Long Term Resource Monitoring Program

Special Report

96-S001

Taxonomic and Distributional Status

of Notropis volucellus

and Notropis wickliffi in the

Mississippi River Drainage:

A Literature Review

This PDF file may appear different from the printed report

because of slight variations incurred by electronic transmission.

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July 1996
Taxonomic and Distributional Status

of Notropis volucellus and Notropis wickliffi

in the Mississippi River Drainage:

A Literature Review

by

Robert A. Hrabik

Missouri Department of Conservation

Open River Field Station

Jackson, Missouri 63755

Prepared for

National Biological Service

Environmental Management Technical Center

575 Lester Avenue

Onalaska, Wisconsin 54650

Project Leader: Steve Gutreuter

July 1996
LTRMP Special Reports provide Long Term Resrouce Program partners

with scientific and technical support. The opinions and conclusions

 in LTRMP Special Reports are those of the author(s) and do not 

necessarily reflect those of the Environmental Management Technical Center.

 All reports in this series receive anonymous peer review.

National Biological Service

Environmental Management

Technical Center

CENTER DIRECTOR

Robert L. Delaney

ECOLOGICAL MONITORING AND

RESEARCH DIRECTOR

Steve Gutreuter

INFORMATION AND TECHNOLOGY

SERVICES DIRECTOR

Norman W. Hildrum

INFORMATION TRANSFER AND

MEDIA SERVICES MANAGER

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REPORT EDITOR

Deborah K. Harris

Cover graphic by Mi Ae Lipe-Butterbrodt

Mention of trade names or commercial products does not constitute endorsement or recommendation

for use by the National Biological Service, U.S. Department of the Interior.

The National Biological Service . . . gathering, analyzing,

and sharing the biological information necessary to support the wise

stewardship of the Nation's natural resources.

Printed on recycled paper
iii

Contents

Page

Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v

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

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

Taxonomic History .......................................... 2

Etymology, Nomenclature, and Type Localities ..................... 2

Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Description of the Taxa and Similar Sympatric Species . . . . . . . . . . . . . . . . . . . 5

Diagnostic Characters ..................................... 5

Similar Sympatric Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Distribution and Habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Biology and Life History ...................................... 9

Notropis wickliffi ........................................ 9

Notropis volucellus ....................................... 9

Habits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Food and Feeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Tables

Table. Selected diagnostic characters of Notropis volucellus and Notropis wickliffi

from the Mississippi River drainage . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Figures

Figure 1. Location of Long Term Resource Monitoring Program field stations

where Program biologists are collecting long-term resource data on the

Upper Mississippi River . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Figure 2. Generalized distribution of Notropis volucellus in North America, after

Gilbert and Burgess (1980) ............................. 8

Figure 3. Generalized distribution of Notropis wickliffi in North America, based on

Etnier and Starnes (1993) and Hrabik (unpublished data) . . . . . . . . . 10
v

Preface

The Long Term Resource Monitoring Program (LTRMP) was authorized under the

Water Resources Development Act of 1986 (Public Law 99-662) as an element of the U.S.

Army Corps of Engineers' Environmental Management Program. The LTRMP is being

implemented by the Environmental Management Technical Center, a National Biological

Service Science Center, in cooperation with the five Upper Mississippi River System

(UMRS) States of Illinois, Iowa, Minnesota, Missouri, and Wisconsin. The U.S. Army

Corps of Engineers provides guidance and has overall Program responsibility. The mode

of operation and respective roles of the agencies are outlined in a 1988 Memorandum of

Agreement.

The UMRS encompasses the commercially navigable reaches of the Upper Mississippi

River, as well as the Illinois River and navigable portions of the Kaskaskia, Black,

St. Croix, and Minnesota Rivers. Congress has declared the UMRS to be both a nationally significant ecosystem and a nationally significant commercial navigation system. The

mission of the LTRMP is to provide decision makers with information for maintaining the

UMRS as a sustainable large river ecosystem given its multiple-use character. The longterm goals of the Program are to understand the system, determine resource trends and

effects, develop management alternatives, manage information, and develop useful

products.

This report was prepared under Strategy 2.2.8, Monitor and Evaluate Fish

Communities, Guilds, and Populations, Task 2.2.8.4, Evaluate and Summarize Annual

Results, as specified in Goal 2 of the Operating Plan of the LTRMP for the Upper

Mississippi River System (USFWS 1993). This report was developed with funding

provided by the Long Term Resource Monitoring Program.

Additional copies of this report may be obtained from the National Technical

Information Service, 5285 Port Royal Road, Springfield, Virginia 22161 (1-800-553-6847

or 703-487-4650).

