ERNEST A. LACHN

and ROBERT E. JENK1N

Systematics, Distribution,

and Evolution of the Chub

Genus Nocomis Girard

{Pisces, Cyprinidae) of

Eastern United States, With

Descriptions of New Species i

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 85
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 85

Ernest A. Lachner Systematics, Distribution,

and Robert E. Jenkins ^ Evolution of the

Chub Genus Nocomis

Girard (Pisces, Gyprinidae)

of Eastern United States,

With Descriptions of

New Species

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Lachner, Ernest A., and Robert E. Jenkins. Systematics, Distribution, and Evolution of the Ghub Genus Nocomis Girard (Pisces, Cyprinidae) of Eastern United

States, with Descriptions of New Species. Smithsonian Contributions to Zoology,

number 85, 97 pages, 1971.—The chub genus Nocomis Girard of North America

is characterized and relationships among three species groups are discussed. Two

new species are described from the central Appalachian region, both in the micropogon species group.

The three species groups are defined and their characters are summarized;

namely, the biguttatus group with three species, the micropogon group with three

species, and the leptocephalus group with three subspecies. The central Appalachian region is inhabited by four species, N. micropogon (river chub), N. platyrhynchus, new species, (bigmouth chub) and N. raneyi, new species (bull chub) of

the micropogon group, all of which are sympatric with N. leptocephalus (bluehead chub) in one or more river drainages. A key to these four species and a

discussion of the nomenclatural history of the genus and nominal species are

included. Important diagnostic characters useful in differentiating among the

specific and several infraspecific populations are discussed, evaluated, and summarized in twenty-seven tables. An account of each species includes a synonymy,

diagnosis, description, and comparison of meristic and morphometric characters,

coloration in life and in preservation, population differentiation, reproduction,

growth and size attained, materials studied and geographic distribution. The

species are illustrated, special body features are drawn, distributions are plotted,

and important character data are shown in thirty figures.

The ecological requirements of the chubs are reviewed and species preferences

discussed. Regional ecology is reviewed in respect to occurrence and abundance

of the species. Associations, interrelationships and frequency of hybridization are

compared among the four species. Differences are observed among the species

of the micropogon group in their frequency of hybridization with N. leptocephalus.

The biological and geological evidence providing an explanation of the dispersal of the chubs and other species of fishes in various interdrainage exchanges

in the central Appalachian region is comprehensively reviewed. Possible routes of

entry into the several drainages are proposed. Present, detailed distributional

patterns of the chubs and other species are related with geologic events, and present and past ecological conditions in an effort to understand the conditions,

barriers, and routes affecting dispersal. Three kinds of geological events that

operated in the dispersal of chubs are considered: stream capture, eustatic changes

of the Atlantic Coastal Plain, and Pleistocene drainage modifications. The discussion on the evolution of the species of Nocomis is based on morphology, coloration, zoogeography, and life history.

Official publication date for each title is handstamped in a limited number of initial copies and

is recorded in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1971

For talc by the Superintendent of Documents, U. S. Government Printing Office

Washington, D. C. 20402 - Price $1.25 (paper cover)
Contents

Page

Preface 1

Introduction 2

Acknowledgments 6

Diagnostic Characters 7

Genus Nocomis Girard 13

Key to the Species of Nocomis 16

The micropogon Group 16

Nocomis raneyi, new species 17

Atypical Nocomis raneyi 32

Nocomis platyrhynchus, new species 37

Nocomis micropogon (Cope) 42

The leptocephalus Group 50

Nocomis leptocephalus (Girard) 50

Ecology, Association, and Frequency of Hybridization 55

Dispersal: The Biological and Geological Evidence 61

Evolution of Nocomis 73

Summary 78

Literature Cited 80

Tables 1-27 85

Index 95

in

Ernest A. Lachner

and Robert E. Jenkins

Systematics, Distribution,

and Evolution of the

Chub Genus Nocomis

Girard (Pisces, Cyprinidae)

of Eastern United States,

With Descriptions of

New Species

Preface

The first author of this series of studies became interested in the systematics and life history of the

North American barbeled cyprinids, Genus Nocomis,

more than twenty years ago. A new species was discovered from the Roanoke River in 1946 based on

one large adult male. Shortly thereafter, a new form

was found in the New-Kanawha River drainage.

At that time three highly differentiated and widely

distributed species in the genus were recognized,

N. biguttatus, N. leptocephalus, and N. micropogon.

Additional collections of the new forms from the

central Appalachians and adequate collections of

the described species accumulated slowly at first,

and little progress was made toward an understanding of the systematics of the group. Relationships

among the species were not clear because the primary

diagnostic characters showed allometric development

and they were also sexually dimorphic. In addition,

some species showed considerable populational divergence in certain river systems on a subspecific or

racial level. It became apparent from preliminary

exploratory studies that new forms were also present

in the Cumberland and Green River drainages, the

Ernest A. Lachner, Department of Vertebrate Zoology, National

Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Robert E. Jenkins, Roanoke College, Salem,

Virginia 24153.

Arkansas River drainage, and in rivers of the southern Appalachian slope.

Solution of the systematic problems necessitated

large series of adults from the major drainages of

eastern United States, westward to the Ozarks and

the Missouri River basin. In recent years extensive

collections became available through the cooperation of many institutional and state workers. Many

tuberculate adult male specimens, which are of

greatest importance for systematic study, were obtained mainly by electro-seining and angling. These

collections and those housed in other American

museums, numbering more than 100,000 specimens,

coupled with recent field work brought the study

to fruition.

The systematic studies revealed that the three

recognized species were members of separate groups,

each group containing three forms. New insight was

obtained on generic relationships among the American cyprinid genera Hybopsis, Nocomis, and Notropis and of the evolution of the nest building

chubs, Nocomis. Field studies contributed comparative data on the ecology, distribution, and dispersal

of the species. Intensive appraisal of important diagnostic characters showed that certain specimens,

identified as "variants" in the early stages of the

study, represented natural interspecific hybrids. Five

interspecific Nocomis hybrid combinations were discovered and seventeen intergeneric combinations in1
2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

volving Nocomis with other cyprinid genera were

described. A population of N. micropogon, inhabiting the Potomac River, approaches characters of the

new species in the New River drainage and may

have resulted from introgressive hybridization from

early and limited dispersal through the headwaters

of the Monongahela River at times when these

waterways (Greenbrier, Monongahela, Potomac)

were variously connected.

