U.S. Department of the Interior

U.S. Geological Survey

Scientific Investigations Report 2010–5103

Prepared in cooperation with Colorado Springs City Engineering

Temporal Change in Biological Community Structure in the 

Fountain Creek Basin, Colorado, 2001–2008
Photo on cover: Upper end of sampling reach looking upstream, Fountain Creek at 

Security, Colorado (07105800). Photo by Erin E. Evans, November 2007.
Temporal Change in Biological Community 

Structure in the Fountain Creek Basin, 

Colorado, 2001–2008

By Robert E. Zuellig, James F. Bruce, and Robert W. Stogner

Prepared in cooperation with Colorado Springs City Engineering

Scientific Investigations Report 2010–5103

U.S. Department of the Interior

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

KEN SALAZAR, Secretary

U.S. Geological Survey

Marcia K. McNutt, Director

U.S. Geological Survey, Reston, Virginia: 2010

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Suggested citation:

Zuellig, R.E., Bruce, J.F., and Stogner, R.W., 2010, Temporal change in biological community structure in the Fountain 

Creek basin, Colorado, 2001–2008: U.S. Geological Survey Scientific Investigations Report 2010–5103, 19 p.
iii

Contents

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

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

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

Study Area..............................................................................................................................................2

Methods...........................................................................................................................................................2

Data Collection......................................................................................................................................2

Data Analysis.........................................................................................................................................7

Determining Site-Specific Temporal Change in Biological Communities among Years ...........7

Relating Environmental Variables to Observed Temporal Change in Biological 

Community Structure..............................................................................................................7

Temporal Change in Biological Community Structure.............................................................................8

Macroinvertebrate Communities .......................................................................................................8

Fish Communities..................................................................................................................................8

Environmental Variables Correlated to Temporal Changes in Biological Community Structure......8

Macroinvertebrate Communities .......................................................................................................8

Fish Communities................................................................................................................................11

Implications for Future Monitoring ...........................................................................................................11

Summary and Conclusions.........................................................................................................................17

Acknowledgments.......................................................................................................................................17

References Cited..........................................................................................................................................17

Figures

1. Map showing locations of sampling sites in the Fountain Creek basin, 

Colorado, 2001–2008.....................................................................................................................3

2. Spatial distribution of sites with and without temporal changes in 

macroinvertebrate communities among years in the Fountain Creek 

basin, Colorado, 2001–2008 .......................................................................................................10

 3. Spatial distribution of sites with and without temporal changes in 

fish communities among years in the Fountain Creek basin, Colorado, 

2003–2008 .....................................................................................................................................12

4. Spatial distribution of sites with and without temporal changes in fish 

(2003–2008) and macroinvertebrate (2001–2008) communities among years 

in the Fountain Creek basin, Colorado ....................................................................................15
iv

Tables

1. Description of sites and period of record for types of data collected in the 

Fountain Creek basin, Colorado, 2001–2008 .............................................................................4

2. Habitat, streamflow, and water-chemistry variables used to relate 

site-specific temporal patterns (seriation) in biological community 

structure, Fountain Creek basin, Colorado, 2001–2008...........................................................6

3. Correlation coefficients, percent significance level, and the number of years 

associated with the strength of multivariate seriation of macroinvertebrate 

communtiy structure among years (RELATE test for seriation)............................................9

4. Correlation coefficients and percent significance level associated with the 

strength of multivariate seriation of fish community structure among 

years (RELATE test for seriation)..............................................................................................11

5. Results of the BIO-ENV procedure determining which environmental 

variables best described observed multivariate temporal change (seriation)

in macroinvertebrate community structure in the Fountain Creek basin, 2001–2008 .....13

6. Results of the BIO-ENV procedure determining which environmental variables 

best described observed multivariate temporal change (serial) in fish 

community structure in the Fountain Creek basin, 2003–2008 ............................................16
v

Conversion Factors

Inch/Pound to SI

Multiply By To obtain

Length

inch (in) 2.54 x 100 centimeter (cm)

inch (in) 2.54 x 101 millimeter (mm)

inch (in) 2.54 x 104 micrometer (mm)

foot (ft) 3.048 x 10-1 meter (m)

mile (mi) 1.609 x 100 kilometer (km)

micrometer (mm) 1.0 x 102 centimeter (cm)

Flow Rate

cubic foot per second (ft3/s) 2.832 x 10-22.590 cubic meter per second (m3/s)

Volume

milliliter (ml) 473.176 pint (pt)

Temperature in degrees Celsius (°C) may be converted to degrees Fahrenheit (°F) as follows:

°F=(1.8×°C)+32

Vertical coordinate information is referenced to the North American Vertical Datum of 1988 

(NAVD 88).

Horizontal coordinate information is referenced to the North American Datum of 1983 (NAD 83).

Altitude, as used in this report, refers to distance above the vertical datum.

Specific conductance is given in microsiemens per centimeter at 25 degrees Celsius (μS/cm at 

25 °C).

Concentrations of chemical constituents in water are given either in milligrams per liter (mg/L) 

or micrograms per liter (μg/L).

Abbreviations

BIO-ENV exploratory multivariate analysis linking biology to environmental variables

CDOW Colorado Division of Wildlife

CDPHE Colorado Department of Public Health and Environment

DC direct current

HIT Hydrologic Index Tool

MDS nonmetric multidimensional scaling

NWIS National Water Information System

NWQL National Water Quality Laboratory

PRIMER Plymouth Routines in Multivariate Ecological Research

QA/QC quality assurance/quality control

QMH qualitative multihabitat

RELATE nonparametric multivariate hypothesis test between two distance matrices

USGS U.S. Geological Survey

Abstract

In 2001, the U.S. Geological Survey, in cooperation with 

Colorado Springs City Engineering, began a study to better 

understand the relations between environmental characteristics and biological communities in the Fountain Creek basin 

in order to aide water-resource management and guide future 

monitoring activities. To accomplish this task, environmental 

(streamflow, habitat, and water chemistry) and biological (fish 

and macroinvertebrate) data were collected annually at 24 

sites over a 6- or 8-year period (fish, 2003 to 2008; macroinvertebrates, 2001 to 2008). For this report, these data were 

first analyzed to determine the presence of temporal change 

in macroinvertebrate and fish community structure among 

years using nonparametric multivariate statistics. Where 

temporal change in the biological communities was found, 

these data were further analyzed using additional nonparametric multivariate techniques to determine which subset of 

selected streamflow, habitat, or water-chemistry variables best 

described site-specific changes in community structure relative 

to a gradient of urbanization. 

This study identified significant directional patterns of 

temporal change in macroinvertebrate and fish community 

structure at 15 of 24 sites in the Fountain Creek basin. At 

four of these sites, changes in environmental variables were 

significantly correlated with the concurrent temporal change 

identified in macroinvertebrate and fish community structure 

(Monument Creek above Woodmen Road at Colorado Springs, 

Colo.; Monument Creek at Bijou Street at Colorado Springs, 

Colo.; Bear Creek near Colorado Springs, Colo.; Fountain 

Creek at Security, Colo.). Combinations of environmental 

variables describing directional temporal change in the biota 

appeared to be site specific as no single variable dominated the 

results; however, substrate composition variables (percent substrate composition composed of sand, gravel, or cobble) collectively were present in 80 percent of the environmental variable subsets that were significantly correlated with temporal 

change in the macroinvertebrate and fish community structure. 

