cyclops Scapholeberis Scapholeberis Cletocamptus Daphnia Lynceus Daphnia Daphnia Sida Ceriodaphnia Ceriodaphnia Diaphanosoma Diaphanosoma Pleuroxus Polyphemus Graptoleberis Moina Eurycercus Moina Chydorus Cyclops Leptodora Family Cyclopidae Cyclopidae Daphnidae Cyclopidae Daphnidae Cyclopidae Daphnidae Daphnidae Canthocamptidae Daphnidae Lynceidae Daphnidae Daphnidae Sididae Daphnidae Daphnidae Sididae Sididae Chydoridae Polyphemidae Chydoridae Moinidae Chydoridae Moinidae Chydoridae Cyclopidae Leptodoridae —Continued

Order Eucopepoda Eucopepoda Cladocera Eucopepoda Cladocera Eucopepoda Cladocera Cladocera Eucopepoda Cyclopoida Cladocera Calanoida Conchostraca Cladocera Conchostraca Cladocera Cladocera Cladocera Cladocera Cladocera Cladocera Cladocera Cladocera Cladocera Cladocera Cladocera Cladocera Cladocera Eucopepoda Cladocera Class Copepoda Branchiopoda Copepoda Branchiopoda Copepoda Branchiopoda Copepoda Branchiopoda Branchiopoda Copepoda Copepoda Branchiopoda Copepoda Copepoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Branchiopoda Copepoda Summary of invertebrate database.

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm Appendix 6. Summary of Invertebrate Database 67 1 7 3 7 3 7 7 7 7 3 5 7 7 5 7 5 5 5 7 7 5 7 3,7 6,7 6,7 3,7 3,7 6,7 6,7 3,7 2,6,7 1,5,6 Source 9.7 9.4 7.9 9.2 9.4 12.9 Max pH 4.9 7.5 8.0 7.8 7.1 7.5 Min ) -1 811 811 960 1,771 4,892 1,568 4,030 3,019 1,568 4,030 3,310 5,800 5,266 3,850 1,493 9,106 9,106 4,030 1,094 3,850 4,378 3,430 1,492 Max 15,524 31,250 46,875 31,250 46,875 , microsiemens per centimeter] -1 192,000 156,250 187,500 99 69 80 57 85 45 45 57 57 45 913 107 913 313 190 105 156 107 156 179 156 650 Min 1,651 6,335 1,094 1,563 3,125 Conductivity (µS cm Species circumcinctus alaskanus filiformis longulus griseomicans sculptilis lindahli griseipennis mohammed falli tenuicaudatus bifarius siciloides antennatus shoshone ajax oregonensis hybridus semisulcatus viridis cordieri Genus Dytiscus Dytiscus Dytiscus Lixellus Coptotomus Litodactylus Coptotomus Caenestheriella Colymbetes Colymbetes Branchinecta Agabus Laophonte Ergasilus Agabus Diaptomus Agabus Diaptomus Agabus Diaptomus Graphoderus Agabus Agabus Diaptomus Dytiscus Acilius Cyclops Acilius Dytiscus Family Dytiscidae Dytiscidae Dytiscidae Dytiscidae Curculionidae Dytiscidae Curculionidae Dytiscidae Caenestheriidae Dytiscidae Dytiscidae Branchinectidae Dytiscidae Laophontidae Ergasilidae Dytiscidae Diaptomidae Dytiscidae Diaptomidae Dytiscidae Diaptomidae Dytiscidae Dytiscidae Dytiscidae Diaptomidae Dytiscidae Dytiscidae Cyclopidae Dytiscidae Dytiscidae —Continued

Order Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Conchostraca Coleoptera Coleoptera Anostraca Coleoptera Eucopepoda Eucopepoda Coleoptera Eucopepoda Coleoptera Eucopepoda Coleoptera Eucopepoda Coleoptera Coleoptera Coleoptera Eucopepoda Coleoptera Coleoptera Eucopepoda Class Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Eubranchiopoda Insecta Insecta Eubranchiopoda Insecta Copepoda Copepoda Insecta Copepoda Insecta Copepoda Insecta Copepoda Insecta Insecta Insecta Copepoda Insecta Insecta Copepoda Summary of invertebrate database.

