Characterization and Analysis of Temporal 

and Spatial Variations in Habitat and 

Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs 

and Vicinity, Colorado, 1998–2001

By James F. Bruce

Denver, Colorado

2002

Prepared in cooperation with the

COLORADO SPRINGS CITY ENGINEERING, and

COLORADO SPRINGS UTILITIES

U.S. GEOLOGICAL SURVEY

Water-Resources Investigations Report 02–4093
District Chief

U.S. Geological Survey

Box 25046, Mail Stop 415

Denver Federal Center

Denver, CO 80225–0046

For additional information write to:

District Chief

U.S. Geological Survey

Box 25046, Mail Stop 415

Denver Federal Center

Denver, CO 80225–0046

U.S. DEPARTMENT OF THE INTERIOR

GALE A. NORTON, Secretary

U.S. GEOLOGICAL SURVEY

Charles G. Groat, Director

The use of firm, trade, and brand names in the report is for identification purposes only and does 

not constitute endorsement by the U.S. Geological Survey.

Copies of this report can be purchased 

from:

U.S. Geological Survey

Information Services

Box 25286

Denver Federal Center

Denver, CO 80225
CONTENTS III

CONTENTS

Abstract.................................................................................................................................................................................. 1

Introduction............................................................................................................................................................................ 1

Purpose and Scope....................................................................................................................................................... 2

Description of Study Area ........................................................................................................................................... 2

Land Use...................................................................................................................................................................... 4

Acknowledgments ....................................................................................................................................................... 5

Methods of Investigation ....................................................................................................................................................... 6

Riparian and Instream Habitat ..................................................................................................................................... 6

Data Collection .................................................................................................................................................. 6

Statistical Analysis of Habitat Data................................................................................................................... 7

Macroinvertebrates ...................................................................................................................................................... 7

Sample Collection.............................................................................................................................................. 7

Sample Processing and Quality Assurance........................................................................................................ 8

Description of Metrics ....................................................................................................................................... 8

Taxa Richness and Total Abundance ....................................................................................................... 9

EPT and Chironomidae............................................................................................................................ 9

Dominant Taxa......................................................................................................................................... 9

Similarity Measures, Cluster Analysis, and Multivariate Analysis ................................................................... 9

Statistical Analysis of Macroinvertebrate Data ................................................................................................. 10

Temporal and Spatial Variation in Habitat and Macroinvertebrates...................................................................................... 10

Habitat.......................................................................................................................................................................... 10

Seasonal Variations............................................................................................................................................ 11

Spatial and Annual Variations ........................................................................................................................... 11

Macroinvertebrates ...................................................................................................................................................... 15

Comparison of Replicate and Composite Samples............................................................................................ 18

Seasonal Variations............................................................................................................................................ 19

Spatial Variation in Invertebrate Community Structure..................................................................................... 21

Environmental Gradient Analysis by Canonical Correspondence Analysis...................................................... 23

Summary................................................................................................................................................................................ 27

Selected References ............................................................................................................................................................... 27

Appendix A. Habitat data on CD-ROM (in pocket)

Appendix B. Macroinvertebrate data on CD-ROM (in pocket)

FIGURES

1. Map showing location of study area and sampling sites in the Fountain Creek Basin............................................... 3

2–6. Graphs showing:

2. Pebble-count data for selected sites in the Fountain Creek Basin, 1998–2001 ................................................. 12

3. Number of macroinvertebrate taxa collected during the fall at selected sites in the Fountain 

Creek Basin, 1998–2000 .................................................................................................................................... 16

4. Total abundance of macroinvertebrates collected during the fall at selected sites in the Fountain 

Creek Basin, 1998–2000 .................................................................................................................................... 16

5. Relative abundance of chironomids collected during the fall at selected sites in the Fountain 

Creek Basin, 1998–2000 .................................................................................................................................... 17

6. Ratio of EPT abundance to chironomid abundance from fall samples collected at selected sites 

in the Fountain Creek Basin, 1998–2000........................................................................................................... 17

7. Dendrogram based upon Jaccard’s Coefficient of Community values summarizing the similarity 

between selected invertebrate communities collected from the Fountain Creek Basin, 1998–2000.......................... 22

8. Dendrogram based upon Sorenson’s Coefficient of Community values summarizing the similarity 

of invertebrate community structure between selected sites in the Fountain Creek Basin, 1998–2000..................... 25

9. Canonical Correspondence Analysis ordination of sampling sites, species, and environmental variables 

for selected sites in the Fountain Creek Basin, 1998–2000 ........................................................................................ 26
IV CONTENTS

TABLES

1. Sampling sites, dates of collection, and types of samples collected in the Fountain Creek Basin, 

1998–2001............................................................................................................................................................... 4

2. Land uses and estimated total impervious and pervious area for the Fountain Creek Basin, 1998–2000.............. 5

3. Summary of two-way analysis of variance (ANOVA) evaluating seasonal and annual differences of 

10 habitat variables measured at selected sites in the Fountain Creek Basin, 1998–2001 ..................................... 13

4. Results of two-way analysis of variance (ANOVA) and Student-Newman-Keuls (SNK) multiple range 

tests for spatial variation in 21 instream and riparian habitat variables measured at selected sites in the 

Fountain Creek Basin, 1998–2000.......................................................................................................................... 14

5. Results of Wilcoxon signed-rank test and Sorenson’s Coefficient of Community for paired replicate 

and composite invertebrate samples collected from selected sites in the Fountain Creek Basin, 1998–2001........ 18

6. Summary of Sorenson’s Coefficient of Community analysis of paired replicate and composite 

invertebrate samples collected from selected sites in the Fountain Creek Basin, 1998–2001................................ 19

7. Mean metric values for fall and spring samples collected from selected sites in the Fountain Creek 

Basin, 1998–2001.................................................................................................................................................... 20

8. Combined results of all fall and spring metric mean comparisons from 10 sites in the Fountain Creek 

Basin, 1998–2001.................................................................................................................................................... 21

9. Seasonal differences in macroinvertebrate samples based upon Jaccard’s Coefficient of Community 

values from selected sites in the Fountain Creek Basin, 1998–2000 ...................................................................... 22

10. Results of Wilcoxon signed-rank test for paired fall and spring metric values from macroinvertebrate 

samples collected from selected sites in the Fountain Creek Basin, 1998–2001.................................................... 23

11. Results of one-way analysis of variance (ANOVA) and Student-Newman-Keuls (SNK) multiple range 

tests for spatial variation in invertebrate community structure from selected sites in the Fountain Creek 

Basin, 1998–2000.................................................................................................................................................... 24

12–18. Tables on CD-ROM (in pocket):

12. Detailed instream and riparian habitat variables measured at selected sites in the Fountain Creek Basin, 

1998–2001

13. Detailed point instream habitat variables measured at selected sites in the Fountain Creek Basin, 1998–2001

14. Pebble-count data for selected sites in the Fountain Creek Basin, 1998–2001

15. Habitat variable scores based upon rapid bioassessment protocols for selected sites in the Fountain Creek

Basin, 1998–2001

16. Geomorphic channel unit measurements and water-surface gradient for selected sites in the Fountain Creek

Basin, 1998–2001

17. Macroinvertebrate taxa collected by kicksweep method at selected sites in the Fountain Creek Basin, 

1998–2001

18. Density of macroinvertebrates collected by Hess sampler at selected sites in the Fountain Creek Basin, 

1998–2001
CONTENTS V

CONVERSION FACTORS, ABBREVIATIONS, AND VERTICAL DATUM

Degree Celsius (C) may be converted to degree Fahrenheit (F) by using the following equation:

F = 9/5 (C) + 32

Degree Fahrenheit (F) may be converted to degree Celsius (C) by using the following equation:

C = 5/9 (F – 32)

Sea level: In this report “sea level” refers to the National Geodetic Vertical Datum of 1929 (NGVD of 1929)—a geodetic datum derived 

from a general adjustment of the first-order level nets of both the United States and Canada, formerly called Sea Level Datum of 1929.

Multiply By To obtain

cubic foot per second (ft3/s) 2.83210–2 cubic meter per second (m3/s)

foot (ft) 3.04810–1 meter (m)

foot per second (ft/s) 3.04810–1 meter per second (m/s)

gallon (gal) 3.785100 liter (L)

gallon (gal) 3.785103 milliliter (mL)

inch 2.54100 centimeter (cm)

inch 2.54101 millimeter (mm)

inch 2.54103 micrometer (m)

mile (mi) 1.609100 kilometer (km)

square foot (ft2) 9.2910–2 square meter (m2)

square mile (mi2) 2.590100 square kilometer (km2)
INTRODUCTION 1

Abstract

The Fountain Creek Basin in and 

around Colorado Springs, Colorado, is affected 

by various land- and water-use activities. Biological, hydrological, water-quality, and land-use 

data were collected at 10 sites in the Fountain 

Creek Basin from April 1998 through April 2001 

to provide a baseline characterization of macroinvertebrate communities and habitat conditions 

for comparison in subsequent studies; and to 

assess variation in macroinvertebrate community 

structure relative to habitat quality. Analysis 

of variance results indicated that instream and 

riparian variables were not affected by season, 

but significant differences were found among 

sites. Nine metrics were used to describe and 

evaluate macroinvertebrate community structure. 

Statistical analysis indicated that for six of the 

nine metrics, significant variability occurred 

between spring and fall seasons for 60 percent 

of the sites. Cluster analysis (unweighted pair 

group method average) using macroinvertebrate 

presence-absence data showed a well-defined 

separation between spring and fall samples. 

Six of the nine metrics had significant spatial 

variation. Cluster analysis using Sorenson’s 

Coefficient of Community values computed from 

macroinvertebrate density (number of organisms 

per square meter) data showed that macroinvertebrate community structure was more similar 

among tributary sites than main-stem sites. 

Canonical correspondence analysis identified a 

substrate particle-size gradient from site-specific 

species-abundance data and environmental correlates that decreased the 10 sites to 5 site clusters 

and their associated taxa.

INTRODUCTION

Biomonitoring uses living organisms to 

evaluate the quality of an aquatic environment. 

Aquatic biological communities integrate acute 

and(or) chronic changes in the physical, chemical, 

and biotic components of their environment (Plafkin 

and others, 1989; Cuffney and others, 1993). Therefore, evaluating the ecological components of an 

aquatic community is useful in assessing the variation 

in macroinvertebrate communities and the effects of 

environmental perturbations on macroinvertebrate 

community structure. Also, programs that are based on 

long-term monitoring are useful in identifying trends 

in biological measurements as well as nonspecific 

water quality. Fitzpatrick and Giddings (1997) have 

also suggested that complementary habitat analysis is 

useful in furthering the understanding of the interactions among chemical, physical, and biological characteristics. When habitat quality among sites is similar, 

differences in macroinvertebrate communities and 

measures of these communities can be attributed to 

water-quality factors. But, when the habitat quality 

differs, evaluation of the habitat is necessary to determine the magnitude that habitat may be a limiting 

factor in biological community structure.

Characterization and Analysis of Temporal and 

Spatial Variations in Habitat and Macroinvertebrate 

Community Structure, Fountain Creek Basin, 

Colorado Springs and Vicinity, Colorado, 1998–2001

By James F. Bruce
2 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

Protocols developed for the U.S. Geological 

Survey (USGS) National Water-Quality Assessment 

Program (NAWQA) by Meador and others (1993) 

and Rapid Bioassessment Protocols (RBP) developed 

by the U.S. Environmental Protection Agency 

(USEPA) (Plafkin and others, 1989; Barbour and 

others, 1997) were used to assess instream and riparian 

habitat conditions. Habitat assessment promotes an 

understanding of the relations among physical factors 

that might be limiting to biological communities and 

provides baseline information necessary to identify 

future changes in habitat conditions. 

