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

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

Literature Review and Database of Relations Between 

Salinity and Aquatic Biota: Applications to Bowdoin National 

Wildlife Refuge, Montana

U.S. Department of the Interior

U.S. Geological Survey

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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Although this report is in the public domain, permission must be secured from the individual copyright owners to 

reproduce any copyrighted materials contained within this report.

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

Long-term accumulation of salts in wetlands at Bowdoin 

National Wildlife Refuge (NWR), Mont., has raised concern 

among wetland managers that increasing salinity may threaten 

plant and invertebrate communities that provide important 

habitat and food resources for migratory waterfowl. Currently, 

the U.S. Fish and Wildlife Service (USFWS) is evaluating 

various water management strategies to help maintain suitable 

ranges of salinity to sustain plant and invertebrate resources of 

importance to wildlife. To support this evaluation, the USFWS 

requested that the U.S. Geological Survey (USGS) provide 

information on salinity ranges of water and soil for common 

plants and invertebrates on Bowdoin NWR lands. To address 

this need, we conducted a search of the literature on occurrences of plants and invertebrates in relation to salinity and 

pH of the water and soil. The compiled literature was used to 

(1) provide a general overview of salinity concepts, (2) document published tolerances and adaptations of biota to salinity, 

(3) develop databases that the USFWS can use to summarize 

the range of reported salinity values associated with plant and 

invertebrate taxa, and (4) perform database summaries that 

describe reported salinity ranges associated with plants and 

invertebrates at Bowdoin NWR. The purpose of this report is 

to synthesize information to facilitate a better understanding 

of the ecological relations between salinity and flora and fauna 

when developing wetland management strategies. A primary 

focus of this report is to provide information to help evaluate 

and address salinity issues at Bowdoin NWR; however, the 

accompanying databases, as well as concepts and information discussed, are applicable to other areas or refuges. The 

accompanying databases include salinity values reported for 

411 plant taxa and 330 invertebrate taxa. The databases are 

available in Microsoft Excel version 2007 (http://pubs.usgs.

gov/sir/2009/5098/downloads/databases_21april2009.xls) and 

contain 27 data fields that include variables such as taxonomic 

identification, values for salinity and pH, wetland classification, location of study, and source of data. The databases are 

not exhaustive of the literature and are biased toward wetland 

habitats located in the glaciated North-Central United States; 

however, the databases do encompass a diversity of biota 

commonly found in brackish and freshwater inland wetland 

habitats.

Introduction

Bowdoin National Wildlife Refuge (NWR) was established in 1936 to provide habitat for breeding and migrating 

waterfowl. The refuge is located about 11 km to the east of 

Malta, in the Milk River Valley of Phillips County, Mont. 

(fig. 1). Characteristic of refuges located in the arid NorthCentral United States, Bowdoin NWR is at risk to increased 

salinization and accumulation of trace elements that can 

threaten wetland biota (DuBois and others, 1992; Nimick, 

1997; Laubhan and others, 2006). Bowdoin NWR has been 

the subject of various investigations and model simulations 

(Lambing and others, 1988; Hamilton and others, 1989; 

Kendy, 1999; Bauder and others, 2007) that suggest that salt 

concentrations in lakes and marshes are increasing and that the 

long-term accumulation of salts could pose a significant threat 

to flora and fauna. The increasing salt concentrations on the 

refuge have been attributed to changes in water availability 

and quality that, in conjunction with water management strategies that stabilize water levels and enhance retention time, 

have amplified the evapoconcentration of salts and reduced the 

effectiveness of natural salt removal processes such as flushing 

and deflation (removal by wind) (Hamilton and others, 1989). 

Further, this increase in salinity has been linked to alteration 

of vegetation communities that provide important wildlife 

habitat. For example, Hamilton and others (1989) observed 

that salt-tolerant plants had replaced cat-tails (Typha sp.) and 

bulrushes (Scirpus sp.), and submerged aquatic species such as 

pondweeds (Potamogeton sp.) and widgeongrass (Ruppia sp.) 

had largely disappeared. Additionally, declines in waterfowl 

use and productivity have been attributed to increased salinity 

and subsequent changes in vegetative composition.

To ensure the long-term success of Bowdoin NWR in 

providing healthy, sustainable habitat for waterfowl and other 

wildlife, the U.S. Fish and Wildlife Service (USFWS) is developing management strategies to maintain salt concentrations 

within ranges suitable for the establishment of historical plant 

and invertebrate communities. Currently, the USFWS is evaluating various water management scenarios that will enhance 

their ability to export salts and manage salinity levels in lakes 

and marshes on the refuge. Water management scenarios 

considered by the USFWS range from no action to controlled 

and uncontrolled (for example, flushing caused by floods) 
ROCKY

MOUNTAINS

River

Fort Peck Missouri

Lake

Milk

River

MONTANA

Study area

107°44′ 107°36′

48°32′

48°24′

48°16′

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)

0 2.5 5 10 KILOMETERS

Lake Bowdoin

Dry Lake

Pond

Dry Lake

Unit

Drumbo Unit

Lakeside Unit

Lakeside Extension

River

Creek

Dodson South Canal

Bowdoin National

Wildlife Refuge

0 1.25 2.5 5 MILES

Nelson

Reservoir

Milk

Beaver

Figure 1. Location of Bowdoin National Wildlife Refuge in relation to 

Beaver Creek, Milk River, Dodson South Canal, and Nelson Reservoir.

2 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.
Description of Bowdoin National Wildlife Refuge 3

exports of saline water to creeks and rivers (oral commun., 

Comprehensive Conservation Plan planning committee). 

Each of these scenarios will result in varying levels of salinity 

control in lakes and marshes that will ultimately affect wildlife 

habitat in terms of plant and invertebrate communities.

To support the evaluation of water management scenarios, the USFWS requested that the U.S. Geological Survey 

(USGS) provide information on suitable ranges of salinity 

for plants and invertebrates of importance to the refuge. To 

address this need, we present summarized data obtained from 

various sources (for example, books, peer-reviewed publications, reports, subject reviews, unpublished data) that relate 

the occurrences of common aquatic plant and invertebrate 

species to various water and soil quality variables such as 

specific conductance and pH. As part of the review, we also 

discuss general physiological adaptations that allow flora and 

fauna to survive in salt-affected environments.

Purpose and Scope

The goal of this report is to provide the USFWS with 

scientifically based information to facilitate a better understanding of the ecological relations between salinity and 

flora and fauna when developing management strategies. 

Although the primary focus is to provide information to help 

address salinity issues at Bowdoin NWR, the databases, as 

well as concepts and information discussed, are intended to 

be applicable to other areas or refuges. To address our goal, 

we conducted an extensive search to locate existing scientific 

literature that reported occurrences of plants and invertebrates 

in relation to salinity and pH of the water and soil. Information 

from this literature search was incorporated into databases and 

used to address the following objectives:

1. Provide a general overview of salinity concepts and the 

tolerances and adaptations of biota to salinity.

2. Develop databases that the USFWS can use to summarize 

the range of reported salinity values associated with plant 

and invertebrate taxa.

3. Perform database summaries that describe reported salinity ranges associated with plants and invertebrates at 

Bowdoin NWR.

Description of Bowdoin National 

Wildlife Refuge

The 6,294-ha refuge is located in the Glaciated Northern 

Grasslands (level IV ecoregion) (Woods and others, 2002) of 

northeastern Mont., which is characterized by rolling hills and 

plains that are the result of glaciation. In general, glacial and 

alluvial deposits overlay bedrock, and the region is characterized by saline-sodic soils (sodium and sulfate are the dominant ions) and grasslands that contain numerous depressional 

wetlands. Terrestrial soils of the refuge consist primarily of 

Bigsag clays and the Phillips-Elloam complex. The portion 

(36 percent) of the refuge inundated by lakes is bordered, and 

likely underlain, by Bigsag and Bowdoin soils (Kendy, 1999), 

which are relatively fine grained with low permeability and a 

high salt content. An inventory of primary soils and complexes 

on the refuge is presented in table 1.

Refuge habitats consist primarily of short- and mixedgrass prairie, planted dense nesting cover, shrublands, and 

both brackish and freshwater wetlands. Wetlands make up 

about 46 percent (2,924 ha) of the refuge, with the majority of this area consisting of five large, shallow lakes: Lake 

Bowdoin, Drumbo Unit, Dry Lake Unit, Dry Lake Pond, and 

Lakeside Unit (fig. 1). Lake Bowdoin, the largest (2,209 ha) 

and most prominent lake, is an oxbow of the preglacial 

Missouri River. The modern Missouri River flows nearly 

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.

Direct precipitation and diverted water from the Milk 

River via the Dodson South Canal are the main sources of 

water entering the refuge, while irrigation drainage returns 

from nearby agricultural lands, overland runoff, floodwaters 

from adjacent drainages (for example, Beaver Creek), and 

groundwater discharges from saline seeps are secondary 

sources. An annual water allocation of 431 ha-m from the 

Milk River Irrigation Project that flows through Dodson South 

Canal provides a relatively reliable source of water to the 

refuge; however, this allocation can fluctuate annually and is 

contingent on the amount of water available for distribution. 

Measured water deliveries from the years 1938 to 2006 ranged 

from 70 to 1,419 ha-m and averaged 597 ha-m. Water losses 

are attributed primarily to evaporation and infrequent surface 

outflows to neighboring Beaver Creek. The average annual 

evaporation (92.71 cm) (Kendy, 1999) greatly exceeds the 

mean annual precipitation (27.94–35.56 cm) (U.S. Department 

of Agriculture, 2004) in this semiarid region; thus, the refuge 

experiences a water deficit for much of the year. Currently, 

USFWS personnel manage refuge wetlands for waterfowl and 

other wildlife by using a system of channels and dikes that 

allows for the capture, storage, and movement of water within 

the refuge.

A total of 210 plant species have been documented on 

the refuge (app. 1). Dominant emergent vegetation includes 

hardstem bulrush (Scirpus acutus), alkali bulrush (S. paludosus), common threesquare (S. pungens), and cat-tail (Typha 

latifolia). Other notable species that occur along the shores 

of lakes and marshes include pickleweed (Salicornia rubra) 

and saltgrass (Distichlis spicata). Common aquatic vegetation 

includes sago pondweed (Potamogeton pectinatus), widgeongrass (Ruppia maritima), and watermilfoil (Myriophyllum 

exalbescens) (Weydemeyer and Marsh, 1936; Hamilton and 

others, 1989; Johnson, 1990). Information on invertebrate 

communities is relatively limited; however, 32 invertebrate 

taxa have been documented within lakes and marshes on the 
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]

Soil Slope range, 

in percent

Area, 

in hectares Percent of refuge Cumulative percent 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

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 (Caenidae), and water fleas (Cladocera) (Johnson, 1990).
Primer on Salinity Concepts 5

Methods

We developed plant and invertebrate databases by 

conducting an extensive search to locate existing scientific 

literature that reported occurrences of plants and invertebrates 

in relation to salinity and pH of the water and soil. Literature 

and keyword (for example, wetlands, salinity, salt, invertebrates, plants) searches were conducted by using various 

electronic databases such as FirstSearch, Cambridge Scientific 

Abstracts, Water Resources Abstracts, Web of Science, Google 

Scholar, and others. Additionally, we used unpublished data 

gathered as part of various investigations conducted primarily 

by the USGS Northern Prairie Wildlife Research Center.

Information obtained from this search was entered 

into spreadsheet databases to facilitate data summaries. The 

detailed databases contain 27 fields that include variables 

such as taxonomic identification, values for salinity and pH, 

wetland classification, location of study, and source of data. 

Appendix 3 provides a detailed description of each variable 

found in the databases, and appendix 4 includes an annotated 

description of the data obtained from each source to provide 

users information on the utility or applicability of the data to 

their region of interest.

Primer on Salinity Concepts

Salinity Definitions and Measurements

Salinity refers to the concentration of dissolved or 

soluble salts in water and soil. Salts are generally defined as 

ionic compounds which in water solution yield a cation other 

than hydrogen (H+

) and an anion other than hydroxyl (OH-) 

or dioxygen (O2

-

). As an example, the following equation 

displays the chemical formula for common table salt, sodium 

chloride:

Na+ + Cl- ↔ NaCl

Common ions (cations and anions) found in northern 

wetlands, including those within Bowdoin NWR (fig. 2), are 

calcium (Ca2+), magnesium (Mg2+), potassium (K+

), sodium 

(Na+

), bicarbonate (HCO3

-

), carbonate (CO3

2-), chloride (Cl-

), 

and sulfate (SO4

2-) (Swanson and others, 1988; Nimick, 1997; 

Bauder and others, 2007). When removed from solution (for 

example, through evaporation), principal salts in water and 

soils at Bowdoin NWR include sodium sulfate (Na2

SO4), 

sodium bicarbonate (NaHCO3

), calcium carbonate (CaCO3), 

calcium sulfate (CaSO4

), and magnesium sulfate (MgSO4); 

minor amounts of chlorine (Cl-

) and fluoride (F-) salts are also 

found (Bauder and others, 2007).

Measures of salinity for both soil and water are usually 

expressed as conductance (that is, specific conductance, electrical conductivity) or total dissolved solids (TDS). Specific 

conductance is a measure of the capacity of an aqueous 

solution to conduct an electrical current and is highly correlated with the concentration of ions in solution. Conductance 

measurements also are highly dependent on temperature 

because an increase in the temperature of a solution will 

decrease viscosity and increase the mobility of ions in solution; a rise in temperature could also increase the number 

of ions in solution because of dissociation of molecules. 

Because of the need to compare conductivity values collected 

at various temperatures by using differing equipment (that is, 

probes that differ with respect to electrode distance and area), 

measurement devices are typically temperature-compensated 

and provide standardized readings at 25°C with an electrode 

distance of 1 cm (for example, 1,000 μS cm-1 at 25°C) (Lind, 

1985). Specific conductance is commonly reported as millisiemens (mS), microsiemens (μS), millimhos (mmhos), or 

micromhos (μmhos). Siemens and mhos are equivalent units 

that are reciprocal of the ohm, the International System (SI) 

unit of electrical resistance.

Total dissolved solids is a measure of all inorganic and 

organic substances dissolved in water and is usually presented 

as parts per thousand (ppt, ‰), percent (%), milligrams per 

liter (mg L-1), or total mass (in grams [g]). The concentration 

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 approximated from specific conductance measurements. For example, 

specific conductance is typically multiplied by an empirical 

factor that generally varies from 0.5 to 1.0 to estimate TDS 

(Lind, 1985). 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). As a reference, in figure 3, three 

common salinity reporting units (μS cm-1, mg L-1, and ppt) are 

compared in relation to plant community salinity categories 

(table 2) described by Stewart and Kantrud (1972).

Sodicity

Sodicity refers to the sodium ion concentration in a soil 

or solution. Substrates classified as sodic have high sodium 

ion concentrations relative to other cations such as calcium or 

magnesium. High sodium concentrations cause soil aggregates to swell and disperse, resulting in destruction of soil 

structure. In contrast, calcium and magnesium ions cause soil 

particles to flocculate, forming aggregates that enhance and 

stabilize soil structure. The extent of soil structure destruction 

or enhancement caused by cations is not only a function of 

concentration but also is related to the organic matter content, 

carbonate and sulfate concentrations (for example, calcite and 

gypsum), leaching potential, and texture of the soil. The loss 

of soil structure can affect plant growth by impairing hydraulic conductivity (water movement through soil), drainage, 

gas exchange, and rooting depth. Additionally, sodicity—and 

salinity to a certain extent—is often accompanied by high 
Figure 2. Concentration of major ions in Lake Bowdoin during April–August 2006 (data provided by the U.S. Fish and Wildlife Service).

0

2,000

4,000

6,000

8,000

10,000

14,000

12,000

APRIL MAY JUNE JULY AUGUST

CONCENTRATION, IN MILLIGRAMS PER LITER

EXPLANATION

Sulfate (SO4

2-) 

Sodium (Na+) 

Bicarbonate (HCO3

-

) 

Magnesium (Mg2+)

Chloride (Cl-) 

Carbonate (CO3

2-) 

Potassium (K+) 

Calcium (Ca2+ ) 

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 

imbalance.

Soils are commonly classified as saline, sodic, or salinesodic on the basis of a combination of three factors: the 

sodium adsorption ratio (SAR), specific conductance, and 

pH (table 3). The SAR is the ratio of sodium to calcium and 

magnesium cations in solution, defined as

In general, when the SAR is above 12, physical properties of 

soil may be altered, and plants can become stressed because 

of ion toxicity and reduced water absorption (osmotic stress). 

We do not have specific field data (see table 3) necessary to 

classify soils found at Bowdoin NWR (table 1); however, on 

the basis of reported ranges of SAR, specific conductance, and 

pH values (table 4) for soils on the refuge, all three soil classes 

likely occur, especially if the classifications were based on the 

upper end of the reported values (table 4).

a relatively wet and dry year. LaBaugh (1989) reviewed data 

from northern prairie wetlands and lakes and reported ranges 

of specific conductance and TDS of 42–472,000 μS cm-1 and 

31–342,000 mg L-1 (approximately 48–534,375 μS cm-1), 

respectively. Additionally, LaBaugh (1989) described studies 

in which specific conductance increased twofold to sixfold and 

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 

fluctuations in water levels that result in the dilution, evapoconcentration, or deflation (blowing of salts from mudflats) 

of salts (Lieffers and Shay, 1983; LaBaugh, 1989; LaBaugh 

and others, 1996). For example, between 1975 and 2007 the 

mean annual salinity in Lake Bowdoin varied from 4,113 (in 

1990) to 28,908 (in 1984) μS cm-1 (fig. 4). During this 33-year 

period, Lake Bowdoin was classified as moderately brackish 9 percent, brackish 79 percent, and subsaline 12 percent 

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 

salinity flux is illustrated when salinity is examined in relation 

to the Palmer Hydrological Drought Index (PHDI), which 

shows long-term cumulative drought and wet conditions 

(fig. 5). Variation in the PHDI over the past 30 years, or since 

salinity has been monitored in Lake Bowdoin, indicates two 

drought periods (1980–85, 1999–2007) and two relatively wet 

periods (1975–80, 1986–98) (fig. 5). During these alternating 

wet and dry periods, salinity levels in Lake Bowdoin tended to 

decrease during wet periods and increase during drought.