It was known that the males of the three previously established species (N. biguttatus, N. micropogon, and N. leptocephalus) build and guard large

gravel nests during the spring reproductive period.

The reproductive behavior, of N. biguttatus and

N. micropogon was reported by several authors as

well as the utilization of the nests of these three

species by other fishes as reproductive sites. Since

the morphology (the number and distribution of

nuptial tubercles on the head and body and the

development of nuptial head crests) differs among

Nocomis species, it was pertinent to investigate

whether all species of Nocomis construct nests, and

to determine possible differences in their breeding

behaviors related to the specific nuptial characters.

It was discovered that the males of all species of

Nocomis build a nest which is guarded by a solitary

male. The interactions between interspecific and

intergeneric breeding populations over the nest

were studied in respect to agonistic and compatible

associations, particularly involving the breeding

male Nocomis. The extent and significance of the

utilization of the nests of Nocomis by other fishes,

especially the cyprinids, were comprehensively evaluated. Since the reproductive activities of chubs necessitates relatively large amounts of water with

moderate to fast flow, beds of small gravel and comparatively large areas (the small creeks are not inhabited by Nocomis, and fighting occurs when the

males have limited nesting space), it is not possible

to experimentally breed the species without elaborate and extensive habitat construction, using large

artificial riffle-pool complexes; consequently stream

sites were sought and selected where observation

and study under natural conditions could be done

for all the species of Nocomis. The various breeding

behavioral patterns and associations of species were

thus evaluated in regard to their functional and/or

evolutionary significance. The impact of these associations in the production of natural hybrids was

appraised. Thus, a study of the evolutionary biology

of the chubs emerged.

Two individuals participated in various portions

of the overall study during their doctoral programs,

Martin L. Wiley and Robert E. Jenkins. One study

in this series was recently published (Lachner and

Jenkins, 1967). A related study was completed by

Wiley (1969) on the comparative morphology and

histology of nuptial tubercles in fishes.

Six parts will complete this series of studies, all

to be published shortly in the Smithsonian Contributions to Zoology:

1. "Systematics, Distribution and Evolution of the

Chub Genus Nocomis (Pisces, Cyprinidae) of Eastern United States, with Descriptions of New

Species," by Ernest A. Lachner and Robert E.

Jenkins. [Number 85]

2. "Populations of the Polytypic Species Nocomis

leptocephalus (Girard) with a Description of a

New Subspecies," by Ernest A. Lachner and Martin L. Wiley. [Number 92]

3. "Systematics, Distribution, and Evolution of the

Nocomis biguttatus species group (Family Cyprinidae: Pisces) with a Description of a New Species from the Ozark Upland," by Ernest A.

Lachner and Robert E. Jenkjns. [Number 91]

4. "Criteria for Analysis and Interpretation of the

American Fish Genera Nocomis Girard and

Hybopsis Agassiz," by Robert E. Jenkins and

Ernest A. Lachner. [Number 90]

5. "Natural Interspecific and Intergeneric Cyprinid

Hybrids Involving the Chub Genus Nocomis,"

by Ernest A. Lachner, Robert E. Jenkins, and

Martin L. Wiley. [In preparation]

6. "Reproduction, Behavior, Nest Associations, Hybridization, and Interrelationships Between the

Chub Genus Nocomis and Other North American

Cyprinid Fishes," by Ernest A. Lachner. [In preparation]

Introduction

During a spring collecting trip in 1946 to the central

Appalachian region of Virginia, Edward C. Raney,

Robert D. Ross, and Ernest A. Lachner examined

the catch of a lone angler on the Roanoke River at

Salem, Virginia. A large cyprinid on his stringer

appeared distinctive from the known members of

the nest-building chubs, genus Nocomis. Subsequent

exploration and study revealed that this specimen
NUMBER 8 5

represented a new species confined to several drainages of the central Atlantic slope and that a related

undescribed form occurred in the New River drainage. Both forms are closely related to the river chub,

N. micropogon, a species widely distributed over

northeastern United States. Another species, N. leptocephalus, sympatric with both species in the area,

was also treated. The large amount of study material obtained in recent years made it possible to

consider the geographic distribution and dispersal

of the species in detail which proved relevant to the

interpretation of their evolution.

The objectives of this paper are to (1) describe

two new species of American barbeled cyprinids

from the central Appalachian region; (2) to discuss

their relationships within the micropogon speciesgroup and to describe the species-groups in the

genus; (3) to summarize the pertinent information

on the ecology of the four species of the central

Appalachian region; and (4) to describe the geographic distributions of the species and to postulate

their dispersal, based on current distribution, ecology, and the geological history of the region.

METHODS.—The methods of counting meristic

characters and measuring body proportions were

taken as outlined by Hubbs and Lagler (1958: 19-

26) in most instances. Other methods were used

when the conventional procedures proved unsatisfactory or when a character was specialized to necessitate exploratory study before a satisfactory procedure was selected. The terminology used is explained

below.

All measurements of length refer to the standard

length (SL) in millimeters (mm) unless otherwise

stated. The width of the gape was measured as the

least distance between the angles of the lips with the

mouth in a normal, closed position. The length of

the head was taken as the greatest distance from the

tip of the snout to the posterior margin of the bony

opercle. The postorbital length is the longest measure from the bony orbit to the posterior margin of

the bony opercle. The total vertebral count includes

the vertebrae in the Weberian complex, presumably

four, and the hypural plate as one. The caudal vertebrae count started with the first, elongate haemal

spine. This distinction was not possible in the small

specimens. The sexes were determined by gonadal

inspection. A binocular microscope was used for

sexing, determining tubercle and scale development

and counting meristic characters in small specimens.

Our tabular data, particularly that referring to

meristic characters, was largely taken from samples

of seven to ten specimens each spread over the range

of a species within the various river systems as the

material permitted. The tabular data is arranged

by drainages and, when possible, in geographical

sequence.

The breast is considered to be the area bounded

by the isthmus, opercular membranes, pectoral fin

bases and a line connecting the posterior pectoral

fin bases. The degree of breast squamation was quantified by employing the lines shown in Figure la

and odd-numbered lines between these (not shown).

To evaluate the degree of squamation (Table 18),

a line was first determined at which the most anterior scales were located. If the breast was fully

scaled up to a certain line, the degree of squamation

was regarded as the number of the line. When the

breast was only partially scaled in areas behind the

most anterior scale, it was given an averaged but

lower rating. For example, the squamation in Figure la is rated as 1. The most anterior scale is at

line 2 but only about one half of the area between

lines 0 and 2 is scaled. Specimens with both an

unsealed breast and an unsealed portion of the area

posterior to line 0 were rated as 0.

