Fish Community Assembly in the Forest River, North Dakota, and resolution of

Campostoma species presence

By

Lucas J. Borgstrom

A thesis submitted in partial fulfillment of the requirements for the

Master of Science

Major in Wildlife and Fisheries Sciences (Fisheries Specialization)

South Dakota State University

2010

11

Fish Community Assembly in t?e Forest River, North Dakota, and resolution of

Campostoma species presence

This thesis is approved as a creditable and independent investigation by a

candidate for the Master of Science degree and is acceptable for meeting the thesis

requirements for this degree. Acceptance of this thesis does not imply that the

conclusions reached by the candidate are necessarily the conclusions of the major

department.

, ~"'", ,..//,. ,/.I /'_'>~ ,! ! .. ~ - ,",./ ~~~1 ' L.'tJ,l, ./ll.'1

~ ..... ' ' _._' .:-/ _·(_· __ · .. ,_'_t.c.../ ..... I<-·'.....:''-, .-.:..' ..=;/-'-"'-' --''--/ --,---,--" ,,-

Dr. Charles R. Berry, Jr. Thesis Advisor

~w,t,/iIk' Dr. David W. Willis Head, Wildlife & Fisheries Sciences

Date

--------

iii

Acknowledgements

A special thanks to Dr. Charles R. Berry, Jr. for providing me with the

opportunity to study eastern North Dakota fishes and for his guidance and patience

through this process. My gratitude is also granted to my committee members, Dr. Brian

D. S. Graeb, Dr. Nels H. Troelstrup, Jr., and Dr. Sandra Bunkers for their assistance with

this thesis. I must thank the landowners who granted permission to sample the streams in

their property, without your cooperation some important stream reaches would not have

been sampled, although one thing that has become evident to me in my work is that all

stream reaches are important. Furthermore, a great deal of gratitude is due to the project

manager, Cari-Ann Hayer, without her efforts this project would not have been as

complete as it is. Using nonmetric multidimensional scaling as a community composition

analysis tool would not have even crossed my mind had it not been for Mark Fincel. My

thanks are also extended to the South Dakota Cooperative Fish and Wildlife Research

Unit for providing vehicles, tools, and sampling gear to complete this project. The Unit is

jointly sponsored by the South Dakota Department of Game, Fish, and Parks, United

States Fish and Wildlife Service, South Dakota State University, and the United States

Geological Survey. A special thanks to the many graduate and undergraduate students for

their assistance in the field and lab, especially Wes Bouska, Dan Dorris, Jake Billings,

Eric Boyda, Scott Sindelar, and Cory R. Stearns.

Beyond my current colleagues, a great deal of my gratitude is extended to the

people who helped me in my undergraduate career, my advisors, Dr. Debbie Guelda, Dr.

Mark Fulton, and Dr. Don Cloutman. Thank you all so much for your guidance. If it were

iv

not for your energy and enthusiasm for this field, I may not have continued to pursue my

interest in it. Also I want to thank Dr. Richard Koch for the internship opportunity he

granted me. This was when I cut my teeth in aquatic biology, and where I learned I want

to deal with fish rather than identifying early instar invertebrates.

Most importantly, I would like to thank my family for their support and

encouragement in my educational endeavors as well as introducing me to the great

outdoors and the educational and sustaining bounty it holds. To my wife Jody, I cannot

thank you enough for having the foresight to follow me to where I needed to be to get the

best education I could. For your persistence in nudging me along to complete this, as well

as your patience when hunting interrupted this writing process, thank you. I do not at all

feel that this is a comprehensive list of those who I want to thank, if I missed anybody, it

was not intentional. If you helped me in some way or somehow, you know it, and I am

grateful to you for that.

Funding for this project was provided by North Dakota Game and Fish

Department as a State Wildlife Grant Project. The state of North Dakota provided many

of the GIS layers used in the creation of maps as well as historic species locations. South

Dakota State University provided part (40%) of the state matching funds for this project.

v

Abstract

Fish Community Assembly in the Forest River, North Dakota, and resolution of

Campostoma sp. presence

Lucas J. Borgstrom

February 5, 2010

I was provided the opportunity to investigate the fish community associated with

the Red River basin of North Dakota. The first aspect I examined was the presence of the

genus Campostoma. In this aspect, I investigated the Campostoma species that were

present historically and documented the species that currently inhabit Red River waters in

North Dakota. The second aspect of my work was to study the fish community patterns

along the longitudinal dimension of the Forest River. I used nonmetric multidimensional

scaling to differentiate the patterns at two scales: level III and level IV ecoregions. The

final aspect of my study was to investigate habitat preferences of fishes in the Forest

River. I sampled fish by macrohabitat and compared the results to a similar study.

In the determination of the Campostoma species present, I investigated the fishes

found in the Red River basin to determine presence of central stoneroller, Campostoma

anomalum, and largescale stoneroller, C. oligolepis because both species have been

reported in the basin historically. Campostoma sp. were found at five sites. These

specimens and museum specimens collected during previous studies were C. oligolepis. I

developed a novel approach, termed the oblique circumferential scale count, to counting

vi

scales around the body to aid in differentiating C. anomalum from C. oligolepis. The sum

of oblique and lateral line scale count differentiates C. oligolepis (n = <84 scales) and C.

anomalum (n = 84 scales or greater).

The second aspect of the study was focused on the Forest River of North Dakota,

which provides a unique situation to assess both environmental and anthropogenic affects

on longitudinal fish assemblage because it flows through several ecoregions and is

divided by four epilimnetic-release dams (4.6–23.2 m high) impounding over 50 hectares

of water each. Sixteen sites were sampled using seining, backpack electrofishing, and

traps. Richness ranged among sites from 5–15 species and from 3–6 families. Shannon‘s

Diversity Index (H’) ranged from 0.77–1.95 and evenness (J’) varied from 0.24–0.81.

Nonmetric multidimensional scaling was conducted using percent similarity index values.

Site groupings were based on dams but not level IV ecoregions. Sites between two dams

were grouped together, excluding the site immediately below dams. The sites directly

below dams separated out closer to the most upstream sites than to the other groups. The

serial discontinuity concept explains this finding, but the process domains concept does

not. When analyzed at the level III ecoregion scale, the site groupings were closely

related to ecoregions. Still, the sites that were directly below a dam were closer to the

upstream sites than were the sites further downstream of the dams or upstream of the

reservoirs. Results from the level III ecoregion scale analysis agree with predictions of

both the serial discontinuity concept and process domains concept. The fish assemblage

in the Forest River is influenced by the anthropogenic forces of dams at the level IV

vii

ecoregion scale. At the level III ecoregion scale, the anthropogenic forces of dams and

natural community breaks based on ecoregions were both evident.

The final aspect of the study was to look into fish habitat preferences. Fish data

were collected at 16 sites on the Forest River, North Dakota and four macrohabitats

(pools, riffles, runs, and backwaters) that were characterized for depth, velocity,

substrate, and cover. Habitat preferences for seven species were in general agreement

with the habitat preferences in a published report on fishes of Minnesota. Hornyhead

chub, common carp, and common shiners were usually found in run macrohabitats that

were 55–85 cm deep. Largescale stoneroller, longnose dace, creek chub, and fathead

minnows were usually found in riffle macrohabitats that was 15–35 cm deep. Water

velocity for the riffle species was generally slower (15–65 cm/s) than for the run species

(65 cm/s). Each species has a specific preference with substrate (sand, gravel, boulders)

and cover (vegetation, boulders) or no cover. None of these species were found to prefer

pool macrohabitats. These findings indicate that the use of stream specific habitat

associations may be better than regional habitat association models when trying to

conserve certain species.

viii

Table of Contents

Page

Acknowledgements…………………………………………………………………...….iii

Abstract…………………………………………………………………………………....v

Table of Contents………………………………………………………………………..viii

List of Tables..………...…………………………………………………………………..x

List of Figures…………………………………………………………………………....xii

Chapter 1: Clarification of the Genus Campostoma in North Dakota and a novel

meristic technique………………………………………………………………………....1

Abstract…………………………………………………………………………………....1

Introduction………………………………………………………………………………..1

Materials and methods…………………………………………………………………….3

Results………………………………………………………………………………….….7

Discussion………………………………………………………………………………....9

Acknowledgements……………………………………………………………………....11

Literature Cited……………………...………………………………………………..….13

Chapter 2: Longitudinal Fish Assemblage in the Forest River, North

Dakota: A Test of Longitudinal Riverine Theory .……………………………….……...26

Abstract…………………………………………………………………………………..26

Introduction……………………………………………………………………………....27

Materials and methods…………………………………………………………………...30

Results……………………………………………………………………………...…….33

ix

Discussion……………………………………………………………………………......34

Acknowledgements…………………………………………………………………...….36

References…………………………………………………………………………...…...37

Chapter 3: Habitat Preferences of Select Fishes of the Forest River in North

Dakota……………………………………………………………………………………50

Summary…………………………………………………………………………….…...50

Introduction……………………………………………………………………………....50

Methods……………...…………………………………………………………………...52

Results……………………………………………………………………………...…….55

Discussion…………………………………………………………………………...…...59

Acknowledgements…………………………………………………………………...….63

References…………………………………………………………………………...…...64

x

List of Tables

Page

Table 1-1. Campostoma oligolepis scale count data for 2006 and 2007

survey with means and standard error (SE) of counts…………………………..........… 17

Table 1-2. Campostoma anomalum from South Dakota State University

fish collection scale count data with means and standard error (SE) of counts………….19

Table 1-3. Campostoma oligolepis scale count data from University of North

Dakota fish collection with means and standard error (SE) of counts .………………….20

Table 1-4. Campostoma oligolepis scale count data from University of

Minnesota Bell Museum with means and standard error (SE) of counts……..………....21

Table 2-1. Total individuals, Shannon‘s index (H’), and evenness (J’) values

for the 16 sampling locations on the Forest River…………………….…………………42

Table 2–2. List of families and associated species of fishes collected from the

Forest River of North Dakota……………………………………………………………43

Table 2-3. Percent similarity matrix for the 16 sites on the Forest River………………. 44

xi

Table 3-1. Total number of macrohabitats sampled in the Forest River, North

Dakota and the number of times the species of concern, invasive species, and other

selected species were present in those macrohabitats……………………………………69

Table 3-2. Dimensions of substrate categories and descriptions of cover

categories from Aadland (1993)…………………………………………………………70

Table 3-3. List of families and associated species of fishes collected from the

Forest River of North Dakota with total individuals and relative abundance for

each species shown………………………………………………………………………71

Table 3-4. Total number of sites in the Forest River, North Dakota where the

species of concern, invasive species, and other selected species were collected

and the optimum depth, velocity, substrate, and cover variables for that species……….72

xii

List of Figures

Page

Figure 1-1. Map of sites sampled in the Red River basin of North Dakota

for the 2006- 2007 survey with the Forest River and Elm River watersheds …………...22

Figure 1-2. Picture depicting the three circumferential scale counts…………………….23

Figure 1-3. Scale count frequency distributions for the University of North

Dakota collection, Bell Museum collection, and our current study……………………...24

Figure 1-4. Oblique and sum of oblique and lateral line scale count frequency

distributions for C. oligolepis and C. anomalum………………………………………...25

Figure 2-1. Map depicting the 16 study sites, the four dams impounding over 50

hectares of water, and the level III and level IV ecoregions along the Forest

River…………………………………………………………………………………...…45

Figure 2-2. Depiction of the species and family richness changes in a

downstream direction along the 16 study sites of the Forest River……………………...46

xiii

Figure 2-3. Two-dimensional nonmetric multidimensional scaling output for

the 16 sites along the Forest River depicting how sites that fall between the

same two dams, excluding the sites directly below a dam, grouped close to each

other ………………………………………………………………….………………….47

Figure 2-4. Two-dimensional nonmetric multidimensional scaling output for

the 16 sites along the Forest River depicting Level IV ecoregions……..……………….48

Figure 2-5. Two-dimensional nonmetric multidimensional scaling output for

the 16 sites along the Forest River depicting Level III ecoregions……...……………….49

Figure 3-1. Map depicting the 16 study sites and the four dams impounding

over 50 hectares of water along the Forest River……………………...………………...73

Figure 3-2. Sampling frequency histograms for depth, velocity, substrate and

cover……………………………………………………………………………………...74

Figure 3-3. Depth, velocity, substrate, and cover preference histograms for

hornyhead chub…………………………………………………………………………..75

Figure 3-4. Depth, velocity, substrate, and cover preference histograms for

largescale stoneroller…………………………………………………………………….76

xiv

Figure 3-5. Depth, velocity, substrate, and cover preference histograms for

common carp……………………………………………………………………………..77

Figure 3-6. Depth, velocity, substrate, and cover preference histograms for

longnose dace…………………………………………………………………………….78

Figure 3-7. Depth, velocity, substrate, and cover preference histograms for

common shiner…………………………………………………………………………...79

Figure 3-8. Depth, velocity, substrate, and cover preference histograms for

creek chub………………………………………………………………………………..80

Figure 3-9. Depth, velocity, substrate, and cover preference histograms for

fathead minnow………………………………………………………………………….81

1

Chapter 1

Clarification of the Genus Campostoma in North Dakota and a novel meristic technique

This chapter is in preparation for submission to the journal Copeia. There will be two co-

authors, Cari- Ann Hayer (2nd

author) and Charles R. Berry, Jr. (3rd author). It is

formatted following Copeia rules.

Abstract.—There are conflicting presence records for Central Stoneroller, Campostoma

anomalum, and the Largescale Stoneroller, C. oligolepis in North Dakota. We sampled

127 sites in the Red River basin of North Dakota to determine presence of Campostoma

anomalum, and C. oligolepis. We collected Campostoma at five sites. We inspected

voucher specimens from two museum collections that identified one or both Campostoma

species as being present. All Campostoma collected historically and by us were C.

oligolepis. Our new method of counting scales was more accurate in differentiating C.

oligolepis and C. anomalum than two traditional methods.

INTRODUCTION

Both Largescale Stoneroller, Campostoma oligolepis, and Central Stoneroller, C.

anomalum, may be present in North Dakota. These species have been collected in the

Lake Agassiz Plain ecoregion, which is in the Red River of the North (hereinafter Red

River) drainage basin that includes portions of North Dakota, Minnesota, and South

Dakota (Hankinson, 1929; Lee et al., 1980; Niemela et al., 1998; Koel and Peterka, 2003;

2

Aadland et al., unpubl.). Most North Dakota records are from the Forest River, a tributary

to the Red River (Fig. 1-1) and the extreme northwest range of the species (Trautman,

1981; Koel and Peterka, 2003). Campostoma anomalum was reported in the Forest River

by Feldmann (1963) and Copes and Tubb (1966).