This report may be cited:

Hrabik, R. 1996. Taxonomic and distributional status of Notropis volucellus and

Notropis wickliffi in the Mississippi River drainage: A literature review. National

Biological Service, Environmental Management Technical Center, Onalaska,

Wisconsin, July 1996. LTRMP 96-S001. 15 pp.
Taxonomic and Distributional Status

of Notropis volucellus and Notropis wickliffi

in the Mississippi River Drainage:

A Literature Review

By Robert A. Hrabik

Abstract

Notropis volucellus was described in 1864 and underwent a confusing taxonomic synonymy with Notropis

blennius. By the 1930s, N. volucellus was recognized as a full species with three subspecies. The subspecies

N. v. wickliffi was described in 1931 but was not generally recognized as a full species until 1991. Notropis

volucellus is widely distributed in the Mississippi River drainage, where it may be found in large creeks and

rivers. Notropis wickliffi inhabits large Midwestern rivers from Louisiana to Wisconsin to Pennsylvania. In

the Upper Mississippi River, N. volucellus and N. wickliffi occur sympatrically. Biologists with the Long Term

Resource Monitoring Program (LTRMP) initially identified N. wickliffi as N. volucellus. I began examining

LTRMP voucher specimens in 1991 and determined that specimens from Pool 13 to the Mississippi River’s

confluence with the Ohio River identified as N. volucellus were actually N. wickliffi. Above Pool 13, cursory

separation of N. wickliffi from N. volucellus becomes increasingly difficult. Specimens from Pool 8 were

mostly N. wickliffi, but several appeared to be intergrades. The purpose of this literature review is to describe

the taxonomic and distributional status of N. volucellus and N. wickliffi in the Mississippi River drainage and

to present supporting biological information that may help to define the taxonomy and distribution of both

species.

Introduction

Biologists working on the Upper Mississippi

River have encountered a significant taxonomic

problem with Notropis volucellus (mimic shiner)

and N. wickliffi (channel shiner), which ichthyologists have attempted to resolve over the years.

For example, in 1931 Trautman described

N. wickliffi from the Ohio and Mississippi Rivers

where it sympatrically occurs with N. volucellus.

Subsequently, Jenkins (1976) and Gilbert (1978)

were the first to elevate N. wickliffi to full species

rank but neither published justification for the action. Trautman (1981) also suggested that

N. wickliffi was specifically distinct from

N. volucellus. Mayden and Kuhajda (1989) elevated N. wickliffi to full species status and gave

morphological and electrophoretic characters.

Robins et al. (1991) recognized N. wickliffi as a

full species, elevating it from a subspecies of

N. volucellus. Several ichthyologists, including

T. M. Cavender, D. A. Etnier, and C. R. Gilbert

(in Robins et al. 1991), have expressed confidence

that these taxa are full species. Etnier and Starnes

(1994) recognized and distinguished N. wickliffi

from N. volucellus in their treatise on Tennessee

fishes.

The Long Term Resource Monitoring Program

(LTRMP) is an element of the U.S. Army Corps

of Engineers' Environmental Management Program for the Upper Mississippi River. Original

authorization for the LTRMP provided for a 10-yr

Program starting in 1987; this authorization was

extended for another 5 years in 1990. Six remote

state-operated field stations have been established

for data collection. Water quality, fish, vegetation, wildlife, and invertebrates are being monitored in five separate reaches of the Mississippi

River and one reach of the Illinois River.

During routine monitoring, LTRMP biologists

collected numerous specimens of the

N. volucellus-N. wickliffi group but all were assigned to N. volucellus. In 1991, I began to examine voucher specimens retained by the six field
2

stations in the Program (Fig. 1). Using meristic

and morphometric information provided in the

literature, I determined that all records from the

open river listed as N. volucellus were actually N.

wickliffi. By 1992, I concluded that N. wickliffi

was the dominant form of the N. volucellus

species-group in the Upper Mississippi River from

Pool 13, to the Ohio River (Fig. 1). However,

cursory separation of N. wickliffi from N.

volucellus becomes increasingly difficult above

Pool 13, where meristic characters and melanophoric patterns begin to overlap. Specimens from

Pool 8 were mostly N. wickliffi but several appeared to be intergrades. Notropis volucelluswickliffi intergrades have also been noted by other

ichthyologists (G. Seegert and D. Fago, personal

communication) in Pools 8 and 9.

Gaps presently exist in the LTRMP database

because biologists are not able to accurately identify members of the N. volucellus species-group.

I prepared this literature review in anticipation of

a morphometric, meristic, and electrophoretic

study to determine the taxonomic validity and distribution of each species in the Mississippi River

drainage. The purpose of the literature review is

to describe the taxonomic and distributional status

of N. volucellus and N. wickliffi in the Mississippi

River drainage and to present supporting biological information that may help to define the taxonomy and distribution of both species.

Taxonomic History

Both N. volucellus and N. wickliffi belong to

the teleost family Cyprinidae (minnows) and the

genus Notropis (shiners, in part). Both are included in the subgenus Notropis and have been

assigned to the volucellus species-group by

Mayden (1989). Other members of this speciesgroup according to Mayden (1989) are the following: Notropis buchanani, N. cahabae,

N. heterolepis, N. maculatus, N. ozarcanus,

N. rupestris, N. spectrunculus, and Opsopeodus

emiliae. However, Amemiya and Gold (1990)

suggested that the Volucellus species-group is not

monophyletic. They demonstrated that O. emiliae

is more closely related to the genus Cyprinella on

the basis of chromosomal nucleolus organizer regions phenotypes. Furthermore, Page and

Johnston (1990) demonstrated that the breeding

behavior of O. emiliae is unique among North

American minnows belonging to a clade comprising the genera Opsopoeodus, Codoma, and

Pimephales.