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm 68 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. 1 3 7 7 7 7 7 7 7 7 7 7 7 3 3 7 7 7 7 3 6,7 3,7 6,7 3,7 3,7 6,7 6,7 3,7 3,7 3,7 3,7 6,7 Source 9.7 9.4 8.7 8.0 10.0 Max pH 7.1 7.5 7.1 8.2 Min ) -1 72 951 324 171 3,822 4,892 3,883 5,266 1,936 4,378 9,106 2,253 2,819 2,575 3,130 9,106 1,602 1,791 4,078 4,210 2,670 3,385 4,892 4,340 4,973 3,310 3,955 4,098 2,046 3,808 Max 15,524 , microsiemens per centimeter] -1 57 45 45 57 45 45 91 45 45 45 45 101 183 238 153 190 228 324 153 279 105 100 153 140 217 168 Min 6,335 Conductivity (µS cm Species subaenus fraterculus undulatus unguicularis tenebrosus turbidus superioris sellatus pervicinus sayi notabilis picatus criniticoxis patruelis modestus masculinus lutescens perplexus impressopunctatus occidentalis maculosus canadensis liberus biguttatus Genus Laccophilus Laccophilus Ilybius Hygrotus Ilybius Ilybius Hydroporus Hygrotus Hydroporus Hygrotus Hydroporus Hygrotus Hydroporus Hygrotus Hydroporus Hygrotus Hydroporus Hydroporus Hygrotus Hydaticus Hygrotus Hydaticus Hygrotus Graphoderus Liodessus Laccornis Hygrotus Graphoderus Laccophilus Hygrotus Graphoderus Family Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae Dytiscidae —Continued

Order Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Class Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Summary of invertebrate database.

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm Appendix 6. Summary of Invertebrate Database 69 1 2 7 7 7 7 7 7 7 7 3 3 3 7 7 3 7 7 7 7 7 6,7 2,7 6,7 6,7 6,7 3,7 2,7 3,7 3,7 6,7 3,7 Source 8.6 9.0 9.7 8.4 9.0 9.4 12.9 10.0 Max pH 7.8 6.1 5.3 7.5 7.5 7.6 6.4 7.3 Min ) -1 72 72 4,720 6,543 5,160 1,714 1,782 3,883 1,682 4,210 2,888 3,720 3,405 4,210 3,356 9,106 4,210 3,706 3,883 3,430 4,744 1,048 3,083 Max 11,532 15,524 18,600 15,524 15,524 15,524 , microsiemens per centimeter] -1 192,000 192,000 91 57 45 45 57 45 45 99 45 99 117 100 400 134 140 153 363 986 153 913 217 991 140 947 Min 1,209 6,335 6,335 1,015 Conductivity (µS cm Species fraternus connexus tortulosis pectoralis edentulus minutus maculiventris subguttatus strigatus frontalis stagninus salinarius striatellus leechi spenceri immaculicollis striatus griseostriatus hoppingi hatchi affinis Genus Berosus Berosus Haliplus Haliplus Peltodytes Gyrinus Peltodytes Gyrinus Peltodytes Gyrinus Haliplus Gyrinus Haliplus Rhantus Haliplus Rhantus Haliplus Potamonectes Haliplus Cercyon Potamonectes Haliplus Berosus Potamonectes Haliplus Berosus Liodessus Family Hydrophilidae Hydrophilidae Hydrophilidae Hydraenidae Haliplidae Helodidae Haliplidae Haliplidae Gyrinidae Haliplidae Gyrinidae Haliplidae Gyrinidae Haliplidae Gyrinidae Gyrinidae Haliplidae Dytiscidae Haliplidae Dytiscidae Haliplidae Dytiscidae Haliplidae Hydrophilidae Dytiscidae Haliplidae Hydrophilidae Dytiscidae Haliplidae Hydrophilidae Dytiscidae —Continued