The first reported C. oligolepis collection was from the Forest River in 1964 by

Copes and Tubb (1966). DeKrey (1998) reported C. oligolepis at three different sites on

the Forest River. One specimen of Campostoma oligolepis was collected in the Elm River

(North Dakota Department of Health, unpubl.; Fig. 1-1).

Campostoma oligolepis was first described by Hubbs and Green (1935) as a

subspecies of C. anomalum; it was classified as a species in 1966 (Pflieger, 1971).

Campostoma oligolepis and C. anomalum can be distinguished by minor, sometimes

overlapping morphometric differences (Burr and Smith, 1976). Genetic analysis revealed

differences between the two species (Buth and Burr, 1978; Blum et al., 2008). Blum et al.

(2008) organized the Campostoma genus by using DNA sequencing. They collected

individuals from 33 sites across the Campostoma range, but did not include the Red River

of the North drainage. In addition to morphological differences, the two species have

slightly different but often overlapping habitat associations. Both adults and juveniles of

C. anomalum and C. oligolepis are classified as ―slow-riffle guild‖ members in

Minnesota (Aadland, 1993). Campostoma anomalum is usually found in small riffles and

slower waters in headwaters (Burr and Smith, 1976; Rakocinski, 1984), whereas C.

oligolepis is usually found in fast, deep rocky riffles with coarse substrates (Mettee et al.,

1996). Campostoma anomalum is turbidity and silt tolerant (Clay, 1975; Lee et al., 1980;

3

Becker, 2001), whereas C. oligolepis is silt intolerant (Page and Burr, 1991; Becker,

2001). Niemela et al. (1998) refers to both Campostoma species as pioneer species, which

are the first to colonize headwater stream sections after desiccation, and species that tend

to predominate in unstable environments affected by anthropogenic stress. The

Campostoma species found in North Dakota are considered a species of conservation

priority. Their rarity is due to being at the edge of the range for these species.

The Campostoma status in North Dakota is uncertain. Records for C. oligolepis

and C. anomalum indicate both species occurring in the same waters but in different

years. The purpose of our study is to investigate the Campostoma species—anomalum,

oligolepis, or both—that were present historically and document the species which

currently inhabit Red River waters in North Dakota.

MATERIALS AND METHODS

Study area and field techniques.—This study was conducted from 2006 to 2007 in the

Red River basin in North Dakota (Fig. 1-1). From its formation at the Otter Tail River

and Bois de Sioux River confluence at Wahpeton, North Dakota the Red River forms the

border between North Dakota and Minnesota as it flows northward to the Canadian

border and on into Manitoba. Fifty-four thousand km2 in eastern North Dakota are

drained by the Red River (Niemela et al., 1998). The largest North Dakota tributary is the

Sheyenne River (Tornes et al., 1997). Eight other tributaries enter the Red River from

North Dakota and include the Wild Rice, Maple, Elm, Goose, Turtle, Forest, Park and

Pembina rivers.

4

Specimens were collected in 2006–2007; four sites were resampled within the

same year and six were resampled between years. This yielded 137 sampling occasions at

127 different sites in Red River tributaries and the Red River mainstem (Fig. 1-1). The

stream was visually assessed for stream macrohabitats prior to sampling. This ensured the

reach encompassed all major macrohabitats within the area. GPS coordinates were

recorded and digital photographs were taken for each site (Borgstrom, 2010).

Depending on habitat conditions, fish were collected at each site with one or a

combination of the following: seining, backpack electrofishing with a Smith-Root LR-24

backpack electrofishing unit, cloverleaf minnow traps, cloverleaf predator traps, or

cylindrical minnow traps. Wadeable sites were sampled for a minimum of ten mean

stream widths or until no new species were collected, whichever was greater. The

distance sampled at some sites was less due to not having permission to sample any more

of the river upstream, and the river being non-wadeable downstream. Sites on the Red

River mainstem were non-wadeable and were sampled with a Smith-Root electrofishing

boat during the last week of August, 2007. Boat electrofishing was conducted in a

downstream direction at each site for five 20 minute intervals.

At wadeable sites fishes were collected with bag seines (1.2 m deep, 9.5-mm²

knotless netting) in 4.6 m and 9.1 m lengths and stretched to cover as much stream

habitat as possible. Seining was used when there was minimal vegetation and boulders.

Seine hauls were conducted in a downstream direction and were made until no new

species were collected or until the water became non-wadeable. Backpack electrofishing

was used in habitats where large boulders or vegetation made seining inefficient.

5

Backpack electrofishing was conducted in an upstream direction and tested prior to

sampling to determine adequate settings, which changed depending on stream water

quality. Backpack electrofishing was also used to collect fish in block nets placed below

waters that were too swift to sample in an upstream direction. Cloverleaf minnow traps

(38.5 cm deep, three 47-cm diameter chambers, 13-mm opening between chambers, 6-

mm mesh), cloverleaf predator traps (47 cm deep, three 47-cm-diameter chambers, 50-

mm opening between chambers, 13-mm mesh) and cylindrical minnow traps (40.6 cm

long, 22-cm-diameter, 19-mm opening, 6-mm mesh) were set overnight in backwater,

deep-water and pool habitats.

After fishes were collected, they were transferred to a live well, identified,

counted and released except for unidentified individuals. Voucher specimens were taken

for each species. Fishes were preserved in a 10% formalin solution and taken to the

laboratory for identification.

Laboratory techniques.—Specimen fixation and preservation followed published

guidelines (Walsh and Meador, 1988). We identified and counted all fixed specimens.

Specimen identification and counts were verified by an ichthyologist. Vouchers are

stored in the South Dakota State University Department of Wildlife and Fisheries fish

collection. Other specimens previously collected by others and stored at the Bell Museum

(University of Minnesota) and the University of North Dakota were used to further

determine the Campostoma species collected in the drainage. We inspected the eight

Campostoma specimens from the Forest River, North Dakota that are held at the Bell

Museum. These eight specimens represented one sampling occasion each at two sites:

6

one from 1993, one from 1994. The Campostoma sp. from the Forest River (n=30) in the

University of North Dakota collection were inspected by others (Kelsch, Professor,

Biology, pers. comm.).

Campostoma species determination depends on scale counts (Burr and Smith,

1976). We conducted five types of scale counts (e.g., circumferential scales, scales above

the lateral line, scales below the lateral line, pre-dorsal scales, and lateral line scales;

Table 1-1) for all Campostoma sp. collected in 2006 and 2007, and for museum

specimens. Scale count methods were those of Hubbs and Lagler (2004) and Pflieger

(1997). When left side lateral line scale deformities were present we used the right side

lateral line scale count. Eighty-three total individuals were used for species

determination.

A combined lateral line scale count plus vertical circumferential scale count with

85 as a break point is 99.8% accurate in determining the species of Campostoma (Burr

and Smith, 1976). Campostoma oligolepis have a sum < 85 whereas C. anomalum have a

sum > 85 (Burr and Smith, 1976).

We used two conventional methods of counting circumferential scales (i.e.

vertical, and Pflieger) and one that we devised and termed the ―oblique‖ count. The

vertical count (Hubbs and Lagler, 2004) counts the scale rows crossed by a line

encompassing the widest point around the body (usually four complete scales anterior to

the dorsal fin origin; personal observation; Fig. 1-2A). This method results in C.

oligolepis having 31–36 scales around the body and C. anomalum having 39–46 (Hubbs

and Lagler, 2004). The Pflieger method (Pflieger 1997) works along the scale diagonals.

7

It starts at the first whole scale anterior to the dorsal fin origin and goes down and back

until the lateral line scale where the count turns forward and down until the ventral

midline (Fig. 1-2B). From here it follows the same pattern up the opposite side to the

starting scale counting each scale row crossed. Campostoma oligolepis usually has 32–39

scale rows around the body whereas C. anomalum usually has 40–55 (Pflieger, 1997).

We experienced difficulties associated with performing the conventional circumferential

scale counts as they pass through the pelvic girdle region.

The oblique count bypasses any difficulties with the scales in the region of the

pelvic girdle and was much easier to perform. The oblique count starts with the first

whole scale anterior to the dorsal fin origin, goes down and back on the diagonal until it

reaches the ventral midline, then comes forward and up back on the other side to the

starting scale (Fig. 1-2C). We conducted the same scale counts on C. anomalum held in

the South Dakota State University (SDSU) fish collection (n=20) to determine the

validity in using the oblique count to distinguish C. anomalum from C. oligolepis.

RESULTS

Current study collection.—We collected 53,069 fish of 52 species in the Red River

mainstem and nine tributaries (Borgstrom et al., unpubl). The Forest River is the only

drainage where Campostoma sp. were collected. Eleven Campostoma sp. were collected

from three sites on the Forest River in 2006; ten were vouchered for identification.

Thirty-five Campostoma sp. were collected from two sites on the Forest River in 2007.

Two sites where Campostoma sp. were collected in 2006 were revisited in 2007;

8

Campostoma sp. were not collected on the return visit. All Campostoma sp. specimens

collected from 2007 were vouchered for identification. Campostoma sp. were not

collected from the six sites on the Elm River.

The scale counts for the oblique method averaged 32.5 ± 0.24 (Table 1-1) for the

45 Campostoma sp. from the Forest River collected in 2006–2007. The conventional

scale counts (Table 1-1) are within published ranges (Burr and Smith, 1976) for C.

oligolepis. Scale count analysis identified all but one (060728_01C; Table 1-1) individual

captured during this study as C. oligolepis. The individual that fell outside the published

ranges had deformed lateral line scales on both sides which increased the scale number.

This is evident in frequency distribution data (Fig. 1-3). However, the predorsal (n=20)

and above lateral line (n=16) scale counts for this individual fell within the published

ranges for C. oligolepis (Burr and Smith, 1976). Therefore, we identified this individual

as C. oligolepis.

Most of the conventional scale counts for C. anomalum from the SDSU collection

fell within the published ranges (Burr and Smith, 1976) for C. anomalum (Table 1-2).

The mean above lateral line count of 17.7 fell just below the published range of 18–20

(Burr and Smith, 1976) and was the only count that was outside of published ranges. The

oblique count for C. anomalum (n=20) had a mean of 38.9 (S.E. = 0.59) compared to

32.5 (S.E. = 0.24) for the 45 C. oligolepis from the Forest River. However, the range of

the count for the two species overlapped somewhat (Fig. 1-4).

Historical collections.—Based on a combined lateral line and circumferential scale count

with 85 as a cut off, all 30 specimens in the University of North Dakota collection fell

9

within the published ranges (Burr and Smith, 1976) for C. oligolepis (Table 1-3). The

average scale counts for the Bell Museum holdings are within the published ranges (Burr

and Smith, 1976) for C. oligolepis (Table 1-4).

The frequency distributions for lateral line scale count, vertical circumferential

scale count, and the lateral line and vertical circumferential scale sum for each collection

are compared (Fig. 1-3). The Bell Museum data and data from our current study follow

the same general curve, but the circumferential and sum counts conducted on the

specimens held at the University of North Dakota are shifted lower by approximately

four, although they also follow the same general curve as the other counts. These lower

counts are still within published ranges for C. oligolepis.

DISCUSSION

All individuals captured during this study were C. oligolepis. Rakocinski (1980)

noted that C. anomalum and C. oligolepis can hybridize, and that hybrids could account

for as much as 4% of the Campostoma spp. population where C. anomalum and C.

oligolepis were sympatric. But the genetic differences between the two species are

maintained even in high hybridization areas. If hybridization between C. oligolepis and

C. anomalum is occurring in the Forest River, we should have collected and identified

some adult C. anomalum, but we did not.

With the lowest standard error (0.24) for circumferential scale counts, the oblique

scale count may be a valid method to aid in future Campostoma species identification.

This count was the easiest circumferential count to use as it avoids the pelvic fin insertion

10

area. This area oftentimes has either embedded scales or is scaleless following the initial

line taken to complete the count. The vertical and Pflieger methods cross through this

area. Therefore the oblique scale count may be a more reliable count. More research

should be conducted using these same scale counts on the other two Campostoma species

(C. ornatum and C. pauciradii).

If we used the combined lateral line and vertical circumferential scale count, we

would have misidentified one fish (2.22% [060728_01C; Table 1]) as C. anomalum. If

we had used the vertical circumferential count we would have misidentified ten fish

(22.22%) as C. anomalum. If we based our identification on the Pflieger circumferential

count we would have misidentified one specimen (2.22% [070807_01Q; Table 1]) as C.

anomalum. It is evident that no single count is the best to differentiate C. anomalum from

C. oligolepis; rather a combination of counts is best. If we used the combined lateral line

and oblique count with 84 as a division between the two species, we would not have

misidentified any individuals.

The oblique count has the lowest standard error of the three circumferential scale

counts for both C. oligolepis and C. anomalum (Tables 1-1, 1-2) and offers a valid

surrogate for the other circumferential scale counts. The oblique count alone does not

differentiate C. oligolepis from C. anomalum (Fig. 1-4), but using the sum of the oblique

and lateral line scale counts differentiates C. oligolepis from C. anomalum with a cutoff

value of 84 (Fig. 1-4) such that a value of 84 or greater indicates an individual is C.

anomalum and a value of <84 indicates an individual is C. oligolepis.

11

The 30 Campostoma sp. specimens at the University of North Dakota and

collected from the Red River basin were initially identified as C. anomalum. These

specimens were reviewed and determined to be C. oligolepis (Kelsch, Professor, Biology,

pers. comm.). The initial identification to C. anomalum is likely due to the collections

having taken place prior to C. oligolepis being elevated to the species rank in 1966

(Pflieger, 1971).

The eight Campostoma sp. individuals from the study area held by the Bell

Museum are C. oligolepis as they were originally identified. The correct initial

identification is understandable as these two samples were collected in 1993 and 1994,

more than twenty years after C. oligolepis was raised to species status. Through this

study, we are confident that the only Campostoma species collected—both presently and

historically—in the Red River basin in North Dakota is C. oligolepis. This represents the

northwestern most collections of C. oligolepis (Trautman, 1981; Koel and Peterka, 2003)

and a new major drainage basin in the range of this species.

The designation of C. oligolepis as a species of conservation priority in the state

of North Dakota is warranted. This species was only collected during five sampling

occasions at five different sites located along the Forest River from a total of 127

different sites throughout the entire Red River basin in North Dakota.