Other important environmental variables related to temporal 

change in the biological community structure included those 

describing channel form (streambank height) and streamflow 

(normalized annual mean daily flow, high flood-pulse count). 

Site-specific results from this study were derived from a 

relatively small number of observations (6 or 8 years of data); 

therefore, additional years of data may reveal other sites with 

temporal change in biological community structure, or could 

define stronger and more consistent linkages between environmental variables and observed temporal change. Likewise 

current variable subsets could become weaker. Nonetheless, 

there were several sites where temporal change was detected 

in this study that could not be explained by the available 

environmental variables studied herein. Modification of 

current data-collection activities may be necessary to better 

understand site-specific temporal relations between biological 

communities and environmental variables. 

Introduction

Fountain and Monument Creeks (hereafter, Fountain 

Creek basin) drain approximately 926 mi2 of the eastern slope 

of the Rocky Mountains in south-central Colorado. Several 

environmental factors that influence stream communities have 

been altered throughout Fountain Creek basin (for example, 

habitat alteration and riparian disturbance), which began in 

the mid-1800s. Several studies of Fountain Creek basin have 

described patterns in surface-water hydrology (Edelmann 

and others, 2002; Stogner, 2000), water chemistry (Mau and 

others, 2007), and sediment transport (von Guerard, 1989), 

and other studies have related some of these characteristics to 

urbanization and biological communities (Zuellig and others, 2007). Continued alteration of environmental factors is 

expected in the future as human population grows, especially 

in regards to surface-water hydrology, which could further 

influence water quality, in-stream habitat, and ultimately, 

biological communities.

Local, State, and Federal agencies are interested in better 

understanding the relations between environmental characteristics and biological communities in the Fountain Creek basin 

in order to aid water-resource management and guide future 

monitoring activities. In response to these needs, the U.S. 

Geological Survey (USGS), in cooperation with Colorado 

Springs City Engineering, began an evaluation of available biological (fish and macroinvertebrates) and associated 

environmental (streamflow, habitat, and water chemistry) data 

Temporal Change in Biological Community Structure in the 

Fountain Creek Basin, Colorado, 2001–2008

By Robert E. Zuellig, James F. Bruce, and Robert W. Stogner
2 Temporal Change in Biological Community Structure in the Fountain Creek Basin, Colorado, 2001–2008

collected in the Fountain Creek basin between 2001 and 2008. 

These data were analyzed to determine the presence of multivariate serial change (hereafter, temporal change) in biological 

community structure. Where temporal change was found, the 

analysis was extended to determine which subset of available 

environmental variables best described observed temporal 

changes in biological communities. Results are intended to 

help water-resource management make informed decisions 

and guide future monitoring activities.

Purpose and Scope

This report describes temporal change in biological 

community structure in the Fountain Creek basin. A compilation of 180 macroinvertebrate and 64 fish community 

samples collected annually for 6 or 8 years from as many as 

24 sites between 2001 and 2008 in the Fountain Creek basin 

is presented. Nine environmental variables (including characteristics of streamflow, habitat, and water chemistry) concurrently collected with the biological data were also compiled. 

Using various combinations of these data, the objectives 

were to (1) determine site-specific temporal changes in fish 

and macroinvertebrate community structure among years; (2) 

where temporal change was detected, determine if changes 

in aquatic communities were related to changes in selected 

environmental variables such as stream hydrology, habitat, 

or water-chemistry; and (3) discuss results in the context of 

long-term biological monitoring of the Fountain Creek basin. 

Habitat, fish, and macroinvertebrate data evaluated in this 

report are available upon request from the USGS Colorado 

Water Science Center (James Bruce, written commun., 2010). 

Stream-discharge (streamflow) and water-chemistry data 

can be obtained online at http://waterdata.usgs.gov/co/nwis

(search by USGS station number; see table 1). 

Study Area

The Fountain Creek basin encompasses approximately 

926 mi2 in south-central Colorado draining the eastern slope 

of the Rocky Mountains (fig. 1). Altitude ranges from 4,700 ft 

at the confluence with the Arkansas River to 14,109 ft at the 

summit of Pikes Peak (source). Fountain and Monument 

Creeks are the two main drainages and are located in the 

transition of two distinctive physiographic landforms—the 

Front Range and the Colorado Piedmont (Hansen and Crosby, 

1982)—which corresponds to two Level III ecoregions: the 

Southern Rockies and the Southwestern Tablelands (Omernik, 

1987). The Front Range, which comprises the western onethird of the study area, is underlain by granite. Soils in this 

area are well drained and occur on steep slopes (Larsen, 1981; 

von Guerard, 1989). The Colorado Piedmont, which comprises the remaining eastern two-thirds of the area, abuts the 

base of the Rampart Range and is underlain by sandstone, 

shale, alluvial, and windlain deposits. Soils in this area are 

generally sandy and well drained with more gentle slopes 

(Larsen, 1981; von Guerard, 1989). The soils and geology on 

the Colorado Piedmont are readily erodible. More details of 

the soils and geology of the study area can be found in Larsen 

(1981) and von Guerard (1989). See Mau and others (2007) 

and Edelman and others (2002) for a more detailed description 

of the study area. Also, Zuellig and others (2007) provide a 

historical perspective of present-day fish and macroinvertebrate communities. 

Methods 

Environmental (streamflow, habitat, and selected water 

chemistry) and macroinvertebrate data were collected annually at 15 sites from 2001 to 2008 and at an additional 9 sites 

between 2003 and 2008 (table 1, fig. 1). Fish data were collected annually at 10 sites between 2003 and 2008 and at one 

site between 2007 and 2008. Macroinvertebrate and fish data 

were analyzed for the presence of temporal change in community structure among years using nonparametric multivariate statistics (RELATE, PRIMER ver. 6.1, Plymouth, United 

Kingdom). Where temporal change was found, the data were 

further analyzed using additional multivariate techniques 

to determine which subset of streamflow, habitat, or waterchemistry variables best described site-specific changes in 

biological community structure (BIO-ENV, PRIMER ver 

6.1, Plymouth, United Kingdom). Discussions of findings are 

focused to help direct future monitoring efforts and needed 

ecological investigations in the Fountain Creek basin. 

Data Collection

Available streamflow, habitat, and water-chemistry 

characteristics as well as macroinvertebrate and fish community structure were quantified at 24 sites (table 1). The list of 

environmental variables chosen for study was, in part, selected 

from the findings of Zuellig and others (2007) where they 

presented a list of strongly correlated variables (Spearman’s 

rho >0.60, p <0.05) with impervious surface coverage in 

the Fountain Creek basin. The variables selected for study 

included percent of the stream reach composed of riffle 

habitat (pctRIF), bank height (BANKHT), Froude number 

(FROUDE), percent sand (pctSAND), percent gravel (pctGRAVEL), percent cobble (pctCOBB), normalized maximum 

daily mean flow (MH20), high flood-pulse count (FH1), and 

dissolved nitrate plus nitrite (DNO23) (table 2). Methods used 

to collect the data addressed in this report are briefly described 

in the sections below. See Zuellig and others (2007), Mau and 

others (2007), and the citations within those two reports for 

further explanation. 