It was necessary to describe the extent of development of head tubercles as well as their distribution

and numbers. For convenience, four stages were

recognized in tubercle development, the spot, bud,

nuptial tubercle, and scar. Spots (tubercle "anlagen")

appear as small, round, light to gray areas in the

deeper epidermis. Spots appear first in juveniles and

with further development they increase in diameter,

become lighter with a darkened center and approach

the epidermal surface. Tubercle buds arise in the

center as small pimples, having rounded or pointed

tips. Nuptial tubercles represent the completed

growth stage following budding (Figure 2, see also

Wiley, 1969, and Wiley and Collette, 1970). They

are hard, cornified structures that attain their greatest development more or less synchronously with

the spring reproductive period of the mature males.

In life the nuptial tubercles reflect a pearlescent hue

particularly at the peak of the reproductive cycle.

Tubercle scars are shallow to moderately deep, discolored cavities resulting from the loss of the nuptial

tubercles. Unless specifically indicated, data refer-
SMITHSONIA

N CONTRIBUTIONS TO ZOOLOGY

SNA SPO

ring to the numbers and distribution of tubercles

may represent any of these four stages.

Tubercle numbers refer, unless otherwise stated,

to the total number of tubercles or their counter-

parts developed on the head. Recently developed

spots often contrast poorly with the head coloration

and may often be difficult to observe. A film of water

applied to the head while counting appreciably

aided in discerning the spots. The absence of an

opening in the spots distinguishes them from ce-

phalic pores which are often superficially similar in

appearance. Tubercles, scars, and buds are easily

observed.

Difficulty was encountered in obtaining accurate

counts on specimens with high numbers of crowded

tubercles. Counting particular tubercles twice or

skipping some was avoided by subdividing the head

into small quadrants. The quadrants were gently

lined with a draftsman pen using a solution of

alizarine in ethyl alcohol after the head was dried

of excess moisture. This stain dissolved from speci-

mens within about one week.

The areas and lines of the head referred to in

discussions of development and distribution of tu-

bercles are indicated in Figure 1 b, c. The interorbi-

tal and occipital areas are subdivided in order to

adequately describe details of tuberculation. The

following areas are designated: snout, anterior to

line AIN; internasal, between lines AIN and PIN; pre-

interorbital, between lines PIN and AIO; anterior

interorbital, between lines AIO and MIO; posterior

interorbital, between lines MIO and PIO; anterior

occipital, between lines PIO and AOC; mid-occipital,

between lines AOC and MOC; posterior occipital, be-

tween lines MOC and POC. The subnasal area is over

the lachrymal bone.

The posterior distribution of tubercles was desig-

nated as the line nearest to the most posterior tu-

bercle along the midline area of the head. The

absence of tubercles from the snout in the approxi-

FIGURE 1.—Heads of Nocomis showing areas and lines pertain-

ing to breast squamation and tuberculation of head: a, degrees

of breast squamation (see page 3, methods); b, head tuber-

culation, dorsal areas; c, head tuberculation, lateral areas.

AIN—anterior internasal line; PIN—posterior internasal line;

AIO, MIO, PIO—anterior, mid and posterior interorbital lines,

respectively; AOC, MOC, POC—anterior, mid and posterior occip-

ital lines, SNA—subnasal [=lachrymal] region; PNA—posterior

nasal region; SPO—suprapostorbital position.
NUMBER 85

mate area from the AIN line to slightly anterior to

this line is regarded as a hiatus.

The terminology "nuptial crest" and "swelling"

is one of degree. A nuptial crest is a large swelling

on the head extending from the internasal area to

the nape, where it often descends abruptly, and resembles a "cocks comb" in profile (Figures 25a,c, 25).

A "swelling" may represent the early stages of a

crest (Figures 10&, 236) or it may remain a slightly

swollen area on the dorsal part of the head. The

swelling generally begins its development between

the internasal and interorbital area.

Life colors were observed in the field from living

specimens and/or from color photographs taken

from freshly killed specimens. These colors are distinguished in the text from colors recorded from

freshly preserved specimens.

The synonymy pertains to those references dealing with the species of Nocomis from the central

Appalachian region and where it is pertinent to an

understanding of the species. A complete synonymy

is not possible, for much of the nomenclature in the

early literature of the area refers to Nocomis (or

Hybopsis) kentuckiensis, an unidentified nominal

form. No specific allocation can be made without

the original material because the descriptions are

not adequate and illustrations are wanting.

The distribution maps (Figures 19, 20, 28, 29),

show all localities involving the new species. Only

those localities of N. micropogon and N. leptocephalus occurring in the drainages with the new species

or in adjacent drainages were plotted. These localities are pertinent to an understanding of the geographic distributions and stream captures, and ecological relationships among populations between

and within respective species. When more than one

collection was taken from a specific locality, only

one was indicated on the map.

Distances pertaining to localities are given in airmiles (airmi) or roadmiles (rdmi), but where it was

not stated by a collector the most likely measure

was determined from maps and/or by some indication of the collector's field route, activities, and

notes. The geographic locations of collections plotted on distribution maps were checked with the

United States Geological Survey topographic and

state maps and route maps of states and counties.

A map tracer was user whenever needed for determining localities given in roadmiles and for measuring stream length.

Descriptive accounts of particular specimens or

populations are included in the text, as necessary

for an understanding of certain special aspects of

FIGURE i.—a, Section of a large tubercle from the nuptial crest

of N. leptocephalus, 147 mm, from the South Fork, Roanoke

River, Montgomery County, Virginia, captured in May 1964.

The section is 2 mm high from basement membrane to tip,

and 2.3 mm wide. The conical keratinized cap is composed of

flattened cells forming laminae that separate in preparation.

The epidermal core of epithelial cells is penetrated by a series

of vascularized dermal papillae. The longer papillae have a

series of whorled, spiraling capillaries supported by collagenous tissue and some germinal epithelial cells. Parts of several

loops are visible in the section. (Section prepared in Mallory's

triple stain.) b, Section of a tubercle from the pectoral fin of

a nuptial N. biguttalus, KU 7295, 122 mm, collected 14 June

1962 from the Pecatorica River. (Both photographs taken by

Martin L. Wiley).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the distribution, dispersal, ecology, or sympatric occurrence of the species. Detailed accounts are given

of variation in morphology and of the coloration

of all size stages, for this information is necessary

for proper identification of living specimens that

are observed in the field in studies of reproduction

and behavior, nest associations and interrelationships, and of hybridization.