ACKNOWLEDGEMENTS

We thank D. Dorris, W. Bouska, S. Sindelar, J. Billings, E. Boyda, and C. Stearns

for their aid in sampling the streams and fishes. The investigation into historical

12

collections could not have been obtained without the help of the staff at the University of

North Dakota and the staff at the Bell Museum of Natural History. Further gratitude is

extended to J. Ladonski for his laboratory assistance in verifying fish identifications. Fish

were obtained legally under the North Dakota Game and Fish Department permitting

guidelines (permit # GNF02362977 and GNF02483803). Funding for this project was

provided in part by the North Dakota Game and Fish Department and South Dakota State

University.

13

LITERATURE CITED

Aadland, L. P. 1993. Stream habitat types: their fish assemblages and relationship to

flow. North American Journal of Fisheries Management 13:790–806.

Becker, G. C. 2001. Fishes of Wisconsin. The University of Wisconsin Press, Madison,

Wisconsin.

Blum, M. A., D. A. Neely, P. M. Harris, and R. L. Mayden. 2008. Molecular

systematics of the cyprinid genus Campostoma (Actinopterygii: Cypriniformes):

Disassociation between morphological and mitochondrial differentiation. Copeia

2008:360–369.

Borgstrom, L. J. 2010. Fish community assembly in the Forest River, North Dakota.

Masters thesis, South Dakota State University, Brookings, South Dakota.

Burr, B. M., and P. W. Smith. 1976. Status of the largescale stoneroller, Campostoma

oligolepis. Copeia 1976:521–531.

Buth, D. G., and B. M. Burr. 1978. Isozyme variability in the cyprinid genus

Campostoma. Copeia 1978:298–311.

Clay, W. M. 1975. The fishes of Kentucky. Kentucky Department of Fish and Wildlife

Resources. Frankfort, Kentucky.

Copes, F. A., and R. A. Tubb. 1966. Fishes of the Red River tributaries in North

Dakota. Contributions of the Institute for Ecological Studies number 1. University

of North Dakota, Grand Forks, ND.

14

DeKrey, D. C. 1998. A comparison of fish community structure in relation to habitat

variation in three North Dakota streams. Masters Thesis. University of North

Dakota, Grand Forks, ND

Feldmann, R. M. 1963. Distribution of fish in the Forest River of North Dakota.

Proceedings of the North Dakota Academy of Science 17: 11–19.

Hankinson, T. L. 1929. Fishes of North Dakota. Papers of the Michigan Academy of

Sciences, Arts, and Letters 10:439–460.

Hubbs, C. L., and C. W. Green. 1935. Two new subspecies of fish from Wisconsin.

Transactions of the Wisconsin Academy of Sciences, Arts and Letters 29:89-101.

Hubbs, C. L., and K. F. Lagler. 2004. Fishes of the Great Lakes Region. Revised

edition. University of Michigan, Ann Arbor, Michigan.

Koel, T. M., and J. J. Peterka. 2003. Stream fish communities and environmental

correlates in the Red River of the North, Minnesota and North Dakota.

Environmental Biology of Fishes 67:137–155.

Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R.

Stauffer. 1980. Atlas of North American Freshwater Fishes. Publication #1980-12

of the North Carolina Biological Survey, North Carolina State Museum of Natural

History, Raleigh, North Carolina

Mettee, M. F., P. E. O’Neil, and J. M. Pierson. 1996. Fishes of Alabama and the

Mobile basin. Oxmoor House, Inc., Birmingham, Alabama.

Niemela, S., E. Pearson, T. P. Simon, R. M. Goldstein, and P. A. Bailey. 1998.

Development of index of biotic integrity expectations for the Lake Agassiz Plain

15

Ecoregion. U.S. Environmental Protection Agency, Report 905/R-96-005, Chicago,

Illinois.

Page, L. M., and B. M. Burr. 1991. A field guide to freshwater fishes of North America

north of Mexico. Houghton Mifflin Company, New York, New York.

Pflieger, W. L. 1971. A distributional study of Missouri fishes. University of Kansas,

Lawrence, Kansas. Museum of Natural History, University of Kansas, Publication

20: 225-570.

Pflieger, W. L. 1997. The fishes of Missouri. Missouri Department of Conservation.

Jefferson City, Missouri.

Rakocinski, C. F. 1980. Hybridization and introgression between Campostoma

oligolepis and C. anomalum pullum (Cypriniformes: Cyprinidae). Copeia

1980:584–594.

Rakocinski, C. F. 1984. Aspects of reproductive isolation between Campostoma

oligolepis and Campostoma anomalum pullum (Cypriniformes: Cyprinidae) in

northern Illinois. American Midland Naturalist 112:138–145.

Tornes, L. H., M. E. Brigham, and D. L. Lorenz. 1997. Nutrients, suspended

sediment, and pesticides in streams in the Red River of the North basin, Minnesota,

North Dakota, and South Dakota, 1993-15. USGS Water-Resources Investigations

Report 97-4053: 70 pp.

Trautman, M. B. 1981. The Fishes of Ohio. Ohio State University Press, Columbus,

Ohio.

16

Walsh, S. J., and M. R. Meador. 1998. Guidelines for quality assurance and quality

control of fish taxonomic data collected as part of the National Water Quality

Assessment Program. US Geological Survey Water Resources Investigations

Report 98-4239.

17

Scale counts

Collection ID

Site

#

Above

lateral

line

Below

lateral

line

Pre-

dorsal

Lateral line Circum-

ferential

(oblique)

Circum-

ferential

(vertical)

Around

body

(Pflieger)

Lat +

Circum-

ferential

(v) L R

060712_A 17 13 15 18 45 43 32 34 35 79

060712_B 17 13 16 19 45 46 31 36 34 81

060712_C 17 13 14 19 43 46 31 35 35 78

060725_01A 58 15 16 20 46 44 34 35 37 81

060725_01B 58 13 16 19 44 44 30 36 36 80

060728_01A 71 15 15 18 46 44 34 37 37 83

060728_01B 71 17 15 19 44 45 35 35 38 79

060728_01C 71 16 15 20 49* 48* 34 41 39 90*

060728_01D 71 15 14 18 43 41 34 37 37 80

060728_01E 71 13 13 20 44 46* 31 35 36 79

070807_01A 761 14 15 21 45 47 32 35 37 80

070807_01B 761 13 15 19 46 46 32 33 34 79

070807_01C 761 13 15 20 43 43 31 36 37 79

070807_01D 761 16 15 20 45 44 36 39 39 84

070807_01E 761 15 16 20 48* 44 35 40 39 84

070807_01F 761 15 15 19 43* 44 34 36 36 80

070807_01G 761 14 14 19 44 45 32 33 34 77

070807_01H 761 14 14 19 44 43 33 36 39 80

070807_01I 761 14 15 18 44 43 31 35 36 79

070807_01J 761 14 15 18 43 43 33 33 34 76

070807_01K 761 14 14 18 42 43 34 35 35 77

070807_01L 761 14 16 19 46* 43 34 38 36 81

070807_01M 761 14 16 19 43 45 32 33 34 76

070807_01N 761 13 15 20 44 44 31 35 35 79

070807_01P 761 13 14 20 46* 43 31 33 34 76

070807_01Q 761 16 14 19 45 44 36 40 42 85

070807_01R 761 16 15 20 43 45 34 36 37 79

070807_01S 761 13 16 20 44 44 31 34 34 78

070807_01T 761 14 16 20 44 44 31 35 35 79

070807_01U 761 13 13 20 42 43 30 33 33 75

070807_01V 761 14 15 21 44 44 32 36 34 80

070807_01W 761 14 15 21 45 47 35 35 38 80

070807_01X 761 14 17 18 45 44 34 37 35 82

070807_01Y 761 13 16 19 44 44 33 36 37 80

070807_01Z 761 13 14 19 44 45 31 36 33 80

Table 1-1. Campostoma oligolepis scale count data for 2006 and 2007 survey of the Red

River of the North with means and standard error (SE) of counts.

* Scale deformities that resulted in smaller scales are included in this count which may bias the count high.

18

Scale counts

Collection ID

Site

#

Above

lateral

line

Below

lateral

line

Pre-

dorsal

Lateral line Circum-

ferential

(oblique)

Circum-

ferential

(vertical)

Around

body

(Pflieger)

Lat +

Circum-

ferential

(v) L R

070807_01AA 761 14 16 19 45* 44 34 39 39 83

070807_01AB 761 14 16 19 45 45 33 37 35 82

070807_01AC 761 14 16 19 43 43 32 34 35 77

070807_01AD 761 14 15 20 44 42 33 36 37 80

070807_01AE 761 13 15 20 45 46* 31 36 35 81

070807_01AF 761 13 14 18 44 44 31 37 36 81

070816_02A 768 14 14 20 45 45 33 35 34 80

070816_02B 768 14 15 18 43 41 31 34 35 77

070816_02C 768 13 15 20 44 43 31 35 35 79

070816_02D 768 13 14 18 43 44 31 34 35 77

Means (SE)

13.98

(0.15)

14.98

(0.13)

19.26

(0.13)

44.36

(0.20)

44.18

(0.22)

32.53

(0.24)

35.69

(0.29)

35.93

(0.29)

79.82

(0.40)

Table 1-1. Continued…

* Scale deformities that resulted in smaller scales are included in this count which may bias the count high.

19

Scale counts

Stream Yr

Above

lateral

line

Below

lateral

line

Pre-

dorsal

Lateral

line Oblique

Circum-

ferential

(vertical)

Around

body

(Pflieger)

Lat +

Circum-

ferential

(vertical)

Lat +

oblique

Clay Creek 2009 16 17 21 49 35 41 40 90 84

Monighan Creek 2009 18 17 23 50 39 45 46 95 89

Monighan Creek 2009 19 18 27 52 42 46 45 98 94

Whetstone Creek 2009 19 18 21 46 40 46 46 92 86

Deer Creek 2000 18 16 23 48 39 44 45 92 87

Deer Creek 2000 19 16 24 52 39 48 47 100 91

Gravel Creek 2000 17 17 24 48 36 44 44 92 84

Camp Creek 2000 16 17 23 51 36 40 46 91 87

Stray Horse Creek 2002 17 18 22 50 38 44 43 94 88

Turkey Ridge Creek 2000 16 15 23 51 36 40 39 91 87

Mahoaney Creek 1995 16 18 23 49 39 42 41 91 88

MN River Basin 1993 18 17 23 48 42 47 46 95 90

MN River Basin 1993 17 19 24 51 39 46 45 97 90

MN River Basin 1993 16 15 23 49 35 42 42 91 84

MN River Basin 1993 18 19 23 49 39 45 46 94 88

MN River Basin 1993 18 15 25 51 39 46 45 97 90

MN River Basin 1993 17 17 21 50 38 44 42 94 88

MN River Basin 1993 22 20 24 48 46 54 55 102 94

MN River Basin 1993 19 18 23 50 41 49 51 99 91

MN River Basin 1993 18 17 25 54 40 47 46 101 94

Mean (SE)

17.7

(0.33)

17.2

(0.30)

23.3

(.032)

49.8

(0.41)

38.9

(0.59)

45.0

(0.74)

45.0

(0.80)

94.8

(0.83)

88.7

(0.70)

Table 1-2. Campostoma anomalum from South Dakota State University fish collection

scale count data with means and standard error (SE) of counts.

20

ID # Year

Lateral

Line Circumferential Lat + Circ

V-4-1952 2240 1952 45 32 77

V-4-1952 2240 1952 44 33 77

V 4-1952-F-88 1952 47 33 80

V 4-1952-F-88 1952 44 32 76

V 4-1952-F-88 1952 44 34 78

V 4-1952-F-88 1952 47 32 79

V 4-1952-F-88 1952 44 30 74

V 4-1952-F-88 1952 47 31 78

IV-23-1962 1169 1962 41 33 74

IV-23-1962 1169 1962 43 32 75

II-15-64-F-23 34 1964 44 32 76

II-15-64-F-23 34 1964 45 31 76

II-15-64-F-23 34 1964 43 31 74

8-5-1977 F-989 1977 46 32 78

8-5-1977 F-989 1977 47 31 78

8-5-1977 F-989 1977 45 33 78

8-5-1977 F-989 1977 44 32 76

46 31 77

970617-F4-E 1997 43 31 74

970617-F4-E 1997 46 31 77

970606-F6-S 1997 41 32 73

970609-F5-E 1997 46 33 79

970714-F6-E 1997 45 31 76

970714-F6-E 1997 45 33 78

970714-F6-E 1997 47 31 78

970714-F6-E 1997 45 31 76

970714-F6-E 1997 42 33 75

970714-F6-E 1997 42 32 74

970606-F6-E 1997 43 30 73

970606-F6-E 1997 42 32 74

Means 44.43 31.83 76.27

(SE) (0.33) (0.18) (0.35)

Table 1-3. Campostoma oligolepis scale count data from University of North Dakota

fish collection (Kelsch, Professor, Biology, personal communication) with means and

standard error (SE) of counts.

21

Scale counts

Collection ID

Site

#

Above

lateral

line

Below

lateral

line

Pre-

dorsal

Lateral line

Circum-

ferential

(oblique)

Circum-

ferential

(vertical)

Around

body

(Pflieger)

Lat +

Circum-

ferential

(v) L R

26871 1 14 14 19 47 45 30 34 34 81

26871 1 13 14 18 45 47 31 35 35 82

26871 1 13 13 18 44 43 30 33 34 77

26871 1 13 15 17 46 44 30 33 32 79

27612 2 14 13 19 46 48* 33 34 35 82

27612 2 15 15 18 45 45 33 35 36 80

27612 2 13 14 18 44 43 30 32 32 76

27612 2 13 14 18 44 44 31 35 34 79

Means (SE)

13.50

(0.76)

14.00

(0.76)

18.13

(0.64)

45.13

(1.13)

44.88

(1.40)

31.00

(1.31)

33.88

(1.13)

34.00

(1.41)

79.50

(2.20)

Table 1-4. Campostoma oligolepis scale count data from University of Minnesota Bell

Museum with means and standard error (SE) of counts.

* Scale deformities that resulted in smaller scales are included in this count which may bias the count high.

22

Fig. 1-1. Map of sites sampled in the Red River basin of North Dakota for the 2006-

2007 survey with the Forest River and Elm River watersheds outlined in dashed lines.

The Forest River is indicated by the northern arrow. The Elm River is indicated by the

southern arrow. These two watersheds have historically been inhabited by Campostoma.

Figure 1-1. Map of sites sampled in the Red River basin of North Dakota for the 2006-

2007 survey.

23

Fig. 1-2. Picture depicting the three circumferential scale counts: A. vertical, B. Pflieger,

and C. oblique.

Figure 1-2. Picture depicting the three circumferential scale counts: A. oblique, B.

vertical, and C. Pflieger.