Hydrology. Instantaneous stream discharge (in ft3/s) and 

computation of mean daily discharge (in ft3/s) were measured 

and computed in accordance with standard USGS procedures described by Rantz and others (1982). Measurements 

of instantaneous stream discharge were made throughout the 
Methods 3

Pueblo

Woodland Park

Palmer Lake

Monument

Manitou Springs

Green Mountain Falls

Pikes Peak

9

8

7 6

5

4 3

2

24

23

22

21

20

19

16 17 18

14

13

12 11

10

15

1

Jimmy Camp

Creek

Cottonwood Creek

Monument Creek

Bear Creek

38°15'

38°30'

38°45'

39°

Fort Carson

Military Reservation

Las Vegas

Street Wastewater

Treatment Plant

US Air Force 

Academy

Peterson Air 

Force Base

0 21⁄2 5 MILES

0 2.5 5 KILOMETERS

Base from U.S. Geological Survey

Albers Equal-area projection

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

Central meridian 105°30' W 

105° 104°45' 104°30'

Fountain Creek

COLORADO

DENVER

Study 

Area

Ecoregion

Southern Rockies

Southwestern Tablelands

Military base or reservation

Municipal area

Sampling site and number

Front Range

Rampart Range

Colorado Piedmont

Colorado Springs

Arkansas River

Basin boundary

(see table 1)

ROCKY MOUNTAINS

7

Arkansas River

Figure 1. Locations of sampling sites in the Fountain Creek basin, Colorado, 2001–2008.
Table 1. Description of sites and period of record for types of data collected in the Fountain Creek basin, Colorado, 2001–2008. [USGS, U.S. Geological Survey; ID, identification; NAVD 88, North American Vertical Datum of 1988; ft, feet; mi2, square miles; X, collected; --, not collected] Site ID1 USGS station number Station name Elevation NAVD 88 (ft) Drainage area (mi2) Streamflow discharge Chemistry Habitat 

1 07103700 Fountain Creek near Colorado Springs, Colo. 6,110 103 X X 2001–2008 2002 07103707 Fountain Creek below 8th Street at Colorado Springs, Colo. 6,000 119 -- X 2001–2008 2003 385854104470100 Kettle Creek above Old Ranch Road near Colorado Springs, Colo. 6,670 15.46 X X 2003–2008 4 07103960 Kettle Creek above U.S. Air Force Academy, Colo. 6,620 16 X X 2003–2008 5 385750104475001 Pine Creek above Highway 83 at Colorado Springs, Colo. 6,540 3.67 X X 2003–2008 6 07103970 Monument Creek above Woodmen Road at Colorado Springs, Colo. 6,270 181 X X 2001–2008 2007 07103977 Cottonwood Creek at Cowpoke Road at Colorado Springs, Colo. 6,875 5.93 X X 2001–2008 8 07103980 Cottonwood Creek at Woodmen Road near Colorado Springs, Colo. 6,680 10.3 X X 2001–2008 9 07103985 Cottonwood Creek Tributary above Rangewood Drive at Colorado Springs, Colo. 6,630 2.81 X X 2001–2008 

10 07103990 Cottonwood Creek at mouth at Pikeview, Colo. 6,265 18.7 X X 2001–2008 

11 385501104483701 Monument Creek Tributary 1 near Pulpit Rock at Colorado Springs, Colo. 6,255 0.697 -- X 2003–2008 

12 07104050 North Rockrimmon Creek above Delmonico Drive at Colorado Springs, Colo. 6,220 1.82 X X 2001–2008 13 385204104510101 Monument Creek Tributary 2 below Fillmore Street at Colorado 

Springs, Colo.

6,270 0.75 X X 2003–2008 

14 385124104501301 Monument Creek Tributary 2 at 

Sondermann Park at Colorado Springs, Colo.

6,060 2.04 X X 2003–2008 

15 07104905 Monument Creek at Bijou Street at 

Colorado Springs, Colo.

5,980 235 X X 2001–2008 200
Table 1. Description of sites and period of record for types of data collected in the Fountain Creek basin, Colorado, 2001–2008.—Continued[USGS, U.S. Geological Survey; ID, identification; NAVD 88, North American Vertical Datum of 1988; ft, feet; mi2, square miles; X, collected; --, not collected] 

Site 

ID1

USGS

station 

number

Station 

name

Elevation 

NAVD 88

(ft)

Drainage 

area

(mi2)

Streamflow

discharge Chemistry Habitat 

16 07105000 Bear Creek near Colorado Springs, 

Colo.

6,520 6.89 X X 2003–2008 

17 384909104504401 Bear Creek above 8th Street at 

Colorado Springs, Colo.

6,037 9.57 X X 2003–2008 

18 07105500 Fountain Creek at Colorado Springs, 

Colo.

5,900 392 X X 2001–2008 200

19 07105530 Fountain Creek below Janitell Road 

below Colorado Springs, Colo.

5,840 413 X X 2001–2008 200

20 07105800 Fountain Creek at Security, Colo. 5,640 495 X X 2001–2008 20021 07105900 Jimmy Camp Creek at Fountain, 

Colo.

5,530 65.6 X X 2001–2008 200

22 07106000 Fountain Creek near Fountain, Colo. 5,355 681 X X 2001–2008 20023 07106300 Fountain Creek near Pinon, Colo. 4,990 849 X X 2001–2008 20024 07106500 Fountain Creek at Pueblo, Colo. 4,705 926 X X 2001–2008 200

1See figure 1.2 Fish data from 07103700 were not used because community composition was restricted to one species, and fish data from 07105900 were not used because there were only 
6 Temporal Change in Biological Community Structure in the Fountain Creek Basin, Colorado, 2001–2008

Table 2. Habitat, streamflow, and water-chemistry variables used to relate site-specific temporal patterns (seriation) in biological 

community structure, Fountain Creek basin, Colorado, 2001–2008. 

[%, percent; m, meters; ft3, cubic feet; /, per; s, second; mi2, square miles; ≥, greater than or equal to; mg/L, milligrams per liter; N, nitrogen]

Code Variable description and unit

Channel form

pctRIF % of reach length composed of riffle habitat (%)

BANKHT1 bank height (m)

FROUDE Froude number (current velocity divided by [{gravitational acceleration multiplied by depth of water} exponent0.5] 

dimensionless)

Substrate composition

pctSAND % occurrence of sand from Wolman pebble count (%)

pctGRAVEL % occurrence of gravel from Wolman pebble count (%)

pctCOBB % occurrence of cobble from Wolman pebble count (%)

Streamflow

MH201 Drainage area normalized annual maximum daily mean flow (ft3/s/mi2)

FH11 High flood-pulse count (number of events /year ≥ 75th percentile)

Water chemistry

DNO231 dissolved nitrite plus nitrate (mg/L as N)

1

Variables from Zuellig and others (2007).

study period at cross sections in the vicinity of the USGS 

streamgages that had the most even distribution of streamflow. 

These measurements of stream discharge were used to develop 

site-specific stage-discharge relations (rating curves) that 

were then used to compute continuous discharge data from 

the stage record associated with the 24 USGS surface-water 

gages or temporary transducers. Mean daily discharge was 

acquired from USGS streamgage records (Crowfoot and others, 2002–2008). The Hydrologic Index Tool (HIT) software 

(Henriksen and others, 2006) was used to calculate annual 

values for normalized maximum daily mean flow (MH20) and 

high flood-pulse count (FH1). 