MATERIALS.—A great portion of the material

examined from the critical area of western Virginia

was taken in recent years during survey studies of

several major drainages by Robert D. Ross, his associates, and students of the Virginia Polytechnic

Institute, Blacksburg, Virginia. Most of these collections were made using an alternate polarity

electroseine with about thirty feet of wire between

terminals and with a 115-230 volt AC 2500 Watt

generator. This device is effective in obtaining large

series of chubs, including the adults from the deeper

waters and beneath boulders and banks, and in

obtaining a more qualitative and quantitative estimate of local populations. Most of our material

from West Virginia was collected or acquired by

Frank J. Schwartz, Institute of Marine Sciences,

University of North Carolina, Morehead City, North

Carolina, and Robert Sumner, Fish Division, Department of Natural Resources, Lewisburg, West

Virginia. The several thousand specimens of Nocomis taken by these individuals in their surveys are

now housed largely at the National Museum of

Natural History.

Other collections in the following institutions

taken mainly over the past half-century by individuals or survey parties using various sized nets, electric shocking devices, poisons particularly at preimpoundment stations, angling, traps, etc., have

been examined: Auburn University; Field Museum

of Natural History, Chicago (CNHM); Cornell University (cv); Duke University (DU); Indiana University (Ind. U.); Kentucky Department of Fish and

Wildlife (KFW); University of Louisville; University

of Michigan Museum of Zoology (UMMZ); University of Minnesota; National Museum of Natural

History (USNM); Oklahoma State University Museum of Zoology (OSUMZ); University of Richmond

(UR); Stanford University, Division of Systematic

Biology (su), recently transferred to the California

Academy of Sciences.

It is estimated that more than 50,000 specimens

have been examined from these institutions, contributing either directly or indirectly to this study.

Locality data for collections pertaining to the new

species are listed in the materials section of the

respective species. Only the institutions and number

of collections from which morphological data were

taken are listed for N. micropogon and N. leptocephalus. Many other collections of these two species

examined in the above institutions provided an

enormous amount of distributional data. These collections are not listed, but the localities are plotted

on the distribution maps when they occur within

the treated area.

Acknowledgments

It would be difficult to list the many individuals

who have contributed to this study through their

field observations, collecting efforts, and museum

aid. We are grateful for this cooperation and we

wish to acknowledge the special effort made by the

following individuals for their prompt assistance in

the loan and exchange of collections or in providing

pertinent information: Joseph R. Bailey, Duke University; Walter R. Courtenay, Jr., Florida Atlantic

University; Reeve M. Bailey and Robert Rush

Miller, Museum of Zoology, University of Michigan;

James E. Bohlke, Academy of Natural Sciences,

Philadelphia; William M. Clay, University of Louisville; Daniel M. Cohen and Bruce B. Collette, Systematics Laboratory, U. S. Fish and Wildlife Service,

Washington, D. C; Royal D. Suttkus and Glenn H.

Clemmer, Tulane University; Edward C. Raney,

Cornell University; John Ramsey, Fisheries Research

Unit, Auburn University; Robert D. Ross, Virginia

Polytechnic Institute; Frank J. Schwartz and Robert

Sumner.

The following present and former staff contributed most of the illustrative material: Carolyn Bartlett Gast and A. M. Awl, detailed drawings of

specimens; Fanny Phillips, diagrammatic figures

and graph; Janet L. Robinson, two scatter diagrams;

Jack Scott and James Griffen, photographs. Douglass

M. Payne, Cornell University, took certain photographs. Glenn H. Clemmer drafted the base map of

the Blackwater River. Other drawings and photographs were made by Martin L. Wiley and the

authors. Martin L. Wiley read the manuscript and

provided valuable suggestions for its improvement.
NUMBER 85

The second author was able to pursue these

studies, including field work, in the fall of 1964 and

through July of 1965 with aid from National Science

Foundation Grant No. 23395 to Dr. Edward C.

Raney, Cornell University. Ernest A. Lachner received support from the Smithsonian Institution

Research Awards, Nos. 3316 and 3347, and the Office

of Environmental Sciences, Smithsonian Institution.

Diagnostic Characters

The important characters useful in differentiating

among the several specific or infraspecific populations are discussed below. Group characters are

largely omitted from this discussion.

HEAD TUBERCULATION, HEAD CRESTS, AND SWELLINGS.—

The distribution, number, size, and developmental

patterns of the head tubercles represent the most

critical characters in the separation of the species,

and they also represent important evolutionary

characters.

Two developmental stages, as defined in the

Methods section (p. 3), precede the attainment

of enlarged head tubercles. The first stage is the

tubercle spot. These first appear, depending on the

species, in juveniles of both sexes from between 30

to 80 mm SL. At first they are barely discernible and

remain so in juvenile males and juvenile to adult

females of some species, while in other species they

become conspicuously enlarged. The second stage is

the tubercle bud, a raised area in the center of a

tubercle spot which occurs in males of all species

and sometimes in females of certain species. The

buds vary in texture and size from relatively small

and moderately soft to large and well cornified, the

latter advanced stage approaching the definitive

adult tubercle in development. Individual specimens are often found with both spots and buds, and

in these cases the buds apparently grow from the

earlier developed spots. Some species develop nearly

their adult complement of tubercle spots or buds

when juveniles. Other species show a progressive

increase in tubercle numbers and distribution over

the head with increase in growth throughout life.

The number and growth patterns of tubercles are

shown in Tables 1-7, 9, 10, 13, 19 and Figure 4. The

incipient species differences in adult tubercle number and distribution are evident from the early developmental stages in juveniles (Figure 3). Many

specimens of both sexes are specifically identifiable

at these stages by their spots and/or buds. Differences also occur in the relative frequency of occurrence of tubercle buds in size classes among the

species (Table 4).

FIGURE 3.—Typical early patterns in the development of tubercles and their size in three species of

Nocomis. a, N. leptocephalus, 66 mm female, James drainage; b. N. micropogon, 85 mm female,

Tennessee drainage; c, N. raneyi, 99 mm male, Roanoke drainage.
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Breeding tubercles develop on the head and the

dorsal surface of the pectoral fin in nuptial males

(Figures 9, 10, 11, 16, 22-27). They attain the height

of their development during or slightly before the

spring spawning period, coincident with the development of nuptial coloration and head crests or

swellings. It is not known if the tubercle "anlagen"

increase in number and size at different rates during

certain seasons in the females and immature males.