Land-use change in the Fountain Creek 

Basin could result in increased storm runoff over 

short periods of time and alter baseflow characteristics 

which might be detrimental to the aquatic community. 

Changes in erosion, sediment transport, and deposition 

could affect channel morphology, aquatic habitat, and 

macroinvertebrate community structure. Little information about the stream-dwelling macroinvertebrates 

or habitat condition of the Fountain Creek Basin has 

been compiled, analyzed, or published. To address 

this concern the USGS, in cooperation with the 

Colorado Springs City Engineering and Colorado 

Springs Utilities, began a study in 1998 to assess 

temporal and spatial variations in habitat and 

macroinvertebrate community structure at selected 

sites in the Fountain Creek Basin in the vicinity of 

Colorado Springs, Colo. The results from this study 

can be used as baseline information to augment 

long-term monitoring of water quality, instream 

and riparian quality and diversity, and biological 

condition.

Purpose and Scope

This report includes an assessment of temporal 

and spatial variations in habitat and macroinvertebrate 

community structure from 10 sites in the Fountain 

Creek Basin in the vicinity of Colorado Springs, Colo. 

(fig. 1, table 1). Macroinvertebrate communities and 

measures of these communities were evaluated, and 

a baseline characterization of habitat and macroinvertebrate community structure were determined for 

the sites. Habitat and macroinvertebrate data were 

collected during spring and fall of 1998 through 

2001 at three sites on Fountain Creek (3700, 5500, 

5800); one site on Monument Creek, site 3970; 

four sites in Cottonwood Creek drainage (3977a, 

3977b, 3980, 3985, and 3990); and one site on 

North Rockrimmon Creek (4050). All habitat data 

collected during 1998–2001 were used in the habitat 

analysis. Macroinvertebrate data collected during 

the fall of 2001 were not available at time of report 

preparation.

Description of Study Area

The study area has a drainage area of about 

495 mi2

 (fig. 1). Elevations in the study area range 

from about 5,460 ft at the southern end of the study 

area to 14,109 ft at the summit of Pikes Peak. The 

Front Range of the southern Rocky Mountains and 

the Colorado Piedmont (Hansen and Crosby, 1982) 

are the two major landforms in the study area. The 

Front Range, which comprises the western onethird of the study area, is underlain by granite. The 

Colorado Piedmont, which comprises the remaining 

two-thirds of the area from about 3 to 5 mi west of 

Interstate Highway 25 to the eastern edge of the 

basin, is underlain by sandstone, shale, and alluvial 

and windlain deposits. Soils in this area are generally 

sandy, well drained, with more gentle slopes (Larsen, 

1981; von Guerard, 1989) than soils developed 

in granite. More details of the soils and geology 

of the study area are contained in von Guerard 

(1989).

Fountain and Monument Creeks are the two 

main drainages in the study area. Fountain Creek 

is a perennial stream that originates near Woodland 

Park, Colo., and flows southeastward through a deeply 

incised canyon to Manitou Springs, Colo. The stream 

channel upstream from Manitou Springs is meandering, has a pool-and-riffle regime, and the bed 

material ranges from sand and gravel to cobbles 

and boulders. From Manitou Springs, Fountain 

Creek flows through alluvial terraces into a wide 

alluvial valley; the channel meanders less due to 

channelization, has a pool-and-riffle-and-run regime, 

and the bed material is predominantly sand, gravel, 

and cobbles. Downstream from the confluence 

with Monument Creek, the channel becomes braided, 

and the streambanks are intermittently lined with 

concrete. The bed material downstream from the 

confluence with Monument Creek is variable. Some 

stream reaches have predominantly cobble beds, 

some are scoured to bedrock, and others are a 

mixture of sand, gravel, and cobbles (von Guerard, 

1989).
INTRODUCTION 3

Albers Equal-area projection

Base from U.S. Geological Survey

Standard parallels 37° 30' N and 40° 30' N

Central Meridan 105° 30' W

Study area

Denver

105° 104° 45'

38° 45'

39°

0 5 KILOMETERS

0 5 MILES

Monument Creek

Cottonwood Creek

Sand Creek

East Fork Sand Creek

Fountain Creek

Fountain Creek

Figure 1. Location of study area and sampling sites in the Fountain Creek Basin.
4 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

Monument Creek, the main tributary to 

Fountain Creek, is a perennial stream that originates 

in the Rampart Range and flows eastward toward 

Palmer Lake, then south to Colorado Springs. 

Upstream from site 3970, Monument Creek is 

meandering, has a pool-and-riffle-and-run regime, 

and the streambed consists of sand, gravel, and 

cobbles. Downstream from site 3970, the channel 

becomes braided, sand and small gravel compose 

the streambed, and the banks are intermittently lined 

with concrete. The braided channel conditions occur 

intermittently throughout the remaining length of the 

channel. 

Cottonwood Creek (fig. 1) has historically been 

an ephemeral stream; however, over the past 15 years, 

flow exists throughout the year. The stream channel is 

meandering, has a riffle-and-run regime, and the bed 

material is predominantly sand.

North Rockrimmon Creek originates in the 

northwest portion of Colorado Springs and flows 

southeast into Monument Creek near site 4050 (fig. 1). 

The stream channel is fairly straight, has a pool-andriffle regime, and the streambed consists of sand, 

gravel, cobbles, and boulders. The flow is affected 

by ground-water discharge and runoff from domestic 

housing, commercial, and industrial developments 

along its route.

Land Use

Land uses within the study area include urban, 

military reservations, agriculture, and undeveloped 

areas. Substantial changes in land use have occurred 

from increased population. Table 2 shows the total 

area for various land-use categories for 1998, 1999, 

and 2000. Estimates of the percentage of impervious 

and pervious material associated with each land use 

were applied to estimate the total impervious and 

pervious area upstream from each sampling site 

(table 2). Estimates of the percentage of impervious 

and pervious material associated with each land use 

Table 1. Sampling sites, dates of collection, and types of samples collected in the Fountain Creek Basin, 1998–2001

[FOCR, Fountain Creek; MOCR, Monument Creek; COCR, Cottonwood Creek; NORO, North Rockrimmon Creek; MS4, municipal stormwater permit 

site; RW, receiving water or main-stem site; B, benthic macroinvertebrate sample; N, National Water-Quality Assessment Program Protocols; R, Rapid 

Bioassessment Protocols; --, not sampled or measured]

Site number and name1

(see fig. 1 for location) Site type

Sample date and types of samples

1998 1999 2000 2001

Spring

B/N/R

Fall

B/N/R

Spring

B/N/R

Fall

B/N/R

Spring

B/N/R

Fall

B/N/R

Spring

B/N/R

Fall

B/N/R

3700,

FOCR @ 33rd

Main stem, RW X/X/X X/--/X X/X/X X/--/X X/X/X X/--/X --/--/-- X/X/X

3970,

MOCR @ Woodmen

Main stem, RW X/X/X X/--/X X/X/X X/--/X X/X/X X/--/X --/--/-- X/X/X

3977a,

COCR @ Cowpoke

Tributary to Monument 

Creek, MS4

X/X/-- X/X/-- --/X/-- X/X/-- X/X/-- X/X/-- X/X/-- X/X/X

3977b,

COCR @ Cowpoke

Tributary to Monument 

Creek, MS4

X/X/-- X/X/-- --/X/-- X/X/-- X/X/-- X/X/-- X/X/-- X/X/X

3980,

COCR @ Woodmen

Tributary to Monument 

Creek, MS4

X/X/-- X/X/-- --/X/-- X/X/-- X/X/-- X/X/-- X/X/-- X/X/X

3985,

COCR Trib @ Rangewood

Tributary to Cottonwood 

Creek, MS4

X/X/-- X/X/-- --/X/-- X/X/-- X/X/-- X/X/-- X/X/-- X/X/X

3990,

COCR @ Mouth @ Vincent

Tributary to Monument 

Creek, MS4

X/X/-- X/X/-- --/X/-- X/X/-- X/X/-- X/X/-- X/X/-- X/X/X

4050,

NORO @ Delmonico

Tributary to Monument 

Creek, MS4

X/X/-- X/X/-- --/X/-- X/X/-- X/X/-- X/X/-- X/X/-- X/X/X

5500,

FOCR @ Nevada

Main stem, RW X/X/X X/--/X X/X/X X/--/X X/X/X X/--/X --/--/-- X/X/X

5800,

FOCR @ Security

Main stem, RW X/X/X X/--/X X/X/X X/--/X X/X/X X/--/X --/--/-- X/X/X

1

Site name used in Colorado Springs Municipal Stormwater Permit.
INTRODUCTION 5

were applied using coefficients adapted from Arnold 

and Gibbons (1996) and the U.S. Environmental 

Protection Agency (1992). The following estimates of 

percent impervious material were used: 88 percent of 

commercial, 75 percent of industrial, 50 percent of residential, 90 percent of streets and easements, 15 percent 

of airports and military reservations, 15 percent of 

agricultural, and 15 percent of undeveloped land-use 

categories. Land-use data for 1998–2000 were provided 

by the City of Colorado Springs. Between 1998 and 

2000, the impervious area upstream from site 3700 

(Fountain Creek upstream from Manitou Springs) 

showed no appreciable change. In Monument Creek, 

drainage upstream from Cottonwood Creek (site 3970), 

no appreciable change in impervious area occurred 

between 1998 and 2000. Overall, the amount of 

estimated impervious area within the 495-mi2

 study 

area upstream from site 5800, increased by an estimated 

8 percent between 1998 and 2000.

Acknowledgments

The author greatly appreciates the efforts of 

Wade J. Walker, Erin E. Evans, Brian G. Scott, and 

Cathy M. Tate, who assisted in the collection of these 

data. Thanks to Bill Van Derveer and Tara Kelley 

of Colorado Springs Utilities for their cooperation 

and for providing habitat characterization and macroinvertebrate data for the main-stem sites. Thanks also 

to Dwight W. Moore for his discussions concerning 

the statistical analysis of these data and to Marianne 

August for her work on figure 1.

Table 2. Land uses and estimated total impervious and pervious area for the Fountain Creek Basin, 1998–2000

[mi2, square miles; --, category not reported]

Site 

(see

table 1)

Drainage

area1

 (mi2)

Commercial

and 

industrial 

(mi2)

Residential 

 (mi2)

Streets 

and 

easements

(mi2)

Airports 

and

military 

 (mi2)

Agriculture

 (mi2)

Undeveloped

(mi2)

Estimated 

total 

impervious 

area

 (mi2)

Estimated

 total 

pervious 

area

 (mi2)

Cumulated

drainage

area2

(mi2)

1998

3700 103 3.91 6.26 2.55 0.00 1.25 88.0 22.2 79.8 102

3970 181 4.72 28.8 7.15 28.9 22.7 87.8 45.8 134 180

3977 5.93 0.06 2.08 -- 0.00 0.94 3.03 1.69 4.42 6.11

3985 2.81 0.17 0.58 -- 0.00 0.96 1.07 0.73 2.05 2.78

3990 18.7 1.19 6.27 -- 0.00 2.44 8.98 5.85 13.0 18.9

5500 392 18.2 55.7 19.1 28.9 32.0 233 105 282 387

5800 495 31.6 82.9 36.8 34.9 59.0 257 154 348 502

1999

3700 103 4.94 7.62 1.66 0.00 1.24 87.1 22.9 79.7 103

3970 181 5.34 29.0 6.76 28.9 25.5 84.5 46.0 134 180

3977 5.93 0.04 2.26 -- 0.00 2.95 .95 1.75 4.45 6.20

3985 2.81 0.17 0.94 -- 0.00 0.46 1.25 0.87 1.95 2.82

3990 18.7 0.91 7.97 -- 0.00 4.57 5.36 6.26 12.6 18.8

5500 392 20.1 57.5 17.7 28.9 34.5 228 106 281 387

5800 495 33.8 85.8 35.4 34.9 60.5 253 155 348 503

2000

3700 103 7.10 5.53 1.79 0.00 1.18 86.4 23.8 78.2 102

3970 181 5.71 30.2 7.44 28.9 22.9 85.3 45.0 135 180

3977 5.93 0.56 2.40 -- 0.00 2.77 0.46 2.18 4.01 6.19

3985 2.81 0.71 0.66 -- 0.00 0.33 1.10 1.16 1.64 2.80

3990 18.7 1.88 7.55 -- 0.00 5.07 4.32 6.81 12.0 18.8

5500 392 23.4 57.7 26.0 28.9 30.6 220 114 273 387

5800 495 38.4 86.2 43.8 34.8 55.1 245 166 337 503

1

Drainage area reported by the U.S. Geological Survey (Crowfoot and others, 2000).