Habitat. Habitat variables were measured after biological sampling each year along a predefined reach defined as 20 

times the mean wetted width (range from 100 to 300 m [328 to 

984 ft]). Bank height and percent riffle were estimated following the methods of Meador and others (1993) where multiple 

measurements were made at six equally spaced transects 

along the reach perpendicular to streamflow. Streambed and 

bank substrate were characterized from a pebble count made 

at approximately 100 points along the stream reach (Wolman, 

1954). Although substrate was characterized into six categories during sampling, only percent sand, percent gravel, and 

percent cobble were considered herein (see table 2; Zuellig 

and others, 2007). 

Water Chemistry. Routine nitrate plus nitrite samples 

were collected during nonstormflow (low summer flows 

between storms) conditions in June and September, bracketing 

the biological sampling using standard equipment with widthand depth-integrating techniques. All samples were collected, 

processed, and preserved in the field according to standard 

methods described in the National Field Manual by the U.S. 

Geological Survey (1997–2008). Samples were analyzed 

at the USGS National Water Quality Laboratory (NWQL) 

Lakewood, Colo. See Mau and others (2007) for detailed 

information concerning water-chemistry collection methods, 

interpretation of water-chemistry data, as well as information 

concerning quality assurance/quality control (QA/QC). Mean 

concentrations were used in this analysis. 

Macroinvertebrates. Macroinvertebrates were sampled 

annually at numerous sites during base-flow conditions (timeof-year when streamflow is dominated by groundwater seepage) in October between 2001 and 2008 (table 1). A dip-net 

equipped with 500-mm mesh was used to collect a qualitative 

multihabitat sample (QMH sample type described by Moulton 

and others, 2000) where organisms were collected from 

all available habitat types within each stream reach. These 

samples often contained a considerable amount of debris that 

was reduced in the field by elutriating and sieving (500-mm 

mesh-metal sieve) until sample volumes were approximately 

500 mL (about 1 pt). Invertebrate samples were preserved in 

the field with either 95-percent ethanol or 10-percent buffered 

formalin, and stored until delivered to the contract laboratory 

(Chadwick and Associates, Littleton, Colo., or Aquatic Associates, Fort Collins, Colo.). All unique taxa were identified from 

each sample and recorded as presence-absence data. A voucher 

collection was prepared and updated annually and verified by 

the USGS NWQL. Before analysis, macroinvertebrate taxonomic names were harmonized and ambiguities were resolved 

for each sample. 

Fish. Fish community data were collected annually at 

nine sites in mid-September during base-flow conditions 

between 2003 and 2008 (table 1). All sites were wadable at the 

time of sampling and electrofished in an upstream direction 
Methods 7

with a single pass (Pusey and others, 1998) using two SmithRoot pulsed direct current (DC) backpack electroshockers 

(model 12B POW and model 15D POW). A concerted effort 

was made to sample all available habitats. After capture, fish 

were maintained in aerated live wells. Fish were identified, 

enumerated, and measured (total length) in the field, tallied in 

the nearest 2-mm (0.79-in.) length class, observed for anomalies, and released. 

Data Analysis

Environmental, macroinvertebrate, and fish data were 

analyzed using two nonparametric multivariate routines available in the PRIMER version 6.1 software package, Plymouth, 

United Kingdom (Clarke and Warwick, 2001). First, the 

RELATE routine was applied to macroinvertebrate and fish 

data to determine the presence of temporal change in community structure among years. Second, at sites where temporal 

change occurred, the BIO-ENV routine was used to determine 

which subset of environmental variables (streamflow, habitat, 

and water chemistry) best described site-specific changes in 

community structure. The underlying ordination method used 

in these multivariate routines is nonmetric multidimensional 

scaling (MDS). The advantages of MDS over other ordination 

techniques is that MDS does not make any assumptions about 

the form of the data or the interrelations of the samples, and it 

has greater ability than other ordination methods to represent 

complex relations in low dimensional space. See Clarke and 

Warwick (2001) and Clarke and Gorley (2006) for details 

regarding the REALTE and BIO-ENV PRIMER routines used 

herein, which are briefly discussed in the sections below. 

Determining Site-Specific Temporal Change in 

Biological Communities among Years

The RELATE routine (test for seriation) was used to 

estimate the degree of site-specific temporal change (Clarke 

and Warwick, 2001) in biological communities among years, 

where temporal change is defined as the extent to which community structure departs from a predefined starting point; in 

this case, the first year a sample was collected. The RELATE 

routine computes the strength between two independently 

derived matrices as the Spearman Rank correlation coefficient (Spearman’s rho; Kendall, 1970). In this case, the two 

matrices were one of biological similarities and the other of 

temporal distances (that is, years) among samples. Under this 

analysis, if changes in community structure exactly match the 

temporal sequence (for example, community composition of 

the 2001 sample is most similar to 2002, less similar to 2003, 

and so on with 2007 being the most dissimilar), Spearman’s 

rho (hereafter, rho) will be close to 1; whereas, it will be near 

zero when there is no apparent temporal pattern. In this report, 

rho greater than 0.9 indicates a very strong temporal change, 

0.8 to 0.9 indicates a strong change, 0.7 to 0.8 indicates a relatively strong change, 0.5 to 0.7 indicates a moderate change, 

and less than 0.5 indicates a weak or no change. Additionally, 

the RELATE routine incorporates a significance test derived 

by permutation to test the null hypothesis of complete absence 

of temporal change (Clarke and Warwick, 2001). Herein, rho 

values were considered significant (that is, the null could not 

be rejected) when less than 5 percent of 999 randomly permutated rho values were greater than the real rho value (p value 

less than 0.05). Because community change was evaluated 

among equally spaced temporal distances (years), the presence 

of a temporal change at a site (significant rho value) indicated 

a linear change in community structure among years. 

Before the analysis, Bray-Curtis similarity values among 

samples were calculated from presence-absence macroinvertebrate data (Clarke and Warwick, 2001). Bray-Curtis similarity values measure the degree of similarity in community 

structure between pairs of samples. Values range between zero 

and one, where one indicates an exact match in community 

composition and zero indicates completely different communities between two samples. A fourth-root transformation 

was applied to the relative abundance fish data in order to 

down weight the most dominant species (that is, to prevent the 

similarity values calculated from these data to be driven by 

mostly the highly abundant species), while retaining information about rare species. The time-sequential matrix among 

samples was calculated directly in the RELATE routine under 

the “result of seriation” option in the Primer software (Clarke 

and Warwick, 2001).

Relating Environmental Variables to Observed 

Temporal Change in Biological Community 

Structure

At sites where temporal patterns were detected based 

on the RELATE test for seriation, the BIO-ENV routine was 

applied to determine which subset of available environmental 

variables best matched temporal change in biological community structure. BIO-ENV also uses rank Spearman’s rho to 

determine the strength of association between two similarity 

matrices (Clarke and Warwick, 2001); in this case, between 

the biota and concurrently collected environmental data. All 

possible subsets of environmental variables were evaluated for 

the best match (that is, highest rho value) with the biological 

data; however, only models with four or fewer variables were 

considered to decrease spurious results. The subset with the 

highest rho value (closest to 1) for each site was determined 

to best describe community change among years, given the 

data. The significance of the best subset (Global BEST test) 

was calculated through permutation as described by Clarke 

and Gorley (2006), testing the null hypothesis of no relation 

between the best subset of environmental variables and the 

associated biota. The best subset of environmental variables 

was considered significant (not due to chance) when less than 

5 percent of 999 randomly permutated rho values were greater 

than the real value (p <0.05). Because the BIO-ENV analysis was specific to sites with temporal change, a significant 
8 Temporal Change in Biological Community Structure in the Fountain Creek Basin, Colorado, 2001–2008

BIO-ENV finding indentified the subset of selected environmental variables that best matched observed linear changes in 

biological community structure among years. 