Factors Influencing Salinity

When evaluating the suitability of habitats for aquatic 

biota, it is important to recognize that water and soil salinity can vary considerably both temporally and spatially. 

Swanson and others (1988) reported that specific conductance 

measurements for 178 prairie lakes in North Dakota ranged 

from 365 to 70,000 μS cm-1 and that measurements from 

1 pothole wetland varied from 522 to 7,700 μS cm-1 between 
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).

40

50

60

MICROSIEMENS PER CENTIMETER

MILLIGRAMS PER LITER

PARTS PER THOUSAND

Slightly brackish

Brackish

Moderately brackish

Subsaline

Saline

Fresh

20,000

40,000

60,000

80,000

100,000

0

20,000

30,000

40,000

50,000

60,000

0

10,000

30

0

20

10

Seawater

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 University Agriculture 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]

Classification Soil condition Specific conductance, 

μS cm-1 pH SAR

Saline Normal1 >4,000 <8.5 <13

Sodic Poor <4,000 >8.5 >13

Saline-sodic Normal1 >4,000 <8.5 >13

1

Soil retains “normal” physical conditions as long as sodium does not dominate the exchange complex of the soil.

Primer on Salinity Concepts 7
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, at http://soils.usda.

gov/technical/classification/osd/index.html. μS cm-1, microsiemens per centimeter; SAR, sodium adsorption ratio; --, no data]

Soil series Total area, 

in hectares1 Taxonomic family Specific conductance, μS cm-1 pH SAR

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 Argiustolls-- 6.6–7.8 --

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 Argiustolls2,000–4,000 6.6–7.8 --

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 Argiustolls0–4,000 6.6–8.4 --

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 

Aridic Ustorthents

-- 7.9–9.0 --

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, superactive, frigid Aridic Argiustolls

-- 6.6–7.8 --

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 -- -- --

1

Total area of all soil complexes that include the soil series (see table 1).

8 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

The increased salinity during drought is attributed to 

evapoconcentration of solutes; however, drought also provides 

mechanisms for removal of salt by deflation and seepage to 

groundwater (Weydemeyer and Marsh, 1936; Hamilton and 

others, 1989; LaBaugh and others, 1996). Superimposed over 

these natural processes of deflation and seepage is the effect 

of water deliveries to the refuge via the Dodson South Canal 

(fig. 6), which can be used to dilute salts or maintain water 

levels during dry periods that would normally result in deflation events. Consequently, manipulating water levels provides 

an opportunity to both trigger and halt deflation episodes.

Decreases in salinity during wet periods are attributed 

to increased freshwater inputs that dilute salt concentrations. 

Further, periodic floodwaters that enter the refuge from Beaver 

Creek not only dilute salts but also flush salts from the system. 

The most notable Beaver Creek flood event that resulted in 

significant salt flushing during the past 30 years occurred 

in 1986 when peak flows exceeded 20,000 ft3/s (fig. 7B); 

this event coincided with a sixfold salinity decrease in Lake 

Bowdoin (fig. 5). There are insufficient data to calculate the 

frequency of Beaver Creek flood events (fig. 7), but refuge 

personnel estimate that the historical average of floodwater 

occurrence on the refuge was once every 3–5 years (Rodney 

and Mohrman, 2006; 2007). More recent observations by 

refuge staff suggest, however, that the frequency of flood 

events has decreased to once every 7–10 years. The reduced 

flood frequency is believed to have been caused by the establishment of small impoundments and irrigation diversions in 
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.

YEAR

1975 1980 1985 1990 1995 2000 2005

SPECIFIC CONDUCTANCE,

IN MICROSIEMENS PER CENTIMETER

0

10,000

20,000

30,000

40,000

50,000

Saline

Subsaline

Brackish

Moderately brackish

Slightly brackish

Fresh

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.
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.

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.

YEAR

1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 

PALMER HYDROLOGICAL DROUGHT INDEX (PHDI)

-10

-5

0

5

10

SPECIFIC CONDUCTANCE,

IN MICROSIEMENS PER CENTIMETER

0

10,000

20,000

30,000

Specific conductance

PHDI

1975

1980

1985

1998

YEAR

PALMER HYDROLOGICAL DROUGHT INDEX (PHDI)

1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 

-10

-5

0

5

10

DELIVERIES FROM MILK RIVER VIA DODSON

SOUTH CANAL, IN HECTARE METERS

1975

1980

1985

1998

0

500

1,000

1,500

Deliveries

PHDI

10 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.
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.

YEAR

PALMER HYDROLOGICAL DROUGHT INDEX (PHDI)

1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 

-10

-5

0

5

10

MONTHLY BEAVER CREEK FLOW,

IN HECTARE METERS

1975

1980

1985

1998

0

10,000

20,000

30,000

Flow

PHDI

A

YEAR

PALMER HYDROLOGICAL DROUGHT INDEX (PHDI)

1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010 

-10

-5

0

5

10

ANNUAL BEAVER CREEK PEAK FLOW,

IN CUBIC FEET PER SECOND

Peak flow

PHDI

20,000

10,000

0

B

Primer on Salinity Concepts 11
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.

YEAR

1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 

CUMULATIVE NUMBER OF RESERVOIRS

All reservoirs

Stock

Wildlife

Irrigation

A

0

200

400

600

800

1,000

1,200

1,400

YEAR

1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 

CUMULATIVE CAPACITY, IN HECTARE METERS

All reservoirs

Stock

Irrigation

Wildlife

1,000

2,000

3,000

4,000

0

B

12 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.
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.

Fresh

Slightly brackish

Moderately brackish

Brackish

Subsaline

Saline

NUMBER OF PLANT TAXA

0

10

20

30

40

50

Tolerances and Adaptations of Biota to Salinity 13

The ability of plants to survive saline environments 

depends on their capacity to tolerate salts throughout various 

life cycle stages. Plants capable of growth, flowering, and 

seed production when rooted in soils with sodium chloride 

concentrations greater than 0.5 percent are commonly referred 

to as halophytes (Baskin and Baskin, 1998). In contrast, plants 

incapable of survival and reproduction in these environments 

are referred to as glycophytes, or salt-sensitive plants.

The ability of halophytes to grow and survive in saline 

environments is possible because of various physiological and 

structural adaptations. Seed germination is affected by many 

abiotic factors such as temperature, photoperiod, soil moisture, and pH; however, salinity is one of the primary factors 

that limits seed germination in both glycophytes and halophytes. Baskin and Baskin (1998) reported that concentrations 

of sodium chloride ranging from about 3,000 (0.3 percent, 

approximately 4,688 μS cm-1) to 100,000 mg L-1 (10 percent, 

approximately 156,250 μS cm-1) were sufficient to reduce 

germination rates of many halophytes to about 10 percent 

Tolerances and Adaptations of Biota to 

Salinity

Plants

Differences in salinity are often reflected in the species 

composition of vegetation; thus, salinity has been incorporated as a modifier in many lake and wetland classification 

systems (Stewart and Kantrud, 1971; Cowardin and others, 

1979). Stewart and Kantrud (1972) related the occurrence 

of 135 plant species in prairie pothole wetlands to six salinity categories (table 2). Of these species, nearly 70 percent 

did not occur in wetlands classified as brackish, subsaline, 

and saline (salinity levels greater than 5,000 μS cm-1), and 

about 35 percent did not occur when salinities exceeded 

2,000 μS cm-1 (moderately brackish, fig. 9). Hence, relatively few inland wetland plant species can persist in saline 

environments.
14 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

(table 5). Reduced germination in halophytes is generally 

attributed to salinity-induced osmotic stress that inhibits water 

uptake and production of compounds (for example, enzymes) 

necessary for germination and embryo development (Baskin 

and Baskin, 1998). The inhibitory effects also vary by species 

of salt; for example, potassium chloride (KCl) or magnesium 

chloride (MgCl2

) may affect the germination of a certain plant 

species more than do other salts, such as sodium chloride or 

calcium chloride (CaCl2

). Additionally, natural ecosystems 

typically contain a mixture of salts (Swanson and others, 

1988), and the presence of one salt may enhance or reduce the 

adverse effects of another (Baskin and Baskin, 1998; Tester 

and Davenport, 2003).

Many halophytes exhibit physiological adaptations that 

allow seeds to remain dormant and not germinate during 

hypersaline conditions. This trait allows seeds to persist in the 

soil seed bank until environmental conditions are favorable 

for germination; for example, seeds will not germinate until 

climatic events lower salinity through dilution or leaching. 

Ultimately, dormancy release mechanisms help improve 

survival during seedling development, which is one of the 

most salt-sensitive periods of the halophyte life cycle.

After germination and early establishment, salinity can 

affect halophyte metabolism (for example, decreased rates of 

photosynthesis and respiration), as well as stimulate anatomical and morphological changes (for example, increased 

succulence, stomata number and size, cuticle thickness, tylose 

(bladder-like growth) development, early onset of lignification, and changes in diameter and number of xylem vessels) 

(Poljakoff-Mayber, 1975). These anatomical adaptations allow 

halophytes to tolerate osmotic gradients associated with saline 

environments.

Mature plants absorb salts from the soil solution through 

their root systems and lose water to the atmosphere via 

transpiration. These processes result in the accumulation of 

salts in plant tissues and often lead to high salt concentrations and death in nonhalophytes. In contrast, halophytes have 

evolved various mechanisms to survive saline environments. 

Generally, two principal types of halophytes are recognized: 

those that tolerate excess salts and those that resist excess 

salts (Ungar, 1974; Baskin and Baskin, 1998). Salt-tolerant 

halophytes either withstand high intracellular salt concentrations (salt-enduring) or secrete excess salts (salt-excluding) by 

means of salt glands, by accumulating salts in special hairs, 

or by transporting salts accumulated in plant structures back 

to the roots. In contrast, salt-resistant halophytes either do not 

absorb salts from the soil solution or do not transport salts to 

sensitive organs or leaves.

High salt concentrations in the soil can also affect plant 

growth by reducing the water potential gradient. For water to 

move from the soil to plant tissues, the water potential of the 

plant must be lower (that is, more negative) than that of the 

soil. The water potential of the soil is related to the concentration of dissolved solids in the soil-water solution; a high 

concentration of salts in the soil reduces the soil water potential and decreases the water potential gradient. Essentially, 

saline soils sequester water, making it unavailable to plants. In 

addition, as water evaporates from shallow, saline lakes (such 

as Lake Bowdoin), salts often accumulate as surficial crusts. 

This crusting can reduce water infiltration, thereby decreasing the availability of water to plants growing on the edges of 

lakes.

Effective management of salt concentrations to promote 

the development of vegetation for wildlife habitat will require 

an understanding of plant adaptations to salinity, which vary 

by life stage. The general overview provided above is intended 

only to highlight some of the more common adaptations and 

life-cycle strategies that should be considered in concert with 

other important determinants of plant community composition. For example, the simple plant/salinity relations discussed 

above should be considered in relation to many other factors 

that significantly influence establishment, growth, and reproduction. Soil pH, texture, temperature, redox potential, moisture, and water depth all influence plant establishment and 

growth in wetland environments (for example, Stewart and 

Kantrud, 1972; van der Valk and Davis, 1978; van der Valk, 

1981; Fredrickson and Laubhan, 1994; Winter, 2003).

Invertebrates

As with that of plants, the composition of aquatic 

invertebrate communities can vary greatly in response to 

changes in salinity. In general, relatively fresh aquatic systems 

contain diverse invertebrate communities, whereas highly 

saline systems tend to be dominated by relatively few taxa. 

It is important to note, however, that even though species 

richness declines in highly saline systems, biomass often 

does not (for example, Euliss and others, 1991). Common 

invertebrates of highly saline systems include copepods and 

rotifers (particularly some species of the genera Brachionus

and Hexarthra), as well as a small number of insects from 

the orders Diptera (typically from the families Chironomidae and Culicidae [genus Aedes]) and Coleoptera (Pennak, 

1989). Shifts in invertebrate community structure that result 

from increased salinities in prairie wetlands typically include 

displacement of amphipods (for example, Hyalella) and 

anostracans (for example, Branchinecta) by more salt-tolerant 

brine shrimp (Artemia). In addition, a narrow range of insect 

taxa such as brine flies (Ephydridae) and salt-tolerant water 

boatmen (Corrixidae) have been found to dominate systems 

with high salt concentrations (Swanson and others, 1988; 

Euliss and others, 1999). For example, the insect community 

of hypersaline (up to 300 mS cm-1) ponds in California was 

dominated by 1 corixid (Trichocorixa reticulate) and 1 midge 

(Tanypus grodhausi) (Euliss and others, 1991), whereas 

1 midge species dominated the benthic community of an 

alkaline lake in central Washington where salinity levels were 

about 17,000 mg L-1 (approximately 26,563 μS cm-1). Species 

richness in this lake increased two to eight times as salinity 

levels were reduced to about 1,500 mg L-1 (approximately 

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 1

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

Genus Species

NaCl concentration

Genus Species

NaC

Molarity mg L-1 Molarity 

Melilotus indica 0.05 2,922.0 Sporobolus virginicus 0.26–>0.60 

Salicornia bigelovii 0.05–0.06 2,922.0–3,506.4 Suaeda nudiflora 0.26 

Sarcobatus vermiculatus 0.06–0.15 3,506.4–8,766.0 Aster tripolium 0.34–0.60 

Zygophyllum qatarense 0.07 4,090.8 Atriplex halimus 0.34 

Distichlis spicata 0.09 5,259.6 Atriplex patula 0.34 

Glaux maritima 0.09 5,259.6 Cotula coronopifolia 0.34 

Plagianthus divaricatus 0.09 5,259.6 Mesembryanthemum australe 0.34 

Scirpus americanus 0.09 5,259.6 Phragmites communis 0.34 

Scirpus olneyi 0.09 5,259.6 Polypogon monspeliensis 0.34 

Selliera radicans 0.09 5,259.6 Prosopis farcta 0.34–0.6 

Spergularia salina 0.09 5,259.6 Salicornia patula 0.34 

Triglochin striatum 0.09–0.17 5,259.6–9,934.8 Atriplex griffithii 0.35 

Iva annua 0.13–0.17 7,597.2–9,934.8 Cochlearia danica 0.43 

Zannichellia pedunculata 0.13 7,597.2 Rumex crispus 0.43 

Ceratoides lanata 0.17–0.34 9,934.8–19,869.6 Puccinellia distans 0.45 

Hordeum jubatum 0.17–0.31 9,934.8–18,116.4 Puccinellia lemmoni 0.45 

Juncus maritimus 0.17 9,934.8 Halopeplis amplexicaulis 0.50 

Melaleuca ericifolia 0.17 9,934.8 Atriplex triangularis 0.51 

Myrica cerifera 0.17 9,934.8 Limonium bellidifolium 0.60 

Schoenus nitens 0.17 9,934.8 Limonium vulgare 0.60 

Scirpus robustus 10.17–0.20 19,934.8–11,688.0 Salsola kali 0.60 

Spergularia marina 0.17–0.34 9,934.8–19,869.6 Salicornia pacifica 0.68 

Suaeda linearis 0.17 9,934.8 Spartina alterniflora 0.68 

Melilotus segetalis 0.20 11,688.0 Puccinellia festucaeformis 0.75 Melilotus messanensis >0.20 >11,688.0 Cressa cretica 0.85 Sesuvium portulacastrum 0.2–>0.60 11,688.0–>35,064.0 Salicornia europaea 0.85 Atriplex canescens 0.21 12,272.4 Suaeda depressa 0.85 

Salicornia brachystachya 10.24 114,025.6 Tamarix pentandra 10.85 

Salicornia dolichostachya 10.24 114,025.6–14,025.6 Suaeda japonica 0.90 

Atriplex prostrata 0.26 15,194.4 Kochia americana 1.02 

Atriplex polycarpa 0.26 15,194.4 Zygophyllum dumosa 1.51 

Plantago coronopus 0.26 15,194.4 Salicornia herbacea 1.70 

Salicornia emerici 0.26 15,194.4

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 

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 

species can survive in highly saline waters. Most invertebrate 

species are unable to maintain their internal salt and water 

balance when placed in a solution that contains greater than 

0.9 percent (9,000 mg L-1 or approximately 14,063 μS cm-1) 

salts, or roughly 25 percent the salt content of seawater 

(Pennak, 1989; Hart and others, 1991). Only a narrow assemblage of euryhaline organisms (for example, inhabitants of 

estuaries and tide pools) can thrive in both freshwater and 

marine environments. In general, habitats that support a large 

number of common freshwater fauna have concentrations of 

TDS that range from about 10 to 1,000 mg L-1 (approximately 

16 to 1,563 μS cm-1) (Pennak, 1989; Hart and others, 1991). 