2Drainage area computed from accumulating individual land uses provided by the City of Colorado Springs.
6 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

METHODS OF INVESTIGATION

Habitat data and benthic macroinvertebrate 

samples were collected during the spring and fall at 

selected sites in the Fountain Creek Basin (table 1). 

Several statistical and ordination approaches were 

used to assess variability between replicate and 

composite Hess stream-bottom samples and temporal 

(seasonal and year) and spatial (between and among 

sites) variability in habitat and macroinvertebrate 

community structure.

Riparian and Instream Habitat

Quantitative and qualitative instream and 

riparian habitat variables were measured following 

the procedures described by Plafkin and others (1989), 

Barbour and others (1997), and Meador and others 

(1993) at four main-stem sites and six tributary sites 

in Fountain Creek Basin (fig. 1, table 1). Colorado 

Springs Utilities (CSU) personnel collected habitat 

data at main-stem sites 3700, 3970, 5500, and 5800. 

USGS personnel collected habitat data at tributary 

sites 3977a, 3977b, 3980, 3985, 3990, and 4050. CSU 

and USGS personnel collaborated on the habitat data 

collection for the main-stem sites during the fall 2001.

Data Collection

 CSU and USGS personnel collected data using 

the same procedures, and, for the most part, the same 

habitat variables were measured. Representative 

reaches were selected on the basis of criteria defined 

by Meador and others (1993) and equally subdivided 

into six transects. All sites had at least one transect 

permanently marked with 4-ft sections of 0.5-inch 

rebar driven to within about 1 ft of the ground surface. 

Habitat assessments were conducted in the spring 

(April or May) and fall (October or November) from 

1998 through 2001.

Two categorical habitat assessments were 

made at each site. The first type (RBP) was a more 

qualitative assessment of instream and riparian 

habitat parameters encountered over the entire reach 

(Plafkin and others, 1989; Barbour and others, 1997). 

Habitat measurements based on RBP were made 

during the spring at sites 3700, 3970, 5500, and 5800 

from 1998–2000, and at all sites during the fall 2001 

(fig. 1, table 1). For each site, the scores for 12 habitat 

parameters were summed to derive a single numeric 

value (maximum possible total score = 180, unitless). 

These habitat parameters were separated into three 

main categories. The primary parameters (scored 

0–20) characterize the instream habitats and have 

the greatest direct influence on biological community 

structure. Secondary parameters (scored 0–15) 

measure the overall channel morphology of the stream 

reach. Tertiary parameters (scored 0–10) characterize 

bank structure and riparian areas. These three categories are weighted according to their influence on the 

biological community, with primary parameters given 

the most weight and secondary parameters having 

more weight than tertiary parameters.

The second type of assessment (NAWQA) 

was a more quantitative study of instream and riparian 

habitat variables (Meador and others, 1993). Habitat 

measurements based on the NAWQA protocols were 

made during the spring for all sites and also in the fall 

for sites 3977a, 3977b, 3980, 3985, 3990, and 4050 

(fig. 1, table 1). These habitat data were collected 

from the main-stem sites by CSU personnel from 

1998 to 2000 and collaboratively by CSU and USGS 

for the fall 2001. All habitat data collection from the 

tributary sites was done by USGS personnel. The 

NAWQA reach characterization included slope, 

channel, substrate, bank, flow, canopy, and riparian 

measurements. For each reach, the water surface 

gradient (dimensionless) was determined by levels, the 

length of each of the geomorphic channel units (pool, 

riffle, run) was measured using a 100-m fiberglass 

tape, and a pebble count (Wolman, 1954) at approximately 100 points was done to characterize stream 

substrate. Wetted channel, streambed, and bankfull 

channel width were measured with a fiberglass tape at 

each transect. If channel bars, shelves, or islands were 

present, their widths were measured in meters. Canopy 

angles, in degrees, were measured with a clinometer. 

Total sun angle was calculated by subtracting the sum 

of the canopy angles from 180 degrees. Bank angles, 

in degrees, were measured with a clinometer. Bank 

substrate was categorized, and erosion was noted 

as present or absent. Qualitative assessment of the 

amount and type of bank and riparian vegetation 

was recorded. Instream habitat features (woody 

debris, overhanging vegetation, undercut bank, 

boulder, aquatic macrophytes, manmade structure, 

or none) were recorded for the left edge of water, 

two stream points (point 1, point 2), thalweg of the 
METHODS OF INVESTIGATION 7

channel, and right edge of water. Stream point 

measurements of depth, in feet, velocity, in feet per 

second, dominant bed substrate, embeddedness, and 

presence or absence of silt were taken in the thalweg 

and two other stream points (point 1, point 2) that 

best described the flow regime in the transect. Froude 

numbers were calculated with depth and velocity point 

measurements. Froude numbers are dimensionless and 

generally increase as velocity increases and depth 

decreases. Froude number (Fr) is calculated by 

equation 1:

(1)

where

Fr is the Froude number, dimensionless;

V is the velocity of water, in feet per second;

g is the gravitational acceleration, in feet per 

second squared; and 

D is the depth of water, in feet.

A Froude number greater than 1 indicates supercritical 

flow.

Statistical Analysis of Habitat Data

All habitat variables were tested for normality 

and homogeneity of variance. Transforming these data 

using logarithm base 10, logarithm base e, arc sin, 

square root, or ranks did not improve normality or 

variance, so raw data were analyzed. Parametric twoway and three-way analysis of variance (ANOVA) 

were used to determine significant differences among 

site, season, and year using the site vegetation data 

collected by USGS personnel. Parametric two-way 

ANOVA was used to determine significant differences 

among sites and spring sample dates for most 

NAWQA variable means and to evaluate seasonal 

differences in the RBP data collected by CSU 

personnel at sites 3700, 3970, 5500, and 5800. Due 

to inconsistencies in the data collection between the 

USGS and CSU, sites 3700, 3970, 5500, and 5800 

were excluded from the two-way ANOVA for the 

percent grass and percent forbs variables. A one-way 

ANOVA was used to determine significant differences 

among all sites for mean streambed width, bank angle, 

percent riparian vegetation, and RBP score. A p < 0.05 

was used to reject the null hypothesis for all tests. The 

Student-Newman-Keuls (SNK) multiple range test 

was used to identify homogeneous group(s) among all 

sites. The SNK multiple range test produced a relative 

ranking of the sites. The site with the greatest mean 

was listed on the left, and the site with the smallest 

mean was listed on the right. Basic statistics, ANOVA, 

and SNK were computed using the computer package 

SigmaStat 2.0 (SPSS, Inc., 1997).

Macroinvertebrates

Benthic invertebrate samples were collected 

from 10 sites in the spring and fall from 1998 to 

2001 that coincided with the stream reaches used 

for the habitat characterization (fig. 1, table 1). USGS 

personnel collected benthic invertebrate samples at 

sites 3977a, 3977b, 3980, 3985, 3990, and 4050. CSU 

personnel collected benthic invertebrate samples at 

sites 3700, 3970, 5500, and 5800. Sites were sampled 

at approximately the same time each year to allow 

seasonal, yearly, and multiple-year comparisons 

within and among sites. Sampling protocols remained 

constant throughout the study to allow for within-site, 

among-site, among-year, and seasonal comparisons.

Sample Collection

Benthic invertebrate samples were collected 

from the richest targeted habitat (RTH) (Cuffney 

and others, 1993) within a transect that was randomly 

chosen along the stream reach. The RTH is habitat 

that is most likely to have the greatest variety and 

density of organisms, usually riffles, where water 

flows over completely or partially submerged coarse 

substrate that produces disruption of the water surface. 

Transects were often located in runs that have less 

RTH than riffles and relatively smooth water surfaces. 

Sample locations within riffles generally had gravel 

or cobble substrate, the highest velocities, sufficient 

depth to remain submerged during periods of low 

flow, and an open canopy. Habitat features associated 

with samples collected from runs usually had sand or 

bedrock substrate, manmade debris, and open canopy. 

No samples were collected from pools.

Abundance data (number of organisms per 

square meter) were collected using Hess streambottom samplers with an area of 0.086 m2

 and 500-m 

mesh. One sample at each site consisted of either a 

composite of three Hess samples or the mean of three 

Fr

V

  gD = ------------
8 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

Hess replicate samples. At selected sites, composite 

and replicate samples were collected to evaluate the 

appropriateness of composite samples. Whenever 

possible, composite samples were used in this report 

for comparisons within and among sites. The Hess 

sampler was placed over the RTH and pushed firmly 

into the substrate to reduce the likelihood of escape 

by benthic invertebrates. Large cobbles were removed 

from the sampler and inspected for invertebrates. 

The streambed area within the Hess sampler was then 

stirred by hand or with rebar to a depth of approximately 10 cm to dislodge invertebrates from smaller 

substrate and those living in the hyporheic zone. 

Forceps and water were used to remove organisms 

clinging to or entwined in the mesh. The Hess sampler 

was rinsed between sample collections to minimize 

cross-contamination between samples and sites. 

Samples were collected from downstream to upstream 

to prevent habitat disruption. Distances between 

transects were rarely greater than 5 m.

A D-frame dip net with a 13-inch by 20-inch

frame and 500-m mesh was used to collect kicksweep samples. These samples provided only qualitative invertebrate data. Kicksweep samples were 

collected downstream or after the Hess samples had 

been collected. Organisms in these samples were 

collected from all available habitat types in the 

stream reach. Infrequently, the entire stream reach 

was sampled, but most often all available habitats 

were found within approximately one-half the reach 

distance. Kicksweep samples contained a considerable 

amount of debris that was reduced in the field by 

elutriating and sieving (500-m mesh metal sieve) 

until sample volumes were less than approximately 

500 mL. All samples were put in separately labeled 

500-mL or 1-L wide-mouth plastic bottles, preserved 

in the field in 95 percent ethanol, and stored until 

they could be delivered to the contract laboratory for 

analysis. Field methods described here were used by 

USGS personnel and field methods similar or compatible were used by CSU personnel.

Sample Processing and Quality Assurance

Benthic invertebrate processing, taxonomic 

identification to the lowest possible level, and enumeration were done by Chadwick and Associates in 

Littleton, Colo., using methods described by Klemm 

and others (1990) and Britton and Greeson (1987). 

Chadwick and Associates also prepared a voucher 

collection of benthic invertebrates collected by the 

USGS. The USGS National Water Quality Laboratory 

(NWQL) in Lakewood, Colo., provided independent 

verification of the voucher collection by a qualified 

taxonomist as a quality-control check. The NWQL 

reported 16 misidentifications from the 127 total taxa. 