They probably do not increase in number in mature

males during the reproductive season since spots

were never seen on tuberculate males or on postnuptial males soon after the reproductive season.

Tubercle size is related to age or maturity since

tubercles average larger in larger males. In N. micropogon some overlap occurs between the size of

small tubercles of breeding males and large buds of

nonbreeding males. There is a consistent difference

in size of tubercles between the species although

considerable variation exists within species.

Tubercles were never found to be fully developed

in females. The female Nocomis is not known to

participate in nest-building nor to share any of the

reproductive activities of the male that may necessitate highly developed tubercles. Ramaswami and

Hasler (1955:65-66) induced growth of tubercles in

females of N. biguttatus to well-developed stages by

testosterone injection.

Tubercles in Nocomis possibly serve several functions. Since these white to gray-colored epidermal

structures are highly conspicuous, they may act as a

warning mechanism to individuals of the same and

other species, including potential egg predators,

that infringe upon the active, nest-building male.

Tubercles provide a formidable armament for the

male in his defense of the nest, accomplished by

butting an intruder with the head. Many breeding

males with well-developed tubercles are found with

fresh, moderately deep cavities on the head, indicating that tubercles were lost by force. In this respect, the large, nuptial crest or swelling, which is

generally best developed in the largest males, may

serve to cushion the brain from such impact. This

is the best use we can presently conceive of as a

possible function of the crest or swelling. The shining tubercles may attract females to the nest, and,

since the patterns in which they develop differ among

the species, they may also function as an aid in

species recognition. The tubercles on the side of the

head, generally all of which are supported beneath

by the lachrymal bone, and possibly those dorsolaterally on the head, may aid in maintaining some

contact with the female during the spawning act, as

do the pectoral fin tubercles. Evidence that male

head tubercles function in the stimulation of the

female through contact was given for a species of

Notropis by Koehn (1965:466) and a species of

Etheostoma (Percidae) by Braasch and Smith (1967:

7, fig. 3). There is no evidence for this function in

Nocomis.

The head tubercles are deciduous on postnuptial

males. Those that are not lost from agonistic behavior apparently fall off shortly after the spawning

period. Some tubercles in preserved specimens appear as if they had crumbled apart. Sites once covered by tubercles are scarred and, being evident,

are of taxonomic importance as are the other stages

of tubercle development. Many of the largest and

oldest males die soon after spawning. These specimens are sometimes seen drifting slowly downstream

before death or are found dead on the stream bottom, with the previously tuberculated areas and

other injuries infected with water fungus, Saprolegnia. Tubercle scars of surviving males apparently

heal during the summer. Some scars may have a

central "core," a remnant of the lost tubercle, or,

sometimes a bud may be present. By late summer,

the larger adults of certain species usually have buds

in varying growth stages and all species show no

evidence of fresh scars.

M ERISTIC CHARACTERS.—The following meristic

characters are tabulated in Tables 8, 11, 12, 14-16:

lateral-line scales, body circumferential scale rows,

rows of scales above and below the lateral line,

caudal peduncle scale rows, pectoral fin rays, and

vertebral numbers.

The species differ little in the above meristic

characters although certain average values and

ranges of counts are particularly useful in distinguishing and relating various populations. The most

useful characters are the circumferential scale rows

around the body, and vertebral numbers. Scale

counts above and below the lateral line and along

the lateral line correlate in numbers within species

with the circumferential count, but the differences

are of somewhat lesser magnitude.

Geographic variation in these meristic characters

is apparent and, in a few cases, we have found
NUMBER 85

2 no

a •••

x -

A _

N. rane/i, Atlantic dr.

N. platyrhynchus, New R.

N. micropogon, Potomac R.

N. micropogon/ James R.

N. micropogon, Tennessee R.

N. leptocephalus, Atlantic dr. and New R.

Standard length in mm.

FIGURE 4.—The relationship between body length and number of head tubercles in four species

of Nocomis of the central Appalachian region. Three populations of the river chub, N. micropogon, are compared from the Potomac, James, and Tennessee Rivers. The curves were fitted to

mean values by inspection. The data for both sexes were combined. The mean values were plotted

from data in Tables 5-7, 9, 10, 13, based on a total of 1712 specimens.

differences among populations within a single drainage. Infraspecific differentiation is discussed in each

species following the descriptive data.

MORPHOMETRIC CHARACTERS—Lachner's (1946:

207-208) key to three species of Nocomis contained

morphometric characters. Raney (1950:161), Moore

(1957:109), and Ross (1959:12) and others employed

a modification of this key. We have reevaluated the
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

characters used by these authors and surveyed additional ones with respect to possible allometric

growth.

Many morphological characteristics of the head

and body among the species were compared in a

preliminary manner. While notable differences were

observed, it soon became apparent that there were

considerable allometric growth changes extending

12.5

120

AON . RANEYI

N. PLATYRHYNCHUS

C • N. MICROPOGO N

150 170

STANDARD LENGTH in mm

FIGURE 5.—The relationship between standard length and gape width in three species of Nocomis.

The plots are based on 146 N. raneyi, 88 N. platyrhynchus and 151 N. micropogon taken from

specimens sampled over the ranges of the species. The regression equations are: N. micropogon,

y = .0294X + 6.24, n = 92; N. platyrhynchus, y = .0247X + 6.07, n = 103; N. raneyi, y =

.OO75X + 6.68, n = 96.

Tests of difference of slopes and heights (intercepts) of pairs of regression lines were performed

by analysis of covariancc (Snedecor, 1956:394-399). F values yielding probabilities greater than .95

were considered significant. The slope of the regression of gape width (as % of SL) on standard

length of N. raneyi is significantly different from N. platyrhynchus (F = 64.18) and N. micropogon (F = 6031). The slope of the regression lines for N. micropogon and N. platyrhynchus

do not differ significantly (F = 2.97) but the heights are different (F = 42.19).
NUMBER 8 5 11

in sizes among the young, juveniles, subadults,

and mature adult males. The range of a character

over these growth stages was so great that they

proved to be of little immediate systematic value.

Exceptions were found, such as the width of gape

and length of snout (Tables 17, 20, 21; Figures

5, 6, 7). These two characters proved useful in

distinguishing the two new allopatric forms when

each character was related with body size and with

each other.