Salinity levels in inland lakes and wetlands, however, can vary 

considerably, and the few species that can tolerate extremely 

high salt concentrations require highly specialized and efficient osmoregulatory systems to maintain a proper internal 

balance of salt and water. Since water will tend to move into 

the hypertonic (high salt) tissues of freshwater invertebrates 

through permeable surfaces such as epithelia, cuticle, chitin, 

and gills (Pennak, 1989), these species have developed various 

adaptations such as contractile vacuoles, flame bulb systems, 

nephridia, and various glandular structures that are capable 

of forming highly dilute urine (hypotonic to body fluids) that 

is excreted in great quantities. As a result, concentrations of 

internal salts and water remain somewhat constant.

Similar to that of plants, the salinity tolerance of aquatic 

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 

others, 2005). Many species of Coleoptera and Hemiptera 

are able to exist in prairie wetlands in spite of highly variable 

salinities because their adult stages are capable of flying to 

suitable habitats. Conversely, many nonflying aquatic invertebrates and immature stages of flying insects must rely on 

various adaptations that allow them to endure highly saline 

habitats, including burrowing into substrates or producing ephippia (resting eggs), cysts, or waterproof secretions 

(Euliss and others, 1999). For example, cladocerans such 

as Daphnia sp. produce ephippia that remain viable in the 

substrate until conditions are favorable for hatching. Because 

of the high mobility of some adult invertebrates and the inability of many immature aquatic invertebrates to cope with high 

salinities, the presence of adults in highly saline wetlands does 

not necessarily indicate the presence of a persistent population 

(Pinder and others, 2005).

Database Summary Statistics

Plants

The plant database (http://pubs.usgs.gov/sir/2009/5098/

downloads/databases_21april2009.xls) contains information 

on 411 taxa obtained from 17 studies conducted in 9 States 

and 3 Canadian Provinces, as well as literature that did not 

specify a location (apps. 4 and 5). Figure 10 is a graphic 

portrayal of the compiled data by the number of plant taxa 

in relation to source (fig. 10A), location (fig. 10B), wetland 

classification (fig. 10C), wetland indicator category (fig. 10D), 

plant physiognomy/life-span (fig. 10E), and salinity categories 

(fig. 10F). We used this database to compute the minimum and 

maximum reported specific conductance and pH by substrate 

(soil, water) for each plant taxon (app. 5).

Invertebrates

The invertebrate database (http://pubs.usgs.gov/

sir/2009/5098/downloads/databases_21april2009.xls) contains 

330 taxa obtained from 7 sources (apps. 4 and 6) that represent 

studies from 3 States, 2 Canadian Provinces, and Australia. 

Figure 11 provides a summary of the number of invertebrate 

taxa by source (fig. 11A), location (fig. 11B), wetland classification (fig. 11C), taxonomic classification (fig. 11D), and 

salinity maxima (fig. 11E). Appendix 6 contains the minimum 

and maximum specific conductance and pH for invertebrate 

taxa included in the database.

Considerations in Using Information from the 

Plant and Invertebrate Databases

The plant and invertebrate databases represent information obtained from an extensive search of the scientific 

literature and unpublished data; however, it is not considered comprehensive. Data are derived primarily from field 

studies conducted in natural wetland ecosystems rather than 

from experimentally derived tolerance values. Users should 

consider the following aspects and limitations when creating 

summaries or conducting analyses:

1. The data represent information from numerous geographic locations (for example, Iowa, North Dakota, 

Prairie Pothole Region, Manitoba) and wetland types (for 

example, palustrine, lacustrine, salt-marsh); users may 

want to focus on, or exclude data from, a certain region or 

wetland type.
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.

Kantrud, 1990

Kantrud, 1991

Kantrud, 1996

Ungar and others, 1979

Grosshans and Kenkel, 1997

Ungar and Riehl, 1980

Ungar and others, 1969

Ungar, 1970

Rawson and Moore, 1944

Ungar, 1967

Johnson, 1990

Ungar, 1966

Hammer and Heseltine, 1988

Ungar, 1974

Kantrud and others, 1989

Stewart and Kantrud, 1972

NUMBER OF PLANT TAXA

0

50

100

150

200

250

300

350

SOURCE OF DATA

A

LOCATION OF STUDY

Manitoba

Ohio

Nebraska

Alberta

Kansas

Bowdoin NWR, Montana

Kansas; Oklahoma

Saskatchewan

Various

Montana

Prairie Pothole Region

Iowa

Minnesota

South Dakota

North Dakota

NUMBER OF PLANT TAXA

0

50

100

150

200

250

300

B

WETLAND CLASSIFICATION

CELL

PEMB

Palustrine

Salt-marsh

Lacustrine

Unknown

PEMA

PEM

PEMC

PEMF

NUMBER OF PLANT TAXA

0

50

100

150

200

250

300

350

C

WETLAND INDICATOR CATEGORY

FACWFACFACUFACW+

N/A

FAC

FACU

UPL

FACW

OBL

NUMBER OF PLANT TAXA

0

20

40

60

80

100

120

140

D

PLANT PHYSIOGNOMY/LIFE-SPAN CATEGORY

H-Vine

A-Sedge

Cryptogam

B-Forb

A-Grass

Tree

Shrub

P-Sedge

P-Grass

A-Forb

P-Forb

NUMBER OF PLANT TAXA

0

20

40

60

80

100

120

140

160

180

200

E

SALINITY CATEGORY

Fresh

Slightly brackish

Moderately brackish

Brackish

Subsaline

Saline

NUMBER OF PLANT TAXA

0

20

40

60

80

100

120

140

160

Water

Soil

F

USGS, unpublished a 

Database Summary Statistics 17
SOURCE OF DATA

NUMBER OF INVERTEBRATE

TAXONOMIC ORDERS

0

5

10

15

20

25

A

McCarraher, 1970

Lancaster and Scudder, 1987

Kay and others, 2001Tangen, unpublished

Rawson and Moore, 1944

USGS, unpublished b

Johnson, 1990

LOCATION OF STUDY

Nebraska

British Columbia

Australia

Bowdoin NWR, Montana

Saskatchewan

North Dakota

NUMBER OF INVERTEBRATE

TAXONOMIC ORDERS

0

5

10

15

20

25

B

WETLAND CLASSIFICATION

Riverine

Lacustrine

PEMF

PEMC

NUMBER OF INVERTEBRATE

TAXONOMIC ORDERS

0

5

10

15

20

25

C

TAXONOMIC CLASSIFICATION

Phylum

Class

Order

Family

Genus

Species

NUMBER OF INVERTEBRATE TAXA

0

20

40

60

80

100

120

140

160

180

D

SALINITY MAXIMA

Fresh

Slightly brackish

Moderately brackish

Brackish

Subsaline

Saline

NUMBER OF INVERTEBRATE FAMILIES

0

5

10

15

20

E

Figure 11. Number of taxonomic orders from the invertebrate 

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 

were reported at each level of taxonomic classification. E, The 

number of taxa from the invertebrate database in relation to plant 

community salinity categories presented in table 2. Invertebrate 

taxa were included in each category on the basis of the maximum 

reported salinity of water.

18 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.
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 11 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

(number of taxa)

FAC (10), FAC- (1), FACU (26), FACU- (2), FACW (9), FACW+ (1), OBL (30), 

UPL (24), n/a (7)

Plant physiognomy/life-span3 category

(number of taxa)

A-forb (19), A-grass (6), A/P-forb (3), B-forb (1), cryptogam (2), H-vine (1), 

P-forb (45), P-grass (17), P-sedge (7), shrub (5), tree (4)

1

See variable “CLASSIFICATION” in app. 3 for definitions.

2

See variable “WIC” in app. 3 for definitions.

3

See variable “PHYSIOGNOMY” in app. 3 for definitions.

Application to Bowdoin National Wildlife Refuge 19

2. To avoid misinterpretation, users should always consider 

the database variables DATA_TYPE and CONDUCTIVITY_CODE. DATA_TYPE defines whether the waterquality value (for example, specific conductance) was 

reported as a single measurement or as a summarized 

value such as a mean, median, minimum, or maximum. 

For consistency, salinity units were converted to microsiemens in the databases. The variable CONDUCTIVITY_CODE identifies whether the original study reported 

the value as microsiemens, or if it was converted (see 

Methods, above).

3. For consistency, taxonomic and common names for plants 

were based on The Great Plains Flora Association (1986) 

when possible. Therefore, the variables GENUS and SPECIES do not necessarily represent the scientific names 

reported by the original source; the variables GENUS_R 

and SPECIES_R present the scientific name reported by 

the original study.

4. Users of the invertebrate database must first determine 

which taxonomic classification is most appropriate since 

individual studies report invertebrate occurrences at varying degrees of resolution (for example, Order, Family, 

Genus, Species). Resolution will be lost when using a 

higher taxonomic classification such as Order, whereas 

using a lower taxonomic classification such as Genus may 

result in the loss of some information on occurrence.

5. Certain taxa in both databases may have only been 

reported in a single or small number of studies. Therefore, salinity values for specific taxa may be limited by 

a paucity of data. Additionally, many studies that report 

salinity values are conducted in highly saline systems and 

focus only on salt-tolerant taxa; therefore, many common 

freshwater taxa may not be well represented.

6. It is important to consider that many abiotic factors that 

control wetland plant and invertebrate community composition are highly dynamic and exhibit seasonal and annual 

fluctuations. Therefore, salinity or other measurements 

recorded on a given day or year may not be representative 

of the site during other time periods. Consequently, the 

occurrence of plant and invertebrate taxa recorded at a site 

may be influenced by factors, or values of factors, other 

than those occurring at the time of sampling.

7. For both plants and invertebrates, tolerance to salinity can 

vary significantly depending on life stage. Mature plants 

may be able to survive in saline conditions as long as the 

proper conditions existed during the period of germination, which is often earlier in the year when snowmelt and 

spring rains dilute waters and result in lower salinities. 

Similarly, presence of mobile adult invertebrates does not 

necessarily indicate the presence of a persistent population.

Application to Bowdoin National 

Wildlife Refuge

Plants

There are about 210 plant species documented on 

Bowdoin NWR according to a vegetation survey conducted 

in the late 1980s and early 1990s (app. 1). Of these plants, 

the database compiled for this report includes information 

for 110 species that was obtained from 13 known locations 

distributed among 9 types of aquatic systems, 8 wetland 

indicator categories, and 11 plant physiognomy/life-span 

categories (table 6). On the basis of this information, at least 

some taxa in each plant physiognomy and wetland indicator 

category can tolerate fresh to moderately brackish habitats 

(fig. 12; based on 10th and 90th percentile ranges); however, 

most taxa within each category (based on mean and median as 

measures of central tendency) tolerate only slightly brackish 
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).

SPECIFIC CONDUCTANCE OF WATER,

IN MICROSIEMENS PER CENTIMETER

PLANT PHYSIOGNOMY/LIFE-SPAN CATEGORY Cryptogam

A-forb

A/P-forb

P-forb

A-grass

P-grass

P-sedge

Shrub

Tree

A

Saline

Subsaline

Brackish

Moderately

brackish

Slightly

Fresh brackish

10 100 1,000 10,000 100,000

SPECIFIC CONDUCTANCE OF SOIL,

IN MICROSIEMENS PER CENTIMETER

PLANT PHYSIOGNOMY/LIFE-SPAN CATEGORY Cryptogam

A-forb

A/P-forb

B-forb

P-forb

A-grass

P-grass

P-sedge

Shrub

Tree

H-vine B

Saline

Subsaline

Brackish

Moderately

brackish

Slightly

Fresh brackish

10 100 1,000 10,000 100,000

20 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.
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

SPECIFIC CONDUCTANCE OF WATER,

IN MICROSIEMENS PER CENTIMETER

WETLAND INDICATOR CATEGORY

OBL

FACW+

FACW

FAC

FACU

FACUUPL

10 100 1,000 10,000 100,000

C

Saline

Subsaline

Brackish

Moderately

brackish

Slightly

Fresh brackish

SPECIFIC CONDUCTANCE OF SOIL,

IN MICROSIEMENS PER CENTIMETER

WETLAND INDICATOR CATEGORY

OBL

FACW+

FACW

FAC

FACU

FACUUPL

Saline

Subsaline

Brackish

Moderately

brackish

Slightly

Fresh brackish

D

10 100 1,000 10,000 100,000

Application to Bowdoin National Wildlife Refuge 21
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 

found on the refuge (see Description of Bowdoin National 

Wildlife Refuge) exhibited the greatest tolerance to salinity. 

Examples include hardstem bulrush (Scirpus acutus), alkali 

bulrush (S. paludosus), common threesquare (S. americanus), 

saltgrass (Distichlis spicata), pickleweed (Salicornia rubra), 

sago pondweed (Potomogeton pectinatus), and widgeongrass (Ruppia maritima). Few of these species, however, 

appear to tolerate salinities above brackish (approximately 

15,000 μS cm-1).

Invertebrates

Most of the taxonomic orders and families of aquatic 

invertebrates known to occur on the refuge (app. 2) are represented in the database (figs. 15, 16). Examination of invertebrates at the taxonomic level of family (fig. 16) indicates that 

most families can occur in fresh to brackish habitats; however, 

the greatest diversity tends to occur within slightly to moderately brackish habitats. Similar to that of plants, information 

on invertebrates compiled in the database suggests that few 

taxa appear capable of tolerating salinities above brackish 

(approximately 15,000 μS cm-1).

Summary

We reviewed and summarized relevant literature to 

assess the relation between salinity and composition of biotic 

communities associated with freshwater ecosystems. This 

is not a comprehensive literature review and should not be 

considered exhaustive. Further, we do not report experimentally derived threshold values (for example, minimum or 

maximum salt tolerance) for individual plant or invertebrate 

species; instead, we present data obtained from field surveys 

that describe species occurrences across naturally fluctuating ranges of salinity and other environmental factors (for 

example, salt composition, temperature, pH). This approach 

allows for the prediction of broad species assemblages that 

have the potential to occur in wetlands with varying ranges of 

salinity that should facilitate evaluation of various management practices and strategies aimed at enhancing wildlife 

habitat. As an example, wetlands on Bowdoin NWR are 

characterized by elevated salinity levels that have been 

exacerbated by changes in land use and past management 

activities that enhance water inputs into a closed basin. The 

high salinity levels have had a negative impact on the ability 

of wetlands to support waterfowl, which is one of the primary 

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 

environmental conditions that favor establishment of plant 

and invertebrate communities that are highly attractive to 

waterfowl. In addition, refuge staff can use this information to help establish specific (for example, obtain a specific 

range of salinity levels) rather than general (for example, 

reduce salinity levels) management goals. A primary focus of 

this report was to provide information to address the salinity 

problem at Bowdoin NWR, but information in the database, 

as well as concepts and information discussed, is intended to 

be applicable to other areas or refuges. Ideally, information in 

the database will be augmented with additional field studies by 

resource managers (for example, salinity monitoring, vegetation inventories) to facilitate a better understanding of the 

ecological relations between salinity and flora and fauna when 

developing management strategies. 

Acknowledgments

Funding for this project was provided by the U.S. Fish 

and Wildlife Service and the U.S. Geological Survey. We 

thank Charles Dahl, Betty Euliss, Rachel Laubhan, Dave 

Mushet, and Kathy Tribby for assisting in the literature search 

and providing data and technical assistance, and Suzanne 

Roberts for the cover art on this report. Additionally, we thank 

the following people for comments on previous versions of 

this report: Mike Artmann, Wayne King, Jana Mohrman, Rick 

Sojda, Amy Symstad, and Kathy Tribby.
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).

SPECIFIC CONDUCTANCE OF WATER,

IN MICROSIEMENS PER CENTIMETER

10 100 1,000 10,000 100,000

Fresh

Slightly

brackish

Moderately

brackish Brackish

Subsaline

Saline

Agrostis stolonifera

Ruppia maritima

Typha latifolia

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

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

Acknowledgments 23
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).

SPECIFIC CONDUCTANCE OF SOIL,

IN MICROSIEMENS PER CENTIMETER

100 1,000 10,000 100,000

Fresh

Slightly

brackish

Moderately

brackish Brackish

Subsaline

Saline

Agrostis stolonifera

Typha latifolia

Sagittaria latifolia

Ranunculus macounii

Phalaris arundinacea

Potamogeton richardsonii

Polygonum lapathifolium

Mentha arvensis

Echinochloa crusgalli

Ceratophyllum demersum

Eleocharis acicularis

Sagittaria cuneata

Potamogeton pusillus

Myriophyllum exalbescens

Utricularia vulgaris

Beckmannia syzigachne

Alisma subcordatum

Rumex maritimus

Scirpus acutus

Salicornia rubra

Potentilla anserina

Juncus balticus

Hordeum jubatum

Triglochin maritima

Scirpus americanus

Distichlis spicata var. stricta

Potamogeton pectinatus

Scirpus paludosus

Solidago gigantea

Lysimachia ciliata

24 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.
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).

SPECIFIC CONDUCTANCE OF WATER,

IN MICROSIEMENS PER CENTIMETER

Diptera

Coleoptera

Eucopepoda

Anostraca

Cladocera

Tricoptera

Amphipoda

Odonata

Hemiptera

Decapoda

Ephemeroptera

Collembola

Rhynchobdellida

Limnophila

Pharyngobdellida

Pelecypoda

Conchostraca

Gnathobdellida

100 1,000 10,000 100,000

Brackish Subsaline Saline Moderately

brackish

Slightly

Fresh brackish

Acknowledgments 25
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).