Etymology, Nomenclature, and

Type Localities

Notropis is a compound word derived from

notus, meaning back, and tropis, meaning keel.

The correct spelling should be Nototropis; however, a ruling by the International Rules of Zoological Nomenclature upholds the current spelling

(Smith 1985). Volucellus is derived from volucer,

meaning winged or swift, and wickliffi is a

patronym for Edward L. Wickliff, an Ohio naturalist and colleague of Milton B. Trautman. The

type locality for N. volucellus is the Detroit River

at Grosse Ile, Wayne County, Michigan (Cope

1864). The type locality for N. wickliffi is at the

mouth of the Miami River, Hamilton County,

Ohio (Trautman 1931).

The vernacular names for N. volucellus and

N. wickliffi are mimic shiner and channel shiner.

The term mimic shiner was applied to

N. volucellus because the species superficially

resembles many North American cyprinids.

Channel shiner was applied to N. wickliffi to describe its primary habitat—the main channel of

large rivers.

Systematics

The phylogenetic relationships of North American cyprinids are poorly understood. Some have

called cyprinid phylogeny "chaotic" (Hubbs and

Miller 1977) because their genetic and ontogenetic

characters are complex and difficult to classify.
3

.

.

.

.

.

.

Lake City

Onalaska

Bellevue

Havana

Alton

Cape Girardeau

MINNESOTA

WISCONSIN

IOWA

MISSOURI

ILLINOIS

Mississippi River

Illinois River

Notropis volucellus

Possible intergrades

Notropis wickliffi

Mississippi River

Figure 1. Location of Long Term Resource Monitoring Program field stations where Program biologists

are collecting long-term resource data on the Upper Mississippi River. Distribution of Notropis volucellus,

Notropis wickliffi, and possible intergrades in the Upper Mississippi River is shown, based on meristic and

morphometric characters.
4

However, Mayden (1985, 1989) conducted

monumental work in this area, setting the course

for clarification of some of the most confusing

relationships in the Cyprinidae. Mayden (1989)

recognized three subgenera and two speciesgroups in the genus Notropis: Notropis,

Alburnops, and Hydrophlox; and volucellus and

texanus, respectively. The relationships between

these groups are unresolved with the exception of

the subgenus Notropis and the species-group

texanus, which are sister clades.

Similarly, the relationships between clades and

among species in the volucellus species-group are

unclear (Mayden 1989). Notropis volucellus is

probably a complex of species, meaning that several closely related sympatrically and

allopatrically occurring species have been assigned to N. volucellus because insufficient data

exist to separate them. I concur with other ichthyologists that this complex exists. For example,

Burr and Warren (1986) stated that "several forms

masquerade under the name N. volucellus."

Etnier and Starnes (1994) summed up the confusion this way: "an understanding of the phylogeny of this group and the number of species or subspecies involved is seen as one of the most interesting, needed, and difficult problems remaining

in the systematics of North American freshwater

fishes."

Notropis volucellus was originally described by

Cope (1864), who assigned it to the genus

Hybognathus (H. volucella). Others placed the

species in Hybopsis, Alburnops, and Notropis

(Nelson 1876; Jordan 1878; Gilbert 1978). The

species epithet deliciosus, an early name for

Notropis stramineus (sand shiner), is believed to

be a senior synonym of volucellus (Hubbs 1926;

Smith 1979). Volucellus and deliciosus have both

been misapplied to various genera (O'Donnell

1935). From its description in 1864 until 1928,

N. volucellus was often in synonymy with the

Notropis blennius (river shiner) complex, which

consisted of the sand, mimic, and ghost shiners

(Hubbs and Greene 1928). Systematics of the

river shiner are also confusing. It was described

by Girard (1856) as Alburnops blennius. Jordan

(1878) redescribed the species and named it

Episema jejuna. Forbes (1884) placed it in

Notropis (jejunus). The sand shiner was named

blennius by Jordan and Evermann (1896–1900)

and by Forbes and Richardson (1920). Essentially, Forbes' jejunus became blennius and Forbes

and Richardson's blennius became stramineus.

Species epithets were resurrected by Hubbs and

Greene (1928), separating this complex into the

sand shiner (deliciosus—i.e., stramineus) and

mimic shiner (volucellus), and recognizing two

subspecies of the mimic shiner, volucellus and

buchanani. Later, Suttkus (1958) relegated

deliciosus to the synonymy of N. texanus. When

Trautman (1931) described the channel shiner, a

third subspecies, wickliffi, was added.