Thirteen additional proportional characters are

presented for adult males in Table 25. Most of

these characters showed allometric growth but the

average values, when based only on adults, provide

useful indices that differ among the species.

INTESTINE.—Raney (1947:129) and Moore (1957:

108-109) reported that the leptocephalus group has

a looped intestine. This feature constitutes a principle diagnostic group character. Miller (1963:19-20)

cited several studies indicating that the Cyprinidae

do not possess a true stomach. We refer to the often

somewhat swollen anterior portion of the digestive

tract (from the posterior end of the esophagus to

the first intestinal bend) as an intestinal bulb. The

leptocephalus group has the ascending and descending portions of the intestine looped or whorled just

posterior to the second intestinal bend and ventral

to the intestinal bulb (Figure 8&). Other species of

Nocomis sometimes have a small to moderate lateral

loop but it is never extensively developed (Figure

8a). Specimens of the leptocephalus complex sometimes lack the looped intestine, particularly in certain southern races, and they thus may resemble

the micropogon group in this character.

13.5

12.5

A • C0 A _•• •

• • AA •

A • •& •* •

A * A

A A

A

• A I

AA

A AA

A A

O O

• O O O <D

o o o o o

o o o o o

• O GO CO OO <D

O O • O O

«• CO O O

CO <D OO O

OD /->

O O O &

• A •

• O» A O

• AO O

A * • O

• o o

OO

(DOO

o o o

OO O 0

o

o

A A

• A

A *

CD

o o o

CO

o

o o

O N. RANEYI

A N. PLATYRHYNCHUS

• N. MICROPOGON

STANDARD LENGTH in mm

FIGURE 6.—The relationship between standard length and snout length in three species of Nocomis.

The plots are based on 151 N. raneyi, 92 N. platyrhynchus, and 159 N. micropogon taken from

specimens sampled over the ranges of the species.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

BREAST SQUAMATION.—A summary of the degree

of breast squamation is given in Table 18 with the

data segregated by species and drainages. This character showed differences among the species, among

some drainage populations within a species, and

within species from various geographical areas.

COLORATION.—Life and alcoholic color patterns

showed important interspecific differences and relationships among the groups of Nocomis. Sexual

dichromatism and color changes associated with

growth and season, and local and geographic variation were studied for each species. Certain color

differences were evident; other color differences

were more subtly developed and noticed only after

handling a large number of specimens.

The brilliant nuptial coloration of the male, strikingly different from that of females and nonbreeding males, is very similar among the species of the

micropogon group. Differences were found between

the micropogon group and the leptocephalus group.

Individual differences in breeding color within each

species, other than that due to geographic variation,

appear to be related to proximity to the peak of

reproductive activities and to age or size. Differences also occur in coloration of females and nonbreeding males, particularly between the species

groups. The color changes with growth are slight

in juveniles, adult females and nonbreeding males

of each species. Geographic variation occurs in nupFIGURE 7.—The width of gape and contour of lower jaw in

three species of Nocomis: a, N. raneyi, 181 mm, James River

drainage; b, N. platyrhynchus, 183 mm, New River drainage;

c, N. micropogon, 185 mm, James River drainage.

FIGURE 8.—Comparison of intestinal looping: a, Nocomis

raneyi, female, 113 mm; b, N. leptocephalus, female, 86 mm.
NUMBER 85 13

tial coloration of N. leptocephalus and perhaps in

fin coloration of N. micropogon.

Sexual dimorphism is pronounced in all species

of Nocomis. The males at maturity, which may be

coincident with the breeding season, or several

months before, develop bright colors on the body

and fins. The young, juveniles, immature adults,

and adult females remain characteristically pale

colored. The nuptial males develop enlarged tubercles on the head and pectoral fins, and laterally on

the body in two species of the biguttatus group.

The adult females develop head spots approaching

the full complement of tubercles of the males. Nuptial males develop large head crests or swellings

(except the biguttatus group) whereas the females

show no trace of tissue enlargement. Adult males

acquire rounded head contours contrasting greatly

with the sharper lines of the young, juveniles, immature adults, and adult females. The solitary

nuptial male constructs a mound nest of gravel

that he transports in his mouth.

Genus Nocomis Girard

Nocomis Girard, 1856:190 (type species: Nocomis nebrascensis

Girard =. Semotilus biguttatus Kirkland by monotypy).

DIAGNOSIS.—Barbled cyprinids, the nuptial males

having the following characters: large breeding tubercules on the head (and on body in two species

of the biguttatus group), that show a phylogeny

among the species in their number, distribution and

size; expansive crest or swelling on head (crests

absent in biguttatus species group); elaborate breeding coloration on body of pink-rosy, orange and/or

blue. Nuptial males construct a mound-nest by

carrying stones in mouth. Body bicolored, dark

above, light below, with heavy dark scale margins

and a dark postopercular bar. Body size large, adult

males 100 to 250 mm SL. Mouth normal, moderate

to large in size, with one terminal maxillary barbel.

Scales large, lateral line almost always fewer than

44, body circumferential scales less than 37. Pharyngeal teeth 1,4-4,1 to 3-3. Scale radii in posterior field

only, total number in adults 23-54. Total vertebrae

number 38 to 43, precaudal vertebrae 21 to 24.

DESCRIPTION.—Body robust to moderately elongate or compressed. Morphometric growth rate generally moderately to markedly allometric. Head

moderate, 25-29 percent of SL, slightly rounded to

flattish above. Snout short to long, 8-14 percent of

SL, acutely pointed to well-rounded or blunt and

declivous. Mouth subterminal, moderately oblique

to nearly horizontal; frenum absent; lower jaw included; gape and upper jaw moderate to large,

7-12 and 7-11 percent of SL, respectively. Lips moderately thick. Single pair of small barbels occasionally absent; origin on flesh over posterior ventral edge of maxilla and sometimes slightly attached

to basal portion of lip angle. Orbit small to moderate, 4-8 percent of SL; position lateral to slightly

dorso-lateral.

Fins moderate in length; dorsal origin about

midlength of body, over or, usually, slightly posterior to pelvic origin; fin margins on dorsal and

anal straight, pectoral and pelvic moderately to

well rounded, caudal deeply to slightly forked.

Caudal fin lobes equal to moderately subequal,

upper lobe generally longest and more pointed.

Dorsal fin rays 8, and 7, pelvic 8, pectoral 15-19.