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm 70 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. 1 3 7 7 6 7 6 7 7 7 7 7 7 7 7 7 6 2 7 7 3 7 7 5 7 6,7 6,7 6,7 6,7 1,6,7 Source 1,2,5,6,7 1,2,5,6,7 9.7 8.9 8.7 8.8 9.2 8.2 7.9 8.4 12.9 12.9 12.9 Max pH 4.6 7.1 7.6 7.5 4.6 8.1 5.9 Min ) -1 273 107 205 4,210 5,266 2,000 4,210 2,000 2,930 4,378 1,080 1,466 6,150 5,554 2,384 2,123 2,000 4,973 3,850 3,946 5,266 2,323 2,888 6,120 Max 11,532 15,000 36,700 15,524 10,938 , microsiemens per centimeter] -1 192,000 192,000 99 57 57 57 57 69 80 57 80 811 118 205 544 134 540 134 455 272 179 100 148 123 313 Min 6,335 3,706 1,682 Conductivity (µS cm Species sp. obtusatus fuscipes oblongus lineatus linearis lateralis diffusus subcupreus dorsalis Genus Paracymus Laccobius Hydrochus Chaoborus Hydrochara Hydrochara Bezzia Hydrobius Hydrobius Helophorus Helophorus Micralymma Helophorus Helophorus Tropisternus Enochrus Tropisternus Enochrus Paracymus Cymbiodyta Family Hydrophilidae Hydrophilidae Hydrophilidae Chironomidae Chaoboridae Hydrophilidae Chaoboridae Hydrophilidae Ceratopogonidae Ceratopogonidae Hydrophilidae Hydrophilidae Sminthuridae Poduridae Hydrophilidae Isotomidae Entomobryidae Hydrophilidae Staphylinidae Hydrophilidae Scirtidae Hydrophilidae Hydrophilidae Hydrophilidae Hydrophilidae Hydrophilidae Hydrophilidae Corethrinae Hydrophilidae —Continued

Order Coleoptera Coleoptera Coleoptera Diptera Diptera Coleoptera Diptera Coleoptera Diptera Diptera Coleoptera Diptera Coleoptera Collembola Collembola Coleoptera Collembola Collembola Coleoptera Collembola Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Coleoptera Diptera Coleoptera Class Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Summary of invertebrate database.

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm Appendix 6. Summary of Invertebrate Database 71 1 7 2 6 6 7 6 7 7 7 7 7 7 7 7 7 7 1,7 1,5 6,7 1,2 6,7 1,5 2,7 5,7 6,7 2,5,7 1,2,7 1,5,7 1,2,6,7 1,2,5,7 1,2,6,7 Source 9.2 9.4 9.7 9.3 9.7 9.1 9.0 8.7 9.3 12.9 12.9 10.0 12.9 10.0 Max pH 7.6 6.1 5.9 4.6 7.1 4.6 6.4 7.1 6.9 5.4 4.6 Min ) -1 637 572 531 4,378 2,245 3,698 2,294 3,536 4,650 2,381 2,123 4,098 2,013 2,930 5,554 5,554 4,210 6,543 2,927 Max 23,100 15,000 14,800 46,875 18,600 19,600 31,250 15,000 , microsiemens per centimeter] -1 156,250 192,000 192,000 192,000 57 69 80 80 85 57 69 91 115 210 200 313 139 185 140 100 153 132 286 185 126 313 159 220 105 Min 2,283 1,512 Conductivity (µS cm Species americanus perturbans Genus Callicorixa Lethocerus Lethocerus Belostoma Pericoma Caenis Callibaetis Baetis Mansonia Culiseta Culex Aedes Odontomyia Family Corixidae Stratiomyidae Corixidae Stratiomyidae Simuliidae Belostomatidae Sciomyzidae Belostomatidae Ptychopteridae Belostomatidae Psychodidae Psychodidae Ephemeroptera Ephydridae Caenidae Dolichopodidae Caenidae Dixidae Baetidae Baetidae Culicidae Baetidae Culicidae Culicidae Tipulidae Culicidae Tabanidae Culicidae Syrphidae —Continued