Although many forms of N. volucellus exist

(e.g., the Neosho River form; in Hubbs and

Bonham 1951), only one subspecies,

N. v. volucellus, is recognized today. Bailey (in

Harlan and Speaker 1951) reelevated the ghost

shiner (Gilbert 1980), and Jenkins (1976), Gilbert

(1978), and finally Robins et al. (1991) elevated

N. wickliffi to full species rank.

Notropis wickliffi was originally described by

Trautman (1931) as a subspecies of N. volucellus.

Trautman (1931, 1981) noted intergrades between

wickliffi and volucellus, especially in tributaries to

large rivers, and seasonally in the Ohio River.

Etnier and Starnes (1994) reported that in some

Tennessee streams, N. wickliffi and N. volucellus

sympatrically occurred and were distinguishable.

Becker (1983), however, expressed reservations

about his ability to distinguish the two taxa from

Wisconsin waters. Overall, there seems to be

little disagreement among ichthyologists that

N. wickliffi warrants full species rank and that

intergrades may occur where they are sympatric

(Trautman 1981; Robins et al. 1991; Etnier and

Starnes 1994).

Taxonomic studies involving N. volucellus and

N. wickliffi are few. Gong and Cavender (1991)

used univariate and multivariate techniques to describe the taxa from Ohio. While finding

N. wickliffi only in the Ohio River, Gong and

Cavender (1991) found a north–south cline in sev-
5

eral morphometric and meristic variables and suggested that if a total geographic analysis was conducted, more taxonomic forms of N. volucellus

could be revealed.

In another taxonomic study, Mayden and

Kuhajda (1989) compared meristic, morphological, and genetic characters of N. wickliffi,

N. volucellus, and N. cahabae. They concluded

that N. volucellus is a complex of more than one

taxon, and N. wickliffi is a distinctive and sister

species to the newly described N. cahabae.

Description of

the Taxa and Similar

Sympatric Species

Diagnostic Characters

Selected diagnostic characters for N. volucellus

and N. wickliffi appear in the Table. Some

N. volucellus characters are ambiguous—possibly

reflecting more than one taxon or geographical

variation. The N. volucellus characters showing

the most variation are lateral-line scales, scales

above and below the lateral line, pectoral fin rays,

anal fin rays, gill rakers, and maximum size.

Lateral-line scales ranged from 32 to 40.

Smaller scaled N. volucellus (35–40) were from

Louisiana (Douglas 1974), New York (Smith

1985), and Illinois (Smith 1979). Scales above

the lateral line were generally 8–10, with 7–9 below. However, Smith (1985) reported that New

York specimens had only about four scales above

and below the lateral line. New York

N. volucellus also had fewer pectoral fin rays

(12–15) and gill rakers (5–6). The largest average

maximum size (65 mm) was reported from Illinois

(Smith 1979) and Tennessee (Etnier and Starnes

1994), but larger specimens have been

documented elsewhere (Trautman 1981). The

smallest N. volucellus specimens were found in

Louisiana (Douglas 1974) and Wisconsin (Becker

1983).

In general, N. wickliffi can be distinguished

from N. volucellus and its forms by having larger

scales (fewer in the lateral line); more scales

above and below the lateral line (less elevated

scales); and a nonexistent to lightly pigmented

predorsal stripe and spot. Mayden and Kuhajda

(1989) noted that the typical N. wickliffi has a

larger eye, longer mouth, more posteriorly placed

anal fin, a longer and more anteriorly placed dorsal fin, shorter predorsal length, and a narrower

caudal peduncle than either N. volucellus or N.

cahabae. In addition, Etnier and Starnes (1994)

noted that N. wickliffi has a less arched back and

more prominent nuptial tubercles on the head.

However, Etnier and Starnes (1994) stated that N.

wickliffi had a deeper caudal peduncle than N.

volucellus, opposite of the observations of

Mayden and Kuhajda (1989). Collections from

the Upper Mississippi River vary in several physical attributes, including body depth, in which

N. wickliffi is sometimes deeper than N. volucellus

(Hrabik, unpublished data). Etnier and Starnes

(1994) noted that N. wickliffi from Tennessee have

a continuous postdorsal dark streak, but that

sympatric N. volucellus have a discontinuous or

no postdorsal streak. Conversely, N. wickliffi

from the Upper Mississippi River rarely have a

postdorsal streak, but N. volucellus I have examined from several upper Midwest localities may

have a continuous postdorsal streak (Hrabik, unpublished data).

Similar Sympatric Species

Notropis volucellus and N. wickliffi are most

often confused with N. stramineus (sand shiner),

N. buchanani (ghost shiner), N. shumardi

(silverband shiner), and N. amnis (pallid shiner).

Notropis volucellus and N. wickliffi can be distinguished from N. stramineus by having eight anal

fin rays (seven in N. stramineus), poorly developed pre- and postdorsal fin stripes (prominent in

N. stramineus), a more rounded snout, and elevated scales in the anterior portion of the lateral

line. They differ from N. buchanani in having
6

Table. Selected diagnostic characters of Notropis volucellus and Notropis wickliffi from the Mississippi River drainage.