24

0

2

4

6

8

10

12

14

16

18

20

41 42 43 44 45 46 47 48 49

0

2

4

6

8

10

12

14

30 31 32 33 34 35 36 37 38 39 40 41

0

2

4

6

8

10

12

73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90

Tesky Bell Museum Current study

Fig. 1-3. Scale count frequency distributions for the University of North Dakota collection

(hollow bars), Bell Museum collection (gray bars), and our current study (black bars): A.

Lateral line scale count. B. Vertical circumferential scale count. C. Vertical

circumferential and lateral line scale count sum. An asterisk (*) indicates an individual

with deformed lateral line scales which increased the scale number.

Figure 1-3. Frequency distributions for the collections reviewed: A. Lateral line scale

count. B. Vertical circumferential scale count. C. Vertical circumferential and lateral line

A

A

B

B

C

C

*

*

25

Fig. 1-4. Oblique (A) and sum of oblique and lateral line (B) scale count frequency

distributions for C. oligolepis and C. anomalum using C. oligolepis from this study and

the Bell Museum at the University of Minnesota and C. anomalum from the South

Dakota State University fish collection. An asterisk (*) indicates an individual with

deformed lateral line scales which increased the scale number.

Figure 1-3. Frequency distributions for the collections reviewed: A. Lateral line scale

count. B. Vertical circumferential scale count. C. Vertical circumferential and lateral

line scale count sum. The differences in B and C may be attributable to the individual

conducting the scale counts.

0

2

4

6

8

10

12

14

16

18

30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46

0

2

4

6

8

10

12

72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 94

C. oligolepis C. anomalum

A

A

B

B

*

26

Chapter 2

Longitudinal Fish Assemblage in the Forest River, North Dakota:

A Test of Longitudinal Riverine Theory

This chapter is in preparation for submission to the journal Aquatic Ecology and was co-

authored with Charles R. Berry, Jr. (2nd

author). It is formatted following Aquatic

Ecology rules.

Abstract.— The Forest River of North Dakota provides a unique situation to assess both

environmental and anthropogenic affects on longitudinal fish assemblage because it flows

through several ecoregions and is divided by four dams (4.6–23.2 m high) impounding

over 50 hectares of water each. Sixteen sites were sampled using seining, backpack

electrofishing, and traps. Richness ranged among sites from 5–15 species and from 3–6

families. Shannon‘s Diversity Index (H’) ranged from 0.77–1.95 and evenness (J’) varied

from 0.24–0.81. Nonmetric multidimensional scaling was conducted using percent

similarity index values. Level IV ecoregions had no effect on the fish assemblage, but

level III ecoregions have an effect on the fish assemblage. Dams had an effect on the fish

assemblage at both scales. The sites directly below a dam were closer on the NMDS plot

to the upstream sites than were the sites further downstream of the dams or upstream of

the reservoirs. Results from the level IV ecoregion scale are supported by the serial

discontinuity concept, but not the process domains concept. Results from the level III

ecoregion scale analysis agree with predictions of both the serial discontinuity concept

27

and process domains concept. The fish assemblage in the Forest River is influenced by

the anthropogenic forces of dams at the level IV ecoregion scale. At the level III

ecoregion scale, the anthropogenic forces of dams and natural community breaks based

on ecoregions are both evident.

Introduction

―From headwaters to mouth, the physical variation within a river

system possesses a continuous gradient of physical conditions. This

gradient should elicit a series of responses within the constituent

populations…‖ Vannote et al. (1980).

This river continuum concept (RCC) statement by Vannote et al. (1980) provided a

synthesis of knowledge pertaining to the linkage of lotic ecosystems to the surrounding

terrestrial land and incorporating the energy cycling and biological community ecology

observed within the lotic system (Minshall et al. 1985) and was a framework that

described how a lotic system functions. Their concept sent researchers to the field for the

better part of the last three decades with a testable hypothesis. It was soon apparent that

the generality of the hypothesis was a handicap (Ward and Stanford 1983; Minshall et al.

1985; Pringle et al. 1988; Junk et al. 1989; Townsend 1989; Ward 1989; Allan 1995;

Montgomery 1999) usually because local controls dominated systemic controls on stream

biota (Vannote et al. 1980; Ward and Stanford 1983; Hughes et al. 1987; Pringle et al.

28

1988; Townsend 1989; Allan 1995; Montgomery 1999). However, the RCC continues to

serve as a useful conceptual framework (Allan 1995; Stanford and Ward 2001).

The suitability of the RCC has been tested in streams that drain agricultural

landscapes and prairie grasslands of the Great Plains. Prairie streams are inverted in

regards to riparian vegetation, stream temperature, and primary productivity when

compared to the prototypical RCC stream (Matthews 1988; Wiley et al. 1990). Light

availability is the driving force behind primary productivity in prairie streams (Wiley et

al. 1990). Hoagstrom et al. (2006) found that fish species composition did not support the

RCC in the Great Plains because species composition was driven by species replacements

rather than additions. A river in semiarid western South Dakota, in contrast to the RCC,

exhibited a random pattern or a pattern of no change in the biological and physical

properties, but a river in the sub-humid eastern South Dakota exhibits gradual and

continuous biological and physical changes as predicted by the RCC (Milewski 2001).

However, research on prairie stream ecology has been meager compared to that on

streams in other landscapes (Matthews 1988; Dodds et al. 2004).

Discontinuities caused by dams have been one of the major findings that have led to

a better understanding of the river continuum (Ward and Stanford 1983; Stanford and

Ward 2001). The Serial Discontinuity Concept (Ward and Stanford 1983; Stanford and

Ward 2001) applies to regulated rivers as a dam is viewed as a discontinuity in the river

continuum that resets the system back to a community similar to one upstream of the dam

in the unregulated reaches. The river does not reach the pre-dam state until sufficiently

downstream from the dam or after enough unaltered tributaries flow into it. Support for

29

this concept can be found from surveys of great rivers (Vinson 2001), and smaller rivers

(Tiemann et al. 2004). Dams alter water chemistry, river temperature regimes, substrate,

turbidity, and fish community (Hannan 1979; Holden 1979; Simons 1979). All of these

variables have distinct breaks at the dam interface, which represent discontinuities in the

river continuum.

The Process Domains Concept (Montgomery 1999) states that there may be breaks

in the downstream continuity of a river based on its geomorphic context, as the stream

travels through a variety of ecoregions. This takes into account the climate, geology and

topography of a stream section. These factors govern the various geomorphological

processes acting on the river, which gives the domain a certain physical habitat and

disturbance regime.

Land forms, processes, geomorphology, and biota can be grouped by ecoregions

(Hughes et al. 1987). Wisconsin ecoregions (Lyons 1996) and Oregon aquatic ecoregions

(Hughes et al. 1987) explain a great deal of variance in fish assemblages. Catchment-

scale habitat variables may account for more variation in fish assemblage than reach- or

site-scale variables (Gido et al. 2006). Where ecoregions abruptly change, so can streams

(Montgomery 1999; Seelbach et al. 2006). Ecoregions can be too large of a scale, and

streams can have breaks at a smaller scale such as where two similar tributaries converge

(Seelbach et al. 2006). This does not hold true for streams in the eastern and central

United States because changes in stream order (i.e. where two similar tributaries

converge) do not provide break points for fish communities (Matthews 1986).

30

The Forest River of North Dakota provides a unique situation to assess both

environmental (i.e. ecoregions) and anthropogenic affects (i.e. dams) on longitudinal fish

assemblage in a prairie stream as predicted by the RCC. It flows through two level III

ecoregions and is divided by four epilimnetic release dams (4.6–23.2 m high)

impounding over 50 hectares of water each. Our objective is to test the Serial

Discontinuity Concept and the Process Domains Concept as they apply to the Forest

River by analyzing fish assemblage data using nonmetric multidimensional analysis.

Materials and methods

Study area and field techniques.— This study was conducted at 16 sites from 1 August

2007 to 17 August 2007 on the Forest River of North Dakota (Fig. 2-1). The Forest River

flows 190-km while draining about 2300-km2. It flows through two level III ecoregions:

the Northern Glaciated Plains and the Lake Agassiz Plain and four level IV ecoregions:

Drift Plains, Glacial Lake Basin, Sand Deltas and Beach Ridges, and Saline Area (Fig. 2-

1; Bryce et al. 1998). About 95% of the native prairie in the Red River basin has been

converted for other uses (North Dakota Parks and Recreation Department, unpubl.). The

Forest River has been the focus of two previous studies in which we could find fish

species reports (Feldmann 1963; Woods 1971). We compared our species accounts to the

accounts from Feldmann (1963) and Woods (1971).

The river was visually assessed at each site to ensure the reach encompassed all

major macrohabitats (i.e. riffle, run, pool) within the area. GPS coordinates were

recorded and photographs were taken for each site (Borgstrom 2010).

31

Depending on habitat conditions within each reach fish were collected with one or

a combination of the following: seining, backpack electrofishing with a Smith-Root LR-

24 backpack electrofishing unit, cloverleaf minnow traps, cloverleaf predator traps, or

cylindrical minnow traps. We used multiple gears to maximize effort and likelihood all

species present would be collected. We sampled reaches ten mean stream widths in

length at each site or until no new species were collected, whichever was greater. One

site we were only able to sample a reach of seven mean stream widths in length due to

not having permission to sample any more upstream, and a reservoir downstream.

Fish were collected with bag seines (1.2 m deep, 9.5-mm² knotless netting) in

lengths of 4.6 m and 9.1 m and stretched to cover as much stream habitat as possible.

Seining was used when there was minimal vegetation and boulders. Seine hauls were

conducted in a downstream direction and made until no new species were collected or

until the water became non-wadeable.

Backpack electrofishing was used in habitats where large boulders or vegetation

made seining inefficient. Backpack electrofishing was conducted in an upstream direction

and tested prior to sampling to determine adequate settings, which changed depending on

stream water quality. Backpack electrofishing was also used to collect fish in block nets

placed below turbulent waters that were too swift to sample in an upstream direction.

Cloverleaf minnow traps (38.5 cm deep, three 47-cm diameter chambers, 13-mm

opening between chambers, 6-mm mesh), cloverleaf predator traps (47 cm deep, three

47-cm-diameter chambers, 50-mm opening between chambers, 13-mm mesh) and

32

cylindrical minnow traps (40.6 cm long, 22-cm-diameter, 19-mm opening, 6-mm mesh)

were set overnight in backwater, deep-water and pool habitats.

After fishes were collected, they were transferred to a live well, identified,

counted and released except for unidentified individuals. Voucher specimens were taken

for each species. Fishes were preserved in a 10% formalin solution and taken to the

laboratory for verification and identification.

Laboratory techniques.—Fixation in the field and specimen preservation in the lab

followed published guidelines (Walsh and Meador 1988). We identified and counted all

fixed specimens. Specimen identification and counts were verified by an ichthyologist.

Vouchers are stored in the South Dakota State University Department of Wildlife and

Fisheries fish collection.

For each site, we determined fish richness values, species diversity (H’), and

percent similarity between all site pairings. Shannon‘s Diversity Index was calculated

using the formula H’ = , where s = number of species and pi = the

proportion of the total sample represented by species i (Kwak and Peterson 2007).

Percent similarity was calculated using the formula Pjk = where Pjk = the

similarity between assemblages j and k, = relative abundance of species i in

assemblage j and = relative abundance of species i in assemblage k (Kwak and

Peterson 2007). We used percent similarity as a measure of community similarity

because it is robust and is insensitive to the number of individuals collected (Kwak and

Peterson 2007). Two-dimensional nonmetric multidimensional scaling (NMDS) was

conducted in Statistical Analysis Software (9.1) using percent similarity index values.

S

i

iei pp1

))(log(

),min( jiki pp

jip

kip

33

Results

Total individuals collected ranged between 54 and 8081 among sites (Table 2-1),

and a total of 17,998 individuals were collected, representing seven families and 25

species (Table 2-2). Species richness ranged from 5–15 among sites. The three most

upstream sites (1, 2, and 3) contained the fewest number of species with 7, 5, and 6

species respectively (Fig. 2-2). The greatest number of species collected (n = 15)

occurred at sites 9 and 11 in the middle reaches of the river (Fig. 2-2). The number of

families ranged from 3–6 with the fewest families (n = 3) collected at site 3 (Fig. 2-2) and

the greatest number of families (n = 6) collected at sites 1, 4, 9, and 11 (Fig. 2-2). Three

of these four sites (1, 4, and 9) are located immediately downstream of dams. There were

no apparent longitudinal trends in Shannon‘s Diversity Index (H’) which ranged from

0.77–1.95 and evenness (J’) which varied from 0.25–0.82 (Table 2-1). Table 2-3 shows

that percent similarity between sites varied from 1.1% similarity (paring sites 5 & 16) to

88.6% similarity (pairing sites 5 & 7).

The two-dimensional NMDS output (Fig. 2-3) yielded one cluster containing ten

sites on the left hand side. The two-dimensional NMDS output yielded a badness-of-fit

value (0.06) that is considered ―good‖ (Kruskal 1964) and was below the threshold value

of 0.15 (Kruskal and Wish 1984). Low values for badness-of-fit indicate a better fit

(Kruskal and Wish 1984) and zero is a ―perfect‖ fit (Kruskal 1964).

The three upstream sites (locations 1, 2, and 3) grouped close to one another on

the right hand side, and the three other sites, which occurred just beyond the plunge pools

of the dams, are scattered between these two groups (Fig. 2-3). This two-dimensional

34

output yielded site groupings based on dams (Fig. 2-3) but not level IV ecoregions (Fig.

2-4). Sites that were grouped close to each other are located within the same between-

dam reaches (Fig. 2-3), excluding the sites immediately below dams. The sites directly

below dams separated out closer to the upstream reaches than to the other groups (Fig. 2-

3). When the two-dimensional output was analyzed at the level III ecoregion scale, the

site groupings were closely related to ecoregions (Fig. 2-5).

Discussion

Temporal analysis of fish presence suggests that the fish assemblage has been

stable over 45 years, although data are meager. Woods (1971) collected 140 individuals

from six sites representing six families and 13 species. We collected all of the 13 species

that Woods (1971) collected. Two of the 23 species captured in 1963 (Feldmann 1963)

but not during this study were Ameiurus nebulosus, and Percopsis omiscomaycus. The

trout-perch, Percopsis omiscomaycus, is listed as a species of conservation priority for

North Dakota (Hagen et al. 2005). We did collect three species (Notropis dorsalis,

Lepomis macrochirus and Ictalurus punctatus) that had never been reported from the

drainage (Hankinson 1929; Feldman 1963; Woods 1971).