Before analysis, the environmental data were log10 transformed to improve normality and then normalized so that each 

variable had a mean of zero and a standard deviation of one. 

This normalization was necessary to ensure that each variable 

took on values over a similar range because different classes 

of environmental variables were measured on different scales. 

After transformation and normalization, Euclidean distance 

values were calculated for the environmental data (Clark and 

Warwick, 2001). Bray-Curtis similarity values were calculated 

from presence-absence macroinvertebrate data and fourth-root 

transformed relative abundance fish data. 

Temporal Change in Biological 

Community Structure

Temporal change in biological community structure was 

defined as the extent to which community structure departed 

from what it was the first year a sample was collected at a 

site (Clarke and Warwick, 2001). The strength of departure 

was measured as Spearman’s rho (Kendall, 1970), where rho 

values were close to one if strong temporal change occurred at 

a site and close to 0 if change was absent. 

Macroinvertebrate Communities

Temporal change in macroinvertebrate community 

structure was present at 10 of the 24 sites evaluated (table 3). 

Of the 10 sites, only site 8 showed relatively strong temporal change (rho = 0.71, p <0.003), whereas the strength of 

temporal change was more moderate at the remaining 9 sites 

(median rho = 0.53; range, 0.42 to 0.59). Sites with significant 

temporal change that were represented by only 6 years of 

macroinvertebrate data (sites 14, 16, and 21) tended to have 

lower rho values (0.49, 0.47, and 0.54) and higher percent 

significance values (4.5, 4.9, and 4.1), which is in part due 

to the smaller number of observations available for analysis. 

Additional years of data are needed to determine how genuine 

these directional changes in community structure are at these 

sites. 

Although temporal change of macroinvertebrate community structure was detected among tributary and main stem 

sites, this change was absent at the three most downstream 

sites as well as at other sites (mostly tributaries) scattered 

throughout the basin (fig. 2). Only 6 years of data were collected at most tributary sites; therefore, additional years of 

data at these sites may uncover other directional changes in 

community structure, especially at those sites with nonsignificant rho values (closest to 0.5). However, it is unlikely 

that temporal change will be detected at the lower sites (22, 

23, and 24) by continuing to collect presence-absence type 

macroinvertebrate data unless major environmental changes 

occur at these sites; for example, changes in stream hydrology 

or water-quality conditions significant enough to restructure 

the current fauna. 

Fish Communities

Significant temporal change in fish community structure 

was detected at five of the nine sites where fish data were 

analyzed (table 4). At three of the five sites, rho values were 

greater than 0.70 (range 0.71 to 0.76), indicating relatively 

strong temporal patterns in fish community structure between 

2003 and 2008. Other temporal changes, although significant, 

were more moderate (rho values 0.46 to 0.51) at the remaining 

two sites. 

Additional years of data are needed at sites with moderately strong rho values (around 0.5) to determine the persistence of temporal changes at these sites. Likewise additional 

years of data may reveal temporal change in community 

structure at site 18 (Fountain Creek at Colorado Springs) as 

rho was closer to 0.50 and percent significance levels were 

relatively low when compared to the other sites with nonsignificant rho values (sites 2, 15, and 22). In contrast to 

macroinvertebrates, fish community temporal changes were 

detected with stronger signatures and at sites distributed along 

the length of the Fountain Creek basin, including the two most 

downstream sites (fig. 3). Unfortunately, little is known about 

the temporal distribution of fish community structure among 

tributary sites as these data were not collected. 

Environmental Variables Correlated 

to Temporal Changes in Biological 

Community Structure

At sites where temporal patterns were detected, additional analyses were completed to determine which subset 

of environmental variables best matched temporal changes 

in biological community structure. The strength of association between the biota and the best subset of environmental 

variables was measured using Spearman’s rho as described by 

Clark and Warwick (2001). 

Macroinvertebrate Communities

Environmental variables significantly described temporal change among macroinvertebrate communities at 4 of the 

10 sites (sites 6, 15, 16, and 20) where seriation was present 

(table 5). At these sites, the strength of association between 

the best subset of environmental variables and community 

structure was moderate (rho values 0.60 and 0.63) to strong 

(rho values 0.81 and 0.84). 

The subsets of environmental variables with significant 

rho values appeared to be site-specific with no single variable 

dominating the models with the highest rho values among 
Environmental Variables Correlated to Temporal Changes in Biological Community Structure 9

Table 3. Correlation coefficients, percent significance level, and the number of years associated with the strength of multivariate 

seriation of macroinvertebrate community structure among years (RELATE test for seriation). 

[Sites with significant correlations between patterns in macroinvertebrate community structure and equally spaced temporal distances (years) are in bold] 

Site 

ID1

Station 

name

Spearman's 

rho

Percent 

significance 

level

Years 

of data

1 Fountain Creek near Colorado Springs, Colo. 0.24 10.7 8

2 Fountain Creek below 8th Street at Colorado Springs, Colo. 0.17 20.4 8

3 Kettle Creek above Old Ranch Road near Colorado Springs, Colo. 0.39 10.7 6

4 Kettle Creek above U.S. Air Force Academy, Colo. 0.28 16.9 6

5 Pine Creek above Highway 83 at Colorado Springs, Colo. 0.39 7.7 6

6 Monument Creek above Woodmen Road at Colorado Springs, Colo. 0.59 0.4 8

7 Cottonwood Creek at Cowpoke Road at Colorado Springs, Colo. 0.29 8.7 8

8 Cottonwood Creek at Woodmen Road near Colorado Springs, Colo. 0.71 0.3 8

9 Cottonwood Creek Tributary above Rangewood Drive at Colorado Springs, Colo. 0.47 1.4 8

10 Cottonwood Creek at mouth at Pikeview, Colo. 0.19 17.9 8

11 Monument Creek Tributary 1 near Pulpit Rock at Colorado Springs, Colo. 0.08 38.2 6

12 North Rockrimmon Creek above Delmonico Drive at Colorado Springs, Colo. 0.30 7.1 8

13 Monument Creek Tributary 2 below Fillmore Street at Colorado Springs, Colo. 0.25 16.2 6

14 Monument Creek Tributary 2 at Sondermann Park at Colorado Springs, Colo. 0.49 4.5 6

15 Monument Creek at Bijou Street at Colorado Springs, Colo. 0.56 0.4 8

16 Bear Creek near Colorado Springs, Colo. 0.47 4.9 6

17 Bear Creek above 8th Street at Colorado Springs, Colo. 0.24 23.0 6

18 Fountain Creek at Colorado Springs, Colo. 0.42 1.8 8

19 Fountain Creek below Janitell Road below Colorado Springs, Colo. 0.53 0.2 8

20 Fountain Creek at Security, Colo. 0.58 0.9 8

21 Jimmy Camp Creek at Fountain, Colo. 0.54 4.1 6

22 Fountain Creek near Fountain, Colo. 0.06 38.3 8

23 Fountain Creek near Pinon, Colo. 0.19 16.3 8

24 Fountain Creek at Pueblo, Colo. 0.25 11.2 8

1

See figure 1. 

sites. For example, percent sand (pctSAND), bank height 

(BANKHT), and normalized annual maximum daily mean 

flow (MH20) best described temporal change at site 6 (Monument Creek above Woodmen Road at Colorado Springs); 

whereas, bank height (BANKHT) and the number of high 

flood-pulse counts (FH1) best described temporal change at 

site 16 (Bear Creek near Colorado Springs, Colo.). 