Pharyngeal teeth, 1,4-4,1 to 3-3, stout, tooth I or

I—III usually with small hook, grinding surface usually developed; tooth arch moderate to stout, pitted

surface well sculptured. Gill rakers usually 5-10,

short to moderate in length, very rarely with bifurcate tips. Pseudobranchae well developed. Intestine simple (S-shaped), or an accessory loop on left

side, or a single but much convoluted ventral whorl.

Peritoneum lightly to heavily pigmented with

melanophores.

Lateral line on body complete, almost straight

and horizontal to moderately curved; cephalic

canals complete, canaliculi well developed (Reno,

1967, 1969b, gives pore counts). Cutaneous taste

buds moderate in size, not densely concentrated,

present on lower head, snout, lips and barbel; compound buds on first pectoral fin membrane.

Scales with sub-basal focus; eccentric circuli

sharply angulate in antero-lateral field; radii numerous limited to posterior field; primary and secondary radii present.

Scales moderately large, the ranges in several

characters follow: lateral line 37-43, body circumference 27-38, caudal peduncle 16-20. Breast completely scaled to naked. Vertebrae 38-42.

Color (in alcohol) dark above, light to colorless

below; midlateral to dorsal scales with posterior and

anterior (scale pockets) margins dark; dark lateral

band and dark caudal spot well developed to absent.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Body color (in life) dorsally blackish, olive or tannish; laterally olive greenish, light tannish or light

yellowish. Fins olivaceous to reddish orange or color

almost entirely absent. Iris partially golden to redorange.

Sexual dimorphism and dichromatism highly developed in adult males; nuptial tubercles moderate

to large, erect, antrorse or curve laterad; tubercles

distributed on dorsal and lateral head, nape and

sometimes on lateral body; tubercles on pectoral

fin rays, 2-9, uniserial, retrorse.

Habitat is streams of sand, gravel, and rocky beds,

living near bottom.

Distribution primarily eastern and central United

States.

Three species groups, seven species, nine forms.

NOMENCLATURAL HISTORY.—The presence of a

barbel at, or very close to, the angle of the lip (and

other gross similarities) was the principle character

involved in the confusion of interpreting several

nominal American cyprinid genera. The taxonomic

rank of Nocomis vascillated between genus, subgenus, and group or section for over 100 years.

Nocomis was recently resubmerged into the synonymy of Hybopsis Agassiz (Bailey, 1951:192) with

seven other nominal genera: Yuriria Jordan and

Evermann, Erimystax Jordan, Platygobio Gill,

Macrhybopsis Cockerell and Allison, Extrarius Jordan, Oregonichthys Hubbs, and Couesius Jordan.

The weakness of this arrangement became evident

when it was learned that the barbel may be a weak,

variable systematic character and that other strong

group characters had been overlooked. Entire populations of the barbeled species Hybopsis (Hybopsis) amblops and Hybopsis (Erimystax) harperi may

lack barbels. Species with barbels also have been

recently assigned to Notropis (N. boucardi for one),

a characteristically nonbarbeled American cyprinid

genus. In barbeled genera not allocated to Hybopsis,

the barbel sometimes may be absent or reduced.

Current studies based on other characters have also

shown a close relationship between Hybopsis amblops winchelli, usually having barbels, and Notropis amnis, lacking barbels. As a consequence, a

number of workers have recently attacked these

generic problems by investigating new character

systems such as tuberculation, pigmentation, brain

patterns, cephalic lateral line sensory canals, and

various osteological features. A review of the nomenclatural history of Hybopsis is given by Reno

(1969a). A critique of the genus Nocomis with a

discussion of other North American barbeled cyprinids, particularly Hybopsis, will be presented in

part 4 of these studies (Smithsonian Contributions

to Zoology, Number 90).

The species of Nocomis, other than our new

forms, were all described by 1864. Species were referred by early workers to the genera Semotilus

Rafinesque, Hybopsis Agassiz, Leuciscus Klein,

Leucosomus Heckel, Nocomis Girard, and Ceratichthys Baird.

The genus Hybopsis and type species Hybopsis

gracilis (=amblops Rafinesque, 1820) were described by Agassiz (1854) but the presence or absence of a barbel was not mentioned. For this reason and because the descriptions were brief, the

identifications of these taxa were long uncertain.

Girard (1856:210-211, 1858:255) recognized Hybopsis and mentioned barbels in his accounts, but

based his generic description on species other than

the type of Hybopsis. Subsequently, such authors

as Cope (1864:279, 1868:230, 1869:379, 1870:460),

Gunther (1868:211), Jordan (1875:38, 1876:94,

1877a:45, 1877b:333, 1877c:27, 1877d:57) and Jordan and Copeland (1876:150) referred certain nonbarbeled cyprinids to Hybopsis. On occasion, the

barbeled species H. storerianus Kirtland was included. Later, however, Jordan (1878b: 426) and

Jordan and Gilbert (1833:212) aligned Hybopsis

with barbeled minnows, although they questioned

their assignment and recognized the junior name

Ceratichthys. Either Ceratichthys or Nocomis were

used rather than Hybopsis in Jordan (1878a:305,

1879:108, 1880:290, 1882:859, 1884:287) and C. H.

Gilbert (1884:203). Ceratichthys Baird and Girard

(1853, type species C. vigilax) was based on a North

American cyprinid currently placed in the genus

Pimephales Rafinesque, 1820. The taxonomic and

nomenclatural histories of Ceratichthys and Pimephales were reviewed by Hubbs and Ortenburger

(1929:35-38), Hubbs and Black (1947), and Cross

(1953).

Close relationships among the forms of Nocomis

were recognized by Cope (1869:364) and Jordan

(1878b: 426-427) who were first to place the nominal forms together and exclude all other groups

and species from one section of their expanded

genus of barbeled minnows. While the forms of
NUMBER 85 15

Nocomis were long thereafter included under a

broadened Nocomis, Ceratichthys, or Hybopsis, the

species of Nocomis apparently were considered to

constitute a distinct group of subgenus. The first

formal opinions that Nocomis is not congeneric

with these groups was based on scale studies by

Cockerell and Allison (1909:159, 161-163), Cockerell and Callaway (1909:189), and Cockerell (1913:

130). These authors apparently examined scales

from only N. effusus (called kentuckiensis) and

were not generally followed by subsequent writers.