SPECIFIC CONDUCTANCE OF WATER,

IN MICROSIEMENS PER CENTIMETER

Libellulidae

Lestidae

Coenagrionidae

Aeshnidae

Velidae

Pleidae

Notonectidae

Mesovelidae

Hebridae

Gerridae

Corixidae

Belostomatidae

Tipulidae

Tabanidae

Syrphidae

Stratiomyidae

Sciomyzidae

Psychodidae

Ephydridae

Dolichopodidae

Dixidae

Culicidae

Corethrinae

Chironomidae

Chaoboridae

Ceratopogonidae

Staphylinidae

Hydrophilidae

Helodidae

Haliplidae

Gyrinidae

Dytiscidae

Curculionidae

Sididae

Polyphemidae

Moinidae

Leptodoridae

Daphnidae

Chydoridae

Bosminidae

Chirocephalidae

Branchinectidae

Artemiidae

Talitridae

Gammaridae Amphipoda

Cladocera

Anostraca

Coleoptera

Diptera

Odonata

Hemiptera

Saline

100 1,000 10,000 100,000

Brackish Subsaline

Moderately

brackish

Slightly

Fresh brackish

26 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.
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28 Database of Relations Between Salinity and Aquatic Biota: Applications to Bowdoin National Wildlife Refuge, Mont.

LaBaugh, J.W., 1989, Chemical characteristics of water in 

northern prairie wetlands, in van der Valk, A., ed., Northern prairie wetlands: Ames, Iowa State University Press, 

p. 56–90.

LaBaugh, J.W., Winter, T.C., Swanson, G.A., Rosenberry, 

D.O., Nelson, R.D., and Euliss, N.H., Jr., 1996, Changes in 

atmospheric circulation patterns affect midcontinent wetlands sensitive to climate: Limnology and Oceanography 

v. 41, p. 864–870.

Lambing, J.H., Jones, W.E., and Sutphin, J.W., 1988, Reconnaissance investigation of water quality, bottom sediment, 

and biota associated with irrigation drainage in Bowdoin 

National Wildlife Refuge and adjacent areas of the Milk 

River Basin, northeastern Montana, 1986-87: Helena, 

Mont., U.S. Geological Survey Water-Resources Investigations Report 87–4243, 71 p.

Laubhan, M.K., Gleason, R.A., Knutsen, G.A., and Euliss, 

N.H., Jr., 2006, A preliminary biological assessment of 

Long Lake National Wildlife Refuge Complex, North 

Dakota: Washington, D.C., U.S. Fish and Wildlife Service 

Biological Technical Publication BTP R6006-2006, 66 p.

Lieffers, V.J., and Shay, J.M., 1983, Ephemeral saline lakes on 

the Canadian prairies—their classification and management 

for emergent macrophyte growth: Hydrobiologia, v. 105, 

p. 85–94.

Lind, O.T., 1985, Handbook of common methods in limnology 

(2d ed.): Dubuque, Iowa, Kendall/Hunt Publishing Company, 199 p.

Moss, B., 1998, Ecology of fresh waters—man and medium, 

past to future (3d ed.): Blackwell Science, Inc., 557 p.

Nimick, D.A., 1997, Hydrology and water chemistry of the 

Benton Lake basin with emphasis on the fate of dissolved 

solids at Benton Lake National Wildlife Refuge, westcentral Montana: Helena, Mont., U.S. Geological Survey 

Water-Resources Investigations Report 97-4100, 79 p.

Parrett, C., 2005, Synthesis of monthly and annual streamflow 

records (water years 1950-2003) for Big Sandy, Clear, Peoples, and Beaver Creeks in the Milk River Basin, Montana: 

Reston, Va., U.S. Geological Survey Scientific Investigations Report 2005–5216, 29 p.

Pennak, R.W., 1989, Fresh-water invertebrates of the United 

States—Protozoa to Mollusca (3d ed.): New York, N.Y., 

John Wiley & Sons, Inc., 628 p.

Pinder, A.M., Halse, S.A., McRae, J.M., and Shiel, R.J., 2005, 

Occurrence of aquatic invertebrates of the wheatbelt region 

of Western Australia in relation to salinity: Hydrobiologia, 

v. 543, p. 1–24.

Poljakoff-Mayber, A., 1975, Morphological and anatomical 

changes in plants as a response to salinity stress, in Poljakoff-Mayber, A., and Gale, J., eds., Plants in saline environments: New York, Heidelberg, Berlin, Springer-Verlag, 

p. 97–117.

Rawson, D.S., and Moore, J.E., 1944, The saline lakes of 

Saskatchewan: Canadian Journal of Research, v. 22, 

p. 141–201.

Rodney, M., and Mohrman, J., 2006, Analysis of stock reservoirs in Beaver Creek Basin, Montana, for downstream 

effects near Bowdoin National Wildlife Refuge: Lakewood, 

Colo., U.S. Fish and Wildlife Service, Mountain-Prairie 

Region, Report dated July 5.

Stewart, R.E., and Kantrud, H.A., 1971, Classification of natural ponds and lakes in the glaciated prairie region: Washington, D.C., U.S. Department of the Interior, Fish and Wildlife 

Service, Resource Publication 92, 57 p.

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.

Swanson, G.A., Winter, T.C., Adomaitis, V.A., and LaBaugh, 

J.W., 1988, Chemical characteristics of prairie lakes in 

south-central North Dakota—their potential for influencing 

use by fish and wildlife: Washington, D.C., U.S. Fish and 

Wildlife Service, Fish and Wildlife Technical Report 18, 

44 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.

Tester, M., and Davenport, R., 2003, Na+

 tolerance and 

Na+

 transport in higher plants: Annals of Botany, v. 91, 

p. 503–527.

The Great Plains Flora Association, 1986, Flora of the Great 

Plains: Lawrence, University Press of Kansas, 1,402 p.

Ungar, I.A., 1966, Salt tolerance of plants growing in saline 

areas of Kansas and Oklahoma: Ecology, v. 47, p. 154–155.

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in northern Kansas: American Midland Naturalist, v. 78, 

p. 98–120.

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v. 83, p. 343–357.

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Reimold, R.J., and Queen, W.H., eds., Ecology of halophytes: New York, N.Y., Academic Press, Inc., p. 235–305.
References

29

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 com-

munities 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.

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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 National Wildlife 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 Engineers, 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

Achillea millefolium; yarrow*

Agropyron cristatum; crested wheatgrass*

Agropyron repens; quackgrass*

Agropyron smithii; western wheatgrass*

Agropyron spicatum; bluebunch wheatgrass

Agrostis stolonifera; redtop*

Alisma subcordatum; common water plantain*

Allium textile; onion

Amaranthus retroflexus; rough pigweed*

Ambrosia artemisiifolia; common ragweed, short ragweed*

Ammannia coccinea; toothcup*

Anemone patens; pasque flower

Apocynum androsaemifolium; spreading dogbane

Arnica fulgens; arnica

Artemisia cana; dwarf sagebrush

Artemisia dracunculus; silky wormwood*

Artemisia frigida; prairie sagewort*

Artemisia ludoviciana; white sage*

Aruncus pubescens; bride’s feathers

Asclepias speciosa; showy milkweed*

Asparagus officinalis; asparagus

Aster ericoides; white aster*

Aster floralvinda; wild aster

Aster puniceus; swamp aster

Aster sp.; wild aster*

Astragalus pectinatus; tine-leaved milk-vetch, narrow-leave

Astragalus racemosus; alkali milk-vetch, creamy poison-vetch

Avena fatua; wild oats*

Beckmannia syzigachne; sloughgrass*

Betula papyrifera; paper birch, canoe birch

Bouteloua gracilis; blue grama

Brassica kaber; charlock*

Bromus inermis; smooth brome*

Bromus japonicus; Japanese brome*

Bromus porteri; nodding brome

Bromus tectorum; downy brome*

Buchloe dactyloides; buffalo grass

Callitriche hermaphroditica; northern water starwort*

Cardaria draba; hoary cress

Carex brevior; fescue sedge*

Carex deweyana; sedge

Carex hookerana; sedge

Carex rostrata; beaked sedge*

Castilleja sulphurea; Indian paintbrush

Centaurea maculosa; spotted knapweed

Cerastium arvense; prairie chickweed

Cerastium brachypodum; nodding chickweed*

Ceratophyllum demersum; hornwort, coontail*

Cheilanthes feei; lip fern

Chenopodium sp.; goosefoot, lamb’s quarter*

Chrysanthemum leucanthemum; ox-eye daisy, marguerite

Chrysopsis villosa; golden aster

Chrysothamnus nauseosus; rabbit brush

Cirsium arvense; Canada thistle, field thistle*

Cirsium undulatum; wavy-leaf thistle

Cirsium vulgare; bull thistle*

Clematis pseudoalpina; virgin’s bower

Clematis terniflora; virgin’s bower

Cleome serrulata; rocky mountain bee plant

Collomia linearis; collomia

Convolvulus arvensis; field bindweed*

Dactylis glomerata; orchard grass*

Distichlis spicata var. stricta; inland saltgrass*

Echinacea angustifolia; purple coneflower

Echinochloa crusgalli; barnyard grass*

Elaeagnus angustifolia; Russian olive*

Eleocharis acicularis; needle spikerush*

Eleocharis macrostachya; common spikerush*

Elodea canadensis; Canadian waterweed*

Elodea nuttallii; waterweed

Elymus canadensis; Canada wild rye*

Elymus glaucus; blue wild rye

Epilobium angustifolium; willow-herb

Equisetum arvense; field horsetail*

Equisetum fluviatile; water horsetail*

Eragrostis cilianensis; stinkgrass

Erigeron glabellus; fleabane

Erigeron sp.; fleabane

Eriogonum flavum; yellow wild buckwheat

Escobaria vivipara; spinystar

Euphorbia esula; leafy spurge*

Euphorbia glyptosperma; ridge-seeded spurge

Fraxinus pennsylvanica; green ash*

Galium boreale; northern bedstraw*

Gaura coccinea; scarlet gaura

Glycyrrhiza lepidota; wild licorice*

Grindelia squarrosa; curly-top gumweed*

Gutierrizia sarothrae; snakeweed

Helianthus laetiflorus; sunflower

Helianthus maximiliani; maximilian sunflower*

Heliotropium curassavicum; seaside heliotrope*

Hordeum jubatum; foxtail barley*

Hydrochloa carolinensis; water grass

Hystrix patula; bottlebrush grass

Iva axillaris; poverty weed

Juncus balticus; baltic rush*

Juncus sp.; rush*

Juniperus communis; common or dwarf juniper

Kochia scoparia; kochia*

Koeleria cristata; junegrass

Lactuca oblongifolia; blue lettuce

Lactuca serriola; prickly lettuce*

Lappula echinata; blue stickseed

Lappula occidentalis; flatspine stickseed

Lappula redowskii; stickseed
Appendix 1. Plant Taxa Identified at Bowdoin National Wildlife Refuge 31

Lappula texana; cupseed stickseed

Lepidium perfoliatum; clasping peppergrass

Lepidium sp.; peppergrass

Liatris punctata; dotted blazing star*

Linaria vulgaris; butter-and-eggs

Lupinius flexuosus; lupine

Lygodesmia juncea; skeletonweed*

Lysimachia ciliata; fringed loosestrife*

Machaeranthera spinulosus; aster

Malva moschata; musk mallow

Malva neglecta; common mallow*

Medicago lupulina; black medick*

Medicago sativa; alfalfa*

Melilotus officinalis; yellow sweet clover*

Mentha arvensis; field mint*

Monarda spicata; horse mint, beebalm

Myriophyllum exalbescens; American milfoil*

Oenothera villosa; common evening primrose

Opuntia polyacantha; plains prickly pear

Oxytropis lambertii; purple locoweed

Oxytropis sericea; white locoweed

Panicum miliaceum; broom-corn millet

Penstemon albidus; white beardtongue

Petalostemum purpurea; purple prairie clover

Phalaris arundinacea; reed canarygrass*

Phleum pratense; timothy*

Phlox douglasii; phlox

Pinus ponderosa; ponderosa pine

Plantago lanceolata; English plantain, buckhorn

Plantago major; common plantain*

Poa pratensis; Kentucky bluegrass*

Poa sandbergii; Sandberg’s bluegrass*

Polanisia dodecandra; clammy-weed

Polemonium haydeni; Jacob’s-ladder

Polygonum aviculare; knotweed*

Polygonum buxiforme; knotweed

Polygonum convolvulus; climbing or wild buckwheat*

Polygonum lapathifolium; pale smartweed*

Polygonum persicaria; lady’s thumb

Polygonum ramosissimum; bushy knotweed*

Populus deltoides; cotton-wood*

Potamogeton filiformis; slender pondweed*

Potamogeton friesii; Fries’ pondweed*

Potamogeton pectinatus; sago pondweed*

Potamogeton pusillus; baby pondweed*

Potamogeton richardsonii; claspingleaf pondweed*

Potentilla anserina; silverweed*

Potentilla argentea; silvery cinquefoil

Potentilla arguta; tall cinquefoil*

Potentilla rivalis; brook conquefoil*

Prunus virginiana; choke cherry

Psoralea argophylla; silver-leaf scurf-pea*

Ranunculus longirostric; white water crowfoot

Ranunculus macounii; Macoun’s buttercup*

Ratibida columnifera; prairie coneflower*

Rhus aromatica; fragrant sumac, polecat bush

Ribes odoratum; buffalo currant

Rosa arkansana; prairie wild rose*

Rosa woodsii; western wild rose*

Rudbeckia sp.; coneflower

Rumex maritimus; golden dock*

Rumex sp.; dock, sorrel*

Rumex venosus; wild begonia

Ruppia maritima; ditchgrass, widgeon grass*

Sagittaria cuneata; arrowhead*

Sagittaria graminea; arrowhead

Sagittaria latifolia; common arrowhead*

Salix exigua subs. interior; sandbar willow*

Salix sp.; willow*

Salsola iberica; russian thistle*

Sarcobatus vermiculatus; greasewood

Scirpus acutus; hardstem bulrush*

Scirpus americanus; chairmaker’s bulrush*

Scirpus paludosus; cosmopolitan bulrush*

Senecio integerrimus; groundsel

Setaria sp.; foxtail*

Setaria viridis; green foxtail

Shepherdia argentea; buffaloberry

Silene cserei; smooth catchfly

Sisymbrium altissimum; tumbling mustard

Solidago gigantea; late goldenrod*

Solidago juncea; early goldenrod

Solidago missouriensis; prairie goldenrod*

Solidago rigida var. rigida; rigid goldenrod*

Sonchus arvensis; field sow thistle*

Sparganium emersum; bur-reed

Sphaeralcea coccinea; red false mallow

Sporobolus airoides; alkali sacaton*

Sporobolus asper; rough dropseed

Stipa comata; needle-and-thread*

Stipa spartea; porcupine-grass

Stipa viridula; green needlegrass*

Symphoricarpos occidentalis; western snowberry*

Symphoricarpos orbiculatus; coralberry, buckbrush

Taraxacum officinale; common dandelion*

Thlaspi arvense; field pennycress*

Tragopogan dubius; goat’s beard, western salsify

Triglochin maritima; seaside arrowgrass*

Typha latifolia; broad-leaved cat-tail*

Utricularia vulgaris; common bladderwort*

Verbena bracteata; prostrate vervain

Vicia americana; American vetch*

Yucca glauca; soapweed, yucca

Zigadenus venenosus; death camass

References

The Great Plains Flora Association, 1986, Flora of the Great 

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 Refuge, 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 L-1 = 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:

• TYPE_CHL (chloride)

• TYPE_MG_L (mg L-1)

• TYPE_PCNT (%)

• TYPE_PH (pH)

• TYPE_PPM (ppm)

• TYPE_SOD (sodium)

• TYPE_SUL (sulfate)

• TYPE_US_CM 

(μS cm-1)

Data type; these variables identify whether the measurement (for example, salinity, pH) was reported as 

a mean, median, range, etc. For example, if TYPE_US_CM = ‘MEAN,’ then the value was reported 

as a mean value by the original source; if TYPE_PH = ‘MIN,’ then the value was reported as a minimum value (for example, smallest value of a range) by the original source, etc.