Order Hemiptera Diptera Hemiptera Diptera Diptera Hemiptera Diptera Hemiptera Diptera Hemiptera Diptera Hemiptera Diptera Ephemeroptera Diptera Ephemeroptera Diptera Ephemeroptera Diptera Ephemeroptera Ephemeroptera Diptera Ephemeroptera Diptera Ephemeroptera Diptera Diptera Diptera Diptera Diptera Diptera Class Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Summary of invertebrate database.

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm 72 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. 1 7 7 6 7 6 6 7 7 7 3 7 7 7 7 7 3 7 3 7 7 3 3 3,7 6,7 6,7 3,7 6,7 6,7 6,7 6,7 1,2,6,7 Source 9.4 8.7 9.7 9.7 10.0 10.0 10.0 10.0 10.0 10.0 Max pH 7.5 5.3 7.3 8.5 7.5 7.5 7.6 7.5 7.5 7.8 Min ) -1 811 811 959 2,745 1,682 4,030 1,583 2,000 2,030 4,210 3,650 4,892 3,755 3,350 4,098 1,679 4,098 3,018 4,210 1,940 5,800 3,850 4,098 4,098 4,892 9,106 Max 26,900 15,524 15,524 15,524 15,524 , microsiemens per centimeter] -1 45 57 80 45 98 99 45 45 57 57 45 308 100 107 665 925 531 194 858 105 274 101 Min 2,014 1,782 1,628 1,608 6,335 1,064 Conductivity (µS cm Species laevigata fusca atopodonta rawsoni americana verticalis interiores naias tarsalis borealis expleta solensis kirbyi bifida vulgaris margaritacea audeni Genus Buenoa Hesperocorixa Ranatra Hesperocorixa Mesovelia Hesperocorixa Hebridae Hespercorixa Rheumatobates Gerris Dasycorixa Cymatia Trichocorixa Cymatia Trichocorixa Corisella Corisella Trichocorixa Trichocorixa Cenocorixa Sigara Notonecta Cenocorixa Sigara Notonecta Cenocorixa Hesperocorixa Buenoa Callicorixa Family Notonectidae Notonectidae Corixidae Nepidae Corixidae Mesovelidae Corixidae Hebridae Corixidae Gerridae Gerridae Corixidae Gerridae Corixidae Corixidae Corixidae Corixidae Corixidae Corixidae Corixidae Corixidae Corixidae Corixidae Notonectidae Corixidae Corixidae Notonectidae Corixidae Corixidae Notonectidae Corixidae —Continued

Order Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Hemiptera Class Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Summary of invertebrate database.

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm Appendix 6. Summary of Invertebrate Database 73 1 7 6 6 7 7 7 7 7 5 7 6 7 7 7 7 2 7 7 6,7 6,7 6,7 6,7 6,7 2,7 3,7 1,2,7 1,2,7 1,6,7 1,2,7 5,6,7 1,2,7 Source 8.6 9.2 9.3 9.0 8.8 9.3 8.3 9.7 9.3 9.7 9.2 9.3 10.0 12.9 10.0 12.9 Max pH 8.1 7.3 7.5 4.9 6.7 7.8 5.9 7.5 7.1 7.3 6.4 7.1 7.1 5.3 5.3 Min ) -1 223 942 978 873 2,725 1,640 4,378 1,030 1,890 1,421 2,012 4,340 4,973 4,210 4,340 2,470 3,350 2,000 4,210 4,098 2,407 2,710 Max 69,100 30,200 15,000 31,250 30,200 46,875 30,200 36,700 33,600 , microsiemens per centimeter] -1 57 98 57 98 80 45 85 117 117 245 845 826 680 100 200 150 313 147 531 147 270 194 286 100 240 240 100 Min 1,454 Conductivity (µS cm Species undulata Genus Triaenodes Enallagma Oecetis Coenagrion Nectopsyche Amphiagrion Brachycentrus Anax Aeshna Sympetrum Libellula Leucorrhinia Plea Neoplea Lestes Limnephilus Notonecta Anabolia Ischnura Family Limnephilidae Leptoceridae Coenagrionidae Leptoceridae Coenagrionidae Leptoceridae Coenagrionidae Leptoceridae Coenagrionidae Brachycentridae Aeshnidae Aeshnidae Aeshnidae Libellulidae Libellulidae Velidae Libellulidae Pleidae Libellulidae Pleidae Lestidae Pleidae Lestidae Limnephilidae Corduliidae Notonectidae Limnephilidae Coenagrionidae —Continued