Based on these results, the overall taxonomic accuracy 

of Chadwick and Associates was considered adequate 

for this study. Seventeen higher taxonomic groups 

(Order or higher) were identified from the benthic 

macroinvertebrates collected by the USGS. The 

Diptera taxa accounted for the largest percentage of 

the total taxa (37 percent) among the 17 higher taxa. 

The NWQL listed three misidentifications from the 

Diptera. Twelve of the 16 misidentifications were from 

the Order Coleoptera. Chadwick and Associates identified 29 Coleoptera taxa compared to 22 taxa identified by the NWQL. Based on NWQL results, the 

coleopterans were revised. Twenty-three Coleoptera 

taxa were used for sites 3977a, 3977b, 3980, 3985, 

3990, and 4050. One misidentification was identified 

by the NWQL from the Heteroptera. In several 

instances, NWQL revisions involved raising the 

taxonomic level of immature or nonideal specimens. 

The revisions made by the NWQL were based on 

the amount of material available in the voucher collection. If a larger series of specimens had been available 

for examination, the original identifications might 

have been endorsed by the NWQL (Brady Richards, 

National Water-Quality Assessment Program, written 

commun., 2001).

Benthic macroinvertebrates collected at 

sites 3700, 3970, 5500, and 5800 by CSU personnel 

are used in this report and were processed by Chadwick 

and Associates. The voucher collection prepared from 

these samples was verified by the laboratory of Boris 

Kondratieff at Colorado State University in Fort 

Collins. Additional taxonomic verifications of selected 

taxa were performed by Len Ferrington, University of 

Kansas in Lawrence. Internal logic checks of the CSU 

database were completed by USGS personnel.

Description of Metrics

Metrics are commonly used to characterize 

the community structure or biological condition of 

a stream (Plafkin and others, 1989). Two types of 

data were used to describe the benthic invertebrate 

communities and measures of these communities. 
METHODS OF INVESTIGATION 9

Qualitative kicksweep samples (collected with a 

D-frame dip net) were intended to provide a list of 

taxa present in the stream reach. Semiquantitative 

samples (collected with a Hess stream-bottom 

sampler) were intended to provide a measure of relative abundance of each taxon present. These samples, 

along with corresponding chemical, physical, and 

land-use data, were used to characterize the aquatic 

community within the sampling reach, compare 

reaches among environmental settings, compare 

changes over time, and couple environmental characteristics with biological characteristics (Cuffney and 

others, 1993). Overall, metrics related to taxonomic 

composition and abundance can be indicative of the 

general condition of the invertebrate community. The 

nine metrics selected to represent biological attributes 

of the aquatic communities sampled and their significance are described in the following section. These 

metrics were not calibrated to a reference or control 

site, but relative comparisons among the sites were 

made to provide assessment of the macroinvertebrate 

communities and measures of these communities. 

Taxa richness was reported from kicksweep data. All 

other metrics were computed with semiquantitative 

data from Hess samples.

Taxa Richness and Total Abundance

 Taxa richness (total taxa) can be useful 

in describing the biological condition of a stream 

(Resh and Grodhaus, 1983). Taxa richness generally 

increases with improving water quality and(or) habitat 

diversity. Total abundance of macroinvertebrates 

across taxa has been used to assess stream quality 

(Plafkin and others, 1989). Generally, invertebrate 

abundances decrease when communities are exposed 

to stresses such as degraded water quality or habitat 

alteration that can result from natural or manmade 

influences or storm runoff.

EPT and Chironomidae

Taxa in the Chironomidae family are generally more tolerant to degraded water quality than 

Ephemeroptera, Plecoptera, and Trichoptera (EPT) 

taxa (Rosenberg and Resh, 1993). Also, the ratio 

of EPT abundance to Chironomidae abundance 

has been used as a stream-water-quality indicator 

(Resh and Grodhaus, 1983). Thus, higher values 

for Total Chironomidae and Percent Chironomidae 

(Chironomidae %) may indicate a degraded 

stream condition, while higher values for Total 

EPT, Percent EPT (EPT %), and EPT:Chironomidae 

Ratio (EPT:C Ratio) may indicate better stream 

condition. EPT:C Ratio was calculated as E/C = 

EPT/(EPT + Chironomidae).

Dominant Taxa 

Total Dominant taxa and Percent Dominant 

taxa (% Dominant taxa) are measures of redundancy 

which assume that a highly redundant community 

reflects an impaired community (Rosenberg and Resh, 

1993). These metrics were calculated as the total or 

percentage of total abundance represented by the two 

most abundant taxa. Invertebrate communities under 

stress are frequently composed of fewer taxa and tend 

to be dominated by a few tolerant species.

Similarity Measures, Cluster Analysis, and 

Multivariate Analysis

Similarity measures and dendrograms were 

used to evaluate the variability between replicate 

and composite Hess samples and the variation of 

community structure between seasons and among 

sites. Sorenson’s Coefficient of Community (Beals, 

1984) used abundance data to determine the variability 

between paired replicate and composite Hess samples 

and to summarize the similarity of invertebrate 

community structure among sites. The Sorenson 

value (C) is calculated by equation 2:

(2)

where

C is the Sorenson’s Coefficient of Community;

w is the sum of the smaller value between 

site A and site B;

a is the sum of the frequencies of each taxon 

at site A; and

b is the sum of the frequencies of each taxon at 

site B.

The Sorenson value (C) can range from 0 (complete 

dissimilarity) to 100 (complete similarity). Values 

above 60 generally indicate agreement between the 

community structure provided by the two sampling 

methods.

C   2w

  a b +  100 = --------------------------------
10 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

The Jaccard Coefficient of Community (Jaccard, 

1912) provided numerical expression of taxonomic 

similarity based on presence or absence of taxa. The 

Jaccard value (J) is calculated by equation 3:

(3)

where

J is the Jaccard’s Coefficient of Community;

a is the number of taxa in common to both 

samples;

b is the number of taxa present in sample B 

but not in sample A; and

c is the number of taxa present in sample A 

but not in sample B.

The Jaccard value can range from 0 (no taxa in 

common) to 1.0 (all taxa shared) and directly 

expresses the percentage of taxa shared between 

two collections. This index is sensitive to variation 

in taxa occurrence, generally unbiased at small 

sample sizes, and interpreted unambiguously 

(Ludwig and Reynolds, 1988).

An unweighted pair group method average 

(UPGMA) cluster analysis was used to determine

 if there was seasonal and(or) spatial separation of 

the samples into discrete groups based on Sorenson 

and Jaccard values. UPGMA clustering refers to 

the measurement of the distance between two clusters 

as measured by the average of all sampling units 

within each group (Pielou, 1984). This agglomerative 

process produced a dendrogram by starting with all 

the samples to be clustered separate, then successively 

combining the most similar samples and(or) clusters 

until all were in a single, hierarchical group.

Principal component analysis (PCA) was used 

to identify environmental variables that accounted for 

the greatest amount of variance in the data and eliminate redundant variables from subsequent analyses. 

The multivariate ordination technique Canonical 

Correspondence Analysis (CCA) was used to identify 

environmental gradients associated with the distribution of aquatic arthropods (ter Braak, 1986; Palmer, 

1993). These analyses were done with the computer 

program MultiVariate Statistical Package 3.1, MVSP 

(Kovach, 1999).

Statistical Analysis of Macroinvertebrate Data

The Wilcoxon signed-rank test, paired t-test, 

and ANOVA were used to examine differences in 

metrics temporally or spatially. A p < 0.05 was used to 

identify significant differences, and because large variance was often associated with benthic invertebrate 

data, a p < 0.10 was used to identify marginally significant differences. The SNK multiple range test was used 

to identify homogeneous group(s) among all sites. Data 

were checked for normality using the KolmogorovSmirnov test and in some cases were log or rank transformed before analysis. These tests were from the 

computer package SigmaStat 2.0 (SPSS, Inc., 1997).

TEMPORAL AND SPATIAL VARIATION IN 

HABITAT AND MACROINVERTEBRATES

Assessments of temporal and spatial variability 

in habitat and macroinvertebrate community structure 

were made for selected sites in the Fountain Creek 

Basin. The limited amount of data collected from 

1998–2001 are not adequate to assess effects from 

stormwater runoff, but results were used to evaluate 

the relative differences among selected sites in the 

basin and provide a baseline characterization of habitat 

and macroinvertebrates for comparison in future 

studies. Natural differences in macroinvertebrate 

communities associated with cold-water (site 3700) 

and warm-water (sites 3970, 3977a, 3977b, 3980, 

3985, 3990, 4050, 5500, 5800) streams complicated 

the analysis of differences among sites. Instream and 

riparian habitat variables are presented in Appendix A 

on CD-ROM in pocket (tables 12–16). Macroinvertebrate data are presented in Appendix B on CD-ROM 

in pocket (tables 17 and 18).

Habitat

Several generalizations concerning the habitat 

data are presented. A direct relation was evident 

between stream discharge (Crowfoot and others, 1999, 

2000, 2001) and 3 of the 10 instream habitat variables 

(wetted channel width, streambed width, and bankfull 

width). These three variables usually had higher values 

for sites with the greatest discharge. The exception is 

site 3990, which ranked higher for these three variables than site 3700. Stream discharge had a less 

J a

abc + + = ---------------------
TEMPORAL AND SPATIAL VARIATION IN HABITAT AND MACROINVERTEBRATES 11

consistent direct relation with bank height and 

stream velocity. Froude number best demonstrated 

an indirect relation between stream discharge and 

any of the instream habitat variables. Froude numbers 

generally increase as velocity increases and depth 

decreases. Relatively large Froude numbers indicated 

that sites 3977a, 3977b, 3980, and 3990 were dominated by fast and shallow flows. The correlation 

between macroinvertebrate community structure and 

Froude number is likely important but is complicated 

by substrate characteristics, water quality, habitat 

quality, and thresholds of these variables that apply 

to distinct segments of an insect’s life cycle (Resh 

and Rosenberg, 1984).

Generally sand and fine gravel dominated 

the streambed substrate within the study-area sites. 

Site 3990 was dominated by bedrock, site 5800 had 

moderate amounts of bedrock, and site 3700 had the 

highest frequencies of substrate with larger particles 

and lower embeddedness (fig. 2, table 14, Appendix A 

on CD-ROM in pocket). Also, personal field observations outside the timeframe of this report identified 

site 3970 as a streambed less dominated by sand 

and gravel with large abundance of cobbles.

Geomorphic channel units were not balanced 

well among riffle, run, and pool. Run dominated every 

site. Pools were identified at sites 3700, 3970, 3977a, 

3977b, 3985, 4050, 3990, and 5500, and only site 3985 

had pools for each year of the study (1998–2001) 

(table 16, Appendix A on CD-ROM in pocket).

The most evident impacts to the instream and 

riparian habitat observed during site visits included 

downcutting (3970, 3977b, 3980, 3985, 3990, 5800); 

deposition (3977a, 3985, 4050); urban development 

(3700, 3970, 3980, 3985); right-of-way construction 

(3980, 3985, 5500); beaver (Castor canadensis) 

activity (3970, 5500); and manmade litter at all sites.

Prior to rigorous statistical analyses, all habitat 

variables were tested for normality and homogeneity 

of variance. All percentage vegetation variables failed 

the normality test, and all passed the equal variance 

test except percentage of forbs on bank. Canopy cover 

and all instream habitat variables failed the tests of 

normality and equal variance except bank angle, which 

passed the test of equal variance. Transforming these 

data using logarithm base 10, logarithm base e, arc sin, 

square root, or ranks did not improve normality or 

variance, so raw data were analyzed. The RBP scores 

were found to be normally distributed with homogeneous variance. Numerous statistical analyses 

presented in the following sections were performed to 

evaluate temporal and spatial effects on instream and 

riparian habitat parameters.