NMDS analysis indicates that at level IV ecoregions, the separation of groups is

based on the local control of dams (Fig. 2-3) and not systemic ecoregional processes (Fig.

2-4). Sites between two dams were grouped together, excluding the site immediately

below dams. The sites directly below dams separated out closer to the most upstream

35

sites than to the other sites. The serial discontinuity concept predicts this finding, but the

process domains concept does not.

When using the larger scale level III ecoregions as a function in the NMDS

analysis, the serial discontinuity concept becomes less evident. The sites that were

directly below a dam were closer to the upstream reaches than were the sites further

downstream of the dams or upstream of the reservoirs (Fig. 2-3). This upholds the serial

discontinuity concept, but the process domains concept is also upheld in this scenario due

to there being two main groups based on level III ecoregions (Fig. 2-5). The fish

assemblage in the Forest River is more strongly influenced by the anthropogenic forces

of dams at the level IV ecoregion scale than any natural community breaks that may

persist (Fig. 2-4), but when analyzed at the larger scale of level III ecoregions (Fig. 2-5),

the anthropogenic forces and natural community breaks are both evident.

Understanding multiple scales and a species relation to habitat at those scales are

important for the conservation of that species. The federally endangered Topeka shiner,

Notropis topeka, presence probability is related to stream condition and land-cover

variables at the valley segment (large) scale (Wall and Berry 2006). Whereas, at the reach

(fine) scale, Notropis topeka presence probability is related to habitat and community

variables (Wall and Berry 2006). Although, the addition of landscape variables negligibly

improved models predicting fish assemblage metrics and fish index of biotic integrity

compared to a model based on physical variables at the site-scale (Rowe et al. 2009).

This is due in part to landscape variables (systemic) influencing stream site (local)

36

variables which have a direct influence on stream biota and assemblages in Iowa (Rowe

et al. 2009).

In conclusion, nonmetric multidimensional scaling is a useful analysis tool for

investigating riverine theory. Furthermore, the importance of scale cannot be overlooked.

One riverine theory may hold up at multiple scales, whereas another theory may only

become evident when studied at a larger scale. The Forest River fish assemblage is

structured by both the serial discontinuity concept and the process domains concept at

differing scales. Future studies should focus on how the reservoirs themselves affect the

fish community assemblage and the contributions reservoir fishes make to the stream fish

community as a source of species and low water refugia.

Acknowledgements

We thank S. Sindelar, J. Billings, E. Boyda, and C. Stearns for their aid in

sampling the Forest River and associated fishes. We would also like to thank Cari-Ann

Hayer for her guidance throughout the sampling for this study. Further gratitude is

extended to J. Ladonski for his laboratory assistance in verifying fish identifications. We

are grateful for the person who suggested we use nonmetric multidimensional analysis to

determine the relationships between sites, Mark Fincel. All fish were obtained legally

under the permitting guidelines of the North Dakota Game and Fish Department (permit

# GNF02362977 and GNF02483803). Funding for this project was provided in part by

the North Dakota Game and Fish Department and South Dakota State University.

37

References

Allan JD (1995) The river continuum concept. In: Allan JD (ed) Stream Ecology:

Structure and Function of Running Waters. Chapman and Hall, London pp 276–281

Borgstrom LJ (2010) Fish community assembly in the Forest River, North Dakota.

Masters thesis, South Dakota State University

Bryce S, Omernik JM, Pater DE, Ulmer M, Schaar J, Freeouf J, Johnson R, Kuck P,

Azevedo SH (1998) Ecoregions of North Dakota and South Dakota. [Two sided

color poster with map, descriptive text, summary tables, and photographs.] Reston,

Virginia: United States Geological Survey (scale 1:1,500,000)

Dodds WK, Gido K, Whiles MR, Fritz KM, Matthews WJ (2004) Life on the edge: the

ecology of Great Plains prairie streams. Biosci 54:205–216

Feldmann RM (1963) Distribution of fish in the Forest River of North Dakota. Proc N D

Acad Sci 17: 11–19

Gido KB, Falke JA, Oakes RM, Hase KJ (2006) Fish-habitat relations across spatial

scales in prairie streams. In: Hughes RM, Wang L, Seelbach PW (eds) Landscape

influences on stream habitats and biological assemblages. American Fisheries

Society, Symposium 48, Bethesda, Maryland, pp 265–285

Hagen SK, Isakson PT, Dyke SR (2005) North Dakota Comprehensive Wildlife

Conservation Strategy. North Dakota Game and Fish Department. Bismarck, ND.

Hankinson TL (1929) Fishes of North Dakota. Pap Mich Acad Sci Arts Lett 10:439–460.

38

Hannan HH (1979) Chemical modifications in reservoir-regulated streams. In: Ward JV,

Stanford JA (eds) The Ecology of Regulated Streams. Plenum Press, New York, pp

75–94

Hoagstrom CW, Wall SS, Duehr JP, Berry CR, Jr (2006) River size and fish assemblages

in southwestern South Dakota. Gt Plains Res 16:117–126

Holden PB (1979) Ecology of riverine fishes in regulated stream systems with emphasis

on the Colorado River. In: Ward JV and Stanford JA (eds) The Ecology of

Regulated Streams. Plenum Press, New York, pp 57–74

Hughes RM, Rexstad E, Bond CE (1987) The relationship of aquatic ecoregions, river

basins and physiographic provinces to the ichthyogeographic regions of Oregon.

Copeia 1987:423–432.

Junk WJ, Bayley PB, Sparks RE (1989) the flood pulse concept in river-floodplain

systems. In: Dodge DP (ed) Proceedings of the International Large River

Symposium, Can Spec Pub Fish Aquat Sci, pp 110–127

Kruskal JB (1964) Multidimensional scaling by optimizing goodness of fit to a nonmetric

hypothesis. Psychometrika 29:1–27

Kruskal JB, Wish M (1978) Multidimensional Scaling. Sage University Paper series on

Quantitative Applications in the Social Sciences, series number 07-011. Beverly

Hills, California: Sage Publications

Kwak TJ, Peterson JT (2007) Community Indices, Parameters, and Comparisons. In: Guy

CS, Brown ML (eds) Analysis and Interpretation of Freshwater Fisheries Data.

American Fisheries Society, Bethesda, Maryland, pp 677–763

39

Lyons J (1996) Patterns in the species composition of fish assemblages among Wisconsin

streams. Envrion Biol Fishes 45:329–341

Matthews WJ (1986) Fish faunal ‗breaks‘ and stream order in the eastern and central

United States. Environ Biol of Fishes 17:81–92

Matthews WJ (1988) North American prairie streams as systems for ecological study. J N

Am Benthol Soc 7:387–409

Milewski CL (2001) Local and systemic controls on fish and fish habitat in South Dakota

rivers and streams: implications for management. Ph.D. dissertation, South Dakota

State University

Minshall GW, Cummins KW, Peterson RC, Cushing CE, Bruns DA, Sedell JR, Vannote

RL (1985) Developments in stream ecosystem theory. Can J Fish Aquat Sci

42:1045–1055

Montgomery DR (1999) Process domains and the river continuum. J Am Water Resour

Assoc 35:397–410

Pringle CM, Naiman RJ, Bretschko G, Karr JR, Oswood MW, Webster JR, Welcomme

RL, Winterbourn MJ (1988) Patch dynamics in lotic systems: the stream as a

mosaic. J N Am Benthol Soc 7:503–524

Rowe DC, Pierce CL, Wilton TF (2009) Physical habitat and fish assemblage

relationships with landscape variables at multiple spatial scales in wadeable Iowa

streams. N Am J Fish Manag 29:1333–1351

Seelbach PW, Wiley MJ, Baker ME, Wehrly KE (2006) Initial classification of river

valley segments across Michigan‘s lower peninsula. In: Hughes RM, Wang L,

40

Seelbach PW (eds) Landscape influences on stream habitats and biological

assemblages. American Fisheries Society, Symposium 48, Bethesda, Maryland, pp

25–48

Simons DB (1979) Effects of stream regulation on channel morphology. In: Ward JV

Stanford JA (eds) The Ecology of Regulated Streams. Plenum Press, New York, pp

95–111

Stanford JA, Ward JV (2001) Revisiting the serial discontinuity concept. Regul Rivers:

Res Manag 17:303–310

Tiemann JS, Gillette DP, Wildhaber ML, Edds DR (2004) Effects of lowhead dams on

riffle-dwelling fishes and macroinvertebrates in a Midwestern river. Trans Am Fish

Soc 133:705–717

Townsend CR (1989) The patch dynamics concept of stream community ecology. J N

Am Benthol Soc 8:36–50

Vannote RL, Minshall GW, Cummins KW, Sedell JR, Cushing CE (1980) The river

continuum concept. Can J Fish Aquat Sci 37:130–137

Vinson MR (2001) Long-term dynamics of an invertebrate assemblage downstream from

a large dam. Ecol Appl 11:711–730

Wall SS, Berry CR, Jr (2006) The importance of multiscale habitat relations and biotic

associations to the conservation of an endangered fish species, the Topeka shiner.

In: Hughes RM, Wang L, Seelbach PW (eds) Landscape influences on stream

habitats and biological assemblages. American Fisheries Society, Symposium 48,

Bethesda, Maryland, pp 305–322

41

Walsh SJ, Meador MR (1998) Guidelines for quality assurance and quality control of fish

taxonomic data collected as part of the National Water Quality Assessment

Program. US Geological Survey Water Resources Investigations Report 98-4239

Ward JV (1989) The four-dimensional nature of lotic ecosystems. J N Am Benthol Soc

8:2–8

Ward JV, Stanford JA (1983) The serial discontinuity concept of lotic ecosystems. In:

Fontaine TD, Bartell SM (eds) Dynamics of lotic ecosystems. Ann Arbor Scientific

Publishers, Ann Arbor, Michigan, pp 29–42

Wiley MJ, Osborne LL, Larimore RW (1990) Longitudinal structure of an agricultural

prairie river system and its relationship to current stream ecosystem theory. Can J

Fish Aquat Sci 47:373–384

Woods CE (1971) Helminth parasites of fishes from the Forest River, North Dakota. Am

Midl Nat 86:212–215

42

Location Dam

Total

individuals

Shannon's index

(H’)

Evenness

(J’)

1 Yes 8081 0.77 0.40

2 No 54 1.11 0.69

3 No 519 0.45 0.25

4 Yes 87 1.63 0.74

5 No 376 1.46 0.59

6 No 269 1.79 0.72

7 No 256 1.34 0.56

8 No 158 1.96 0.82

9 Yes 1269 1.82 0.67

10 No 883 1.39 0.67

11 No 2166 1.67 0.62

12 No 1633 1.30 0.52

13 No 681 1.25 0.57

14 No 737 1.52 0.61

15 No 466 1.70 0.77

16 Yes 363 1.37 0.66

Table 2-1. Total individuals, Shannon‘s index (H’), and evenness (J’) values for the 16

sampling locations on the Forest River and whether or not they fell downstream of dams.

Locations correspond to the downstream direction (1 is most upstream location).

Table 2-2. Total individual, Shannon‘s index (H‘), and evenness (J’) values for the 16

sampling locations on the Forest River. A yes (Y) for dam indicates the location was

located directly downstream from a dam outlet. Locations correspond to the downstream

direction (1 is most upstream location).

43

Family Species Common name # of Individuals

Esocidae Esox lucius northern pike 53

Cyprinidae Campostoma oligolepis largescale stoneroller 35

Cyprinella spiloptera spotfin shiner 15

Cyprinus carpio common carp 134

Luxilus cornutus common shiner 3715

Nocomis biguttatus hornyhead chub 346

Notropis dorsalis bigmouth shiner 1093

Pimephales notatus bluntnose minnow 1170

Pimephales promelas fathead minnow 6403

Rhinichthys cataractae longnose dace 143

Rhinichthys obtusus western blacknose dace 123

Semotilus atromaculatus creek chub 1193

Catostomidae Carpiodes cyprinus quillback 7

Catostomus commersonii white sucker 2304

Ictaluridae Ameiurus melas black bullhead 26

Ictalurus punctatus channel catfish 1

Noturus gyrinus tadpole madtom 206

Gasterosteidae Culaea inconstans brook stickleback 2

Centrarchidae Ambloplites rupestris rock bass 4

Lepomis macrochirus bluegill 577

Pomoxis nigromaculatus black crappie 5

Percidae Etheostoma nigrum johnny darter 284

Perca flavescens yellow perch 89

Percina maculata blackside darter 69

Sander vitreus walleye 1

Table 2–2. List of families and associated species of fishes collected from the Forest

River of North Dakota.

Table 2–1. List of families and associated species of fishes collected from the Forest

River of North Dakota.

44

16 100

15 100

6.34

14 100

48.7

8

6.34

13 100

49.7

5

68.9

7

6.34

12 100

77.3

9

67.5

6

62.5

2

6.34

11 100

67.4

6

76.9

1

43.7

5

68.6

5

1.75

10 100

68.7

4

56.2

7

60.0

1

31.6

4

74.2

8

1.81

9 100

18.6

5

29.6

4

18.2

1

21.7

5

29.0

4

26.7

0

9.60

8 100

32.7

7

67.4

1

65.1

8

49.6

9

55.7

3

43.9

1

67.7

5

1.10

7 100

62.5

0

28.6

1

62.3

1

81.6

5

69.2

8

88.3

5

45.1

3

65.8

5

4.62

6 100

55.3

8

60.9

0

41.0

5

52.3

6

59.9

2

47.1

3

46.4

6

40.0

2

48.6

9

1.47

5 100

59.1

9

88.6

3

64.1

3

33.1

4

64.9

5

83.9

0

66.8

0

84.2

9

40.0

5

61.8

1

1.10

4 100

19.4

9

18.3

2

12.8

0

19.9

5

34.2

4

9.47

14.2

5

4.10

9.84

8.34

14.8

2

8.05

3 100

10.4

0

9.16

1.13

6.64

7.71

9.39

3.56

6.16

1.98

6.55

5.68

7.37

42.1

5

2 100

30.3

9

38.4

4

12.5

0

11.8

5

12..8

9

19.5

3

26.4

0

3.28

10.7

6

5.33

8.52

14.5

2

15.6

7

21.4

7

1 100

48.8

7

78.0

4

21.2

2

10.1

7

8.02

12.5

4

15.8

5

20.3

3

2.63

10.2

2

5.05

8.52

12.3

2

15.2

6

43.2

5

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

Loc

atio

n

Tab

le 2

-3. P

erce

nt s

imila

rity

mat

rix

for

the

16 s

ites

on th

e F

ores

t Riv

er. L

ocat

ion

1 is

the

mos

t ups

trea

m s

ite a

nd 1

6 is

the

mos

t

dow

nstr

eam

site

.