Order Tricoptera Odonata Tricoptera Odonata Tricoptera Odonata Tricoptera Odonata Tricoptera Odonata Tricoptera Odonata Tricoptera Odonata Trichoptera Odonata Odonata Odonata Odonata Hemiptera Odonata Hemiptera Odonata Hemiptera Odonata Hemiptera Odonata Tricoptera Odonata Hemiptera Tricoptera Class Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Insecta Summary of invertebrate database.

Phylum Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm 74 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont. 1 7 7 7 7 7 5 5 1 7 2 7 1 2 7 5 5 6 5 2 7 7 2 7 5 1,7 6,7 6,7 6,7 5,7 1,7 1,7 Source 9.0 8.0 8.8 9.4 8.8 8.8 10.0 12.9 Max pH 7.1 5.4 4.9 7.6 7.3 5.9 6.1 Min ) -1 469 185 1,416 4,214 1,618 4,098 3,658 6,543 1,608 7,600 4,214 2,594 1,542 3,125 1,875 1,640 1,563 6,150 3,808 1,985 3,906 Max 15,000 15,625 15,000 15,000 69,100 46,875 15,000 15,000 29,200 34,100 , microsiemens per centimeter] -1 57 98 119 118 911 324 510 313 254 196 313 313 500 217 200 168 500 100 313 540 162 363 313 910 313 150 100 101 200 767 Min Conductivity (µS cm Species reflexa elodes caparata azteca lacustre nitidum casertanum flavicornis stagnalis Genus Lymnaea Lymnaea Lymnaea Cambarus Lymnaea Lymnaea Hyalella Hyalella Anodonta Sphaerium Gammarus Pisidium Pisidium Cernotina Hydra Fabria Plumatella Molanna Family Lymnaeidae Lymnaeidae Lymnaeidae Cambaridae Lymnaeidae Lymnaeidae Talitridae Lymnaeidae Talitridae Unionidae Talitridae Perthiidae Sphaeriidae Gammaridae Gammaridae Sphaeriidae Ceinidae Sphaeriidae Polycentropodidae Sphaeriidae Polycentropodidae Hydridae Phryganeidae Plumatellidae Phryganeidae Parastacidae Palaemonidae Molannidae —Continued

Order Limnophila Limnophila Limnophila Decapoda Limnophila Limnophila Amphipoda Limnophila Amphipoda Pelecypoda Amphipoda Amphipoda Pelecypoda Amphipoda Amphipoda Pelecypoda Amphipoda Amphipoda Pelecypoda Tricoptera Pelecypoda Tricoptera Hydroida Tricoptera Plumatellida Tricoptera Decapoda Decapoda Tricoptera Class Gastropoda Gastropoda Gastropoda Malacostraca Gastropoda Gastropoda Malacostraca Gastropoda Gastropoda Malacostraca Bivalvia Malacostraca Malacostraca Bivalvia Malacostraca Malacostraca Bivalvia Malacostraca Malacostraca Bivalvia Insecta Bivalvia Insecta Hydrozoa Insecta Ostracoda Phylactolaemata Insecta Malacostraca Malacostraca Insecta Summary of invertebrate database.