 Character Notropis volucellus Notropis wickliffi

Lateral-line scales 32–40 32–34

Scales above lateral line 4–10 10–11

Scales below lateral line 3–9 9–11

Circumferential scales 17–22 21–24

Predorsal scales 13–16 12–15

Breast squamation None to fully scaled Usually none or a few scales

Scale form Elevated anteriorly 2–4 times height

over width

Elevated anteriorly 2–3 times

height over width

Pectoral fin rays 12–17 14–17

Pelvic fin rays 8–9 8

Dorsal fin rays 8 8

Anal fin rays 7–9 8

Gill rakers 5–10 5–8

Teeth 2,4–4,2 and 4–4 Not known

Maximum size 50–76 mm •65 mm

Body depth—greatest point 20%–25% of standard length About 20% standard length

Head length 3.5–4.0 times into standard length 3–4 times into standard length

Eye diameter 2.6–3.5 times into head length 2–3 times into head length

Lateral stripe Variable; well developed to vague; usually more intense posteriorly; expanded

on caudal peduncle usually with ventral

lobe; spot reported on some specimens

Less variable; moderately developed and more intense posteriorly; expanded on caudal

peduncle usually with ventral

lobe; no spot

Predorsal stripe and spot Heavily pigmented and evident Nonexistent to lightly pigmented

Caudal peduncle scales More heavily pigmented posteriorly Usually uniformly pigmented

Preoperculo-mandibular canal Usually complete or interrupted Complete

Infraorbital canal Uninterrupted and complete Uninterrupted and complete

Vertebrae 32–37 Unknown

Peritoneum Silvery to scattered dark speckles Lightly scattered dark speckles

Tubercles Absent on fins, except dorsal surface of

pectorals; well developed on snout;

noted on specimens 40 mm or larger

Same except less developed on

snout and not found on specimens less than 50 mm

Data summarized from: Becker (1983); Burr and Warren (1986); Cross and Collins (1975); Douglas (1974); Etnier and

Starnes (1993); Hrabik (unpublished data); Mayden and Kuhajda (1989); Pflieger (1975); Robison and Buchanan (1988);

Smith (1979); Smith (1985); and Trautman (1981).
7

more pigment on the body; a less pointed dorsal

fin; moderately elevated scales in the anterior portion of the lateral line; pelvic fins not reaching the

anal fins when depressed; a dusky stripe on the

caudal peduncle; speckled peritoneum; less deep,

slab-sided body; and a complete infraorbital canal.

They differ from N. shumardi in having elevated

scales, eight anal fin rays (nine in N. shumardi),

a more terminal mouth (as opposed to oblique),

and no dark pigment along the lateral line. They

differ from N. amnis in pharyngeal tooth count

(1,4–4,1 in N. amnis); and in the mouth (upper

lip) extending to the eye, being slightly more

oblique, and level with the lower margin of the

eye.

Distribution and Habitat

Notropis volucellus and its forms are widely

distributed from southern Canada to Texas; they

are widespread in the eastern drainages of the

Mississippi River (Fig. 2) and extend westward

through the Guadalupe and San Antonio river basins in southeastern Texas (Gilbert and Burgess

1980). Buchanan (1976) reported N. volucellus to

be one of the most abundant minnows in the lower

White River system of Arkansas. This conclusion

is questionable because the lower White River

flows through the Mississippi River Alluvial

Plain, where habitat seems more suited to N.

wickliffi. Notropis volucellus is also found in the

Red River of the North, the Great Lakes, and St.

Lawrence drainages and Lake Champlain system

(Gilbert and Burgess 1980). This species has a

curious distribution in Tennessee, where it is

largely absent from direct tributaries to the Mississippi River (Etnier and Starnes 1994), and in

southern Michigan where it is absent (Gilbert and

Burgess 1980). The absence of N. volucellus

from southern Michigan may be an artifact, as

University of Michigan Museum of Zoology fish

records may not have been available to all authors

in the Atlas of North American Freshwater Fishes

(D. Etnier, personal communication). Disjunct

populations of N. volucellus occur in Missouri

(Pflieger 1975), North Carolina, and Virginia

(Gilbert and Burgess 1980). Notropis volucellus

has been introduced to the Susquehanna River

basin where it is rapidly spreading (Malick 1978;

Gilbert and Burgess 1980; Cooper 1983), and has

been reported to be declining in some Midwestern

states, especially in Illinois (Smith 1979), Ohio

(Trautman 1981), and southern Wisconsin (Becker

1983).

Because of the confusing taxonomy, the definition of N. volucellus and N. wickliffi zoogeography is tenuous. However, it seems plausible that

both species evolved from stock in the Teays-Mississippi River system (Pflieger 1971).

Notropis volucellus avoids small headwater

streams but may be found in a variety of other

aquatic environments. In northern parts of its

range, it is found in vegetated pools or backwaters

of small streams and rivers (Black 1945; Becker

1983; Smith 1985). In Missouri, it is locally

abundant in clear, vegetated, and flowing

Bootheel ditches (Pflieger 1975; Hrabik, unpublished data). In Louisiana, it is more common in

low gradient streams, especially of the Mississippi

River Alluvial Plain, than in higher gradient

streams of the surrounding Coastal Plain (Douglas

1974). Elsewhere, it is found in clear to moderately clear medium-sized streams to highly turbid,

large rivers (Pflieger 1975; Trautman 1981; Burr

and Warren 1986; Etnier and Starnes 1994). The

small stream forms are generally found in clear

pools over gravel substrate in moderate current.