• MAXIMUM = reported as maximum value, typically as part of a range (for example, 

1–10)

• MEAN = reported as a calculated mean

• MEDIAN = reported as a calculated median

• MINIMUM = reported as minimum value, typically as part of a range (for example, 

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 Conservation 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. Salinity 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

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 Refuge: 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 Wildlife 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 Service, 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: Washington, D.C., National Biological Service Information and Technology Report 6, 77 p.
Appendix 4. Description of Each Data Source Used to Develop the Plant and Invertebrate Databases 41

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 macroinvertebrate 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 Naturalist, 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 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.
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Achillea millefolium yarrow S 400 3,330 

Acorus calamus sweet flag S 600 

Agropyron caninum slender wheatgrass S 640 3,800 

Agropyron cristatum crested wheatgrass S 400 1,050 

Agropyron dasystachyum thickspike wheatgrass S 20,313 

Agropyron elongatum tall wheatgrass S 340 5,500 

Agropyron intermedium intermediate wheatgrass S 345 5,500 

Agropyron intermedium intermediate wheatgrass W 1,623 7.7 

Agropyron repens quackgrass S 140 6,460 

Agropyron repens quackgrass W 40 5,000 7.2 8.8 

Agropyron smithii western wheatgrass S 400 7,000 7.7 8.1 

Agropyron smithii western wheatgrass W 307 1,623 7.7 8.0 

Agropyron sp. wheatgrass S 350 2,790 

Agrostis hyemalis ticklegrass S 490 

Agrostis scabra ticklegrass S 345 4,688 

Agrostis scabra ticklegrass W 1,301 7.9 

Agrostis stolonifera redtop S 290 5,500 

Agrostis stolonifera redtop W 200 630 7.2 8.3 

Alisma gramineum narrowleaf water plantain S 400 1,850 

Alisma gramineum narrowleaf water plantain W 300 15,000 7.4 8.6 

Alisma subcordatum common water plantain S 300 4,550 

Alisma subcordatum common water plantain W 113 9,500 6.7 9.8 

Alisma triviale water plantain W 40 5,000 

Allium canadense wild onion S 400 620 

Allium stellatum pink wild onion S 650 2,200 

Alopecurus aequalis short-awn foxtail S 300 1,850 

Alopecurus aequalis short-awn foxtail W 40 5,000 7.0 9.7 

Amaranthus retroflexus rough pigweed S 730 2,450 

Ambrosia artemisiifolia common ragweed, short ragweed S 300 20,313 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Ambrosia artemisiifolia common ragweed, short ragweed W 326 1,582 7.3 8.6 

Ambrosia psilostachya western ragweed S 140 4,000 

Ambrosia psilostachya western ragweed W 560 729 8.0 8.6 

Ambrosia sp. ragweed S 590 

Ammannia coccinea toothcup W 40 2,000 

Amorpha canescens lead plant S 400 850 

Amorpha fruticosa false indigo S 525 850 

Andropogon gerardii big bluestem S 350 2,620 

Andropogon scoparius little bluestem S 1,500 

Anemone canadensis meadow anemone S 140 2,940 

Anemone canadensis meadow anemone W 355 429 7.6 8.4 

Apocynum cannabinum indian hemp dogbane, prairie 

dogbane

S 310 1,975 

Apocynum cannabinum indian hemp dogbane, prairie 

dogbane

W 40 5,000 7.2 8.6 

Artemisia absinthium wormwood S 350 5,500 

Artemisia absinthium wormwood W 1,623 7.7 

Artemisia biennis biennial wormwood S 290 3,970 

Artemisia biennis biennial wormwood W 40 15,000 9.7 

Artemisia dracunculus silky wormwood S 290 2,380 

Artemisia frigida prairie sagewort S 900 3,330 

Artemisia ludoviciana white sage S 290 2,200 

Asclepias incarnata swamp milkweed S 295 1,725 

Asclepias incarnata swamp milkweed W 274 900 8.1 9.6 

Asclepias ovalifolia ovalleaf milkweed S 300 1,340 

Asclepias sp. milkweed S 550 

Asclepias speciosa showy milkweed S 300 1,730 

Asclepias speciosa showy milkweed W 40 2,000 8.5 8.6 

Asclepias syriaca common milkweed S 140 2,830 

Asclepias syriaca common milkweed W 326 774 7.2 9.5 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Asclepias verticillata whorled milkweed S 620 700 

Aster brachyactis rayless aster S 620 2,930 

Aster brachyactis rayless aster W 500 15,000 8.2 8.4 

Aster ericoides white aster S 380 4,700 

Aster ericoides white aster W 492 630 7.8 8.1 

Aster falcatus white prairie aster S 400 3,970 

Aster falcatus white prairie aster W 630 8.1 

Aster hesperius panicled aster W 400 9,800 

Aster novae-angliae new england aster S 590 

Aster sagittifolius arrow-leaved aster W 1,000 

Aster simplex panicled aster S 295 5,500 

Aster simplex panicled aster W 40 16,100 6.7 9.7 

Aster sp. wild aster S 300 850 

Aster sp. wild aster W 391 590 7.4 8.3 

Aster subulatus saltmarsh aster S 1,875 9,844 

Astragalus canadensis Canada milk-vetch S 530 540 

Atriplex argentea silver-scale saltbush S 3,594 15,625 

Atriplex subspicata spearscale S 1,250 55,000 7.0 8.7 

Atriplex subspicata spearscale W 500 76,400 8.4 

Avena fatua wild oats S 820 1,470 

Azolla mexicana water fern, mosquito fern S 330 

Azolla mexicana water fern, mosquito fern W 471 7.3 

Baccharis salicina willow baccharis S 313 9,844 

Bacopa rotundifolia water hyssop S 410 

Bacopa rotundifolia water hyssop W 40 2,000 8.2 

Beckmannia syzigachne sloughgrass S 300 4,550 

Beckmannia syzigachne sloughgrass W 40 9,500 6.7 9.7 

Bidens cernua nodding beggar-ticks S 290 5,500 

Bidens cernua nodding beggar-ticks W 40 2,500 8.0 8.4 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Bidens frondosa devil’s beggar-ticks S 290 3,465 

Bidens frondosa devil’s beggar-ticks W 40 6,673 6.7 8.4 

Bidens sp. beggar-ticks S 820 

Bidens vulgata big devil’s beggar-ticks W 40 2,000 

Boltonia asteroides boltonia S 290 3,480 

Boltonia asteroides boltonia W 100 6,800 7.4 8.2 

Boltonia asteroides var. latisquama violet boltonia W 40 2,000 

Bouteloua curtipendula sideoats grama S 770 1,150 

Brassica kaber charlock S 1,320 1,430 

Bromus inermis smooth brome S 140 4,000 

Bromus inermis smooth brome W 256 5,387 7.1 9.5 

Bromus japonicus japanese brome S 400 900 

Bromus japonicus japanese brome W 1,623 7.7 

Bromus tectorum downy brome S 400 650 

Bromus tectorum downy brome W 326 7.8 

Calamagrostis canadensis bluejoint S 350 3,480 

Calamagrostis canadensis bluejoint W 40 3,800 6.9 9.5 

Calamagrostis stricta slimstem reedgrass S 140 4,700 

Calamagrostis stricta slimstem reedgrass W 40 17,600 7.3 9.8 

Callitriche hermaphroditica northern water starwort W 40 2,500 

Callitriche verna vernal water starwort W 40 2,000 

Calystegia sepium hedge bindweed S 430 5,156 

Carduus nutans musk thistle, nodding thistle S 500 

Carex alopecoidea foxtail sedge S 400 

Carex aquatilis water sedge S 330 1,475 

Carex aquatilis water sedge W 170 3,800 6.9 9.6 

Carex atherodes slough sedge S 295 3,075 

Carex atherodes slough sedge W 40 8,500 6.6 9.9 

Carex bebbii bebb’s sedge S 340 530 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Carex brevior fescue sedge S 400 700 

Carex buxbaumii brown bog sedge S 450 

Carex buxbaumii brown bog sedge W 391 1,400 8.0 

Carex granularis meadow sedge S 410 720 

Carex granularis meadow sedge W 113 6.9 

Carex hallii deer sedge S 630 

Carex hallii deer sedge W 412 7.9 

Carex interior interior sedge S 480 

Carex lacustris hairy sedge W 900 1,700 

Carex laeviconica smoothcone sedge W 40 3,200 

Carex lanuginosa woolly sedge S 290 2,615 

Carex lanuginosa woolly sedge W 40 32,600 7.2 9.5 

Carex praegracilis clustered-field sedge S 620 3,800 

Carex praegracilis clustered-field sedge W 40 5,000 

Carex rostrata beaked sedge S 305 

Carex rostrata beaked sedge W 200 2,600 

Carex sartwellii sartwell’s sedge S 450 900 

Carex sartwellii sartwell’s sedge W 40 5,000 8.0 

Carex sp. sedge S 290 4,700 

Carex sp. sedge W 192 1,487 6.9 9.9 

Carex stipata sawbeak sedge W 400 

Carex stricta tussock sedge W 100 9,400 

Carex sychnocephala manyhead sedge S 340 

Carex tetanica rigid sedge W 900 5,500 

Carex vulpinoidea fox sedge S 300 3,120 

Carex vulpinoidea fox sedge W 40 2,000 7.5 8.6 

Carum carvi caraway S 850 

Centaurium pulchellum branched centaury S 350 

Cerastium brachypodum nodding chickweed S 450 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Ceratophyllum demersum hornwort, coontail S 300 3,300 

Ceratophyllum demersum hornwort, coontail W 40 5,100 7.0 9.9 

Chenopodium album lamb’s quarters S 300 4,000 

Chenopodium album lamb’s quarters W 40 500 

Chenopodium glaucum oak-leaved goosefoot S 39,800 

Chenopodium glaucum oak-leaved goosefoot W 500 15,000 

Chenopodium rubrum alkali blite S 350 39,800 

Chenopodium rubrum alkali blite W 40 15,000 7.8 9.7 

Chenopodium sp. goosefoot, lamb’s quarter S 860 1,870 

Chenopodium sp. goosefoot, lamb’s quarter W 311 8.6 

Cicuta maculata common water hemlock S 450 1,720 

Cicuta maculata common water hemlock W 274 2,200 7.4 9.1 

Cicuta maculata angustifolia common water hemlock W 3,500 7,500 8.3 8.7 

Cirsium arvense Canada thistle, field thistle S 140 5,500 

Cirsium arvense Canada thistle, field thistle W 40 5,387 7.2 9.6 

Cirsium flodmanii flodman’s thistle S 480 3,330 

Cirsium vulgare bull thistle S 380 1,700 

Cirsium vulgare bull thistle W 1,623 7.7 

Convolvulus arvensis field bindweed S 140 2,830 

Convolvulus arvensis field bindweed W 450 729 8.2 8.6 

Conyza canadensis horse-weed S 300 4,000 

Coreopsis tinctoria plains coreopsis S 345 615 

Cornus sericea redosier dogwood S 550 575 

Cornus sericea redosier dogwood W 537 8.5 

Crataegus rotundifolia northern hawthorn S 550 

Crataegus rotundifolia northern hawthorn W 450 8.6 

Crepis runcinata hawk’s-beard S 140 1,260 

Crepis runcinata hawk’s-beard W 326 1,263 7.2 8.6 

Cyperus acuminatus tapeleaf flatsedge S 720 850 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Cyperus acuminatus tapeleaf flatsedge W 40 2,000 

Cyperus erythrorhizos redrooted cyperus S 900 

Cyperus odoratus fragrant flatsedge S 480 1,975 

Cyperus sp. umbrella sedge S 310 1,470 

Cypripedium sp. lady’s-slipper S 1,730 

Dactylis glomerata orchard grass S 395 770 

Dalea purpurea purple prairie clover S 290 850 

Descurainia sophia flixweed S 560 2,380 

Dichanthelium oligosanthes dichanthelium S 530 

Distichlis spicata var. stricta inland saltgrass S 400 104,000 6.8 10.0 

Distichlis spicata var. stricta inland saltgrass W 500 120,000 7.9 9.1 

Drepanocladus sp. drepanocladus moss W 40 5,000 

Echinochloa crusgalli barnyard grass S 290 5,500 

Echinochloa crusgalli barnyard grass W 40 5,000 6.7 9.1 

Elaeagnus angustifolia russian olive S 550 1,150 

Elaeagnus commutata silverberry S 900 6,460 

Eleocharis acicularis needle spikerush S 350 2,000 

Eleocharis acicularis needle spikerush W 40 5,800 7.2 9.8 

Eleocharis compressa flatstem spikerush S 500 2,400 

Eleocharis compressa flatstem spikerush W 400 5,000 8.2 

Eleocharis macrostachya common spikerush W 100 41,000 

Eleocharis obtusa var. ovata blunt spikerush S 500 900 

Eleocharis obtusa var. ovata blunt spikerush W 40 2,000 7.8 9.1 

Eleocharis palustris common spikerush S 305 10,938 

Eleocharis palustris common spikerush W 40 15,000 6.8 9.8 

Eleocharis sp. spikerush S 290 3,160 

Eleocharis sp. spikerush W 113 2,597 6.6 9.5 

Ellisia nyctelea waterpod S 620 

Elodea canadensis canadian waterweed W 40 2,000 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Elymus canadensis Canada wild rye S 400 3,800 

Epilobium ciliatum willow-herb S 500 2,930 

Epilobium ciliatum willow-herb W 40 5,000 

Epilobium leptophyllum narrow-leaved willow-herb S 290 4,000 

Epilobium leptophyllum narrow-leaved willow-herb W 1,232 8.5 

Epilobium sp. willow-herb, fireweed S 460 

Equisetum arvense field horsetail S 140 1,520 

Equisetum arvense field horsetail W 400 450 8.6 

Equisetum fluviatile water horsetail W 300 

Equisetum hyemale common scouring rush S 470 2,100 

Equisetum laevigatum smooth scouring rush S 140 3,800 

Equisetum laevigatum smooth scouring rush W 192 412 7.9 9.7 

Erigeron philadelphicus Philadelphia fleabane S 340 1,613 

Erigeron philadelphicus Philadelphia fleabane W 401 729 8.0 8.6 

Erigeron strigosus daisy fleabane S 300 1,260 

Eriophorum polystachion narrowleaf cottongrass W 500 2,200 

Eupatorium maculatum joe-pye weed, spotted joe-pye weed S 400 760 

Eupatorium maculatum joe-pye weed, spotted joe-pye weed W 148 700 7.3 

Euphorbia esula leafy spurge S 600 1,880 

Euphorbia maculata spotted spurge S 570 2,570 

Euphorbia maculata spotted spurge W 148 7.3 

Eustoma grandiflorum showy prairie gentian S 938 15,156 

Euthamia graminifolia var. graminif flat-top goldentop S 450 1,730 

Euthamia graminifolia var. graminif flat-top goldentop W 100 2,100 8.0 

Fragaria virginiana wild strawberry S 140 3,800 

Fragaria virginiana wild strawberry W 450 8.6 

Fraxinus pennsylvanica green ash S 345 2,050 

Galium boreale northern bedstraw S 400 1,270 

Galium sp. bedstraw, cleavers S 540 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Galium sp. bedstraw, cleavers W 355 7.6 

Galium trifidum small bedstraw S 400 630 

Galium trifidum small bedstraw W 300 590 7.4 7.6 

Glaux maritima sea milkwort S 1,290 4,000 

Glaux maritima sea milkwort W 288 5,000 7.6 

Glyceria borealis northern mannagrass W 40 1,000 

Glyceria grandis tall mannagrass S 330 1,290 

Glyceria grandis tall mannagrass W 40 4,000 6.7 9.9 

Glyceria striata fowl mannagrass S 350 2,200 

Glyceria striata fowl mannagrass W 260 800 9.8 

Glycyrrhiza lepidota wild licorice S 300 4,000 

Glycyrrhiza lepidota wild licorice W 450 1,487 8.1 8.6 

Gratiola neglecta hedge hyssop W 40 2,000 

Grindelia squarrosa curly-top gumweed S 450 20,313 

Hedeoma hispidum rough false pennyroyal S 300 630 

Helenium autumnale sneezeweed W 40 2,500 

Helianthus annuus common sunflower S 340 3,480 

Helianthus annuus common sunflower W 326 7.8 

Helianthus maximiliani maximilian sunflower S 140 3,800 

Helianthus maximiliani maximilian sunflower W 450 590 7.4 8.6 

Helianthus nuttallii nuttall’s sunflower S 400 3,800 

Helianthus nuttallii nuttall’s sunflower W 590 856 7.4 8.1 

Helianthus petiolaris plains sunflower S 400 760 

Helianthus rigidus stiff sunflower S 400 2,200 

Helianthus sp. sunflower S 140 450 

Heliopsis helianthoides false sunflower, ox-eye S 500 1,730 

Heliotropium curassavicum seaside heliotrope W 2,000 15,000 

Hesperis matronalis dame’s rocket S 450 2,615 

Hesperis matronalis dame’s rocket W 355 7.6 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Hierochloe odorata sweetgrass S 760 

Hierochloe odorata sweetgrass W 40 2,000 

Hippuris vulgaris mare’s tail S 850 

Hippuris vulgaris mare’s tail W 267 4,100 8.1 8.6 

Hordeum jubatum foxtail barley S 290 39,800 6.4 9.5 10,

Hordeum jubatum foxtail barley W 40 48,600 7.3 9.7 

Hypoxis hirsuta yellow stargrass S 530 640 

Impatiens capensis spotted touch-me-not W 400 

Iris sp. iris, flag S 270 890 

Iris sp. iris, flag W 274 729 7.2 9.0 

Iris versicolor blue flag S 600 750 

Iris versicolor blue flag W 302 729 8.6 9.1 

Iva annua marsh elder S 1,875 24,000 6.9 8.2 

Iva xanthifolia marsh elder S 340 2,930 

Iva xanthifolia marsh elder W 1,582 7.3 

Juncus alpinus richardson’s rush S 850 2,570 

Juncus alpinus richardson’s rush W 964 1,947 7.3 8.2 

Juncus balticus baltic rush S 290 43,750 

Juncus balticus baltic rush W 40 45,000 7.8 8.1 

Juncus bufonius toad rush S 450 

Juncus bufonius toad rush W 40 2,300 

Juncus dudleyi dudley rush W 40 2,000 

Juncus interior inland rush S 380 4,000 

Juncus interior inland rush W 40 2,000 7.8 

Juncus sp. rush S 1,225 2,790 

Juncus torreyi torrey’s rush S 490 5,500 

Juncus torreyi torrey’s rush W 40 10,000 7.3 

Kochia scoparia kochia S 300 20,313 

Kochia scoparia kochia W 40 45,000 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Lactuca serriola prickly lettuce S 340 2,930 