Phylum Mollusca Mollusca Mollusca Arthropoda Mollusca Mollusca Arthropoda Mollusca Mollusca Arthropoda Mollusca Arthropoda Arthropoda Mollusca Arthropoda Arthropoda Mollusca Arthropoda Arthropoda Mollusca Arthropoda Mollusca Arthropoda Cnidaria Arthropoda Arthropoda Bryozoa Arthropoda Arthropoda Arthropoda Arthropoda Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm Appendix 6. Summary of Invertebrate Database 75 1 7 7 7 7 7 7 5 5 7 7 7 7 7 7 7 7 6 1,5 1,6,7 1,6,7 Source 9.4 10.0 10.0 Max pH 7.1 7.1 7.9 Min ) -1 7, USGS, unpublished b. 469 286 261

3,456 1,264 4,650 4,378 2,407 4,250 1,094 5,266 1,985 3,880 3,680 4,340 3,480 1,579 Max 15,000 15,000 , microsiemens per centimeter] -1 156,250 98 57 85 57 98 69 98 119 334 168 313 105 313 122 313 351 188 220 500 540 Min Conductivity (µS cm 6, Tangen, unpublished; Tangen, 6,

Species trivolvis anceps circumstriatus parvus crista jennessi lewesi gyrina umbilicatellus exacuous hypnorum campestris 5, Rawson and Moore, 1944;

Genus Helisoma Helisoma Gyraulus Gyralus Gyralus Armiger Planaria Gordius Physa Valvata Physa Promenetus Physa Promenetus Aplexa Planorbula 4, McCarraher, 1970; 4, McCarraher,

Family Planorbidae Planorbidae Planorbidae Planorbidae Planorbidae Planorbidae Planariidae Planorbidae Gordiidae Physidae Valvatidae Physidae Planorbidae Physidae Planorbidae Physidae Physidae Planorbidae Lymnaidae —Continued

3, Lancaster and Scudder, 1987; 3, Lancaster and Scudder,

Order Limnophila Limnophila Limnophila Limnophila Limnophila Limnophila Tricladida Limnophila Gordea Limnophila Mesogastropoda Limnophila Limnophila Limnophila Limnophila Limnophila Limnophila Limnophila Limnophila 2, Kay and others, 2001;

Class Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Turbellaria Gastropoda Gordioida Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Summary of invertebrate database.

- - 1, Johnson, 1990;

minthes morpha Phylum Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Platyhel Mollusca Nematoda Nemato Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Source: Appendix 6. [The minimum and maximum values represent all data types (that is, reported as minimum, maximum, mean, median, or single occurrence). µS cm 1 76 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

References

Johnson, K.M., 1990, Aquatic vegetation, salinity, aquatic invertebrates, and duck brood use at Bowdoin National Wildlife Refuge: Bozeman, Montana State University, master’s thesis. Kay, W.R., Halse, S.A., Scanlon, M.D., and Smith, M.J., 2001, Distribution and environmental tolerances of aquatic macro- invertebrate families in the agricultural zone of southwestern Australia: Journal of the North American Benthological Society, v. 20, p. 182–199. Lancaster, J, and Scudder, G.G.E., 1987, Aquatic Coleoptera and Hemiptera in some Canadian saline lakes—patterns in community structure: Canadian Journal of Zoology, v. 65, p. 1383–1389. McCarraher, D.B., 1970, Some ecological relations of fairy shrimps in alkaline habitats of Nebraska: American Midland Natu- ralist, v. 84, p. 59–68. Rawson, D.S., and Moore, J.E., 1944, The saline lakes of Saskatchewan: Canadian Journal of Research, v. 22, p. 141–201. Publishing support provided by: Helena and Lafayette Publishing Service Centers

For more information concerning this publication, contact: Director, Northern Prairie Wildlife Research Center U.S. Geological Survey 8711 37th Street Southeast Jamestown, North Dakota 58401 (701) 253-5500

Or visit the Northern Prairie Wildlife Research Center Web site at: http://www.npwrc.usgs.gov/ Gleason and others—Literature Review and Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Montana—Scientific Investigations Report 2009–5098 Printed on recycled paper