Seasonal Variations

A three-way ANOVA was used to analyze 

10 vegetation variables for seasonal differences. 

These results were from data measured using NAWQA 

protocols from the six tributary sites (3777a, 3977b, 

3980, 3985, 3990, and 4050). Site, season, and year 

were the factors used in the ANOVA. Nine of the 

10 variables had significant differences (p < 0.05) 

among sites. Season and year were significantly 

different for six and seven of the variables, respectively. Due to the high percentage of significant differences among sites and nondefinitive percentage of 

significant differences attributed to season, an unambiguous statement regarding seasonal effects on vegetation based on the three-way ANOVA could not be 

made.

Table 3 summarizes the results, from the 

preceding data set, of 60 two-way ANOVA’s that 

analyzed the factors season and year for each site 

separately. Only 16 of the possible 60 ANOVA’s 

(27 percent) indicated a significant seasonal difference, and three of these tests had p-values of 0.049. 

Differences were proportioned almost equally between 

bank (7/16) and riparian variables (9/16). Nineteen 

significant differences for the variables based on factor 

year were identified. All sites accounted for at least 

one significant difference due to season or year. Also, 

results from a two-way ANOVA on RBP scores for 

main-stem sites on Fountain and Monument Creek 

(from 1998 to 2001) indicated no significant differences due to season with a p-value of 0.301. The 

measured habitat variables among the sites in this 

study did not vary seasonally to the extent that requires 

measurements each season.

Spatial and Annual Variations

Two-way ANOVA results indicated that all 

instream and vegetative habitat variables, except 

bank forbs, had significant differences among sites 

(table 4). The habitat variables streambed width, 

bank angle, riparian vegetation, and RBP score were 

analyzed with a one-way ANOVA because not all 

these data were collected at each site or were not 

collected consistently with other sites. Percent grass 

and percent forbs results were not available for
12 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

0

102030405060708090

100

PERCENT OCCURRENCE

3700

SPRING 1998

3970

SPRING 1998

5500

SPRING 1998

5800

SPRING 1998

3700

SPRING 1999

3970

SPRING 1999

5500

SPRING 1999

5800

SPRING 1999

3700

SPRING 2000

3970

SPRING 2000

5500

SPRING 2000

5800

SPRING 2000

3700

FALL 2001

3970

FALL 2001

3977a

FALL 2001

3977b

FALL 2001

3980

FALL 2001

3985

FALL 2001

3990

FALL 2001

4050

FALL 2001

5500

FALL 2001

SITE NUMBER AND DATE

Figure 2. Pebble-count data for selected sites in the Fountain Creek Basin, 1998–2001.
TEMPORAL AND SPATIAL VARIATION IN HABITAT AND MACROINVERTEBRATES 13 Table 3. Summary of two-way analysis of variance (ANOVA) evaluating seasonal and annual differences of 10 habitat variables measur

Fountain Creek Basin, 1998–2001 (site locations shown in fig. 1) [S, seasonal p-value; Y, annual p-value; --, not significantly different (p > 0.05)] Site number (fig. 1) Bank vegetation Bank grass Bank forbs Bank shrubs Bank trees Riparian vegetation Riparian grass Riparian forbs 

S Y S Y S Y S Y S Y S Y S Y S Y 

3977a -- 0.016 0.009 -- <0.001 -- -- -- -- -- -- -- 0.033 -- -- -- 3977b -- -- -- 0.042 -- -- 0.005 -- -- -- 0.027 -- -- -- -- <0.001 3980 -- -- -- -- -- -- 0.026 -- -- -- -- -- 0.049 -- 0.009 <0.001 3985 -- -- -- -- -- 0.016 -- -- -- 0.041 0.004 <0.001 <0.001 0.010 <0.001 <0.001 3990 -- -- -- -- -- <0.001 -- -- 0.049 -- -- -- -- 0.023 -- <0.001 4050 -- -- -- -- -- 0.033 0.003 0.044 0.006 0.002 -- <0.001 -- -- -- <0.001 
14 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

Table 4. Results of two-way analysis of variance (ANOVA) and Student-Newman-Keuls (SNK) multiple range tests for spatial 

variation in 21 instream and riparian habitat variables measured at selected sites in the Fountain Creek Basin, 1998–2000

[Instream habitat variables, 1–10; vegetation habitat variables, 11–20; qualitative habitat variable, 21; RBP, Rapid Bioassessment Protocol; blue shading 

indicates variable with at least one exclusive group; sites joined by a line were not significantly different; %, percent; no., number; site locations shown 

in fig. 1; <, less than]

Habitat variable Unit SNK multiple range test

ANOVA

p-value Test 

power

1 Wetted channel 

width

Meters Mean 24.4 12.6 10.0 5.8 5.7 3.7 2.5 1.8 1.4 1.1

Site no. 5800 5500 3970 3990 3700 3980 3977a 3985 3977b 4050 <0.001 1.000

2a Streambed width Meters Mean 27.5 15.3 10.8 7.9 7.6 7.0 3.0 2.9 2.6 2.1

Site no. 5800 5500 3970 3990 3700 3980 3977a 4050 3977b 3985 <0.001 1.000

3 Bankfull width Meters Mean 30.0 22.0 13.5 11.0 8.8 8.6 6.2 5.3 4.9 4.9

Site no. 5800 5500 3970 3990 3700 3980 3977a 3977b 3985 4050 <0.001 1.000

4 Bank height Meters 6.8 3.7 3.0 1.5 1.2 1.1 0.9 0.8 0.6 0.4

5800 3970 5500 3977b 3700 3990 3980 4050 3977a 3985 <0.001 1.000

5a Bank angle Degrees Mean 58 49 48 47 46 44 38 35 29 29

Site no. 3977b 5500 5800 4050 3990 3980 3970 3985 3977a 3700 <0.001 1.000

6 Total sun angle Degrees Mean 154 140 135 121 118 113 100 96 85 41

Site no. 3985 3980 3977a 5500 3990 3970 3977b 4050 5800 3700 <0.001 1.000

7 Embeddedness % Mean 97 95 88 86 86 79 78 77 72 49

Site no. 3985 3977a 3977b 5500 3970 5800 3700 4050 3980 3990 <0.001 1.000

8 Stream depth Feet Mean 1.7 1.3 1.3 1.0 0.2 0.2 0.2 0.2 0.2 0.2

Site no. 3970 5500 5800 3700 3977b 3990 3985 3980 4050 3977a <0.001 1.000

9 Stream velocity Feet per Mean 3.2 3.0 2.9 2.9 2.3 2.2 2.0 1.9 1.3 0.9

second Site no. 5500 3970 5800 3990 3700 3980 3977b 3977a 3985 4050 <0.001 1.000

10 Froude number None Mean 1.1 0.9 0.8 0.7 0.5 0.5 0.5 0.4 0.4 0.4

Site no. 3990 3980 3977a 3977b 3985 5500 5800 3970 3700 4050 <0.001 1.000

11 Bank vegetation % Mean 63 49 46 44 41 39 28 26 24 20

Site no. 3985 3970 3977a 5500 3700 4050 3980 3990 5800 3977b <0.001 1.000

12b Bank grass % Mean 88 84 60 49 28 27

Site no. 3977a 3985 4050 3980 3977b 3990 0.001 1.000

13b Bank forbs % Mean

Site no. not significant 0.052 0.432

14 Bank shrubs % Mean 14.2 14.2 13.0 8.3 5.7 4.9 4.7 4.6 2.8 0.6

Site no. 3970 5500 3700 4050 3977b 3980 5800 3990 3985 3977a <0.001 1.000

15 Bank trees % Mean 12.8 12.8 6.4 5.3 5.0 3.0 2.8 0.8 0.0 0.0

Site no. 5500 3700 4050 5800 3990 3985 3970 3977b 3977a 3980 0.011 0.683

16a Riparian 

vegetation

% Mean 75.8 75.7 71.9 67.9 66.7 65.1 59.2 58.8 56.7 27.5

Site no. 3700 3977b 3985 4050 3990 3980 5800 3977a 3970 5500 <0.001 1.000

17b Riparian grass % Mean 68 66 64 49 48 44

Site no. 3985 3977a 3980 4050 3990 3977b <0.001 1.000

18b Riparian forbs % Mean 19 16 15 12 10 8

Site no. 3985 3980 3977a 4050 3990 3977b 0.004 0.819

19 Riparian shrubs % Mean 30.4 27.5 17.1 15.6 13.9 12.4 10.3 9.7 6.7 2.8

Site no. 3977b 4050 3990 3980 3970 3985 3977a 5500 3700 5800 <0.001 1.000

20 Riparian trees % Mean 19.0 17.5 13.3 11.9 11.8 10.0 4.3 3.9 0.8 0.0

Site no. 3990 3977b 5500 5800 4050 3700 3980 3970 3985 3977a <0.001 1.000

21a RBP score None Mean 114 87 75 67 56 53 50 49 28 22

Site no. 3700 4050 3970 3985 3977b 3990 5500 3980 3977a 5800 <0.001 1.000

a

Results from one-way ANOVA.

b

Results unavailable for sites 3700, 3970, 5500, and 5800.
TEMPORAL AND SPATIAL VARIATION IN HABITAT AND MACROINVERTEBRATES 15

sites 3700, 3970, 5500, and 5800 because these data 

were collected inconsistently with the remaining sites. 

Measurements for site 3977a were omitted from the 

spring 1998 and 1999 analysis of bankfull width due 

to error in data collection or transcription.

Instream habitat variables, which tend to 

describe channel morphology features, provided 

the best means of discriminating the differences 

among sites. The SNK multiple range tests identified 

11 habitat variables with at least one exclusive group 

(table 4, highlighted in blue). An exclusive group 

includes only one site. Nine of these 11 habitat variables (82 percent) were instream habitat variables. 

Sites 3970, 5500, and 5800 were found in four exclusive groups, while sites 3977a and 3977b were not 

isolated into an exclusive group by the SNK method. 

The sites with the largest drainage area (table 2) had 

the greatest means for the instream habitat variables 

wetted channel width, streambed width, and bankfull 

width. Also, the embeddedness results indicated that 

site 3990 provided preferred macroinvertebrate habitat 

with the lowest embeddedness mean, but this measurement was heavily influenced by bedrock (defined as 

0 percent embeddedness), which commonly limits the 

macroinvertebrate community structure. The overall 

habitat site similarity was greater among the tributary 

sites compared to the site similarity among main-stem 

sites.

Two-way ANOVA results indicated that 8 of 

13 habitat variables had significant differences due 

to year, and that during the study, the instream and 

riparian habitat in the Fountain Creek Basin were not 

stable. The year 1998 was isolated four times, and 

1999 and 2000 were isolated from all other years twice 

and once, respectively. The year pairs 1998, 1999; 

1998, 2000; and 1999, 2000 were found in SNK 

homogeneous subsets twice, once, and five times, 

respectively. Although based on a limited amount of 

data, the observed yearly pattern in habitat variation 

might be due to the pre- and postconditions of the 

1999 flood (Crowfoot and others, 2000). Stogner 

(2000) found that the high daily-mean streamflows 

that occurred during the spring 1999 flood remained 

elevated for a longer period of time than any other 

event for the entire period of record for six sites in the 

Fountain Creek Basin. Continued collection of habitat 

data would be necessary to identify trends in habitat 

stability at the study sites. This information could 

be used to evaluate the relation between streamflow 

patterns and rates of change in measured habitat 

variables.