Large river forms are found along shorelines over

gravel, sand, and mud substrates usually in slow

to moderate current. Becker (1983) reported the

presence of N. volucellus in the shallow bays of

the Great Lakes, and Smith (1979) and Becker

(1983) noted this species from glacial lakes in Illinois and Wisconsin. Etnier and Starnes (1994)

noted that it has a "tolerance" for large reservoirs

in Tennessee.

In comparison, N. wickliffi is seemingly restricted to large rivers, the lower reaches of direct

tributaries, and some large creeks (Trautman

1981; Etnier and Starnes 1994). Cross (1967)

referred to the Neosho River form as N. wickliffi,

 but I doubt the accuracy of that identification.
8

M I S S O U R I R I V E R

A R K A N S A S R I V E R 

R E D R I V E R 

O H I O R I V E R 

T E N N E S S E E R I V E R 

Range of Notropis volucellus

. .

.

.

. . .

. . . .

.

. .

. .

.

.

.

.

. San Antonio, TX Houston, TX

Dallas, TX

Alexandria, LA

Jackson,MS

Texarkana, TX

Tulsa, OK

Springfield, MO

St. Louis, MO

Memphis, TN

Indianapolis, IN

Nashville, TN

Green Bay, WI

Minneapolis, MN

Fargo, ND

Toledo, OH

Roanoke, VA

Pittsburgh, PA

Syracuse, NY

Montreal, Canada

Birmingham, AL

Figure 2. Generalized distribution of Notropis volucellus in North America, after Gilbert and Burgess

(1980). Distribution extends into southern Canada and may include large rivers within its range.
9

The Neosho-Spring River form is probably an

undescribed species or subspecies of N. volucellus

(Pflieger 1975). Etnier and Starnes (1994) show

N. wickliffi to be distributed in the Mississippi

River from the mouth of the Missouri River to

northeastern Louisiana, the Ohio River from

Pennsylvania to its confluence with the Mississippi River, the Tennessee River, and the Wabash

River (Fig. 3).

In Missouri, N. wickliffi has been collected in

the lower Missouri River (Pflieger 1975), but is

seemingly rare in the pooled portion of the Mississippi River above St. Louis (Pflieger 1975;

Hrabik, unpublished data). However, N. wickliffi

was collected in the lower Fabius River in northeastern Missouri in 1989 (Hrabik, unpublished

data). Unfortunately, voucher specimens were

mistaken for N. stramineus and discarded.

Biologists from the LTRMP field station at

Bellevue, Iowa (Upper Mississippi River, Pool

13), commonly collect N. wickliffi during routine

monitoring (S. Gritters, personal communication)

and have retained numerous voucher specimens

verified by B. Kuhajda and me. Specimens collected by biologists from the Onalaska, Wisconsin, LTRMP field station (Upper Mississippi

River, Pool 8) have been identified as N. wickliffi

and N. wickliffi-volucellus intergrades (Hrabik,

unpublished data). G. Seegert (personal communication) has also collected N. wickliffi and

intergrades from Upper Mississippi River pools in

Illinois and Wisconsin.

Notropis wickliffi is found along shoreline and

shoal habitats, with or without current, over mud,

sand, and gravel substrates (Etnier and Starnes

1994; Hrabik, unpublished data; LTRMP, unpublished data). The species seems to tolerate high

turbidity but is also found in moderately clear

large creeks and rivers. Trautman (1981) noted

that when the Ohio River becomes very clear during droughts and winter, N. wickliffi numbers decrease and intergrades increase.

Biology and Life History

Notropis wickliffi

Because N. wickliffi has only recently been recognized as a species, little is known about its biology and habits. In the Middle Mississippi River

(Pflieger 1975; Hrabik, unpublished data), it is

most often found in association with Notropis

buchanani, N. atherinoides, N. shumardi, and

N. blennius. Trautman (1957) reported spawning

from June to August in Ohio. Etnier and Starnes

(1994) collected tuberculate specimens during the

same months in Tennessee, the nuptial tubercles

verifying that the species were in a stage of breeding. In contrast with these few facts, much more

is known about the biology and life history of N.

volucellus.

Notropis volucellus

Habits

In Missouri streams, N. volucellus is most often found with N. greenei, N. texanus,

N. rubellus, Luxilus zonatus, and Lythrurus

fumeus (Pflieger 1975). In one Wisconsin stream,

it was associated with relatively large populations

of Luxilus cornutus, Moxostoma erythrurum,

Nocomis biguttatus, Pimephales notatus, Percina

maculata, and Etheostoma nigrum (Becker 1983).