Although the best subsets were not dominated by single 

variables, groups of similar variables were present at the four 

sites with significant BIO-ENV results (fig. 4). For example, 

substrate composition variables were present in three of the 

four best subsets. 

Although approached differently, others have found 

substrate particle size to be an important indicator of macroinvertebrate community structure in the Fountain Creek basin 

(von Guerard, 1989; Bruce, 2002). Both the von Guerard 

(1989) and Bruce (2002) studies looked at macroinvertebrate 

and environmental data from several sites collected over 

a few years that were longitudinally arranged in the basin. 

Under such longitudinal designs (especially when inclusive 

of transitional areas between mountains and plains), macroinvertebrate communities are expected to change predictably 

along the naturally occurring environmental gradients (for 

example, substrate particle size decreases in a downstream 

direction). Integrating the results from this study with those of 

von Guerard (1989) and Bruce (2002) indicates that substrate 

characteristics of the basin, as well as site-specific changes 

in substrate composition over time, influenced macroinvertebrate community structure—a finding well represented in the 

literature (for example, see Waters, 1995). Additionally, von 

Guerard (1989) found that flooding 30 days prior to invertebrate sampling explained as much as 77 percent of variation in 

macroinvertebrate densities. Herein, three of the four best subsets of environmental variables included similar streamflow 
10 Temporal Change in Biological Community Structure in the Fountain Creek Basin, Colorado, 2001–2008

EXPLANATION

Sites without macroinvertebrate temporal changes

! Sites with macroinvertebrate temporal changes

!

!

!

!

!

!!

!

!

!

!

! !

!

!

!

!

9

8 7

6

4

3

5

2

1

24

23

23

22

21

20

19

18

16

15

12 11

13

17

134

110

!

Number corresponds to site identification numbers 

 in table 1

Stream or canal

105o

105o45’

39o

39o45’

39o30’

!

!

!

!!

!

!

0 5 10 20 MILES

0 5 10 20 KILOMETERS

Figure 2. Spatial distribution of sites with and without temporal changes in macroinvertebrate 

communities among years in the Fountain Creek basin, Colorado, 2001–2008.
Implications for Future Monitoring 11

Table 4. Correlation coefficients and percent significance level associated with the strength of multivariate seriation of fish 

communtiy structure among years (RELATE test for seriation). 

[Sites with significant correlations between patterns in fish community structure and equally spaced temporal distances (years) are in bold] 

Site ID1 Station name Spearman's 

rho

Percent 

significance level

2 Fountain Creek below 8th Street at Colorado Springs, Colo. –0.19 72.7

6 Monument Creek above Woodmen Road at Colorado Springs, Colo. 0.76 0.6

15 Monument Creek at Bijou Street at Colorado Springs, Colo. 0.10 34.4

18 Fountain Creek at Colorado Springs, Colo. 0.36 9.5

19 Fountain Creek below Janitell Road below Colorado Springs, Colo. 0.76 0.5

20 Fountain Creek at Security, Colo. 0.71 0.8

22 Fountain Creek near Fountain, Colo. –0.25 72.7

23 Fountain Creek near Pinon, Colo. 0.46 2.5

24 Fountain Creek at Pueblo, Colo. 0.51 3.0

1

See figure 1.

characteristics that were previously linked to impervious surface coverage in the Fountain Creek basin (Zuellig and others, 

2007). Channel form characteristics (for example, BANKHT 

and pctRIF) were also included in some of the best subsets 

of environmental variables; however, nutrient concentrations 

(DNO23) were not found important among the best subsets 

of environmental variables (table 5, fig. 4) as was reported by 

Zuellig and others (2007).

Although environmental variables described temporal 

change in macroinvertebrate community structure at some 

sites (four), there were several sites (six) where temporal 

change was present but unexplained by the available variables 

(table 5, fig. 4). In most cases (sites 8, 9, 14, 18, and 19), rho 

values were greater than 0.4 (near moderate) and percent significance levels were weak (about 19 to 73 percent). Because 

the significance level of the Global BEST test in the BIO-ENV 

procedure is strongly related to the number of data points 

being fit (Clarke and Warwick, 2001), additional years of data 

may reveal more conclusive results, as long as the descriptive variables of temporal change in community structure are 

measured. 

Fish Communities

Although temporal change among fish communities 

was detected at many sites, they were explained by environmental variables at only one site (site 6, table 6). At this 

site, the strength of association between the best subset of 

environmental variables (substrate composition—pctSAND, 

pctGRAVEL; channel form—BANKHT) and temporal change 

in fish community structure was very strong (rho 0.94; percent 

significance 0.30). 

Additionally, environmental variable subset three (table 

6; site 6, subset 3) for this site was limited to pctSAND and 

BANKHT, which described patterns almost equally as well 

(rho = 0.90) as the subset with the highest rho value for that 

site (table 6; site 6, subset 1). These two variables (pctSAND, 

BANKHT) were also included in the best subset that described 

temporal change in macroinvertebrate community structure at 

this site (table 5; site 6, subset 1). 

As with macroinvertebrate communities, there were 

several sites where temporal change in fish community 

structure was present, but these changes were not described 

by the environmental variables available for analysis. All fish 

analyses were limited to 6 years of data (2003 to 2008) at each 

site, which likely influenced RELATE and BIO-ENV results 

as described in the previous sections. Additional years of data 

may reveal more conclusive BIO-ENV results, if the most 

influential environmental variables are measured. 

Implications for Future Monitoring

Results from this study were derived from a relatively 

small number of observations (6 or 8 years of data); therefore, 

additional years of data may reveal other sites with temporal 

change or possibly define stronger linkages between environmental variables and observed temporal change. Likewise the 

temporal changes that were detected could become weaker. 

Nonetheless, there were several sites where temporal changes 

in community structure were detected in this study that could 

not be explained by the selected environmental variables 

studied herein. At some of these sites, it is possible that the 

important environmental variables necessary to describe these 

temporal changes in aquatic communities were not available 

for analysis, because they were not consistently collected 

(see tables 5 and 6) or not part of the current study. Either 

way, modification of future data-collection activities is likely 
12 Temporal Change in Biological Community Structure in the Fountain Creek Basin, Colorado, 2001–2008

! Sites without fish temporal changes

! Sites with fish temporal changes

!

!

!

6

2

24

23 23

22

20

19

18

15

Number corresponds to site identification numbers 

 in table 1

Stream or canal

105o

105o45’

39o

39o45’

39o30’

!

!

!

!

!

!

EXPLANATION

0 5 10 20 MILES

0 5 10 20 KILOMETERS

Figure 3. Spatial distribution of sites with and without temporal changes in fish communities among 

years in the Fountain Creek basin, Colorado, 2003–2008. 
Table 5. Results of the BIO-ENV procedure determining which environmental variables best described observed multivariate temporal change (sermacroinvertebrate community structure in the Fountain Creek basin, 2001–2008. [Three subsets for each site with the highest Spearman's rho are presented, those that were significant (that is < 5 percent of 999 permutated rho values were greater than the reaidentification; %, percent; /, per; mg/L, milligrams per liter]

Site

ID1 Site name Subset number Number of variables Spearman's rho Percent significance level Variables included in model2 

6 Monument Creek above Woodmen Road at 

Colorado Springs, Colo.