Overall, sand and fine gravel dominated the 

substrate, and runs were the prevalent geomorphic 

channel unit found at the selected sites in the study 

area. The range in streamflow among the sites varied 

greatly, but the prevalent velocity-depth regime was 

fast and shallow flows. Readily observed deleterious 

effects to the instream and riparian habitats at the 

study sites included downcutting, deposition, scour, 

urban development, right-of-way construction, beaver 

activity, and manmade litter. Seasonal variation of the 

habitat variables measured was generally not found. 

Statistically significant spatial variation in habitat 

variables was found among the sites, and channel 

morphology features provided the best discrimination. 

Habitat similarity was greater within the tributary sites 

compared to the main-stem sites. Effects on the habitat 

from the spring 1999 flood might be demonstrated by 

the yearly grouping pattern that appears to separate 

pre- and postflood conditions.

Macroinvertebrates

Benthic macroinvertebrate taxa collected 

from selected sites in the Fountain Creek Basin are 

listed in Appendix B on CD-ROM in pocket. Hess 

and kicksweep sampling yielded the identification of 

218 macroinvertebrate taxa that were represented by 

9 insect orders and 35 noninsect taxa. As a percentage 

of taxa richness, dipterans accounted for 40 percent 

of the taxa. Comparisons among the sites showed that 

site 3700 had consistent and relatively high taxa diversity (number of macroinvertebrate taxa, fig. 3). Taxa 

diversity was inconsistent at sites 3970, 4050, and 

5500. The remaining sites were depauperate in terms 

of taxa diversity. Total abundance (number per square 

meter) followed a pattern similar to taxa diversity, 

with site 3700 having the highest yearly values and 

sites 3970, 4050, and 5500 having at least 1 year with 

high total abundance (fig. 4). Only site 3700 had a 

consistently low relative abundance of chironomids 

compared to the other sites (fig. 5). Site 3700 was the 

only site that had the EPT:C ratio skewed in favor of 

the EPT taxa for each year of the study (fig. 6).

In general, site 3700 was the relatively least 

impaired stream segment as indicated by the structure 

of its benthic macroinvertebrate community, described 

previously by the four metrics, with sites 3970, 4050, 

and 5500 forming a relatively moderately impaired 

group. At site 4050, lack of an adequate amount of 

streamflow could be an important limiting factor of
16 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

0

5

10

15

20

25

30

35

FALL

3700

FALL

3970

FALL

3977a

FALL

3977b

FALL

3980

FALL

3985

FALL

3990

FALL

4050

FALL

5500

FALL

5800

1998 1999 2000

NUMBER OF MACROINVERTEBRATE TAXA

0

1,000

2,000

3,000

4,000

5,000

6,000

FALL

3700

FALL

3970

FALL

3977a

FALL

3977b

FALL

3980

FALL

3985

FALL

3990

FALL

4050

FALL

5500

FALL

5800

1998 1999 2000

NUMBER OF ORGANISMS PER SQUARE METER

Figure 4. Total abundance of macroinvertebrates collected during the fall at selected sites in the Fountain Creek Basin, 

1998–2000.

Figure 3. Number of macroinvertebrate taxa collected during the fall at selected sites in the Fountain Creek Basin, 

1998–2000.
TEMPORAL AND SPATIAL VARIATION IN HABITAT AND MACROINVERTEBRATES 17

0

10

20

30

40

50

60

70

80

90

100

FALL

3700

FALL

3970

FALL

3977a

FALL

3977b

FALL

3980

FALL

3985

FALL

3990

FALL

4050

FALL

5500

FALL

5800

1998 1999 2000

PERCENT CHIRONOMIDS

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

1.0

FALL

3700

FALL

3970

FALL

3977a

FALL

3977b

FALL

3980

FALL

3985

FALL

3990

FALL

4050

FALL

5500

FALL

5800

1998 1999 2000

RATIO EPT:CHIRONOMIDAE

Figure 5. Relative abundance of chironomids collected during the fall at selected sites in the Fountain Creek Basin, 

1998–2000.

Figure 6. Ratio of EPT abundance to chironomid abundance from fall samples collected at selected sites in the 

Fountain Creek Basin, 1998–2000.
18 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

macroinvertebrate populations. The remaining sites 

might be termed relatively severely impaired, with 

inadequate streamflow likely the primary limiting 

factor of macroinvertebrate populations at sites 3977a 

and 3977b. Identifying the hydrological, environmental, biological, water-quality, or climatic factors 

that influenced the macroinvertebrate communities 

and measures of these communities was not straightforward due to the lack of reference or control sites 

and qualitative and quantitative differences between 

and among sites. For example, the difference in baseflow stream discharge (sustained ground-water inflow 

rather than surface runoff) was up to four orders of 

magnitude (sites 3977a, 3977b, 4050 < 0.1 ft3

/s; 

site 5800 > 100 ft3

/s) (Crowfoot and others, 1999, 

2000, 2001; field observations, 1998, 1999, 2000), 

size of drainage basin ranged from 1.8 mi2 (site 4050) 

to 495 mi2 (site 5800) (table 2), land-use types and 

proportions differed greatly among the sites (table 2), 

natural differences between cold-water and warmwater stream-dwelling biota, and field observations 

of the effects of point and nonpoint sources of waterquality degradation were not evenly distributed among 

the sites. Also, biological data that would best determine possible effects from wastewater-treatment 

facilities and assess the further relative degradation 

or recovery of macroinvertebrate communities that 

inhabit Fountain Creek to its confluence with the 

Arkansas River were not available.

Comparison of Replicate and Composite Samples

Results from Sorenson’s Coefficient of 

Community (C) analysis and from the Wilcoxon signedrank test are given in table 5. Six of 11 of the signedrank tests indicated no significant differences (p > 0.05) 

between the replicate and composite samples. Further, 

6 of the 11 C values from log-transformed data were 

greater than 60, indicating relatively high similarity 

between the samples, and only two of these C values 

(site 3970 in fall 1998, and site 3985 in spring 2000) 

indicated relatively low similarity between the sample 

types. Differences in results between the tributary and 

main-stem sites was not apparent, as the nonsignificant 

signed-rank test-result frequencies are almost equal 

between stream types, with nonsignificant differences 

occurring in 50 percent of the tributary sites and 

57 percent of the main-stem sites. Table 6 summarizes 

the Sorenson’s Coefficient of Community (C) analysis. 

All mean and median similarity values from logtransformed data were higher than those from raw 

data, possibly due to the dampening of differences 

between rare and dominant taxa. Mean similarity 

values based on log-transformed data were equal 

between stream types, with the median value for the 

main-stem sites slightly higher. These results were not 

definitive but indicated that composite sampling at these 

sites is an appropriate alternative to replicate sampling 

and that evaluations among and within sites can be 

made with data from either sampling method.

Table 5. Results of Wilcoxon signed-rank test and Sorenson's Coefficient of Community1 for paired replicate and composite 

invertebrate samples collected from selected sites in the Fountain Creek Basin, 1998–2001

[x, integer]

Site number

(fig. 1) Date Signed-ranktest p-value

Sorenson’s

Coefficient of

Community

Sorenson’s

Coefficient of

Community

on log(x+1)

3700 Spring 1999 0.456 72.2 65.9

3700 Spring 2000 0.003 48.0 62.2

3970 Spring 1998 0.001 42.7 62.0

3970 Fall 1998 <0.001 12.7 25.7

3970 Spring 2000 0.216 53.2 62.7

3977b Fall 1999 0.241 56.4 72.0

3980 Fall 2000 <0.001 15.0 54.0

3985 Spring 2000 <0.001 4.1 18.2

4050 Spring 2001 0.561 63.2 63.2

5500 Fall 1999 0.194 45.0 54.9

5800 Fall 1999 1.000 40.0 55.6

1Values above 60 generally indicate agreement between the community structure provided by the two sampling methods.
TEMPORAL AND SPATIAL VARIATION IN HABITAT AND MACROINVERTEBRATES 19

Seasonal Variations

The number of invertebrate species inhabiting 

the sites ranged from very few to possibly 50 or more 

from a wide variety of taxonomic orders. Each of these 

species has evolved unique life history strategies or 

adaptive traits to minimize competition and predation, 

exploit seasonally available food sources, avoid 

unfavorable environmental conditions, and synchronize reproduction with favorable environmental conditions. The sum of all species’ life cycles at a given 

location can potentially cause the community structure 

to be a continually changing aggregate of populations. 

Therefore, metrics or indices computed from species 

assemblages collected in different seasons can reflect 

natural variation and make comparisons among and 

within sites more difficult and complicate the assessment of biological condition.

Table 7 lists the seasonal mean for each 

metric by site and can be used to determine if 

seasonal differences in metrics occurred at any particular site and compare the behavior of a metric between 

sites. For example, percent EPT (EPT %) indicated 

seasonal variation at site 3700 (fall 72 percent, spring 

55 percent) but was essentially unchanged at site 3970 

(fall 26 percent, spring 25 percent). Preliminary analysis of these data indicated that among all sites, eight 

out of nine metric values calculated from fall samples 

were greater than those from spring samples (table 8). 

These observations coupled with field experience were 

indications that seasonal differences in community 

structure existed for at least some of the study sites.

A between-season comparison was performed using a one-way ANOVA on metrics listed 

in table 7 and, when appropriate, ANOVA on ranks, 

by combining metric values from like stream types 

(that is, tests were made on samples collected at tributary sites, n = 18; and main-stem sites, n = 12). Significant seasonal variation (p < 0.05) was identified for 

all non-percentage metrics from the tributary sites, 

with the fall values indicating a more robust biological 

condition. However, results for the main-stem sites did 

not show significant seasonal differences for any individual metric. The results from the ANOVA test for 

the main-stem sites might have been complicated by 

analyzing metric values from a cold-water stream 

(site 3700) with metrics associated with warm-water 

streams (sites 3970, 5500, 5800), and further analysis 

would be necessary to determine if seasonal variation 

affected the community structure of the main-stem 

sites. Additional tests also would be necessary to evaluate the components of seasonal and yearly variation 

at individual sites.

Three cluster analyses (UPGMA), based on 

Jaccard (J) values, were used to determine if seasonal 

variation was present at the sites. The first analysis 

was based on a matrix of J values computed from all 

kicksweep samples collected from sites 3700, 3970, 

5500, and 5800. The results from this analysis demonstrated that site 3700 had higher within-site similarity 

by forming two-sample fall/spring clusters (nodes) 

in the dendrogram. Site 3700 was on a cold-water 

stream dominated by cobbles, whereas the remaining 

main-stem sites are designated as warm-water streams 

that were dominated by shifting sand channels. The 

clustering of site 3700 showed that site differences 

were more important than seasonal differences to the 

ordered arrangement of the cold-water site. Eleven of 

the first 13 clusters were composed of samples from 

the same season. Differences between the spring and 

fall samples existed for the warm-water sites, and 

these seasonal differences override site similarity in 

the clustering of the warm-water sites.

The second cluster analysis that excluded data 

from site 3700 was done on the remaining 18 samples. 