Lactuca serriola prickly lettuce W 2,000 45,000 

Lactuca tatarica blue lettuce S 290 3,970 

Lactuca tatarica blue lettuce W 307 7.8 

Lathyrus palustris marsh vetchling S 340 1,730 

Lathyrus palustris marsh vetchling W 590 7.4 

Leersia oryzoides rice cutgrass S 380 1,475 

Leersia oryzoides rice cutgrass W 192 326 7.8 9.7 

Lemna minor common duckweed S 350 4,000 

Lemna minor common duckweed W 40 15,000 6.6 9.9 

Lemna trisulca star duckweed S 350 4,000 

Lemna trisulca star duckweed W 40 13,900 7.0 9.9 

Lepidium densiflorum peppergrass S 300 

Leptochloa fascicularis bearded sprangletop S 469 7,656 

Leptochloa fascicularis bearded sprangletop W 577 7.9 

Liatris ligulistylis rocky mountain blazing star S 3,330 3,800 

Liatris punctata dotted blazing star S 650 

Liatris pycnostachya prairie blazing star S 1,730 

Lilium philadelphicum wild lily S 540 

Limosella aquatica mudwort W 40 2,000 

Lindernia dubia false pimpernel W 40 2,000 

Linum perenne blue flax S 450 

Lithospermum canescens hoary puccoon S 550 

Lithospermum canescens hoary puccoon W 450 8.6 

Lobelia spicata palespike lobelia S 770 2,620 

Lotus unifoliolatus Amercian bird’s-foot trefoil S 670 3,330 

Lycopus americanus Amercian bugleweed S 300 4,000 

Lycopus americanus Amercian bugleweed W 274 24,000 7.2 9.6 

Lycopus asper rough bugleweed S 330 4,700 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Lycopus asper rough bugleweed W 40 32,600 6.8 9.8 

Lygodesmia juncea skeletonweed S 690 

Lysimachia ciliata fringed loosestrife S 500 2,780 

Lysimachia ciliata fringed loosestrife W 166 2,653 6.7 8.3 

Lysimachia hybrida loosestrife S 330 860 

Lysimachia hybrida loosestrife W 40 2,000 7.7 9.0 

Lysimachia thyrsiflora tufted loosestrife S 370 735 

Lysimachia thyrsiflora tufted loosestrife W 192 3,800 7.4 9.7 

Malva neglecta common mallow S 1,320 

Marsilea vestita western water clover W 40 2,000 

Medicago lupulina black medick S 300 2,500 

Medicago sativa alfalfa S 300 5,500 

Medicago sativa alfalfa W 629 7.2 

Melilotus officinalis yellow sweet clover S 140 6,460 

Melilotus officinalis yellow sweet clover W 326 1,582 7.3 8.6 

Melilotus sp. sweet clover S 375 5,500 

Melilotus sp. sweet clover W 362 7.0 

Mentha arvensis field mint S 290 2,380 

Mentha arvensis field mint W 40 5,000 6.7 9.0 

Mentha arvensis villosa field mint W 3,500 4,100 8.3 

Mimulus ringens alleghany monkey-flower W 600 

Monarda fistulosa wild bergamot S 450 550 

Muhlenbergia asperifolia scratchgrass S 880 4,688 

Muhlenbergia asperifolia scratchgrass W 700 45,000 

Muhlenbergia racemosa marsh muhly S 1,725 

Muhlenbergia richardsonis mat muhly S 400 3,800 

Myriophyllum exalbescens American milfoil S 450 4,740 

Myriophyllum exalbescens American milfoil W 40 8,000 7.2 9.8 

Myriophyllum heterophyllum water milfoil W 40 2,000 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Myriophyllum pinnatum green parrot’s feather W 100 

Myriophyllum verticillatum whorl-leaf watermilfoil W 400 5,000 

Najas flexilis naiad W 300 700 

Nepeta cataria catnip S 380 

Nuphar lutea var. variegatum yellow water lily W 400 

Oxalis stricta yellow wood sorrel S 140 1,700 

Oxalis stricta yellow wood sorrel W 450 8.6 

Oxalis violacea violet wood sorrel S 450 620 

Panicum americanum pearl millet S 290 5,500 

Panicum americanum pearl millet W 275 7.9 

Panicum capillare common witchgrass S 290 2,500 

Panicum capillare common witchgrass W 40 15,000 

Panicum virgatum switchgrass S 300 2,750 

Panicum virgatum switchgrass W 260 856 7.8 9.8 

Parietaria pensylvanica Pennsylvania pellitory S 900 

Parnassia glauca grass-of-parnassus W 900 

Pedicularis canadensis common lousewort, wood betony S 500 1,730 

Phalaris arundinacea reed canarygrass S 300 3,210 

Phalaris arundinacea reed canarygrass W 40 3,800 6.9 9.9 

Phleum pratense timothy S 400 2,050 

Phleum pratense timothy W 600 729 8.3 8.6 

Phragmites australis common reed S 580 4,660 7.3 

Phragmites australis common reed W 100 32,600 7.6 9.0 

Physalis virginiana Virginia ground cherry S 350 1,700 

Plagiobothrys scouleri popcorn-flower W 40 2,000 

Plantago eriopoda alkali plantain W 500 20,100 

Plantago major common plantain S 400 5,500 

Plantago major common plantain W 40 5,000 7.8 

Poa arida plains bluegrass S 1,094 20,313 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Poa compressa Canada bluegrass S 1,250 5,156 

Poa palustris fowl bluegrass S 140 5,500 

Poa palustris fowl bluegrass W 40 5,000 7.4 9.7 

Poa pratensis Kentucky bluegrass S 140 5,500 

Poa pratensis Kentucky bluegrass W 192 1,623 7.4 9.7 

Poa sandbergii Sandberg’s bluegrass S 400 

Poa sp. bluegrass S 550 

Poa sp. bluegrass W 632 7.4 

Polygonum amphibium water smartweed S 270 3,020 

Polygonum amphibium water smartweed W 40 5,000 6.6 9.9 

Polygonum amphibium var. emersum swamp smartweed W 40 5,000 

Polygonum aviculare knotweed S 630 

Polygonum aviculare knotweed W 355 7.6 

Polygonum convolvulus climbing or wild buckwheat S 300 2,380 

Polygonum erectum erect knotweed S 380 1,560 

Polygonum lapathifolium pale smartweed S 310 2,450 

Polygonum lapathifolium pale smartweed W 40 5,000 7.8 8.6 

Polygonum pensylvanicum Pennsylvania smartweed S 1,145 2,450 

Polygonum pensylvanicum Pennsylvania smartweed W 629 7.2 

Polygonum ramosissimum bushy knotweed S 410 54,063 

Polygonum ramosissimum bushy knotweed W 2,000 45,000 

Populus deltoides cotton-wood S 300 1,850 

Populus deltoides cotton-wood W 362 1,385 7.0 7.9 

Populus tremuloides quaking aspen S 300 2,620 

Populus tremuloides quaking aspen W 192 429 7.8 9.7 

Potamogeton diversifolius waterthread pondweed W 40 500 

Potamogeton filiformis slender pondweed W 560 8,000 

Potamogeton friesii Fries’ pondweed W 300 4,100 8.3 

Potamogeton gramineus variable pondweed S 430 660 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Potamogeton gramineus variable pondweed W 40 2,000 6.7 8.3 

Potamogeton natans floatingleaf pondweed S 500 

Potamogeton natans floatingleaf pondweed W 777 7.7 

Potamogeton nodosus longleaf pondweed S 410 535 

Potamogeton nodosus longleaf pondweed W 401 496 8.0 8.2 

Potamogeton pectinatus sago pondweed S 300 31,406 8.4 8.9 

Potamogeton pectinatus sago pondweed W 38 162,500 6.3 10.8 2Potamogeton pusillus baby pondweed S 330 3,000 

Potamogeton pusillus baby pondweed W 40 6,700 6.9 9.9 

Potamogeton richardsonii claspingleaf pondweed S 450 520 

Potamogeton richardsonii claspingleaf pondweed W 40 5,000 8.0 9.5 

Potamogeton sp. pondweed W 148 7.3 

Potamogeton strictifolius narrowleaf pondweed W 267 

Potamogeton vaginatus sheathed pondweed W 267 15,000 8.9 

Potamogeton zosteriformis flatstem pondweed S 380 630 

Potamogeton zosteriformis flatstem pondweed W 260 5,000 7.3 9.9 

Potentilla anserina silverweed S 420 4,700 

Potentilla anserina silverweed W 100 45,000 7.2 

Potentilla arguta tall cinquefoil S 530 1,520 

Potentilla norvegica Norwegian cinquefoil S 400 2,940 

Potentilla norvegica Norwegian cinquefoil W 40 2,000 8.6 

Potentilla pensylvanica Pennsylvania cinquefoil S 400 1,990 

Potentilla rivalis brook conquefoil W 300 

Psoralea argophylla silver-leaf scurf-pea S 290 1,870 

Puccinellia nuttalliana nuttall’s alkaligrass S 880 39,800 7.7 8.6 

Puccinellia nuttalliana nuttall’s alkaligrass W 700 120,000 7.9 9.6 

Ranunculus cymbalaria shore buttercup S 760 4,688 

Ranunculus cymbalaria shore buttercup W 500 34,000 8.0 9.6 

Ranunculus flabellaris threadleaf buttercup S 450 890 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Ranunculus flabellaris threadleaf buttercup W 40 2,500 7.4 9.6 

Ranunculus gmelinii small yellow buttercup S 410 2,100 

Ranunculus gmelinii small yellow buttercup W 113 2,867 6.9 9.9 

Ranunculus longirostris white water crowfoot S 530 1,520 

Ranunculus longirostris white water crowfoot W 40 5,000 7.5 9.7 

Ranunculus macounii macoun’s buttercup S 450 1,530 

Ranunculus macounii macoun’s buttercup W 40 2,100 7.4 7.8 

Ranunculus pensylvanicus bristly crowfoot S 1,005 1,440 

Ranunculus pensylvanicus bristly crowfoot W 1,263 8.6 

Ranunculus sceleratus cursed crowfoot S 550 

Ranunculus sceleratus cursed crowfoot W 40 8,500 

Ranunculus sp. buttercup, crowfoot S 1,130 2,790 

Ranunculus sp. buttercup, crowfoot W 729 8.7 

Ranunculus subrigidus white water crowfoot W 200 4,500 

Ratibida columnifera prairie coneflower S 400 2,270 

Rhus glabra smooth sumac S 550 

Ribes americanum wild black currant S 535 630 

Ribes americanum wild black currant W 355 391 7.6 8.0 

Riccia fluitans slender riccia S 400 740 

Riccia fluitans slender riccia W 40 4,700 6.9 7.9 

Riccia sp. liverwort S 375 4,000 

Riccia sp. liverwort W 307 6,673 7.0 8.4 

Ricciocarpus natans purple-fringed riccia S 1,120 

Ricciocarpus natans purple-fringed riccia W 40 3,200 7.9 

Rorippa palustris bog yellow cress S 290 2,930 

Rorippa palustris bog yellow cress W 40 3,200 7.6 9.0 

Rosa arkansana prairie wild rose S 300 3,800 

Rosa blanda smooth wild rose S 500 740 

Rosa woodsii western wild rose S 140 900 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Rosa woodsii western wild rose W 113 6.9 

Rudbeckia hirta black-eyed susan S 400 3,800 

Rudbeckia hirta black-eyed susan W 1,623 7.7 

Rumex crispus curly dock S 290 11,250 

Rumex crispus curly dock W 326 1,623 7.2 9.7 

Rumex maritimus golden dock S 270 4,550 

Rumex maritimus golden dock W 40 15,000 7.1 9.9 

Rumex mexicanus willow-leaved dock W 40 5,000 

Rumex occidentalis western dock S 300 2,930 

Rumex occidentalis western dock W 40 5,000 7.9 8.8 

Rumex orbiculatus great water dock S 885 

Rumex salicifolius willow dock S 300 5,500 

Rumex salicifolius willow dock W 307 6,673 7.2 9.5 

Rumex sp. dock, sorrel S 420 2,700 

Rumex sp. dock, sorrel W 113 1,582 6.7 8.5 

Rumex stenophyllus Eurasian dock S 700 

Ruppia maritima ditchgrass, widgeon grass S 31,406 3.1 8.8 

Ruppia maritima ditchgrass, widgeon grass W 94 609,375 6.0 10.8 Ruppia maritima var. occidentalis ditchgrass, widgeon grass W 600 14,200 

Ruppia maritima var. rostrata ditchgrass, widgeon grass W 5,500 66,000 

Sagittaria cuneata arrowhead S 270 1,400 

Sagittaria cuneata arrowhead W 40 6,700 6.8 9.5 

Sagittaria latifolia common arrowhead S 400 900 

Sagittaria latifolia common arrowhead W 148 1,444 6.7 9.8 

Sagittaria sp. arrowhead S 400 650 

Sagittaria sp. arrowhead W 256 782 7.2 8.6 

Salicornia rubra saltwort S 2,344 142,813 7.0 9.0 

Salicornia rubra saltwort W 5,000 45,000 

Salix amygdaloides peachleaf willow S 350 2,620 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Salix amygdaloides peachleaf willow W 192 1,385 7.4 9.7 

Salix exigua sandbar willow, coyote willow S 300 1,655 

Salix exigua sandbar willow, coyote willow W 192 600 8.3 9.7 

Salix exigua subs. interior sandbar willow W 300 1,700 

Salix nigra black willow S 620 

Salix sp. willow S 410 1,850 

Salix sp. willow W 201 1,896 7.7 9.8 

Salsola iberica Russian thistle S 39,800 

Scirpus acutus hardstem bulrush S 270 5,500 

Scirpus acutus hardstem bulrush W 40 37,000 6.6 9.9 

Scirpus americanus chairmaker’s bulrush S 1,719 11,200 7.9 8.5 

Scirpus americanus chairmaker’s bulrush W 500 120,000 8.3 9.6 

Scirpus atrovirens darkgreen bulrush S 580 2,270 

Scirpus atrovirens darkgreen bulrush W 500 2,200 7.5 

Scirpus fluviatilis river bulrush S 330 4,550 

Scirpus fluviatilis river bulrush W 40 6,700 6.6 9.9 

Scirpus heterochaetus slender bulrush S 300 2,470 

Scirpus heterochaetus slender bulrush W 40 4,200 6.9 9.8 

Scirpus microcarpus panicled bulrush W 300 900 

Scirpus nevadensis Nevada bulrush W 12,000 45,000 

Scirpus pallidus cloaked bulrush S 340 2,050 

Scirpus pallidus cloaked bulrush W 565 1,809 7.2 8.6 

Scirpus paludosus cosmopolitan bulrush S 410 120,000 5.2 8.9 7

Scirpus paludosus cosmopolitan bulrush W 113 481,250 6.4 9.8 

Scirpus pungens common threesquare S 465 5,500 

Scirpus pungens common threesquare W 100 70,000 7.2 8.4 

Scirpus robustus sturdy bulrush S 2,600 111,000 3.1 6.6 

Scirpus robustus sturdy bulrush W 313 60,938 4.0 8.3 

Scirpus sp. bulrush S 450 975 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Scirpus sp. bulrush W 192 629 7.2 9.7 

Scirpus validus softstem bulrush S 280 4,550 

Scirpus validus softstem bulrush W 40 6,673 7.0 9.7 

Scolochloa festucacea sprangletop S 400 4,660 

Scolochloa festucacea sprangletop W 40 15,000 6.8 9.8 

Scrophularia lanceolata figwort S 340 

Scutellaria galericulata marsh skullcap W 300 

Scutellaria sp. skullcap S 380 760 

Scutellaria sp. skullcap W 256 661 7.2 9.1 

Senecio congestus swamp ragwort S 620 2,450 

Senecio congestus swamp ragwort W 40 5,000 

Senecio pseudaureus falsegold groundsel S 610 1,260 

Sesuvium verrucosum sea purslane S 3,594 81,500 7.0 9.7 

Setaria sp. foxtail S 300 2,500 

Setaria sp. foxtail W 192 964 7.3 9.7 

Silene sp. catchfly, campion S 340 500 

Sisyrinchium campestre white-eyed grass S 530 

Sium suave water parsnip S 400 3,800 

Sium suave water parsnip W 40 5,000 7.2 9.9 

Solidago canadensis Canada goldenrod S 140 4,700 

Solidago canadensis Canada goldenrod W 590 725 7.4 8.6 

Solidago gigantea late goldenrod S 140 2,620 

Solidago gigantea late goldenrod W 629 7.2 

Solidago missouriensis prairie goldenrod S 140 1,220 

Solidago mollis soft goldenrod S 500 6,460 

Solidago rigida var. rigida rigid goldenrod S 140 4,700 

Sonchus arvensis field sow thistle S 290 20,313 

Sonchus arvensis field sow thistle W 40 20,800 7.2 9.6 

Sonchus asper prickly sow thistle S 490 800 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Sorghastrum nutans Indian grass S 530 