1 3 0.63 1.3 pctSAND, BANKHT, MH20 2 4 0.62 pctSAND, pctCOBB, BANKHT, MH3 2 0.60 pctSAND, MH20

8 Cottonwood Creek at Woodmen Road near 

Colorado Springs, Colo.

1 3 0.43 31.5 pctCOBB, FROUDE, FH1 2 2 0.42 pctCOBB, FROUDE, FH13 3 0.39 pctGRAVEL, pctCOBB, FROUDE

9 Cottonwood Creek tributary above Rangewood 

Drive at Colorado Springs, Colo.

1 2 0.45 19.4 pctCOBB, FROUDE 2 3 0.44 pctCOBB, FROUDE, BANKHT3 1 0.44 FROUDE

14 Monument Creek Tributary 2 at Sondermann 

Park at Colorado Springs, Colo.

1 2 0.42 73.2 pctGRAVEL, pctCOBBLE 2 2 0.35 pctGRAVEL, BANKHT3 3 0.35 pctSAND, pctGRAVEL, BANKHT

15 Monument Creek at Bijou Street at Colorado 

Springs, Colo.

1 2 0.60 3.2 pctGRAVEL, pctCOBB 2 3 0.58 pctGRAVEL, pctCOBB, FROUDE3 4 0.53 pctGRAVEL, pctCOBB, DNO23, 

FROUDE

16 Bear Creek near Colorado Springs, Colo. 1 2 0.81 4.2 BANKHT, FH1 

2 3 0.80 DNO23, BANKHT, FH13 4 0.78 pctSAND, pctRIF, BANKHT, FH1
Table 5. Results of the BIO-ENV procedure determining which environmental variables best described observed multivariate temporal change (sermacroinvertebrate community structure in the Fountain Creek basin, 2001–2008.—Continued [Three subsets for each site with the highest Spearman's rho are presented, those that were significant (that is < 5 percent of 999 permutated rho values were greater than the reaidentification; %, percent; /, per; mg/L, milligrams per liter] Site ID1 Site name Subset number Number of variables Spearman's rho Percent significance level Variables included in model2 

18 Fountain Creek at Colorado Springs, Colo. 1 4 0.47 28.7 pctGRAVEL, pctRIF, BANKHT 

2 3 0.47 pctGRAVEL, pctCOBBLE, BANKH3 4 0.44 pctGRAVEL, pctCOBBLE, BANKH

MH20 19 Fountain Creek below Janitell Road below Colorado Springs, Colo. 1 2 0.44 49.0 DNO23, BANKHT 

2 1 0.39 BANKHT 3 2 0.35 pctSAND, BANKHT 20 Fountain Creek at Security, Colo. 1 4 0.84 0.9 pctSAND, pctRIF, FH1, MH20 2 4 0.82 pctSAND, pctCOBB, FH1, MH20 3 3 0.80 pctSAND, FH1, MH20 21 Jimmy Camp Creek at Fountain, Colo. 1 1 0.10 90.1 pctGRAVEL 2 2 0.08 pctSAND MH20 3 4 0.02 pctSAND, pctGRAVEL, BANKHT, MH20 1See figure 1. 2pctSAND, % occurrence of sand from Wolman pebble count (Wolman, 1954); BANKHT, bank height; MH20, drainage area normalized annual maximum daily mean flow; 

cobble from Wolman pebble count; FH1, high flood-pulse count (number of events per year with flows above the 75th percentile); FROUDE, (Froude number, current velocity acceleration multiplied by depth of water} exponent0.5] dimensionless); DNO23, dissolved nitrite plus nitrate (mg/L as N); pctGRAVEL, % occurrence of gravel from Wolman reach composed of riffle habitat.
Implications for Future Monitoring 15

! Sites without temporal changes

! Sites with macroinvertebrate temporal changes

! Sites with fish temporal changes

! Sites with both macroinvertebrate and fish temporal 

 changes

!

!

!

!

!

!

!!

!

!

! !

!

!

! !

!

!

!

!

!

07103970–Monument Creek above Woodmen Rd. 

at Colorado Springs, Colo. 

07104905–Monument Creek 

at Bijou St. at Colorado Springs, Colo.

07105000–Bear Creek near

 Colorado Springs, Colo. 

07105800–Fountain Creek 

at Security, Colo.

9

8 7

6

4 3

5

2

1

24

23

22

21

20

19

18

16

15

12 11

13

17

134

110

!

15 Number corresponds to site identification numbers 

 in table 1

Stream or canal

!

!

105o

105o45’

39o

39o45’

39o30’

EXPLANATION

0 5 10 20 MILES

0 5 10 20 KILOMETERS

Macroinvertebrates–Substrate composition

Macroinvertebrates–Substrate

composition, channel form, 

streamflow

Macroivertebrates–Channel

form, streamflow

Fish–Substrate composition, 

channel form

Macroinvertebrates–Substrate composition, 

channel form, streamflow

Figure 4. Spatial distribution of sites with and without temporal changes in fish (2003–2008) and macroinvertebrate 

(2001–2008) communities among years in the Fountain Creek basin, Colorado.
Table 6. Results of the BIO-ENV procedure determining which environmental variables best described observed multivariate temporal change (serstructure in the Fountain Creek basin, 2003–2008. [Three subsets for each site with the highest Spearman's rho are present, those that were significant (that is, < 5 percent of 999 permutated rho values greater than observed rho cation; %, percent; /, per; mg/L, milligrams per liter] Site ID1 Site name Subset number Number of variables Spearman's rho Percent significance level Variables included in model2 

6 Monument Creek above Woodmen Road at Colorado Springs, Colo. 1 3 0.94 0.30 pctSAND, pctGRAVEL, BAN

2 4 0.92 pctSAND, pctGRAVEL, DNO

BANKHT 3 2 0.90 pctSAND, BANKHT 19 Fountain Creek below Janitell Road below Colorado Springs, Colo. 1 1 0.72 25.20 pctRIF 

2 3 0.70 pctGRAVEL, pctCOBB, pctRI3 4 0.69 pctGRAVEL, DNO23, pctRIF, 

20 Fountain Creek at Security, Colo. 1 1 0.72 28.40 pctCOBB 

2 2 0.70 pctCOBB, DNO23 3 4 0.70 pctCOBB, DNO23, BANKHT

23 Fountain Creek near Pinon, Colo. 1 3 0.66 27.10 pctSAND, pctGRAVEL, MH2

2 3 0.58 pctGRAVEL, BANKHT, MH23 2 0.58 pctGRAVEL, MH20 24 Fountain Creek at Pueblo, Colo. 1 3 0.57 49.10 pctSAND, DNO23, FROUDE 2 2 0.50 DNO23, FROUDE 3 4 0.50 pctSAND, DNO23, FROUDE

BANKHT 1See figure 1. 2pctSAND, % occurrence of sand from Wolman pebble count (Wolman, 1954); pctGRAVEL, % occurrence of gravel from Wolman pebble count; BANKHT, bank height; D

nitrate (mg/L as N); pctRIF, % of reach composed of riffle habitat; pctCOBB, % occurrence of cobble from Wolman pebble count; FH1, high flood pulse count (number of evethe 75th percentile); MH20, drainage are normalized annaul maximum daily mean flow; FROUDE, (Froude number, current velocity divided by [{gravitational acceleration muexponent0.5] dimensionless). 
References Cited 17

necessary to better understand site-specific relations between 

biological communities and selected environmental variables. 