The results of this analysis are reported in a dendrogram (fig. 7) and in table 9. For this data set, the 

minimum and maximum number of agglomerations 

to place all samples within a first-order cluster was 9 

and 17, respectively. A first-order cluster is one that 

includes a novel sample. Twelve agglomerations were 

needed to place each sample within a group. Eleven of

Table 6. Summary of Sorenson's Coefficient of Community 

analysis of paired replicate and composite invertebrate 

samples collected from selected sites in the Fountain Creek 

Basin, 1998–2001

[x, integer; MS4, municipal stormwater permit or tributary sites are 

sites 3977a, 3977b, 3980, 3985, 3990, 4050; RW, receiving water 

or main-stem sites are sites 3700, 3970, 5500, 5800]

Sites

(fig. 1;

table 1)

Data

type

MinimumMaximum Mean Median

All MS4 raw 4.12 63.17 34.70 35.70

log(x+1) 18.18 71.96 51.80 58.60

All RW raw 12.74 72.24 44.80 45.00

log(x+1) 25.68 65.86 51.80 62.00

All sites raw 4.12 72.24 41.10 45.00

log(x+1) 18.18 71.96 54.20 62.00
20 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001 Table 7. Mean metric values for fall and spring samples collected from selected sites in the Fountain Creek Basin, 1998–2001 [F, fall mean; S, spring mean; >, greater than; EPT, Ephemeroptera, Plecoptera, Trichoptera; C, Chironomidae; %, percent] Metric Site number (see fig. 1 for locations) 3700 3970 3977a 3977b 3980 3985 3990 4050 

F S F S F S F S F S F S F S F S 

Total taxa 24 18 18 15 7 2 11 2 11 4 13 5 5 4 20 10 Total 

abundance

3,924 4,741 1,990 1,799 547 12 313 16 1,518 60 1,339 211 442 55 1,764 464 1

Total EPT 2,760 2,959 332 323 42 0 86 3 96 9 56 26 12 8 280 4 Total 

Chironomidae

510 1,257 1,367 1,249 274 7 93 9 701 38 539 34 420 39 776 373 1

EPT % 72 55 26 25 6 0 33 10 17 10 9 43 5 7 13 2 Chironomidae % 11 37 63 62 27 42 26 69 50 74 34 8 92 83 55 77 EPT:C ratio 0.88 0.6 0.3 0.28 0.3 0 0.59 0.11 0.19 0.12 0.22 0.65 0.05 0.08 0.22 0.03 Total dominant 

taxa

2,391 2,620 1,002 1,168 446 9 195 13 1,361 47 974 185 407 39 828 318 1

% Dominant 

taxa

64 59 61 56 86 83 65 87 89 84 64 90 91 83 53 65 

Number times 

F > S

4 8 8 7 8 6 7 7 

Percentage of 

times F > S

44 89 89 78 89 67 78 78 
TEMPORAL AND SPATIAL VARIATION IN HABITAT AND MACROINVERTEBRATES 21

these 12 (92 percent) first-order clusters placed 

samples into groups of like season. The 1998 spring 

sample from site 3970 was placed into an existing 

cluster of fall samples (node 11). This analysis 

strongly indicated that seasonal differences occurred 

among the warm-water main-stem sites. The third 

cluster analysis of the J values from the kicksweep 

samples collected at the tributary sites (3977a, 

3977b, 3980, 3985, 3990, 4050) also indicated 

strong seasonal differences with 23 of 25 firstorder clusters composing like seasons.

Site-specific seasonality was evaluated using 

two-way ANOVA (season and year) and the Wilcoxon 

signed-rank test. Two-way ANOVA’s were performed 

on the nine metrics for each site, which produced 

a total of 180 test results. The powers of these tests 

were extremely low and, therefore, the results were not 

interpreted. For the 90 tests based on season, none had 

a power greater than or equal to the desired value of 

0.80, and 93 percent had powers less than 0.40. Two 

tests for the season factor had powers greater than 

or equal to 0.80, but 87 percent had powers less than 

0.20.

Results of the Wilcoxon signed-rank test to 

evaluate seasonal difference between metric values for 

each site are presented in table 10. Results indicated 

that significant and marginally significant seasonal 

variation was found in 67 percent of the tributary sites 

and 25 percent of the main-stem sites, respectively. 

Paired t-test results also were not reported or interpreted in this report because the powers of these tests 

were low (below 0.80).

Natural seasonal variation was found in both 

stream types, with fall samples more robust, as indicated by the one-way ANOVA, cluster analysis, and at 

specific sites with the signed-rank test. Therefore, the 

best evaluation of temporal and spatial variation in the 

aquatic arthropod community structure within and 

among sites would be achieved by comparing fall 

samples.

Spatial Variation in Invertebrate Community 

Structure

A two-way ANOVA was initially used to evaluate annual and spatial variation within and among 

sites, respectively. In this analysis, metric data from 

main-stem and tributary sites were analyzed separately. Metric values from fall samples collected 

between 1998 and 2000 were used to determine if 

invertebrate community structure varied significantly 

among years and(or) sites. All metrics were tested 

for differences in factors, site and year, resulting in 

18 tests for each stream type (4 by 9 matrix). The 

average power of the 36 tests was 0.31, with only 

three tests having a power greater than 0.80. Test 

power ranges from 0 to 1.0, and a test power equal to 

or greater than 0.80 is desired. Results from individual 

tests that did not meet this criterion were not reported, 

but a summary of the results was included. Marginally 

significant (p < 0.10) or significant (p < 0.05) annual 

differences were found for two metrics from each of 

the stream types. The average power of these tests 

was 0.22, with a range of 0.05–0.72. For the tributary 

sites, three metrics varied significantly among sites, 

but the power of these tests was low with a range of 

0.05–0.64 and a mean of 0.27. Total EPT, percent EPT, 

and EPT:Chironomidae ratio (mean power 0.94, range 

0.93–0.95) were significantly different (p < 0.05) 

among the main-stem sites. Additionally, marginal 

or significant differences may have existed for three 

other metrics among the main-stem sites, but these 

results had a mean power of 0.45 and a range of 

0.35–0.64. Overall, these results tentatively indicated 

that invertebrate community structure did not vary 

significantly between years for either main-stem 

or tributary stream types. In addition, the benthic 

invertebrate community structure did vary significantly among sites, with the main-stem sites showing 

more variation.

Because annual variation in the invertebrate 

community structure appeared negligible, a one-way 

ANOVA or ANOVA on ranks of the nine metrics from 

fall samples was used to examine spatial variation in

Table 8. Combined results of all fall and spring metric 

mean comparisons from 10 sites in the Fountain Creek 

Basin, 1998–2001

[F, mean fall value; S, mean spring value; >, greater than; EPT, 

Ephemeroptera, Plecoptera, Trichoptera; C, Chironomidae; %, percent]

Metric

Combined sites

Percentage of times F > S

Total taxa 90

Total abundance 70

Total EPT 90

Total Chironomidae 70

EPT % 70

Chironomidae % 40

EPT:C ratio 70

Total dominant taxa 60

% Dominant taxa 60
22 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001

Table 9. Seasonal differences in macroinvertebrate samples based upon Jaccard's Coefficient of Community values from 

selected sites in the Fountain Creek Basin, 1998–2000

[F, fall; S, spring; Sample, site number and date or node; *, not a first-order cluster; site locations shown in fig. 1]

Node

(fig. 7) Sample 1 Sample 2 Samples in group Seasons in group

1 3970, F 2000 5500, F 2000 2 F-F

2 3970, F 1998 5800, F 1998 2 F-F

3 Node 1 5800, F 2000 3 F-F-F

4 3970, S 1999 5500, S 1999 2 S-S

5 3970, F 1999 5500, F 1999 2 F-F

6 3970, S 2000 5500, S 2000 2 S-S

7 Node 2 5500, F 1998 3 F-F-F

8 Node 4 5800, S 1999 3 S-S-S

9 Node 5 5800, F 1999 3 F-F-F

10 Node 7 Node 9 6 *

11 3970, S 1998 Node 10 7 S-F-F-F-F-F-F

12 Node 11 Node 8 10 *

13 Node 6 5800, S 2000 3 S-S-S

14 Node 13 Node 3 6 *

15 Node 12 Node 14 16 *

16 5500, S 1998 5800, S 1998 2 S-S

17 Node 15 Node 16 18 *

1

4

3

2

5

6

7

8

9

10

12

13

14

15

16

17

11

5800 SPRING 1998

5500 SPRING 1998

5800 FALL 2000

5500 FALL 2000

3970 FALL 2000

5800 SPRING 2000

5500 SPRING 2000

3970 SPRING 2000

5800 SPRING 1999

5500 SPRING 1999

3970 SPRING 1999

5800 FALL 1999

5500 FALL 1999

3970 FALL 1999

5500 FALL 1998

5800 FALL 1998

3970 FALL 1998

3970 SPRING 1998

0.04 0.20 0.36 0.52 0.68 0.84 1.00

JACCARD'S COEFFICIENT

SITE DATE

Figure 7. Dendrogram (unweighted pair group method average, UPGMA) based upon Jaccard’s 

Coefficient of Community values summarizing the similarity between selected invertebrate communities collected from the Fountain Creek Basin, 1998–2000. Values approaching 1 are more 

similar. Nodes on the dendrogram are indicated by highlighted numbers. Site locations shown 

in figure 1.
TEMPORAL AND SPATIAL VARIATION IN HABITAT AND MACROINVERTEBRATES 23

macroinvertebrate community structure among all 

sites. The test power is an important consideration when 

interpreting these results. Significant differences among 

sites were indicated by five metrics, and the results for 

percent Chironomidae indicated a marginally significant difference (table 11). Results of the SNK multiple 

range tests showed a fairly consistent pattern for site 

pairs 3700/5500 and 3990/5800 arranged on opposite 

ends of the continuum (table 11). Total taxa, Total EPT, 

percent EPT, and EPT:Chironomidae ratio are positive 

indicators that tend to increase under improving 

water quality and(or) habitat conditions. The remaining 

significant metrics in table 11 are usually considered 

negative indicators that tend to increase in response 

to decreasing water quality and(or) habitat conditions. 

Based on these metrics, the biological condition 

of site pair 3700/5500 was always better than site 

pair 3990/5800. The mean metric value (n = 3) was 

presented above each site number in table 11 to demonstrate that substantial differences among the sites were 

evident, even though the multiple range tests could only 

accomplish gross separation of statistically significant 

groups. For example, only two homogeneous groups 

were identified for the metric Total EPT, but the metric 

mean values suggested that four exclusive and probably 

overlapping groups could be formed with sites 3700, 

5500/3970/4050, 3980/3977b/3985/3977a, and 

3990/5800. The power of all the tests for assessing 

the variation in invertebrate community structure was 

adversely affected by sample size and high variance 

within sites. Sample variance will likely continue to be 

high, but as more data are collected, the confidence in 

the test results can improve to an acceptable level.

Table 10. Results of Wilcoxon signed-rank test for paired 

fall and spring metric values from macroinvertebrate 

samples collected from selected sites in the Fountain Creek 

Basin, 1998–2001

[*, marginally significant difference; **, significant difference]

Site number

(fig. 1) p-value

3700 0.164

3970 0.098*

3977a 0.039**

3977b 0.098*

3980 0.039**

3985 0.164

3990 0.027**

4050 0.129

5500 0.426

5800 0.426

The similarity of invertebrate community 

structure among sites also was examined by cluster 

analysis because the SNK multiple range tests 

provided little delineation among sites. This cluster 

analysis used the mean taxa abundances from fall 

samples to compute a matrix of Sorenson’s Coefficient 

of Community (C) values. Figure 8 is a dendrogram 

that summarizes and illustrates the similarity of invertebrate community structure provided by the matrix. 

The clustering of sites in figure 8 does not have statistical significance but furthers the understanding of 

which sites and clusters are more similar in terms of 

community composition. Node 1 indicated that the 

sites with the least spatial separation (sites 3977a and 

3977b) had the highest similarity. Node 3 might indicate that community structure is more similar within 

tributary than main-stem stream types. However, 

nodes 4, 5, and 6 had sites 3980, 3990, and 5800 

forming heterogeneous stream-type clusters, which 

indicated that site clustering was not completely 

dependent on stream type. Sites 3700 and 5800 

were the last sites to be included in a cluster, and 

the percent similarity of the nodes that included 

these sites was small at 23.9 percent and 22.7 percent, 

respectively. The last observation confirmed results 

in the multiple range test from the one-way ANOVA 

of spatial variation that sites 3700 and 5800 were 

not similar. Beyond node 2, similarity in invertebrate 

community structure was relatively low between 

sites.