Sparganium eurycarpum giant burreed S 270 4,550 

Sparganium eurycarpum giant burreed W 40 5,000 6.6 9.6 

Spartina gracilis alkali cordgrass S 4,688 7,813 

Spartina gracilis alkali cordgrass W 700 45,000 

Spartina pectinata prairie cordgrass S 290 20,313 

Spartina pectinata prairie cordgrass W 40 33,500 6.6 9.5 

Spergularia marina salt-marsh sand spurry S 2,344 39,800 

Spergularia marina salt-marsh sand spurry W 2,000 15,000 

Sphenopholis obtusa wedgegrass S 17,188 20,313 

Sphenopholis obtusata prairie wedgegrass S 1,900 

Spiraea alba meadow-sweet S 525 740 

Spiraea alba meadow-sweet W 588 774 8.9 9.6 

Spirodela polyrhiza duckmeat, greater duckweed S 330 730 

Spirodela polyrhiza duckmeat, greater duckweed W 40 3,000 6.9 9.0 

Sporobolus airoides alkali sacaton S 156 42,656 6.5 10.0 

Sporobolus texanus Texas dropseed S 1,875 72,000 7.4 8.3 

Stachys palustris hedge-nettle, marsh betony S 290 3,480 

Stachys palustris hedge-nettle, marsh betony W 40 5,000 6.9 9.8 

Stipa comata needle-and-thread S 620 1,150 

Stipa viridula green needlegrass S 400 1,500 

Stipa viridula green needlegrass W 725 8.6 

Suaeda depressa sea blite S 5,781 110,400 6.8 10.0 

Suaeda depressa sea blite W 5,000 66,000 

Symphoricarpos occidentalis western snowberry S 350 4,000 

Symphoricarpos occidentalis western snowberry W 725 3,480 8.1 8.6 

Tamarix ramosissima salt cedar S 2,188 96,000 7.4 8.7 

Taraxacum officinale common dandelion S 300 4,688 

Taraxacum officinale common dandelion W 326 630 7.8 8.4 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Teucrium canadense wood sage S 450 1,720 

Teucrium canadense wood sage W 391 856 7.4 9.5 

Teucrium canadense var. boreale wood sage W 40 9,100 

Thalictrum dasycarpum purple meadow rue S 550 860 

Thalictrum dasycarpum purple meadow rue W 450 8.6 

Thalictrum venulosum early meadow rue S 525 1,730 

Thalictrum venulosum early meadow rue W 590 7.4 

Thlaspi arvense field pennycress S 470 2,380 

Tradescantia bracteata spiderwort S 400 740 

Tragopogon dubius goat’s beard S 380 2,000 

Trifolium hybridum alsike clover S 450 520 

Trifolium pratense red clover S 400 930 

Trifolium pratense red clover W 326 7.8 

Trifolium repens white clover S 520 640 

Triglochin maritima seaside arrowgrass S 900 72,000 8.1 8.2 

Triglochin maritima seaside arrowgrass W 700 120,000 7.9 9.6 

Triglochin palustris marsh arrowgrass W 9.6 

Typha angustifolia narrow-leaved cat-tail S 280 4,550 

Typha angustifolia narrow-leaved cat-tail W 192 15,000 7.0 9.9 

Typha latifolia broad-leaved cat-tail S 270 5,500 

Typha latifolia broad-leaved cat-tail W 40 13,600 6.2 9.9 

Typha sp. cat-tail S 420 4,660 

Typha sp. cat-tail W 195 5,387 6.9 8.5 

Typha x glauca hybrid cat-tail S 330 4,550 

Typha x glauca hybrid cat-tail W 100 6,600 6.7 9.9 

Urtica dioica stinging nettle S 400 3,800 

Urtica dioica stinging nettle W 274 5,387 7.1 9.0 

Utricularia vulgaris common bladderwort S 330 3,370 

Utricularia vulgaris common bladderwort W 40 8,100 6.8 9.9 
Appendix 5. Summary of plant database.—Continued[Scientific and common names primarily follow The Great Plains Flora Association (1986), and the minimum and maximum values represent all data types (that is, values repormaximum, mean, median, or single occurrence). S, soil; W, water; μS cm-1, microsiemens per centimeter]

Genus Species Common name Substrate

Conductivity

(μS cm-1) pH 

Min Max Min Max

Vallisneria americana tapegrass S 410 880 

Vallisneria americana tapegrass W 192 852 7.6 9.8 

Verbena hastata blue vervain S 415 1,690 

Verbena hastata blue vervain W 355 632 7.4 7.6 

Verbena sp. vervain S 590 

Vernonia fasciculata ironweed S 310 1,400 

Vernonia fasciculata ironweed W 40 500 7.2 7.8 

Veronica anagallis-aquatica water speedwell S 500 

Veronica peregrina purslane speedwell S 4,000 

Veronica peregrina purslane speedwell W 40 2,000 

Vicia americana American vetch S 400 1,720 

Vicia sp. vetch S 140 770 

Vicia sp. vetch W 450 8.6 

Viola nuttallii yellow prairie violet S 580 650 

Viola sororia downy blue violet S 535 860 

Viola sororia downy blue violet W 391 590 7.4 8.6 

Xanthium strumarium cocklebur S 300 5,500 

Xanthium strumarium cocklebur W 40 5,000 7.7 9.7 

Zannichellia palustris horned pondweed W 300 45,000 

Zigadenus elegans white camass S 530 760 

Zizia aptera meadow parsnip S 400 620 

Zizia aptera meadow parsnip W 192 450 8.6 9.7 

Zizia aurea golden alexanders S 400 1,725 

Zizia aurea golden alexanders W 590 856 7.4 8.1 

1Source: 1, Grosshans and Kenkel, 1997; 2, Hammer and Heseltine, 1988; 3, Johnson, 1990; 4, Kantrud and others, 1989; 5, Kantrud, 1990; 6, Kantrud, 1991; 7, Kantrud, 199

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, Ungar17, Ungar, 1974.
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 Refuge: 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 Wildlife 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 Service, 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: Washington, 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 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.
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Annelida Hirudinea 270 15,625 7.1 

Annelida Hirudinea Gnathobdellida Hirudinidae 540 1,997 7.1 

Annelida Hirudinea Pharyngobdellida Erpobdellidae 531 2,000 7.1 

Annelida Hirudinea Pharyngobdellida Erpobdellidae Erpobdella 99 4,340 

Annelida Hirudinea Rhynchobdellida Glossiphoniidae 531 2,000 7.3 

Annelida Hirudinea Rhynchobdellida Glossiphoniidae Glossiphonia 105 2,930 

Annelida Hirudinea Rhynchobdellida Glossiphoniidae Helobdella 69 5,554 7.1 

Annelida Hirudinea Rhynchobdellida Glossiphoniidae Helobdella stagnalis 2,644 

Annelida Hirudinea Rhynchobdellida Glossiphoniidae Placobdella 168 1,243 

Annelida Hirudinea Rhynchobdellida Glossiphoniidae Theromyzon 217 4,378 

Annelida Oligochaeta 57 69,100 5.4 

Arthropoda Arachnida Acarina 100 33,600 4.6 

Arthropoda Arachnoidea Hydracarina 57 15,625 7.1 

Arthropoda Branchiopoda Anastraca Streptocephalidae Streptucephalus texanus 101 982 

Arthropoda Branchiopoda Anostraca 80 2,178 

Arthropoda Branchiopoda Anostraca Artemiidae Artemia salina 26,622 187,500 10.0 

Arthropoda Branchiopoda Anostraca Branchinectidae Branchinecta campestris 5,130 5,557 9.8 

Arthropoda Branchiopoda Anostraca Branchinectidae Branchinecta lindahli 1,312 36,342 9.3 

Arthropoda Branchiopoda Anostraca Branchinectidae Branchinecta mackini 2,394 

Arthropoda Branchiopoda Anostraca Chirocephalidae Chirocephalopsis bundyi 680 1,250 8.8 

Arthropoda Branchiopoda Anostraca Chirocephalidae Eubranchipus 188 892 

Arthropoda Branchiopoda Anostraca Chirocephalidae Eubranchipus bundyi 91 388 

Arthropoda Branchiopoda Anostraca Chirocephalidae Eubranchipus ornatus 80 892 

Arthropoda Branchiopoda Anostraca Streptocephalidae Streptocephalus seali 136 

Arthropoda Branchiopoda Cladocera 99 15,000 

Arthropoda Branchiopoda Cladocera Bosminidae Bosmina obtusirostris 156 31,250 

Arthropoda Branchiopoda Cladocera Chydoridae Alona costata 156 7,813 

Arthropoda Branchiopoda Cladocera Chydoridae Alona rectangula 313 1,563 

Arthropoda Branchiopoda Cladocera Chydoridae Alonella 1,879 

Arthropoda Branchiopoda Cladocera Chydoridae Chydorus 185 3,658 

Arthropoda Branchiopoda Cladocera Chydoridae Chydorus gibbus 313 1,563 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Branchiopoda Cladocera Chydoridae Chydorus sphaericus 156 31,250 

Arthropoda Branchiopoda Cladocera Chydoridae Eurycercus lamellatus 156 2,497 

Arthropoda Branchiopoda Cladocera Chydoridae Graptoleberis testudinaria 313 1,563 

Arthropoda Branchiopoda Cladocera Chydoridae Pleuroxus denticulatus 156 7,813 

Arthropoda Branchiopoda Cladocera Daphnidae Ceriodaphnia 91 4,650 

Arthropoda Branchiopoda Cladocera Daphnidae Ceriodaphnia quadrangula 313 15,625 

Arthropoda Branchiopoda Cladocera Daphnidae Daphnia 80 6,150 

Arthropoda Branchiopoda Cladocera Daphnidae Daphnia longispina 156 187,500 

Arthropoda Branchiopoda Cladocera Daphnidae Daphnia pulex 313 31,250 

Arthropoda Branchiopoda Cladocera Daphnidae Scapholeberis 196 3,698 

Arthropoda Branchiopoda Cladocera Daphnidae Scapholeberis mucronata 156 31,250 

Arthropoda Branchiopoda Cladocera Daphnidae Simocephalus 57 5,266 

Arthropoda Branchiopoda Cladocera Daphnidae Simocephalus vetulus 156 15,625 

Arthropoda Branchiopoda Cladocera Leptodoridae Leptodora kindtii 156 7,813 

Arthropoda Branchiopoda Cladocera Moinidae Moina 2,295 

Arthropoda Branchiopoda Cladocera Moinidae Moina macrocopa 3,706 

Arthropoda Branchiopoda Cladocera Polyphemidae Polyphemus pediculus 156 1,563 

Arthropoda Branchiopoda Cladocera Sididae Diaphanosoma 1,104 4,073 

Arthropoda Branchiopoda Cladocera Sididae Diaphanosoma leuchtenbergianum 156 31,250 

Arthropoda Branchiopoda Cladocera Sididae Sida crystallina 313 1,563 

Arthropoda Branchiopoda Conchostraca 119 2,407 

Arthropoda Branchiopoda Conchostraca Lynceidae Lynceus 140 2,930 

Arthropoda Copepoda 500 15,000 

Arthropoda Copepoda Calanoida 105 6,150 

Arthropoda Copepoda Cyclopoida 69 6,543 

Arthropoda Copepoda Eucopepoda Canthocamptidae Cletocamptus albuquerquensis 31,250 187,500 

Arthropoda Copepoda Eucopepoda Cyclopidae Cyclops albidus 313 7,813 

Arthropoda Copepoda Eucopepoda Cyclopidae Cyclops bicuspidatus 313 15,625 

Arthropoda Copepoda Eucopepoda Cyclopidae Cyclops fimbriatus 313 1,563 

Arthropoda Copepoda Eucopepoda Cyclopidae Cyclops leuckarti 313 3,125 

Arthropoda Copepoda Eucopepoda Cyclopidae Cyclops serrulatus 313 31,250 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Copepoda Eucopepoda Cyclopidae Cyclops viridis 156 46,875 

Arthropoda Copepoda Eucopepoda Diaptomidae Diaptomus oregonensis 156 1,094 

Arthropoda Copepoda Eucopepoda Diaptomidae Diaptomus shoshone 3,125 31,250 

Arthropoda Copepoda Eucopepoda Diaptomidae Diaptomus siciloides 1,563 187,500 

Arthropoda Copepoda Eucopepoda Diaptomidae Diaptomus tenuicaudatus 156 46,875 

Arthropoda Copepoda Eucopepoda Ergasilidae Ergasilus 1,493 

Arthropoda Copepoda Eucopepoda Laophontidae Laophonte mohammed 1,094 31,250 

Arthropoda Eubranchiopoda Anostraca Branchinectidae Branchinecta lindahli 105 5,800 

Arthropoda Eubranchiopoda Conchostraca Caenestheriidae Caenestheriella 960 

Arthropoda Insecta Coleoptera 313 156,250 7.8 

Arthropoda Insecta Coleoptera Curculionidae Litodactylus griseomicans 913 1,568 

Arthropoda Insecta Coleoptera Curculionidae Lixellus filiformis 913 1,568 

Arthropoda Insecta Coleoptera Dytiscidae 69 192,000 4.9 

Arthropoda Insecta Coleoptera Dytiscidae Acilius 650 1,492 

Arthropoda Insecta Coleoptera Dytiscidae Acilius semisulcatus 811 

Arthropoda Insecta Coleoptera Dytiscidae Agabus 57 4,378 7.5 

Arthropoda Insecta Coleoptera Dytiscidae Agabus ajax 45 9,106 

Arthropoda Insecta Coleoptera Dytiscidae Agabus antennatus 45 9,106 

Arthropoda Insecta Coleoptera Dytiscidae Agabus bifarius 107 811 

Arthropoda Insecta Coleoptera Dytiscidae Agabus falli 3,850 

Arthropoda Insecta Coleoptera Dytiscidae Agabus griseipennis 6,335 15,524 

Arthropoda Insecta Coleoptera Dytiscidae Colymbetes 85 5,266 

Arthropoda Insecta Coleoptera Dytiscidae Colymbetes sculptilis 190 3,310 

Arthropoda Insecta Coleoptera Dytiscidae Coptotomus 57 4,030 8.0 

Arthropoda Insecta Coleoptera Dytiscidae Coptotomus longulus 107 3,019 

Arthropoda Insecta Coleoptera Dytiscidae Dytiscus 80 4,030 7.5 

Arthropoda Insecta Coleoptera Dytiscidae Dytiscus alaskanus 99 4,892 

Arthropoda Insecta Coleoptera Dytiscidae Dytiscus circumcinctus 1,651 1,771 

Arthropoda Insecta Coleoptera Dytiscidae Dytiscus cordieri 45 3,430 

Arthropoda Insecta Coleoptera Dytiscidae Dytiscus hybridus 179 3,850 

Arthropoda Insecta Coleoptera Dytiscidae Graphoderus 57 4,030 7.1 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Insecta Coleoptera Dytiscidae Graphoderus liberus 45 72 

Arthropoda Insecta Coleoptera Dytiscidae Graphoderus occidentalis 140 3,955 

Arthropoda Insecta Coleoptera Dytiscidae Graphoderus perplexus 45 4,340 

Arthropoda Insecta Coleoptera Dytiscidae Hydaticus 105 3,385 

Arthropoda Insecta Coleoptera Dytiscidae Hydaticus modestus 279 2,670 

Arthropoda Insecta Coleoptera Dytiscidae Hydroporus 91 4,210 8.2 

Arthropoda Insecta Coleoptera Dytiscidae Hydroporus criniticoxis 324 

Arthropoda Insecta Coleoptera Dytiscidae Hydroporus notabilis 1,602 

Arthropoda Insecta Coleoptera Dytiscidae Hydroporus pervicinus 228 951 

Arthropoda Insecta Coleoptera Dytiscidae Hydroporus superioris 153 2,575 

Arthropoda Insecta Coleoptera Dytiscidae Hydroporus tenebrosus 183 2,253 

Arthropoda Insecta Coleoptera Dytiscidae Hydroporus undulatus 1,936 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus 57 5,266 7.5 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus canadensis 168 2,046 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus impressopunctatus 153 3,310 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus lutescens 45 4,892 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus masculinus 6,335 15,524 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus patruelis 153 4,078 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus picatus 324 1,791 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus sayi 45 9,106 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus sellatus 190 3,130 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus turbidus 238 2,819 

Arthropoda Insecta Coleoptera Dytiscidae Hygrotus unguicularis 45 9,106 

Arthropoda Insecta Coleoptera Dytiscidae Ilybius 101 4,378 7.1 

Arthropoda Insecta Coleoptera Dytiscidae Ilybius fraterculus 45 3,883 

Arthropoda Insecta Coleoptera Dytiscidae Ilybius subaenus 45 4,892 

Arthropoda Insecta Coleoptera Dytiscidae Laccophilus 57 3,822 7.1 

Arthropoda Insecta Coleoptera Dytiscidae Laccophilus biguttatus 45 3,808 

Arthropoda Insecta Coleoptera Dytiscidae Laccophilus maculosus 217 4,098 

Arthropoda Insecta Coleoptera Dytiscidae Laccornis 171 

Arthropoda Insecta Coleoptera Dytiscidae Liodessus 100 4,973 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Insecta Coleoptera Dytiscidae Liodessus affinis 140 4,744 