In Ontario, Hallam (1959) found it in warmwater

sections of streams with Ambloplites rupestris and

Micropterus dolomieu. In a Minnesota lake,

Moyle (1973) found it at night with Pimephales

notatus.

Notropis volucellus is a schooling fish usually

found at the surface or in midwater (Moyle 1973;

Robison and Buchanan 1988). Moyle (1973) reported finding schools with as many as 20,000

individuals in a Minnesota lake.
10

M I S S O U R I R I V E R

A R K A N S A S R I V E R

R E D R I V E R

O H I O R I V E R

T E N N E S S E E R I V E R

Etnier and Starnes

Hrabik and others

Possible range

.

.

.

.

.

.

.

.

.

.

.

.

Baton Rouge, LA Alexandria, LA

Vicksburg, MS

Florence, AL Little Rock, AR

Jonesboro, AR

Louisville, KY St. Louis, MO

Kansas City, MO

Davenport, IA

La Crosse, WI

Wheeling, PA

Figure 3. Generalized distribution of Notropis wickliffi in North America, based on Etnier and Starnes

(1993) and Hrabik (unpublished data).
11

Diurnal movements have been observed by

several researchers (Black 1945; Moyle 1973;

Gascon and Leggett 1977; Becker 1983; Hanych

et al. 1983). During the day, Notropis volucellus

was found in large schools inshore. These

schools break into smaller groups offshore at night

in Indiana and Minnesota lakes (Black 1945;

Moyle 1973). However, Hanych et al. (1983)

found the reverse, with N. volucellus making

movements inshore at sunset, nearshore at night,

and offshore at sunrise. Hanych et al. (1983) explained this behavior as a predator-response mechanism.

Although N. volucellus has been observed in

habitats containing aquatic vegetation (Black

1945; Becker 1983; Smith 1985; Etnier and

Starnes 1994), it is often found in areas without

cover and, in one Alabama study, did not seek

cover at all (Kinsolving and Bain 1990).

Reproduction

Spawning takes place as early as June (Black

1945; Pflieger 1975; Becker 1983) and may continue into August (Moyle 1969; Cross and Collins

1975; Becker 1983). In Wisconsin, Becker

(1983) found gravid females in June and ripe eggs

in July.

Weather may influence spawning activity.

Moyle (1969) found that N. volucellus did not

spawn in a Minnesota lake exposed to low summer temperatures because these temperatures possibly affected food web dynamics and energetics.

Spawning activity has never been observed.

Black (1945) suggested that spawning takes place

at night in deep, vegetated areas in lakes. Females outnumber males two to one at this time

(Moyle 1969), and nesting or parental care do not

seem to exist (Black 1945).

Sexual maturity is reached in its first year

(Black 1945; Moyle 1973; Etnier and Starnes

1994). The average number of eggs per female in

an Indiana lake was 367 (Black 1945), although

Becker (1983) reported as many as 960. A description of the protolarvae is given by Potter and

Potter (1981). Notropis dorsalis, N. stramineus

(Becker 1983), and N. atherinoides (Mayhew

1983) have been documented to hybridize with N.

volucellus.

Food and Feeding

Notropis volucellus has a varied diet generally

consisting of small crustaceans, insects, algae, and

detritus (Black 1945; Moyle 1973; Gascon and

Leggett 1977; Magnin et al. 1978; Becker 1983).

Johnson and Dropkin (1992) found it will also eat

larval Alosa sapidissima (American shad) when

this species is abundant.

Moyle (1973) observed N. volucellus in large

schools feeding in midwater or at the surface of a

Minnesota lake but saw no evidence of resource

(food) competition with other cyprinids in that

system. Black (1945) and Moyle (1969) observed

similar diel feeding patterns in lakes they studied.

Notropis volucellus fed on zooplankton in the

early morning, switched to diptera and terrestrial

insects during the day, and ate mayflies and amphipods in the evening. Strangely, copepods were

ignored even when abundant. Some feeding may

occur at night (Black 1945).

Gascon and Leggett (1977) studied the response of several littoral zone fishes to nutrientproduction gradients in Lake Memphremagog

(northern Vermont and southern Quebec). They

found that N. volucellus had a rather specialized

diet consisting mostly of cladocerans and diptera

larvae, but would eat detritus when resource-limited. The distribution of N. volucellus in Lake

Memphremagog was determined by the relative

abundance of preferred food resources.

However, Olmsted et al. (1979) found

N. volucellus to be less than particular in its food

preferences. They presented two prey types in

five densities and found large variations in prey

preference. Notropis volucellus did not exhibit a

"precise preference tactic based on prey availability."
12

In a Susquehanna River study, Johnson and

Dropkin (1991) found that N. volucellus fed

heavily on aerial aquatic and terrestrial insects.

They concluded that dietary overlap was high with

Cyprinella spiloptera (spotfin shiner) and that

drifting prey was an important food item.

Growth

Females grow faster and attain a larger size

than males (Black 1945). Length at age I was 48

mm and at age II was 55 mm in an Indiana lake

(Black 1945), and 43–52 and 52–65 mm in Wisconsin (Becker 1983). Sexual maturity was

reached at 42 mm in a Minnesota lake (Moyle

1973). Maximum age is 2–3 years (Black 1945;

Moyle 1973).