Location

45

Fig. 2-1. Map depicting the 16 study sites, the four dams (4.6–23.2 m high) impounding

over 50 hectares of water, and the level III and IV ecoregions (Bryce et al. 1998) along

the Forest River. Sites are numbered in a downstream manner with site one being the

most headwater site sampled, and site 16 being the most downstream site sampled. The

Drift Plain is a level IV ecoregion that is part of the Northern Glaciated Plains level III

ecoregion. The remaining level IV ecoregions are part of the larger Lake Agassiz Plain

level III ecoregion.

Fig. 2-3. Map depicting the 16 study sites, the 4 large dams, and the level IV ecoregions

(Bryce et al. 1998) along the Forest River.

46

Fig. 2-2. Depiction of the species and family richness changes in a downstream direction

along the 16 study sites of the Forest River. Sites immediately downstream of a dam are

marked with an arrow.

Fig. 2-4. Depiction of the species and family richness changes in a downstream direction

along the 16 study sites of the Forest River.

0

2

4

6

8

10

12

14

16

0 2 4 6 8 10 12 14 16 18

Ric

hn

ess

Site

Number of species

Number of families

47

Fig. 2-3. Two-dimensional nonmetric multidimensional scaling (NMDS) output for the

16 sites along the Forest River. Sites in the right-hand group are all sites directly below

dams and two sites below the first dam in the upstream portion of the watershed. A circle

( ) indicates a site in the upstream portion of the watershed below the first dam. A

diamond ( ) indicates a site below the second dam. And a hollow circle ( ) indicates a

site below the third dam. A hollow square ( ) indicates a site that occurred directly

downstream from a dam.

48

Fig. 2-4. Two-dimensional nonmetric multidimensional scaling output for the 16 sites

along the Forest River depicting Level IV ecoregions. A circle ( or ) indicates a site

in the Drift Plains. A diamond ( ) indicates a site in the Glacial Lake Basin. A star ( or

) indicates a site in the Sand Deltas and Beach Ridges. A square ( ) indicates a site

in the Saline Area. A hollow shape indicates a site that occurred directly downstream

from a dam. A solid shape indicates a site that is not below a dam.

49

Fig. 2-5. Two-dimensional nonmetric multidimensional scaling output for the 16 sites

along the Forest River depicting Level III ecoregions. A square ( or ) indicates a site

in the Northern Glaciated Plains. A circle ( or ) indicates a site in the Lake Agassiz

Plain. A hollow shape indicates a site that occurred directly downstream from a dam. A

solid shape indicates a site that is not below a dam.

50

Chapter 3

Habitat Preferences of Select Fishes of the Forest River in North Dakota

This chapter is in preparation for submission to the journal Freshwater Biology and was

co-authored with Charles R. Berry, Jr. (2nd

author) It is formatted following

Freshwater Biology rules.

Summary—

1. To better understand fish habitat preferences in the Forest River of North

Dakota, we collected fish data at 16 sites each with four macrohabitats (pool,

riffle, run, backwater) that were characterized for depth, velocity, substrate, and

cover.

2. Habitat preferences for seven species were in general agreement with published

data from Minnesota rivers. Two species agreed wholly, four species varied on

one habitat variable, commonly substrate or velocity, and one species varied on

two variables. Combining these habitat preferences with other knowledge can

aid in better conservation of the habitat necessary for these species.

Introduction

The demands on our limited freshwater resources are increasing as the global

population increases (Vörösmarty, 2000). Some regions have such high demands for

water (i.e. irrigation, industrial and domestic use, reservoirs) that some rivers, like the

Colorado River, USA, no longer flow to their mouth (Mueller et al., 2005). This demand

51

impacts the fish assemblage by changing the thermal regime and hydrologic cycle as well

as permitting nonnative species to become established (Mueller et al., 2005). Physical

habitat characteristics influence fish occurrence and abundance, although the associations

are poorly understood for many species (Hubert & Rahel, 1989). Recognizing habitat use

patterns of stream fishes is a major goal of fisheries biologists because this information

helps to conserve species as river ecology is altered and lotic ecosystems change.

To better predict the responses of fish communities to the demands for freshwater,

fisheries biologists have used a number of assessment techniques such as the habitat guild

approach (Aadland, 1993), the community approach (Gerhard et al., 2004), and native

and introduced groups (Gido & Propst, 1999). Another technique used to better predict a

fish community response to the increasing and varying demands on freshwater resources

is to quantify habitat use patterns of individual species that are considered rare,

threatened, or endangered (Converse et al., 1998; Dunham et al., 2003; Dieterman &

Galat, 2004; Grabowski & Isely; 2006; Hoagstrom et al., 2008). By trying to conserve

these less common species, the more common species will also be conserved.

Fish communities in Great Plains lotic systems are strongly influenced by abiotic

factors (Power et al., 1988). Fish distribution is related to instream habitat and

environmental conditions (Walters et al., 2003). Different flow levels (Kessler et al.,

1995; Braaten & Berry, 1997) and seasonal and daily variations in habitat use

(Grabowski & Isely, 2006) can alter fish distribution and abundance.

Seven species of conservation priority in North Dakota have been documented from

the Forest River (Kelsch, Alm & Tesky, unpubl.) a tributary to the Red River of the

52

North. An aquatic nuisance species, the non-native common carp (Cyprinus carpio

Linnaeus, 1758) has been reported from the Forest River (Kelsch, Alm & Tesky,

unpubl.). This is the only non-native fish in North Dakota that is detrimental to other

species (Panek, 1987). Management of both rare and exotic fishes depends on an

understanding of their habitat associations (Hagen et al., 2005)

The object of this study was to investigate the habitat preferences of the species of

conservation priority (Hagen et al., 2005), invasive species, and selected abundant

species in the Forest River. The findings also have implications for the management of

riverine fishes in other streams in the northern Great Plains agricultural setting.

Methods

Study area and field techniques.— This study was conducted at 16 sites from 1 August

2007 to 17 August 2007 on the Forest River of North Dakota (Fig. 3-1). The Forest River

flows 190-km while draining about 2300-km2 of the North Dakota portion of the Red

River basin. The Forest River is dominated by sand substrates and runs (Table 3-1; Fig.

3-2).

Sixteen sites from headwaters to terminus were chosen to represent river

conditions in major ecoregions and above and below four epilimnetic release dams (4.6–

23.2 m high) impounding over 50 hectares of water each. The river was visually assessed

at each site to ensure the reach encompassed all major macrohabitats (i.e. riffle, run, pool)

for that river segment. GPS coordinates were recorded and photographs were taken for

each site (Borgstrom, 2010).

53

Fish were collected with one or a combination of the following: seining, backpack

electrofishing with a Smith-Root LR-24 backpack electrofishing unit, cloverleaf minnow

traps, cloverleaf predator traps, or cylindrical minnow traps. We used multiple gears at a

site but only one gear per macrohabitat to maximize effort and likelihood all species

present would be collected depending on habitat conditions within each macrohabitat.

Fish were collected with bag seines (1.2 m deep, 9.5-mm² knotless netting) in two

lengths: 4.6 m and 9.1 m and stretched to cover as much stream habitat as possible. Seine

hauls were made to encompass a macrohabitat unit and conducted in a downstream

direction. Seining was used when there was minimal vegetation and boulders.

Backpack electrofishing was used in habitats where large boulders or vegetation

made seining inefficient. Backpack electrofishing was conducted in an upstream direction

and tested prior to sampling to determine adequate settings, which changed depending on

stream water quality. Backpack electrofishing was also used to collect fish in block nets

placed below turbulent waters that were too swift to sample in an upstream direction.

Cloverleaf minnow traps (38.5 cm deep, three 47-cm diameter chambers, 13-mm

opening between chambers, 6-mm mesh), cloverleaf predator traps (47 cm deep, three

47-cm-diameter chambers, 50-mm opening between chambers, 13-mm mesh) and

cylindrical minnow traps (40.6 cm long, 22-cm-diameter, 19-mm opening, 6-mm mesh)

were set overnight in backwater, deep-water and pool habitats.

After fishes were collected, they were transferred to a live well, identified,

counted and released except for unidentified individuals. Voucher specimens were taken

54

for each species. Fishes were preserved in a 10% formalin solution and taken to the

laboratory for verification and identification.

Once the reach was sampled for fishes by macrohabitats, we then measured the

habitat variables for each macrohabitat. Depth and velocity were collected at three points

representative of the macrohabitat using a top setting wading staff. Velocity was

determined using a Marsh-McBirney model 2000 flow meter (Frederick, Maryland,

USA) and taking the average of three cycles at 60% depth. Substrate and cover were

recorded at each velocity and depth location with classifications as described (Aadland,

1993; Table 3-2).

Laboratory techniques.—Fixation in the field and specimen preservation in the lab

followed published guidelines (Walsh & Meador, 1998). We identified and counted all

fixed specimens. Specimen identification and counts were verified by an ichthyologist.

Vouchers are stored in the South Dakota State University Department of Wildlife and

Fisheries fish collection.

Data analysis.— Data analysis follows Aadland & Kuitunen (2006) and is based on the

premise fish do not avoid cover. Fish with higher densities in areas of no cover are

assumed to be there based on other variables, and not the lack of cover. The preference

value for no cover is then assigned to all covers that have a lower preference value. An

example is the fathead minnow (Pimephales promelas) which has a preference value of

1.00 for edge, 0.40 for vegetation, 0.18 for boulder, and 0.05 for no cover. The

remaining cover types had preferences less than 0.05 but were assigned the preference

value of 0.05.

55

Habitat preference values were calculated for velocity, depth, substrate, and

cover. Depth and velocity variables were broken into 10-cm intervals with the midpoint

as a class value (e.g., 5-cm, 15-cm, etc.). The velocity class also contained a zero class as

some backwaters had no flow. The number of habitat measures within each habitat

interval were counted. This was the available habitat. The total number of individuals

sampled within each interval was summed to get the habitat use. Habitat preference was

determined by dividing habitat use for each interval by the habitat available for the same

interval. Preference values were determined by standardizing the habitat preference by

dividing by the maximum habitat preference, thus yielding a 0.0 to 1.0 scale, with 0.0

being the least preferred habitat and 1.0 being the most preferred habitat. Data was

visually inspected through the use of histograms and compared to values for the same

species found in Minnesota (Aadland & Kuitunen, 2006). Some of the classes for each

habitat variable had low sampling occurrences (Fig. 3-2), and could skew the visual

interpretation without taking into account the low sample size.

Results

We collected 17,998 individuals among seven families (Table 3-3). Ten

backwaters (7.3%), 13 pools (9.5%), 16 riffles (11.7%), and 98 runs (71.5%) were

sampled. Table 3-1 shows three species are run species, but these three most prevalent

species were found in macrohabitats in proportion to the macrohabitat occurrence (Table

3-1).

56

Depth classes ranged from 15 cm to 115 cm with a mode in the 35-cm class

(n=31). The velocities sampled were as low as 0 cm/s and extended up to the 65 cm/s

class (Fig. 3-2). The mode for velocities was the 15 cm/s class (n=43). Bedrock was the

only substrate type not sampled and was excluded from all analyses (Fig. 3-2). The most

abundant substrate sampled was sand (n=64). Flotsam was not sampled as a cover type

and was excluded from all analyses (Fig. 3-2). The most commonly sampled cover class

was ―none‖ (n=52; Fig. 3-2).

Two species of conservation priority were collected from the Forest River:

hornyhead chub (Nocomis biguttatus (Kirtland, 1840)) and largescale stoneroller

(Campostoma oligolepis Hubbs & Greene, 1935). Hornyhead chubs were collected from

63 macrohabitats at 12 sites (46%; Tables 3-1, 3-4): three backwaters (30% of all

backwaters sampled), seven pools (54% of all pools sampled), nine riffles (56% of all

riffles sampled), and 44 runs (45% of all runs sampled). They exhibited a preference for

deeper water with the 75-cm class being preferred (Fig. 3-3A). Hornyhead chub showed a

preference for faster flowing waters with the optimum being the 65 cm/s class (Fig. 3-

3B). Gravel was the preferred substrate for hornyhead chub (Fig. 3-3C) followed by

small boulder, large boulder, and sand with preference values of 0.64, 0.50, 0.42

respectively. Vegetation was the preferred cover type for hornyhead chub (Fig. 3-3D).

Largescale stonerollers were collected at five macrohabitats (3.6%) at two sites

(Tables 3-1, 3-4): one pool (8%), three riffles (19%), and one run (1%). Largescale

stoneroller exhibited a preference for shallower water with the 35-cm class being

preferred (Fig. 3-4A). They also preferred faster water with the 65-cm/s class being

57

preferred followed by the 55-cm/s class with a preference value of 0.03 (Fig. 3-4B). They

exhibited the highest preference for large boulder substrates and small boulder substrates

had a preference value of 0.22 (Fig. 3-4C). They showed highest preference for boulder

cover types (Fig. 3-4D).

We collected common carp at four macrohabitats at one site (2.9%; Tables 3-1, 3-

4). All of these macrohabitats were at the same downstream site below a dam. Common

carp were sampled in one backwater (10%) and three runs (3%). They showed a

preference for mid depth ranges and preferred the 55-cm class (Fig. 3-5A). They also

exhibited a preference for slower water with the 15-cm/s class being preferred followed

by the 5-cm/s class with a preference value of 0.25 (Fig. 3-5B). Sand was the preferred

substrate for common carp followed by silt (preference value of 0.35; Fig. 3-5C). Carp

were most commonly sampled from habitats without cover (Fig. 3-5D).

Longnose dace, Rhinichthys cataractae (Valenciennes, 1842), were found at 13

macrohabitats (9.5%; Tables 3-1, 3-4) amongst seven sites. They were located in two

pools (15%), six riffles (38%), and five runs (5%). Longnose dace exhibited a preference

for shallow waters with the three most preferred being the 15-, 25-, and 35-cm classes

with preference values of 0.12, 0.35 and 1.00 respectively (Fig. 3-6A). The velocity

preferred by longnose dace was the 65-cm/s class (Fig. 3-6B). They showed a high

preference for large boulder substrates followed by small boulder substrates with a

preference value of 0.28 (Fig. 3-6C). This boulder preference is further evident because

boulder is the preferred cover (Fig. 3-6D).