Arthropoda Insecta Coleoptera Dytiscidae Potamonectes griseostriatus 45 72 

Arthropoda Insecta Coleoptera Dytiscidae Potamonectes spenceri 6,335 15,524 

Arthropoda Insecta Coleoptera Dytiscidae Potamonectes striatellus 6,335 9,106 

Arthropoda Insecta Coleoptera Dytiscidae Rhantus 57 4,210 7.3 

Arthropoda Insecta Coleoptera Dytiscidae Rhantus frontalis 45 15,524 

Arthropoda Insecta Coleoptera Gyrinidae 153 18,600 6.4 

Arthropoda Insecta Coleoptera Gyrinidae Gyrinus 117 3,405 7.6 

Arthropoda Insecta Coleoptera Gyrinidae Gyrinus maculiventris 363 2,888 

Arthropoda Insecta Coleoptera Gyrinidae Gyrinus minutus 1,682 

Arthropoda Insecta Coleoptera Gyrinidae Gyrinus pectoralis 1,782 

Arthropoda Insecta Coleoptera Haliplidae Haliplus 57 5,160 7.5 

Arthropoda Insecta Coleoptera Haliplidae Haliplus connexus 947 1,048 

Arthropoda Insecta Coleoptera Haliplidae Haliplus hoppingi 217 3,883 

Arthropoda Insecta Coleoptera Haliplidae Haliplus immaculicollis 45 15,524 

Arthropoda Insecta Coleoptera Haliplidae Haliplus leechi 45 72 

Arthropoda Insecta Coleoptera Haliplidae Haliplus salinarius 1,209 3,356 

Arthropoda Insecta Coleoptera Haliplidae Haliplus stagninus 913 11,532 

Arthropoda Insecta Coleoptera Haliplidae Haliplus strigatus 45 15,524 

Arthropoda Insecta Coleoptera Haliplidae Haliplus subguttatus 986 3,720 

Arthropoda Insecta Coleoptera Haliplidae Peltodytes 153 4,210 7.5 

Arthropoda Insecta Coleoptera Haliplidae Peltodytes edentulus 140 3,883 

Arthropoda Insecta Coleoptera Haliplidae Peltodytes tortulosis 1,714 

Arthropoda Insecta Coleoptera Helodidae 134 6,543 

Arthropoda Insecta Coleoptera Hydraenidae 400 192,000 5.3 

Arthropoda Insecta Coleoptera Hydrophilidae 100 192,000 6.1 

Arthropoda Insecta Coleoptera Hydrophilidae Berosus 91 4,720 7.8 

Arthropoda Insecta Coleoptera Hydrophilidae Berosus fraternus 99 3,083 

Arthropoda Insecta Coleoptera Hydrophilidae Berosus hatchi 991 3,430 

Arthropoda Insecta Coleoptera Hydrophilidae Berosus striatus 99 3,706 

Arthropoda Insecta Coleoptera Hydrophilidae Cercyon 1,015 4,210 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Insecta Coleoptera Hydrophilidae Cymbiodyta dorsalis 2,888 

Arthropoda Insecta Coleoptera Hydrophilidae Enochrus 80 5,266 

Arthropoda Insecta Coleoptera Hydrophilidae Enochrus diffusus 3,706 15,524 

Arthropoda Insecta Coleoptera Hydrophilidae Helophorus 57 4,973 

Arthropoda Insecta Coleoptera Hydrophilidae Helophorus linearis 179 2,123 

Arthropoda Insecta Coleoptera Hydrophilidae Helophorus lineatus 205 

Arthropoda Insecta Coleoptera Hydrophilidae Helophorus oblongus 811 1,080 

Arthropoda Insecta Coleoptera Hydrophilidae Hydrobius 69 2,930 

Arthropoda Insecta Coleoptera Hydrophilidae Hydrobius fuscipes 107 

Arthropoda Insecta Coleoptera Hydrophilidae Hydrochara 134 4,210 

Arthropoda Insecta Coleoptera Hydrophilidae Hydrochara obtusatus 273 

Arthropoda Insecta Coleoptera Hydrophilidae Hydrochus 99 4,210 

Arthropoda Insecta Coleoptera Hydrophilidae Laccobius sp. 6,335 11,532 

Arthropoda Insecta Coleoptera Hydrophilidae Paracymus 205 6,120 

Arthropoda Insecta Coleoptera Hydrophilidae Paracymus subcupreus 1,682 2,323 

Arthropoda Insecta Coleoptera Hydrophilidae Tropisternus 123 3,946 

Arthropoda Insecta Coleoptera Hydrophilidae Tropisternus lateralis 148 3,850 

Arthropoda Insecta Coleoptera Scirtidae 100 36,700 5.9 

Arthropoda Insecta Coleoptera Staphylinidae Micralymma 2,000 

Arthropoda Insecta Collembola 272 2,384 8.1 

Arthropoda Insecta Collembola Entomobryidae 118 5,554 

Arthropoda Insecta Collembola Isotomidae 80 6,150 

Arthropoda Insecta Collembola Poduridae 455 1,466 

Arthropoda Insecta Collembola Sminthuridae 134 4,378 

Arthropoda Insecta Diptera 57 15,000 

Arthropoda Insecta Diptera Ceratopogonidae 57 192,000 4.6 

Arthropoda Insecta Diptera Ceratopogonidae Bezzia 540 2,000 7.5 

Arthropoda Insecta Diptera Chaoboridae 544 2,000 7.6 

Arthropoda Insecta Diptera Chaoboridae Chaoborus 57 5,266 7.1 

Arthropoda Insecta Diptera Chironomidae 57 192,000 4.6 

Arthropoda Insecta Diptera Corethrinae 313 10,938 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Insecta Diptera Culicidae 91 192,000 4.6 

Arthropoda Insecta Diptera Culicidae Aedes 69 4,210 

Arthropoda Insecta Diptera Culicidae Culex 159 5,554 

Arthropoda Insecta Diptera Culicidae Culiseta 126 5,554 

Arthropoda Insecta Diptera Culicidae Mansonia perturbans 1,512 2,013 

Arthropoda Insecta Diptera Dixidae 153 2,123 

Arthropoda Insecta Diptera Dolichopodidae 85 192,000 4.6 

Arthropoda Insecta Diptera Ephydridae 140 156,250 4.6 

Arthropoda Insecta Diptera Psychodidae 139 3,536 

Arthropoda Insecta Diptera Psychodidae Pericoma 2,283 2,294 

Arthropoda Insecta Diptera Ptychopteridae 3,698 

Arthropoda Insecta Diptera Sciomyzidae 637 

Arthropoda Insecta Diptera Simuliidae 200 14,800 5.9 

Arthropoda Insecta Diptera Stratiomyidae 80 15,000 

Arthropoda Insecta Diptera Stratiomyidae Odontomyia 105 2,927 

Arthropoda Insecta Diptera Syrphidae 220 6,543 

Arthropoda Insecta Diptera Tabanidae 115 15,000 

Arthropoda Insecta Diptera Tipulidae 57 192,000 5.4 

Arthropoda Insecta Ephemeroptera 313 31,250 

Arthropoda Insecta Ephemeroptera Baetidae 185 19,600 6.9 

Arthropoda Insecta Ephemeroptera Baetidae Baetis 286 2,930 

Arthropoda Insecta Ephemeroptera Baetidae Callibaetis 132 4,098 7.1 

Arthropoda Insecta Ephemeroptera Caenidae 100 18,600 6.4 

Arthropoda Insecta Ephemeroptera Caenidae Caenis 80 2,381 7.1 

Arthropoda Insecta Ephemeroptera Ephemeroptera 185 4,650 

Arthropoda Insecta Hemiptera 313 46,875 

Arthropoda Insecta Hemiptera Belostomatidae Belostoma 531 

Arthropoda Insecta Hemiptera Belostomatidae Lethocerus 572 

Arthropoda Insecta Hemiptera Belostomatidae Lethocerus americanus 210 2,245 

Arthropoda Insecta Hemiptera Corixidae 69 23,100 6.1 

Arthropoda Insecta Hemiptera Corixidae Callicorixa 57 4,378 7.6 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Insecta Hemiptera Corixidae Callicorixa audeni 45 4,892 

Arthropoda Insecta Hemiptera Corixidae Cenocorixa 57 4,098 7.8 

Arthropoda Insecta Hemiptera Corixidae Cenocorixa bifida 45 15,524 

Arthropoda Insecta Hemiptera Corixidae Cenocorixa expleta 6,335 15,524 

Arthropoda Insecta Hemiptera Corixidae Corisella 3,018 

Arthropoda Insecta Hemiptera Corixidae Corisella tarsalis 1,628 1,679 

Arthropoda Insecta Hemiptera Corixidae Cymatia 98 3,350 7.5 

Arthropoda Insecta Hemiptera Corixidae Cymatia americana 45 4,892 

Arthropoda Insecta Hemiptera Corixidae Dasycorixa rawsoni 858 15,524 

Arthropoda Insecta Hemiptera Corixidae Hespercorixa 531 2,000 7.5 

Arthropoda Insecta Hemiptera Corixidae Hesperocorixa 57 4,030 

Arthropoda Insecta Hemiptera Corixidae Hesperocorixa atopodonta 107 811 

Arthropoda Insecta Hemiptera Corixidae Hesperocorixa laevigata 45 9,106 

Arthropoda Insecta Hemiptera Corixidae Hesperocorixa vulgaris 811 

Arthropoda Insecta Hemiptera Corixidae Sigara 57 5,800 7.5 

Arthropoda Insecta Hemiptera Corixidae Sigara solensis 99 1,940 

Arthropoda Insecta Hemiptera Corixidae Trichocorixa 274 4,210 7.6 

Arthropoda Insecta Hemiptera Corixidae Trichocorixa borealis 1,608 4,098 

Arthropoda Insecta Hemiptera Corixidae Trichocorixa naias 1,782 4,098 

Arthropoda Insecta Hemiptera Corixidae Trichocorixa verticalis interiores 2,014 3,755 

Arthropoda Insecta Hemiptera Gerridae 105 3,650 

Arthropoda Insecta Hemiptera Gerridae Gerris 80 4,210 

Arthropoda Insecta Hemiptera Gerridae Rheumatobates 194 2,030 

Arthropoda Insecta Hemiptera Hebridae Hebridae 925 1,583 8.5 

Arthropoda Insecta Hemiptera Mesovelidae Mesovelia 665 959 7.3 

Arthropoda Insecta Hemiptera Nepidae Ranatra fusca 1,682 

Arthropoda Insecta Hemiptera Notonectidae 100 26,900 5.3 

Arthropoda Insecta Hemiptera Notonectidae Buenoa 308 2,745 7.5 

Arthropoda Insecta Hemiptera Notonectidae Buenoa margaritacea 1,064 4,098 

Arthropoda Insecta Hemiptera Notonectidae Notonecta 101 3,850 7.5 

Arthropoda Insecta Hemiptera Notonectidae Notonecta kirbyi 45 15,524 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Insecta Hemiptera Notonectidae Notonecta undulata 45 4,098 

Arthropoda Insecta Hemiptera Pleidae 240 978 

Arthropoda Insecta Hemiptera Pleidae Neoplea 286 2,000 7.1 

Arthropoda Insecta Hemiptera Pleidae Plea 270 3,350 

Arthropoda Insecta Hemiptera Velidae 531 942 7.3 

Arthropoda Insecta Odonata 80 46,875 7.1 

Arthropoda Insecta Odonata Aeshnidae 117 30,200 5.9 

Arthropoda Insecta Odonata Aeshnidae Aeshna 57 4,973 7.8 

Arthropoda Insecta Odonata Aeshnidae Anax 150 4,340 

Arthropoda Insecta Odonata Coenagrionidae 200 30,200 6.7 

Arthropoda Insecta Odonata Coenagrionidae Amphiagrion 223 

Arthropoda Insecta Odonata Coenagrionidae Coenagrion 680 1,890 

Arthropoda Insecta Odonata Coenagrionidae Enallagma 57 4,378 7.3 

Arthropoda Insecta Odonata Coenagrionidae Ischnura 85 2,710 

Arthropoda Insecta Odonata Corduliidae 100 33,600 5.3 

Arthropoda Insecta Odonata Lestidae 117 36,700 5.3 

Arthropoda Insecta Odonata Lestidae Lestes 100 4,210 7.1 

Arthropoda Insecta Odonata Libellulidae 194 30,200 6.4 

Arthropoda Insecta Odonata Libellulidae Leucorrhinia 147 2,470 

Arthropoda Insecta Odonata Libellulidae Libellula 147 4,340 

Arthropoda Insecta Odonata Libellulidae Sympetrum 98 4,210 7.5 

Arthropoda Insecta Trichoptera 313 31,250 

Arthropoda Insecta Tricoptera 98 15,000 

Arthropoda Insecta Tricoptera Brachycentridae Brachycentrus 1,454 2,012 

Arthropoda Insecta Tricoptera Leptoceridae 100 69,100 4.9 

Arthropoda Insecta Tricoptera Leptoceridae Nectopsyche 1,421 

Arthropoda Insecta Tricoptera Leptoceridae Oecetis 826 1,030 7.5 

Arthropoda Insecta Tricoptera Leptoceridae Triaenodes 845 1,640 8.1 

Arthropoda Insecta Tricoptera Limnephilidae 245 2,725 

Arthropoda Insecta Tricoptera Limnephilidae Anabolia 873 

Arthropoda Insecta Tricoptera Limnephilidae Limnephilus 240 2,407 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Arthropoda Insecta Tricoptera Molannidae Molanna flavicornis 911 3,808 

Arthropoda Insecta Tricoptera Phryganeidae 150 15,000 

Arthropoda Insecta Tricoptera Phryganeidae Fabria 910 1,640 7.3 

Arthropoda Insecta Tricoptera Polycentropodidae 363 15,000 

Arthropoda Insecta Tricoptera Polycentropodidae Cernotina 540 1,542 7.6 

Arthropoda Malacostraca Amphipoda 313 46,875 

Arthropoda Malacostraca Amphipoda Ceinidae 100 69,100 4.9 

Arthropoda Malacostraca Amphipoda Gammaridae 500 15,000 

Arthropoda Malacostraca Amphipoda Gammaridae Gammarus 118 4,214 

Arthropoda Malacostraca Amphipoda Perthiidae 200 7,600 5.4 

Arthropoda Malacostraca Amphipoda Talitridae 500 15,000 

Arthropoda Malacostraca Amphipoda Talitridae Hyalella 98 6,543 7.1 

Arthropoda Malacostraca Amphipoda Talitridae Hyalella azteca 254 4,098 

Arthropoda Malacostraca Decapoda Cambaridae Cambarus 313 3,906 

Arthropoda Malacostraca Decapoda Palaemonidae 200 34,100 6.1 

Arthropoda Malacostraca Decapoda Parastacidae 100 29,200 5.9 

Arthropoda Ostracoda 101 6,150 Bryozoa Phylactolaemata Plumatellida Plumatellidae Plumatella 313 1,563 

Cnidaria Hydrozoa Hydroida Hydridae Hydra 313 1,875 

Mollusca Bivalvia Pelecypoda Sphaeriidae 162 3,125 

Mollusca Bivalvia Pelecypoda Sphaeriidae Pisidium casertanum 185 

Mollusca Bivalvia Pelecypoda Sphaeriidae Pisidium nitidum 168 2,594 

Mollusca Bivalvia Pelecypoda Sphaeriidae Sphaerium lacustre 217 1,608 

Mollusca Bivalvia Pelecypoda Unionidae Anodonta 313 469 

Mollusca Gastropoda 313 15,625 

Mollusca Gastropoda Limnophila Lymnaeidae 119 15,000 

Mollusca Gastropoda Limnophila Lymnaeidae Lymnaea 196 3,658 

Mollusca Gastropoda Limnophila Lymnaeidae Lymnaea caparata 510 1,618 

Mollusca Gastropoda Limnophila Lymnaeidae Lymnaea elodes 57 4,214 

Mollusca Gastropoda Limnophila Lymnaeidae Lymnaea reflexa 324 1,416 

Mollusca Gastropoda Limnophila Lymnaeidae Lymnaea stagnalis 767 1,985 
Appendix 6. Summary of invertebrate database.—Continued[The minimum and maximum values represent all data types (that is, values reported as minimum, maximum, mean, median, or single occurrence). μS cm-1, microsiemens per c

Phylum Class Order Family Genus Species Conductivity (μS cm-1) p

Min Max Min 

Mollusca Gastropoda Limnophila Lymnaidae 540 1,579 7.9 

Mollusca Gastropoda Limnophila Physidae 500 15,000 7.1 

Mollusca Gastropoda Limnophila Physidae Aplexa hypnorum 188 3,480 

Mollusca Gastropoda Limnophila Physidae Physa 119 286 

Mollusca Gastropoda Limnophila Physidae Physa gyrina 351 3,880 

Mollusca Gastropoda Limnophila Physidae Physa jennessi 98 5,266 

Mollusca Gastropoda Limnophila Planorbidae 105 15,000 7.1 

Mollusca Gastropoda Limnophila Planorbidae Armiger crista 57 4,250 

Mollusca Gastropoda Limnophila Planorbidae Gyralus 168 2,407 

Mollusca Gastropoda Limnophila Planorbidae Gyralus parvus 85 4,378 

Mollusca Gastropoda Limnophila Planorbidae Gyraulus circumstriatus 57 4,650 

Mollusca Gastropoda Limnophila Planorbidae Helisoma anceps 334 1,264 

Mollusca Gastropoda Limnophila Planorbidae Helisoma trivolvis 98 3,456 

Mollusca Gastropoda Limnophila Planorbidae Planorbula campestris 220 261 

Mollusca Gastropoda Limnophila Planorbidae Promenetus exacuous 98 4,340 

Mollusca Gastropoda Limnophila Planorbidae Promenetus umbilicatellus 69 3,680 

Mollusca Gastropoda Mesogastropoda Valvatidae Valvata lewesi 122 1,985 

Nematoda 313 156,250 Nematomorpha Gordioida Gordea Gordiidae Gordius 313 1,094 

Platyhelminthes Turbellaria Tricladida Planariidae Planaria 313 469 

1Source: 1, Johnson, 1990; 2, Kay and others, 2001; 3, Lancaster and Scudder, 1987; 4, McCarraher, 1970; 5, Rawson and Moore, 1944; 6, Tangen, unpublished; 7, USGS, unpubl
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 macroinvertebrate 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 Naturalist, 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.
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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

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