Summary

Notropis volucellus is probably a complex of

many forms. This may explain the variability in

morphological and meristic characters, phenotypic

expression, and the wide distribution across several North American drainages observed by many

ichthyologists. On the basis of recent morphological and genetic evidence and known habitat preference, I conclude that Notropis wickliffi is probably a distinct species.

Because N. wickliffi has only recently been

given specific status, its distributional limits and

life history are poorly known. And because of the

magnitude of the taxonomic confusion in this complex, it may be some time before new taxa are

described after geographic-wide systematics studies.

In the Mississippi River drainage, the distributional limits and habitat requirements for N.

volucellus (and its forms) and N. wickliffi are virtually unknown. Because biologists are not confident in their ability to identify these species, they

can make no progress toward monitoring

trends and understanding the ecology of these species. These constraints render decision makers

powerless to pursue economic progress without

the danger of destroying critical habitat.

The need for a definitive systematics analysis

of the Mississippi River drainage complex is evident, and the ramifications of such a project are

important. In this age of rapidly changing landscapes and reductions in species ranges, the time

is right to solve this problem.

Acknowledgments

I thank the administration and S. Gutreuter of

the Long Term Resource Monitoring Program for

their support of this project. D. A. Etnier and

B. R. Kuhajda reviewed the manuscript and provided helpful comments.

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14

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the genus Cyprinella (Teleostei:

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fish endemic to the Cahaba River of the Mobile basin. Bulletin of the Alabama Museum

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15

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among three species of minnows (Cyprinidae)

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wickliffi, a new subspecies of cyprinid fish

from the Ohio and Upper Mississippi Rivers.

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Plan for the Upper Mississippi River System

Long Term Resource Monitoring Program.

Environmental Management Technical Center, Onalaska, Wisconsin, Revised September

1993. EMTC 91-P002R. 179 pp. (NTIS

#PB94-160199)
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1. AGENCY USE ONLY (Leave blank) 2. REPORT DATE

July 1996

3. REPORT TYPE AND DATES COVERED

4. TITLE AND SUBTITLE

Taxonomic and Distributional Status of Notropis volucellus and Notropis wickliffi in the Mississippi River Drainage: 

A Literature Review

5. FUNDING NUMBERS

6. AUTHOR(S)

Robert A. Hrabik

7. PERFORMING ORGANIZATION NAME AND ADDRESS

Missouri Department of Conservation

Open River Field Station

Jackson, Missouri 63766

8. PERFORMING ORGANIZATION

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9. SPONSORING/MONITORING AGENCY NAME(S) AND ADDRESS(ES)

National Biological Service 

Environmental Management Technical Center

575 Lester Avenue

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96-S001

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Release unlimited. Available from National Technical Information Service, 5285 Port Royal Road, Springfield, VA 22161 

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13. ABSTRACT (Maximum 200 words)

Notropis volucellus was described in 1864 and underwent a confusing taxonomic synonymy with Notropis blennius. By the 1930s, N. volucellus was recognized as a

full species with three subspecies. The subspecies N. v. wickliffi was described in 1931 but was not generally recognized as a full species until 1991. Notropis

volucellus is widely distributed in the Mississippi River drainage, where it may be found in large creeks and rivers. Notropis wickliffi inhabits large Midwestern rivers

from Louisiana to Wisconsin to Pennsylvania. In the Upper Mississippi River, N. volucellus and N. wickliffi occur sympatrically. Biologists with the Long Term

Resource Monitoring Program (LTRMP) initially identified N. wickliffi as N. volucellus. I began examining LTRMP voucher specimens in 1991 and determined that

specimens from Pool 13 to the Mississippi River’s confluence with the Ohio River identified as N. volucellus were actually N. wickliffi. Above Pool 13, cursory separation of N. wickliffi from N. volucellus becomes increasingly difficult. Specimens from Pool 8 were mostly N. wickliffi, but several appeared to be intergrades. The

purpose of this literature review is to describe the taxonomic and distributional status of N. volucellus and N. wickliffi in the Mississippi River drainage and to present

supporting biological information that may help to define the taxonomy and distribution of both species.

14. SUBJECT TERMS

biology, distribution, life history, Long Term Resource Monitoring Program, Mississippi River, Notropis volucellus,

Notropis wickliffi, systematics, taxonomy

15. NUMBER OF PAGES

15 pp.

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20. LIMITATION OF ABSTRACT
The Long Term Resource Monitoring Program (LTRMP) for the Upper

Mississippi River System was authorized under the Water Resources

Development Act of 1986 as an element of the Environmental Management

Program. The mission of the LTRMP is to provide river managers with

information for maintaining the Upper Mississippi River System as a sustainable

large river ecosystem given its multiple-use character. The LTRMP is a

cooperative effort by the National Biological Service, the U.S. Army Corps of

Engineers, and the States of Illinois, Iowa, Minnesota, Missouri, and Wisconsin.