Environmental Gradient Analysis by Canonical 

Correspondence Analysis

The statistical analysis of geographic variation 

in habitat variables and macroinvertebrate community 

structure identified several patterns and relations 

among the sites, but consistent separation of the 

sites into recognizable groups did not occur. Canonical 

Correspondence Analysis (CCA) was used to determine if distinct species assemblages were associated 

with specific site(s) or groups of sites. This technique 

performed a constrained ordination using a matrix of 

species abundances from each site and an environmental data matrix that corresponded to each site. 

Seventy-four environmental variables were initially 

used in the ordination. The type and number of environmental variables used was: stream discharge, 9; 

water quality, 16; instream habitat, 21; riparian habitat, 

6; land use, 22. Taxa that had mean abundances less
24 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–2001 Table 11. Results of one-way analysis of variance (ANOVA) and Student-Newman-Keuls (SNK) multiple range tests for spatial variat

community structure from selected sites in the Fountain Creek Basin, 1998–2000 [Sites joined by a line were not significantly different; sample size was three (n = 3) for metric means listed above the site numbers; no., number; --, not applicable; site 

less than] Metric SNK Multiple range test 

Total taxa Mean 24 22 20 18 13 11 11 7 5 4 

Site no. 3700 5500 4050 3970 3985 3980 3977b 3977a 3990 5800 Total abundance Mean not significant Site no. Total EPT Mean 2760 387 332 280 96 86 56 42 12 9 Site no. 3700 5500 3970 4050 3980 3977b 3985 3977a 3990 5800 Total Chironomidaea Mean not significant Site no. Percent EPT Mean 72 40 33 26 17 13 9 6 5 0.5 Site no. 3700 5500 3977b 3970 3980 4050 3985 3977a 3990 5800 Percent Chironomidae Mean 92 85 63 55 50 42 34 27 26 11 Site no. 3990 5800 3970 4050 3980 5500 3985 3977a 3977b 3700

EPT:Chironomidae ratio Mean 0.88 0.59 0.50 0.30 0.30 0.22 0.22 0.19 0.05 0.01 

Site no. 3700 3977b 5500 3970 3977a 4050 3985 3980 3990 5800

Total dominant taxa Mean not significant 

Site no.

Percent dominant taxa Mean 96 91 89 86 65 64 63 61 57 53 

Site no. 5800 3990 3980 3977a 3977b 3700 3985 3970 5500 4050

aResults based upon ANOVA of ranks.
TEMPORAL AND SPATIAL VARIATION IN HABITAT AND MACROINVERTEBRATES 25

 than 5 percent of the average taxa mean were removed 

from the species matrix to reduce the number of nondifferential taxa. The final ordination (fig. 9) contained 

59 taxa, 10 sites, and three environmental variables. 

To limit the noise in the ordination, mean species 

abundance values from fall samples were used 

along with means calculated for the environmental 

variables.

The CCA identified a particle-size gradient 

(abscissa, CCA 1) that produced a joint plot with 

five recognizable site clusters and their associated 

taxa (1, site 3700; 2, sites 3970, 5500; 3, site 3990; 

4, sites 3977a, 3977b, 3980, 3985, 4050; and 5, 

site 5800) (fig. 9). The direction of the solid arrows 

(vectors) in figure 9 indicates where the largest 

values for that variable are located, and the length 

of the solid arrows indicates the relative change 

in the magnitude of the variables. In figure 9, the 

vectors for very coarse gravel and sand are highly 

correlated with the first ordination axis, and their 

relative lengths indicate large changes in the values 

for these variables. Therefore, the macroinvertebrate 

community variation partitioned in figure 9 is 

strongly correlated with substrate size. The position 

of site 3990 along CCA 1 was unaffected by the 

particle-size gradient and can be explained by 

bedrock dominating the substrate at this site. The 

positions of site clusters 1 and 4 were most influenced 

by the particle-size gradient. Cluster 5 is the most 

downstream site, and its position in the ordination 

was likely influenced more by stream discharge than 

by the particle-size gradient. The position of cluster 5 

indicated a discontinuity in the geographic distribution 

of the sites in the network and that environmental 

gradients other than substrate size are likely important 

in constraining species assemblages persistent in the 

Fountain Creek Basin.

Metric values computed from the presence 

or absence of taxa associated with the clusters indicated disturbance- or pollution-intolerant taxa were 

associated with cluster 1 (fig. 9). The remaining 

site clusters were composed primarily of disturbanceor pollution-tolerant taxa and formed a less than 

strict continuum from the relatively least impaired 

to most impaired with the site cluster sequence of 

2, 4, 3, 5.

1

2

5

6

7

9

8

4

3

3700

5500

3990

3970

3980

5800

4050

3985

3977b

3977a

4 20 36 52 68 84 100

PERCENT SIMILARITY

SITE NUMBER

Figure 8. Dendrogram (unweighted pair group method average, UPGMA) based upon Sorenson’s Coefficient 

of Community values summarizing the similarity of invertebrate community structure between selected sites 

in the Fountain Creek Basin, 1998–2000. Values approaching 100 are more similar. Nodes on the dendrogram 

are indicated by highlighted numbers. Site locations shown in figure 1.
26 Characterization and Analysis of Temporal and Spatial Variations in Habitat and Macroinvertebrate Community Structure, 

Fountain Creek Basin, Colorado Springs and Vicinity, Colorado, 1998–20013700 3970

3977a

3977b

3980

3985 

3990 

4050 

5500

5800 

–0.7

–1.4

–2.1–2.7

0.71.42.12.73.4

–2.7 –2.1 –1.4 –0.7 0.7 1.4 2.1 

 

wetted_channel

Cluster 2, sites 3970, 5500Ttaxa = 14EPT % = 21EPT:C = 0.43C % = 28taxa: Ace ins, Tri min, Tri spp, Opt spp, Cri tri, Mic spp, Ort spp, Pag spp, Nem spp, Lum spp, Hom nai,  Nai bre, Nai com, Nai spp 

Cluster 1, site 3700Ttaxa = 22EPT % = 73EPT:C % = 0.89C % = 9taxa: Bae tri, Dru dod, Eph ine, Cap spp, Swe spp, Pro spp, Iso spp, Skw ame, Pte bad, Ami asp, Bra ame, Arc gra, Cer osl, Hyd spp, Rhy bru, Rhy rot, Mic pus, Bri spp, Pse spp, Ant spp, Dic spp, Pri lon 

Cluster 5, site 5800Ttaxa = 1EPT % = 0EPT:C = 0C % = 100taxa: Sae spp

Cluster 3, site 3990Ttaxa = 5EPT % = 0EPT:C = 0C % = 60taxa: Euk spp, Ort Cri, Hem spp, Tub wcc, Chi sp

Cluster 4, sites 3977a, 3977b, 3980, 3985, 4050Ttaxa = 20EPT % = 20EPT:C = 0.40C % = 30taxa: Bae mag, Cal spp, Fal qui, Arg viv, Arg spp, Hyd spp, Aga  Cri spp, Cry spp, Het spp, Mac spp, Lim Lis, Gon spp, Tub 

VC gravel

EXPLANATION

SITESTAXA

sand

Figure 9. Canonical Correspondence Analysis (CCA) ordination of sampling sites, species, and environmental variables for selected sit

Basin, 1998–2000. Site cluster metric values were computed from presence-absence of taxa associated with the clusters. Taxa are indicof their genus and species name (tables 17 and 18), and environmental variables are represented by solid arrows. Site locations shown coarse gravel was abbreviated VC gravel, and spp represents species.
SUMMARY 27

SUMMARY

Little information concerning the streamdwelling macroinvertebrates or habitat condition 

of the Fountain Creek Basin has been compiled, 

analyzed, or published. In the spring of 1998, the 

USGS began a noncomprehensive study to characterize and analyze temporal and spatial variations in 

habitat and macroinvertebrate community structure 

by using biological, hydrological, water-quality, and 

land-use data collected at 10 sites in the Fountain 

Creek Basin. The results from this study can be used 

as baseline information to augment long-term monitoring of water quality, instream and riparian habitat 

quality and diversity, and biological condition.

Generally, sand and fine gravel dominated the 

substrate, and runs were the prevalent geomorphic 

channel unit found at the selected sites in the study 

area. The range in discharge among the sites varied 

greatly, but the prevalent velocity-depth regime was 

fast and shallow flows. Site 3700 was a cold-water 

stream, whereas the remaining sites were designated 

as warm-water streams. Readily observed deleterious 

effects to the instream and riparian habitats at the 

study sites included downcutting, deposition, scour, 

urban development, right-of-way construction, beaver 

activity, and manmade litter. Seasonal variation of the 

habitat variables measured was generally not found. 

Statistically significant spatial variation in habitat 

variables was found among the sites, and channel 

morphology features provided the best discrimination. 

Habitat similarity was greater within the tributary sites 

compared to the main-stem sites.

Hess and kicksweep sampling yielded the identification of 218 macroinvertebrate taxa that were 

represented by 9 insect orders and 35 noninsect taxa. 

As a percentage of taxa richness, dipterans accounted 

for 40 percent of the taxa. Site 3700 was the relatively 

least impaired site based on the metrics used in this 

study. Lack of adequate streamflow likely had the 

greatest influence on macroinvertebrate populations at 

sites 3977a, 3977b, and 4050. Statistical analyses indicated that variability in macroinvertebrate populations 

between seasons was significant and that fall samples 

generally had diversity and abundances greater than 

spring samples. Clustering methods provided strong 

evidence that seasonal differences in macroinvertebrate populations existed. ANOVA results indicated 

that annual variation of invertebrate community structure within a given site was negligible and that significant geographic variation for 5 of the 9 metrics 

existed. Mean metric values indicated substantial 

biological differences between sites or groups of 

sites that SNK multiple range tests did not delineate. 

Because the SNK multiple range tests provided little 

delineation among the sites, a dendrogram based on 

the Sorenson’s Coefficient of Community was used 

to examine geographic similarity among the sites. 

The structure of the dendrogram from this analysis 

indicated that invertebrate community structure was 

more similar among tributary sites than main-stem 

sites. CCA ordination identified a substrate particlesize gradient from site-specific species-abundance 

data and environmental correlates that reduced the 

10 sites to 5 site clusters and their associated taxa. 

A continuum of the study sites based on metrics 

computed from the presence or absence of taxa associated with the five CCA clusters from the relatively 

least impaired to most impaired was 3700, 3970/5500, 

3977a/3977b/3980/3985/4050, 3990, 5800.

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Appendix A. Habitat Data (available in Excel format)

Table 12. Detailed instream and riparian habitat variables measured at selected sites in the Fountain Creek Basin, 

1998–2001

Table 13. Detailed point instream habitat variables measured at selected sites in the Fountain Creek Basin, 

1998–2001

Table 14. Pebble-count data for selected sites in the Fountain Creek Basin, 1998–2001

Table 15. Habitat variable scores based upon rapid bioassessment protocols for selected sites in the Fountain 

Creek Basin, 1998–2001

Table 16. Geomorphic channel unit measurements and water-surface gradient for selected sites in the Fountain 

Creek Basin, 1998–2001

Tables 12 and 13

Table 14, 15, and 16

Appendix B. Macroinvertebrate Data (available in Excel format)

Table 17. Macroinvertebrate taxa collected by kicksweep method at selected sites in the Fountain Creek Basin, 

1998–2001

Table 18. Density of macroinvertebrates collected by Hess sampler at selected sites in the Fountain Creek Basin, 

1998–2001

Table 17

Table 18