58

The common shiner, Luxilus cornutus (Mitchill, 1817), was collected in 88 of the

137 sampled macrohabitats (64.2%; Tables 3-1, 3-4) and 11 sites. This species was the

most numerous species in the Forest River. They were collected in two backwaters

(20%), six pools (46%), nine riffles (56%), and 71 runs (72%). They elicited a preference

for greater depths with the 75-cm class being preferred and the 85-cm class having a

preference value of 0.73 (Fig. 3-7A). The preferred velocity for common shiner was the

65-cm/s class (Fig. 3-7B). Gravel was the preferred substrate followed by sand and silt

with preference values of 0.87 and 0.61 respectively (Fig. 3-7C). The cover where

common shiner was sampled most was none (Fig. 3-7D).

Creek chub, Semotilus atromaculatus (Mitchill, 1818), were collected at 73

macrohabitats (53.3%; Tables 3-1, 3-4) and 14 sites. We sampled creek chub from seven

backwaters (70%), seven pools (54%), ten riffles (63%), and 49 runs (50%). They exhibit

a preference for shallower waters with the 25-cm class being preferred (Fig. 3-8A). Creek

chub display a preference for moderate flows with 35 cm/s being preferred (Fig. 3-8B).

They also show a preference for moderate sized substrate with gravel being preferred

(Fig. 3-8C). Vegetation is the preferred cover type for creek chub followed by edge with

a preference value of 0.69 (Fig. 3-8D).

The fathead minnow, Pimephales promelas Rafinesque, 1820, was collected from

23 macrohabitats (16.8%; Tables 3-1, 3-4) and five sites. They were collected from three

backwaters (30%), two riffles (13%), and 18 runs (18%). They exhibited a preference for

shallow water with the 25-cm class being preferred (Fig. 3-9A). Fathead minnows show a

preference for the 35-cm/s velocity class (Fig. 3-9B), and slower, with no individuals

59

captured in the 45-cm/s class or faster. Rubble is the substrate preferred by fathead

minnows followed by gravel with a preference value of 0.29 (Fig. 3-9C). The preferred

cover type for fathead minnows is edge followed by vegetation with a preference value of

0.40 (Fig. 3-9D).

A synthesis of these findings indicates that hornyhead chub, common carp, and

common shiners were usually found in run macrohabitats that were 55–85 cm deep.

Largescale stoneroller, longnose dace, creek chub, and fathead minnows were usually

found in riffle macrohabitats that was 15–35 cm deep. Water velocity for the riffle

species was generally slower (15–65 cm/s) than for the run species (65 cm/s). Each

species has a specific preference with substrate (sand, gravel, boulders) and cover

(vegetation, boulders) or no cover. None of these species were found to prefer pool

macrohabitats

Discussion

Our habitat preference results generally agree with those of Aadland & Kuitunen

(2006). The best example was the longnose dace, which was in agreement on all the four

habitat variables. Other species (hornyhead chub, common shiner, creek chub, and

fathead minnow) were in agreement on three of four habitat variables that Aadland and

Kuitunen (2006) reported as optimal. Substrate or velocity was the variable where our

findings for these species did not agree with those of Aadland & Kuitunen (2006). Our

analysis demonstrated a preference between longnose dace and shallow riffles and runs

that have high flow velocities and boulder substrates. This agrees with the findings in a

60

similar sized river (Mullen & Burton, 1995) and habitat suitability index variables for this

species (Edwards et al., 1983). In contrast, Hubert & Rahel (1989) found longnose dace

were associated with overhead cover, aquatic vegetation, and backwater pools.

Our findings for largescale stoneroller agreed on two of the four variables (depth

and velocity) identified by Aadland & Kuitunen (2006). The discrepancy was perhaps

because our study was limited to conditions in one river with low individual sample size

(n=35; Table 3-3) and macrohabitats (n=5; Table 3-1). If the differences are not

attributable to the low sample size, then the differences in optimal cover type and

substrate class for largescale stonerollers between our study and that of Aadland &

Kuitunen (2006) are likely due to characteristics of the Forest River in the reaches where

largescale stonerollers were collected. The river substrate was mainly sand or silt (Fig. 3-

2). The few areas with gravel were located among boulders in riffles. Therefore, boulders

were classified as the dominate substrate and cover even if the fish were seeking gravel,

as indicated in Minnesota (Aadland & Kuitunen, 2006).

Extreme high and low depth and velocity classes are always going to be under

sampled because they are extremes. This will lead to bias (Hansen et al., 2007). Many of

the species investigated here were affected to some degree by this sample size bias,

although we also investigated the habitat preferences with the undersampled habitats

removed.

For example, the preferred velocity for common shiner is biased high (Fig. 3-7B)

due to the low sample size for that velocity (Fig. 3-2), but with that class removed, the

species shows a bimodal velocity preference. This bimodal preference for velocity may

61

be indicating the preferences of different size classes of fish as common shiner adults and

juveniles are found in very different velocities in Minnesota (Aadland & Kuitunen,

2006). The preference of the common shiner to silt, sand, and gravel substrates agrees

with the substrate requirement previously established for this species (Trail et al. 1983),

but the preference for no cover contrasts to a study where common shiners were

associated with submerged aquatic vegetation (Hubert & Rahel, 1983). Common shiners

have been reported to inhabit main-channel areas (Hubert & Rahel, 1983). Our findings

agree with this because 97.7% of common shiners collected were from main-channel

areas (Table 3-1).

Some differences between the findings of our study and those of Aadland &

Kuitunen (2006) are not attributable to low sample size bias. For example, creek chub

associated with gravel substrates in the Forest River, but in Minnesota the optimal

substrate for creek chub is small boulder (Aadland & Kuitunen, 2006). Our finding of

creek chub associating with vegetative cover agrees with previous studies (McMahon,

1982; Hubert & Rahel 1989). The creek chub preference for moderate velocity (35 cm/s)

agrees with the habitat suitability index variable (McMahon, 1982) but not the findings of

Hubert & Rahel (1989). The findings of creek chub associating with runs and shallower

water disagree with those of McMahon (1982) and Hubert & Rahel (1989). Fathead

minnows associated with gravel or rubble substrates whereas in Minnesota they were

substrate generalists (Aadland & Kuitunen, 2006). For both fathead minnows and creek

chub, a high number of individuals were captured immediately below a dam where much

62

of the finer substrates had been washed away. This could be a case of an impoundment

and habitat interaction confounding the interpretation of the results.

Dams create barriers to fish movement and can affect the fish community found

upstream of the dam. Common carp in the Forest River were only collected below a dam

near the mouth of the watershed. This dam creates a barrier to upstream invasion, thereby

serving to benefit the upper reaches of the Forest River.

The species investigated during this study show similar results to those in

Minnesota streams (Aadland & Kuitunen, 2006). Some of the differences between these

species preferences in the Forest River and species preferences in streams in Minnesota

can be attributed to the relatively narrow scope of this study, 137 macrohabitats at 16

sites on one waterway. This limited the possible habitats to sample and we did not have

the range of environmental conditions (45 sites, 4375 macrohabitats) as Aadland &

Kuitunen (2006). Future research should focus on smaller and more quantified habitats

with a more standardized sampling protocol so that fish habitat relationships and

preferences can have more supporting evidence.

Recognizing the habitat use patterns of stream fishes can be used to facilitate

management of species of conservation priority. Although site specific river restoration

projects are the building blocks for watershed restoration (Ziemer, 1997), knowledge of

how the watershed affects the aquatic habitat is also important or some site specific

projects might fail (Frissell, 1997). For example, restoring the riparian ecosystem resulted

in a dramatic improvement in the aquatic physical habitat of a midwestern stream

(Isenhart et al., 1997). Our findings have immediate practical applications for species

63

conservation and river restoration in North Dakota. Future researchers can increase

sampling efficiency by focusing on certain habitats associated with target species.

Furthermore, local habitat restoration projects could create specific habitats associated

with habitat specialist species which would benefit the entire fish community.

Acknowledgements

We thank S. Sindelar, J. Billings, E. Boyda, and C. Stearns for their aid in

sampling the Forest River and associated fishes. We would also like to thank Cari-Ann

Hayer for her guidance throughout the sampling for this study. Further gratitude is

extended to J. Ladonski for his laboratory assistance in verifying fish identifications. All

fish were obtained legally under the permitting guidelines of the North Dakota Game and

Fish Department (permit # GNF02362977 and GNF02483803). Funding for this project

was provided in part by the North Dakota Game and Fish Department and South Dakota

State University.

64

References

Aadland L.P. (1993) Stream habitat types: their fish assemblages and relationship to

flow. North American Journal of Fisheries Management, 13, 790–806.

Aadland L.P. & Kuitunen A. (2006) Habitat suitability criteria for stream fishes and

mussels of Minnesota. Minnesota Department of Natural Resources, Special

Publication #162. 167pp.

Borgstrom L.J. (2010) Fish community assembly in the Forest River, North Dakota.

Masters thesis, South Dakota State University, Brookings, South Dakota

Braaten P.J. & Berry C.R. Jr. (1997) Fish associations with four habitat types in a South

Dakota prairie stream. Journal of Freshwater Ecology, 12, 477–489.

Converse Y.K., Hawkins C.P. & Valdez R.A. (1998) Habitat relationships of subadult

humpback chub in the Colorado River through Grand Canyon: spatial variability

and implications of flow regulation. Regulated Rivers: Research and Management,

14, 267–284.

Dieterman D.J. & Galat D.L. (2004) Large-scale factors associated with sicklefin chub

distribution in the Missouri and lower Yellowstone Rivers. Transactions of the

American Fisheries Society, 133, 577–587.

Dunham J., Rieman B. & Chandler G. (2003) Influences of temperature and

environmental variables on the distribution of bull trout within streams at the

southern margin of its range. North American Journal of Fisheries Management,

23, 894–904.

65

Edwards E.A., Li H., Schreck C.B. (1983) Habitat suitability index models: longnose

dace. United States Department of the Interior, Fish and Wildlife Service,

FWS/OBS-82/10.33.

Frissell C.A. (1997) Ecological principles. In: Watershed restoration: principles and

practices (Eds J.E. Williams, C.A. Wood, M.P. Dombeck), pp. 96–115. American

Fisheries Society, Bethesda, Maryland.

Gerhard P., Moraes R. & Molander S. (2004) Stream fish communities and their

associations to habitat variables in a rain forest reserve in southeastern Brazil.

Environmental Biology of Fishes, 71, 321–340.

Gido K.B. & Propst D.L. (1999) Habitat use and association of native and nonnative

fishes in the San Juan River, New Mexico and Utah. Copeia, 1999, 321–332.

Grabowski T.B. & Isely J.J. (2006) Seasonal and diel movements and habitat use of

robust redhorses in the lower Savannah River, Georgia and South Carolina.

Transactions of the American Fisheries Society, 135, 1145–1155.

Hagen S.K., Isakson P.T. & Dyke S.R. (2005) North Dakota Comprehensive Wildlife

Conservation Strategy. North Dakota Game and Fish Department. Bismarck, ND.

454pp. http://gf.nd.gov/conservation/cwcs.html.

Hansen M.J., Beard T.D. Jr., & Hayes D.B. (2007) Sampling and Experimental Design.

In: Analysis and Interpretation of Freshwater Fisheries Data (Eds C.S. Guy, M.L.

Brown), pp 51–120. American Fisheries Society, Bethesda, Maryland,.

66

Hoagstrom C.W., Brooks J.E. & Davenport S.R. (2008) Recent habitat association and

the historical decline of Notropis simus pecoensis. River Research and

Applications, 24, 789–803.

Hubert W.A. & Rahel F.J. (1989) Relations of physical habitat to abundance of four

nongame fishes in high-plains streams: a test of habitat suitability index models.

North American Journal of Fisheries Management, 9, 332–340.

Isenhart T.M., Schultz R.C., Colletti J.P. (1997) Watershed restoration and agricultural

practices in the Midwest: Bear Creek of Iowa. In: Watershed restoration: principles

and practices (Eds J.E. Williams, C.A. Wood, M.P. Dombeck), pp. 318–334.

American Fisheries Society, Bethesda, Maryland.

Kessler R.K., Casper A.F. & Weddle G.K. (1995) Temporal variation in microhabitat use

and spatial relations in the benthic fish community of a stream. American Midland

Naturalist, 134, 361–370.

McMahon T.E. (1982) Habitat suitability index models: creek chub. United States

Department of the Interior, Fish and Wildlife Service, FWS/OBS-82/10.4.

Mueller G.A., Marsh P.C. & Minckley W.L. (2005) A legacy of change; the lower

Colorado River, Arizona–California–Nevada, USA, and Sonora–Baja California

Norte, Mexico. In: Historical changes in large river fish assemblages of the

Americas (Eds J.N. Rinne, R.M. Hughes & B. Calamusso), pp. 139-156. American

Fisheries Society, Symposium 45, Bethesda, Maryland.

Mullen D.M. & Burton T.M. (1995) Size-related habitat use by longnose dace

(Rhinichthys cataractae). American Midland Naturalist, 133, 177–183.

67

Panek F.M. 1987. Biology and ecology of carp. In: Carp in North America (Ed E.L.

Cooper), pp 1–15. American Fisheries Society, Bethesda, Maryland.

Power M.E., Stout R.J., Cushing C.E., Harper P.P., Hauer F.R., Matthews W.J., Moyle

P.B., Statzner B. Wais De Badgen I. (1988) Biotic and abiotic controls in river and

stream communities. Journal of the North American Benthological Society, 7, 456–

479.

Quist M.C. & Guy C.S. (2001) Growth and mortality of prairie stream fishes: relations

with fish community and instream habitat characteristics. Ecology of Freshwater

Fish, 10, 158–163.

Trail J.G., Wade C.S., Stanley J.G., Nelson P.C. (1983) Habitat suitability information:

common shiner. United States Department of the Interior, Fish and Wildlife

Service, FWS/OBS-82/10.40.

Vörösmarty C.J., Green P., Salisbury J. & Lammers R.B. (2000) Global water resources:

vulnerability from climate change and population growth. Science, 289, 284–288.

Walsh S.J. & Meador M.R. (1998) Guidelines for quality assurance and quality control of

fish taxonomic data collected as part of the National Water Quality Assessment

Program. US Geological Survey Water Resources Investigations Report 98-4239.

Walters D.M., Leight D.S., Freeman M.C., Freeman B.J. & Pringle C.M. (2003)

Geomorphology and fish assemblages in a Piedmont river basin, U.S.A. Freshwater

Biology, 48, 1950–1970.

68

Ziemer R.R. (1997) Temporal and spatial scales. In: Watershed restoration: principles

and practices (Eds J.E. Williams, C.A. Wood, M.P. Dombeck), pp. 318–334.

American Fisheries Society, Bethesda, Maryland.