Previous biological studies in the Fountain Creek basin 

were observational in nature. Although these studies have 

done well at characterizing the environment and biology, they 

have difficulty linking human-induced environmental alterations to changes in aquatic community structure. To develop 

causal linkages between individual stressors and changes in 

aquatic communities in the Fountain Creek basin, it will be 

necessary to conduct observational studies (such as this one) 

along with field and laboratory experiments (Clements and 

others, 2002). Information generated from such integrative 

studies are powerful for developing causal relations and may 

be imperative for well-targeted mitigation and/or management 

efforts in the Fountain Creek basin.

Summary and Conclusions

Fountain and Monument Creeks (hereafter, Fountain 

Creek basin) drain approximately 926 mi2 of the eastern slope 

of the Rocky Mountains in south-central Colorado. In 2001, 

the U.S. Geological Survey, in cooperation with Colorado 

Springs City Engineering, began a study to better understand 

the relations between environmental characteristics and 

biological communities in the Fountain Creek basin in order 

to aid water-resource management and guide future monitoring activities. Environmental (streamflow, habitat, and water 

chemistry) and biological (fish and macroinvertebrates) data 

were collected annually at 24 sites between 2001 and 2008 

(fish data collected 2003–2008 only). For this report, these 

data were first analyzed to determine the presence of temporal 

change in macroinvertebrate and fish community structure 

among years using nonparametric multivariate statistics 

(RELATE, PRIMER ver. 6.1, Plymouth, United Kingdom). 

Where temporal change in the biological communities was 

found, these data were further analyzed using additional nonparametric multivariate techniques to determine which subset 

of selected streamflow, habitat, or water chemistry variables 

best described site-specific changes in community structure 

relative to a gradient of urbanization (BIO-ENV, PRIMER ver. 

6.1, Plymouth, United Kingdom). 

This study found the presence of temporal change in 

community structure at 10 sites for macroinvertebrates and 

5 sites for fish in the Fountain Creek basin. Of these sites, 

environmental variables significantly described temporal 

change among macroinvertebrate communities at four sites 

(Monument Creek above Woodmen Road at Colorado Springs, 

Colo.; Monument Creek at Bijou Street at Colorado Springs, 

Colo.; Bear Creek near Colorado Springs, Colo.; Fountain 

Creek at Security, Colo.) and at one site for fish (Monument 

Creek above Woodmen Road at Colorado Springs, Colo.). 

Combinations of environmental variables describing directional temporal change in the biota appeared to be site specific 

as no single variable dominated the results; however, substrate 

composition variables (percent substrate composition composed of sand, gravel, or cobble) collectively were present in 

almost 80 percent of the environmental variable subsets that 

were significantly correlated with temporal change in the macroinvertebrate and fish community structure. Other important 

environmental variables related to temporal change in the biological community structure included those describing channel 

form (streambank height) and streamflow (normalized annual 

mean daily flow, high flood-pulse count).

Site-specific results from this study were derived from 

a relatively small number of observations (6 or 8 years of 

data). Additional years of monitoring data will likely improve 

statistical interpretations and provide more insight into the 

observations described in this report. There were several sites 

where temporal change was detected in this study that could 

not be explained by the selected environmental variables studied herein. Modification of current data-collection activities is 

necessary to better understand site-specific relations between 

biological communities and environmental variables. 

Acknowledgments

Gary Dowler, Paul Foutz, Dan Frankowski, Londell Ackley, and Harry Crockett of the Colorado Division of Wildlife 

(CDOW) provided backpack electrofishing equipment and 

contributed greatly to the fish-sampling efforts. Thanks to Lisa 

Ross, Colorado Springs City Engineering, for her support of 

the program. Thanks also to Colorado Springs Utilities for 

their support. Harry Crockett and Travis Schmidt (USGS) 

provided thoughtful reviews and comments that improved the 

report. The authors also thank the USGS hydrographers that 

collected, compiled, and analyzed streamflow measurement 

data.

References Cited

Bruce, J.F., 2002, Characterization and analysis of temporal 

and spatial variations in habitat and macroinvertebrate community structure, Fountain Creek basin, Colorado Springs 

and vicinity, Colorado, 1998–2001: U.S. Geological Survey 

Water-Resources Investigations Report 02–4093, 28 p.

Clarke, K.R., and Gorley, R.N., 2006, Primer version 6—User 

manual/tutorial, Primer-E: Plymouth, United Kingdom, 

190 p.

Clarke, K.R., and Warwick, R.M., 2001, Change in marine 

communities—An approach to statistical analysis 

and interpretation (1st ed.): Plymouth, United Kingdom, 

Plymouth Marine Laboratory, 172 p.
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Clements, W.H., Carlisle, D.M., Courtney, L.A., and Harrahy, 

E.A., 2002, Integrating observational and experimental 

approaches to demonstrate causation in stream biomonitoring studies: Environmental Toxicology and Chemistry, 

v. 21, p. 1138–1146.

Crowfoot, R.M., Steger, R.D., Payne, W.F., and O’Neill, G.B., 

2002, Water resources data, Colorado, water year 2001, v. 1: 

U.S. Geological Survey Water-Data Report CO 01–1, 539 p.

Crowfoot, R.M., Payne, W.F., and O’Neill, G.B., 2003, Water 

resources data, Colorado, water year 2002, v. 1: U.S. 

Geological Survey Water-Data Report CO 02–1, 609 p.

Crowfoot, R.M., Boulger, R.W., Payne, W.F., and O’Neill, 

G.B., 2004, Water resources data, Colorado, water year 

2003, v. 1: U.S. Geological Survey Water-Data Report 

CO 03–01, 577 p.

Crowfoot, R.M., Payne, W.F., O’Neill, G.B., and Boulger, 

R.W., 2005, Water resources data, Colorado, water year 

2004, v. 1: U.S. Geological Survey Water-Data Report 

CO 04–01, 344 p.

Crowfoot, R.M., Payne, W.F., O’Neill, G.B., Boulger, R.W., 

and Sullivan, J.R., 2005, Water resources data, Colorado, 

water year 2005, accessed March 2008, available online at 

http://wdr.water.usgs.gov.

Crowfoot, R.M., Payne, W.F., O’Neill, G.B., Boulger, R.W., 

Sullivan, J.R., and Lewis, M.E., 2006, Water resources data, 

Colorado, water year 2006, accessed March 2008, available 

online at http://wdr.water.usgs.gov.

Crowfoot, R.M., Payne, W.F., O’Neill, G.B., Sullivan, J.R., 

and Lewis, M.E., 2007, Water resources data, Colorado, 

water year 2007, accessed March 2008, available online at 

http://wdr.water.usgs.gov.

Crowfoot, R.M., Payne, W.F., O’Neill, G.B., Sullivan, J.R., 

Smith, G.J., and Lewis, M. E., 2008, Water resources data, 

Colorado, water year 2008, accessed January 15, 2008, 

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Publishing support provided by: 

Denver Publishing Service Center

For more information concerning this publication, contact:

Director, USGS Colorado Water Science Center

Box 25046, Mail Stop 415

Denver, CO 80225

(303) 236-4882

Or visit the Colorado Water Science Center Web site at:

http://co.water.usgs.gov/
Zuellig and others—Temporal Change in Biological Community Structure in the Fountain Creek Basin, Colorado, 2001–2008—SIR 2010-5103