69

Macrohabitat

Number

Sampled

Hornyhead

chub

N. biguttatus

Largescale

stoneroller

C. oligolepis

Common

carp

C. carpio

Longnose

dace

R. cataractae

Common

shiner

L. cornutus

Creek chub

S. atromaculatus

Fathead

minnow

P. promelas

Backwater 10 3 (7) 0 1 (13) 0 2 (122) 7 (25) 3 (66)

Pool 13 7 (39) 1 (1) 0 2 (23) 6 (129) 7 (64) 0

Riffle 16 9 (53) 3 (33) 0 6 (112) 9 (382) 10 (124) 2 (565)

Run 98 44 (247) 1 (1) 3 (120) 5 (8) 71 (3082) 49 (972) 18 (5608)

Total 137 63 (346) 5 (35) 4 (133) 13 (143) 88 (3715) 73 (1185) 23 (6239)

Table 3-1. Total number of macrohabitats sampled in the Forest River, North Dakota and

the number of times the selected species were present in those macrohabitats. The

numbers of individuals are in parentheses.

Macrohabitat

Number

Sampled

Hornyhead

chub

N. biguttatus

Largescale

stoneroller

C. oligolepis

Common

carp

C. carpio

Longnose

dace

R. cataractae

Common

shiner

L. cornutus

Creek chub

S. atromaculatus

Fathead

minnow

P. promelas

Backwater 10 3 (7) 0 1 (13) 0 2 (122) 7 (25) 3 (66)

Pool 13 7 (39) 1 (1) 0 2 (23) 6 (129) 7 (64) 0

Riffle 16 9 (53) 3 (33) 0 6 (112) 9 (382) 10 (124) 2 (565)

Run 98 44 (247) 1 (1) 3 (120) 5 (8) 71 (3082) 49 (972) 18 (5608)

Total 137 63 (346) 5 (35) 4 (133) 13 (143) 88 (3715) 73 (1185) 23 (6239)

Table 3-1. Total number of macrohabitats sampled in the Forest River, North Dakota

and the number of times the species of concern (Horny head chub Nocomis biguttatus,

Largescale stoneroller Campostoma oligolepis), invasive species (common carp Cyprinus

carpio), and other selected species (Longnose dace Rhinichthys cataractae, Common

shiner Luxilus cornutus, Creek chub Semotilus atromaculatus, and Fathead minnow

Pimephales promelas) were present in those macrohabitats. The number of individuals is

in parenthesis.

70

Substrate

Dimension

(mm) Cover Description

Silt <0.062 Undercut Undercut bank

Sand 0.062–3.2 Vegetation Rooted or unrooted plants

Gravel 3.2–64 Wood Woody matter

Cobble 64–128 Boulder Boulders > 10 cm above streambed

Rubble 128–256 Flotsam Thick foam on water surface Small

Boulder 256–508 Canopy Canopy or overhead structure

Large Boulder 508–1016 Edge A break from high to low velocities

Bedrock >1016

Table 3-2. Dimensions of substrate categories and descriptions of cover categories from

Aadland (1993).

Table 3-1. Dimensions of substrate categories and descriptions of cover categories from

Aadland (1993).

71

Family Species Common name

Number of

Individuals

Relative

Abundance

Esocidae Esox lucius northern pike 53 0.003

Cyprinidae Campostoma oligolepis largescale stoneroller 35 0.002

Cyprinella spiloptera spotfin shiner 15 0.001

Cyprinus carpio common carp 134 0.007

Luxilus cornutus common shiner 3715 0.206

Nocomis biguttatus hornyhead chub 346 0.019

Notropis dorsalis bigmouth shiner 1093 0.061

Pimephales notatus bluntnose minnow 1170 0.065

Pimephales promelas fathead minnow 6403 0.355

Rhinichthys cataractae longnose dace 143 0.008

Rhinichthys obtusus

western blacknose

dace 123 0.007

Semotilus atromaculatus creek chub 1193 0.066

Catostomidae Carpiodes cyprinus quillback 7 0.000

Catostomus commersonii white sucker 2304 0.128

Ictaluridae Ameiurus melas black bullhead 26 0.001

Ictalurus punctatus channel catfish 1 <0.001

Noturus gyrinus tadpole madtom 206 0.011

Gasterosteidae Culaea inconstans brook stickleback 2 <0.001

Centrarchidae Ambloplites rupestris rock bass 4 <0.001

Lepomis macrochirus bluegill 577 0.032

Pomoxis nigromaculatus black crappie 5 <0.001

Percidae Etheostoma nigrum johnny darter 284 0.016

Perca flavescens yellow perch 89 0.005

Percina maculata blackside darter 69 0.004

Sander vitreus walleye 1 <0.001

Table 3–3. List of families and associated species of fishes collected from the Forest

River of North Dakota showing total individuals and relative abundance for each species.

Table 2–1. List of families and associated species of fishes collected from the Forest

River of North Dakota.

Table 3–4. List of families and associated species of fishes collected from the Forest

River of North Dakota with total individuals and relative abundance for each species

shown.

Table 2–1. List of families and associated species of fishes collected from the Forest

River of North Dakota.

72

Macrohabitat Microhabitat

Species

Backwater

n = 10

Pool

n = 13

Riffle

n = 16

Run

n = 98

Depth

(cm)

Velocity

(cm/s) Substrate Cover

Hornyhead chub

N. biguttatus 30 54 56 45 75 65 G Veg

Largescale

stoneroller

C. oligolepis

0 8 19 1 35 65 LB B

Common carp

C. carpio 10 0 0 3 55 15 S None

Longnose dace

R. cataractae 0 15 38 5 15 - 35 65 LB B

Common shiner

L. cornutus 20 46 56 72 75 - 85 65 G - S None

Creek chub

S. atromaculatus 70 54 63 50 25 35 G Veg

Fathead minnow

P. promelas 30 0 13 18 25 35 Rub-G Veg

Table 3-4. Preferred macrohabitat and microhabitat metrics for the species of concern,

invasive species, and other selected species collected from in the Forest River, North

Dakota. The macrohabitat columns (backwater, pool, riffle, and run) refer to the

percentage of that habitat in which the species was collected. For the substrate column,

S= sand, G = gravel, Rub = rubble, LB = large boulder. For the cover column, None = no

cover, Veg = vegetation, B = boulders.

73

Fig. 3-1. Map depicting the 16 study sites and the four dams impounding over 50 hectares

of water along the Forest River. Sites are numbered in a downstream manner with site one

being the most headwater site sampled, and site 16 being the most downstream site

sampled.

74

52

3

9 8

39

04

22

0

10

20

30

40

50

60

Fre

qu

ency

25

64

21

3 1

1013

00

10

20

30

40

50

60

70

Fre

qu

nce

y

7

19

43

23

28

14

2 1

0

5

10

15

20

25

30

35

40

45

50

0 5 15 25 35 45 55 65

Fre

qu

en

cy

Velocity (cm/s)

0 0

7

21

31 30

14 13

7 8

3 2 1

0

5

10

15

20

25

30

35

0 5 15 25 35 45 55 65 75 85 9510

511

5

Fre

qu

en

cy

Depth (cm)

A B

C D

Fig. 3-2. Sampling frequency distributions for depth (A), velocity (B), substrate (C) and

cover (D). Depth and velocity axis values are based on the center point for the 10–cm

class. The number of times each habitat class was sampled is above the corresponding

bar.

75

Fig. 3-3. Depth (A), velocity (B), substrate (C) and cover (D) preference histograms for

hornyhead chub with standardized preference value above the bar. Depth and velocity

axis values are based on the center point for the 10–cm class. The value of 0.75 was

assigned to all covers that had a preference value less than the preference value for no

cover based on the assumption that fish will not choose to avoid cover if all other habitat

requirements are met.

0.22

0.42

1.00

0.26

0.00

0.64

0.50

0.0

0.2

0.4

0.6

0.8

1.0

Pre

fere

nce

val

ue 0.75 0.75

1.00

0.75 0.75 0.75 0.75

0.0

0.2

0.4

0.6

0.8

1.0

Pre

fere

nce

val

ue

0.17

0.280.32

0.15

0.34

0.07

1.00

0.210.26

0.66

0.11

0

0.2

0.4

0.6

0.8

1

15

25

35

45

55

65

75

85

95

105

115

Pre

fere

nce

val

ue

Depth (cm)

0.01

0.21

0.35

0.080.14

0.17

0.33

1.00

0

0.2

0.4

0.6

0.8

1

0 5 15 25 35 45 55 65

Pre

fere

nce

va

lue

Velocity (cm/s)

A B

C D

76

Fig. 3-4. Depth (A), velocity (B), substrate (C) and cover (D) preference histograms for

largescale stoneroller with standardized preference value above the bar. Depth and

velocity axis values are based on the center point for the 10–cm class. The value of 0.02

was assigned to all covers that had a preference value less than the preference value for

no cover based on the assumption that fish will not choose to avoid cover if all other

habitat requirements are met.

.

0.02 0.02 0.02 0.02

1.00

0.02 0.02

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.00 0.00 0.02 0.00 0.00

0.22

1.00

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.000.05

1.00

0.070.000.000.000.000.00

0.50

0.000

0.2

0.4

0.6

0.8

1

15 25 35 45 55 65 75 85 95 105

115

Pre

fere

nce

val

ue

Depth (cm)

0.00 0.00 0.00 0.00 0.00 0.000.03

1.00

0

0.2

0.4

0.6

0.8

1

0 5 15 25 35 45 55 65

Pre

fere

nce

val

ue

Velocity (cm/s)

A B

C D

77

Fig. 3-5. Depth (A), velocity (B), substrate (C) and cover (D) preference histograms for

common carp with standardized preference value above the bar. Depth and velocity axis

values are based on the center point for the 10–cm class. The value of 1.00 was assigned

to all covers that had a preference value less than the preference value for no cover based

on the assumption that fish will not choose to avoid cover if all other habitat

requirements are met.

.

1.00 1.00 1.00 1.00 1.00 1.00 1.00

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.35

1.00

0.00 0.00 0.00 0.00 0.000

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.000.000.00

0.18

1.00

0.000.000.00

0.89

0.000.000

0.2

0.4

0.6

0.8

1

15

25

35

45

55

65

75

85

95

105

115

Pre

fere

nce

val

ue

Depth (cm)

0.00

0.25

1.00

0.00 0.00 0.00 0.00 0.000

0.2

0.4

0.6

0.8

1

0 5 15 25 35 45 55 65

Pre

fere

nce

va

lue

Velocity (cm/s)

A B

C D

78

Fig. 3-6. Depth (A), velocity (B), substrate (C) and cover (D) preference histograms for

longnose dace with standardized preference value above the bar. Depth and velocity axis

values are based on the center point for the 10–cm class. The value of 0.01 was assigned

to all covers that had a preference value less than the preference value for no cover based

on the assumption that fish will not choose to avoid cover if all other habitat

requirements are met.

.

0.01 0.010.06 0.04

1.00

0.01 0.03

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.000.04 0.02

0.09

0.00

0.28

1.00

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.12

0.35

1.00

0.050.000.000.000.000.000.000.00

0

0.2

0.4

0.6

0.8

1

15

25

35

45

55

65

75

85

95

105

115

Pre

fere

nce

val

ue

Depth (cm)

0.00 0.00 0.01 0.00 0.01 0.00 0.02

1.00

0

0.2

0.4

0.6

0.8

1

0 5 15 25 35 45 55 65

Pre

fere

nce

va

lue

Velocity (cm/s)

A B

C D

79

Fig. 3-7. Depth (A), velocity (B), substrate (C) and cover (D) preference histograms for

common shiner with standardized preference value above the bar. Depth and velocity

axis values are based on the center point for the 10–cm class. The value of 1.00 was

assigned to all covers that had a preference value less than the preference value for no

cover based on the assumption that fish will not choose to avoid cover if all other habitat

requirements are met.

.

1.00 1.00 1.00 1.00 1.00 1.00 1.00

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.61

0.871.00

0.00 0.00

0.37

0.54

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.280.23

0.56

0.36

0.250.28

1.00

0.73

0.36

0.74

0.000

0.2

0.4

0.6

0.8

1

15

25

35

45

55

65

75

85

95

105

115

Pre

fere

nce

val

ue

Depth (cm)

0.06

0.22

0.06 0.040.10

0.19

0.07

1.00

0

0.2

0.4

0.6

0.8

1

0 5 15 25 35 45 55 65

Pre

fere

nce

va

lue

Velocity (cm/s)

A B

C D

80

Fig. 3-8. Depth (A), velocity (B), substrate (C) and cover (D) preference histograms for

creek chub with standardized preference value above the bar. Depth and velocity axis

values are based on the center point for the 10–cm class. The value of 0.13 was assigned

to all covers that had a preference value less than the preference value for no cover based

on the assumption that fish will not choose to avoid cover if all other habitat

requirements are met.

.

0.13 0.13

1.00

0.16

0.26

0.13

0.69

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.05

0.34

1.00

0.01

0.71

0.47

0.29

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.17

1.00

0.300.24

0.35

0.040.030.06

0.80

0.10

0.000

0.2

0.4

0.6

0.8

1

15

25

35

45

55

65

75

85

95

105

115

Pre

fere

nce

val

ue

Depth (cm)

0.180.25

0.340.39

1.00

0.300.24 0.27

0

0.2

0.4

0.6

0.8

1

0 5 15 25 35 45 55 65

Pre

fere

nce

val

ue

Velocity (cm/s)

A B

C D

81

Fig. 3-9. Depth (A), velocity (B), substrate (C) and cover (D) preference histograms for

fathead minnows with standardized preference value above the bar. Depth and velocity

axis values are based on the center point for the 10–cm class. The value of 0.05 was

assigned to all covers that had a preference value less than the preference value for no

cover based on the assumption that fish will not choose to avoid cover if all other habitat

requirements are met.

.

0.05 0.05

0.40

0.05

0.18

0.05

1.00

0.0

0.2

0.4

0.6

0.8

1.0

Pre

fere

nce

val

ue

0.02 0.01

0.29

0.00

1.00

0.02 0.05

0

0.2

0.4

0.6

0.8

1

Pre

fere

nce

val

ue

0.00

1.00

0.04

0.12

0.000.000.000.00

0.12

0.000.000

0.2

0.4

0.6

0.8

1

15 25 35 45 55 65 75 85 95 105

115

Pre

fere

nce

val

ue

Depth (cm)

0.00 0.020.09

0.30

1.00

0.00 0.00 0.000

0.2

0.4

0.6

0.8

1

0 5 15 25 35 45 55 65

Pre

fere

nce

va

lue

Velocity (cm/s)

A B

C D