We do, however, agree that the scales of Nocomis

are distinctive from those species once thought to be

close relatives. Nocomis and other groups with

which it was regarded as congeneric were elevated

to separate generic status by Jordan (1924) when

Hybopsis was (temporarily) reidentified with species

of Notropis. Hubbs (1926:27) and others also recognized Nocomis as a genus. Nocomis held generic

status until Bailey (1951:192) grouped it, with the

nominal genera listed above, under Hybopsis. Platygobio and Couesius had not been previously associated with the other seven taxa. There was discontent with this arrangement and since 1966 at

least six publications include Nocomis at the generic

level.

The taxonomic and nomenclatural confusion in

the early history of the species of Nocomis was

largely a result of the very close species relationships, their relatively similar general morphology,

and the failure by systematists to recognize the value

of certain secondary sexual characters, such as the

development of tubercles.

Although seven specific names were proposed

clearly for Nocomis by the end of 1864, three of

which are now applied to species and one to a subspecies, only one, Luxilus kentuckiensis Rafinesque,

1820, was generally recognized up until 1926, as a

senior synonym for all species of Nocomis. The

earliest alignments of kentuckiensis were with

species of the subgenus Cyprinella of Notropis

(analostanus, see Gibbs, 1963:521) and at least one

other species, Semotilus biguttatus Kirtland (1840a,

b). Jordan (1876) identified it with Notropis cornntus but later (1877c, 1882) stated that it was still

not satisfactorily identified. Soon after, Jordan

(1885b: 185) curiously stated in his Errata section

that (N.) biguttatus "seems to be the Luxilus kentuckiensis Raf." Thereafter, kentuckiensis was used

for species of Nocomis until Hubbs (1926:28) considered it to be unidentfiable. Both Hubbs (loc.

cit.) and Lachner and Jenkins (1967:558) gave reasons for regarding kentuckiensis as unrecognizable.

Hubbs (1926:27-29) recognized Nocomis as a complex of three species, biguttatus, leptocephalus and

micropogon, illustrated each, including features of

the nuptial males, and clarified their synonymy.

Semotilus biguttatus Kirtland (1840a,b) was described and figured from nuptial males, leaving no

doubt as to its identity. Its synonyms are Ceratichthys nebracensis Girard (1856), Ceratichthys stigmaticus Cope (1864), and Ceratichthys cyclotis

Cope (1864). The species leptocephalus and one of

its subspecies, bellicus, were described by Girard in

1856 under Ceratichthys and Nocomis respectively.

Ceratichthys micropogon dates from Cope (1864).

It is interesting to note that the supposed single

species of Nocomis (kentuckiensis) was frequently

stated to be the, or one of the, most widely distributed of North American cyprinids.

Catostomus melanotus Rafinesque (1820:302) as

with Luxilus kentuckiensis was briefly and poorly

described, based apparently on specimens from the

Ohio River basin. It was referred to Nocomis with

uncertainty, or in part, by Jordan (1876, 1877c,

1882, 1885a, 1885b), Jordan and Copeland (1876),

Jordan and Evermann (1896), and others. Hubbs

(1926:28) regarded melanotus as unidentifiable and

as likely a catostomid as a cyprinid. We concur with

Hubbs' decision. We have also examined the unpublished rough sketch of melanotus by Rafinesque

(Smithsonian Institution Library) but this afforded

no additional clue.

Recently, additional species and forms have been

discovered or recognized, and considerable information on the evolution and interrelationships with

the genus is known. Our studies indicate that

Nocomis is clearly divisible into three species

groups, each partly comprised of one of the species

recognized by Hubbs (1926). The differences

among the species groups are compared in Table 22.

Some characters in which they differ only slightly

are included since these appear to have some biological significance. The major group differences

and aspects of their evolution are discussed by

Lachner and Jenkins (1967) and herein in the

section on evolution.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species of Nocomis of the Central Appalachian Region

1. Tubercles present on snout anterior to internasal area and on subnasal area in one to three

rows in adults and larger juveniles; many tubercles on head, numbering more than 25 in

adults; intestine not whorled, occasionally with, at most, a small dorsad or laterad loop

along the anterior half of descending portion; scales on body smaller, circumferential scale

rows number 30 or more in about 94 percent of specimens sampled 2

Tubercles always absent on snout anterior to internasal area and always absent on subnasal

area; head tubercles greatly reduced in numbers, almost always fewer than 25; intestine

usually whorled or with an anterior loop extending to right side covering part of alimentary

tract; scales on body larger, circumferential scale rows number less than 30 in 72 percent

of specimens sampled. Bluehead chub, N. leptocephalus (Girard)

2. Adults typically with smaller and more numerous head tubercles extending in a relatively

uniform pattern from the snout along the midline to the occipital area; head tubercles extend

posteriorly on midline beyond the anterior interorbital area (AIO) in almost all specimens

over 100 mm; hiatus in tuberculation on snout in adults only occasionally present; tubercles

number about 60 to 200 in subadults to adults; scales smaller, average values of circumferential scale rows segregated by drainage populations are higher than 32.6 and for lateral

line scale rows higher than 40.1; more vertebrae, average values for drainage populations

range from 40.5 to 41.6 3

Adults typically with larger and fewer head tubercles, grouped in the area from the snout

to the interorbital area; head tubercles do not extend posteriorly on midline beyond the

anterior interorbital area (except some Potomac and Monongahela drainage specimens in

which a few tubercles extend to the posterior interorbital area) and almost always occur in

a V-shaped, anteriorly directed pattern; a hiatus in tuberculation almost always occurring

along midline of snout leaving a small isolated patch near tip of snout; tubercles number

fewer than 60 in almost all specimens, and appreciably fewer in the subadults (except the

Potomac River population, where slightly higher numbers occur) ; scales larger, averages

for circumferential scale rows range below 32.3 and for lateral line scale rows from 39.9 or

lower, when segregated by drainage populations; fewer vertebrae, average values for drainage populations range from 39.3 to 40.4 River chub, W. micropogon (Cope)

3. Mouth narrow (Figure 5), gape width almost always less than 95 percent of standard length;

snout short (Figure 6); body more compressed and deeper in the adults; dark lateral band

on body inconspicuous in preserved specimens of juveniles and absent in adults; breast usually naked, or partially scaled; tubercles on forehead crowded, those posterior to midinterorbital line equally or slightly more widely spaced than those anterior to line; pharyngeal

arch stout, the limbs short and broad Bull chub, IS. raneyi, new species

Mouth wide, gape width almost always greater than 9.5 percent of standard length in adults

and subadults larger than 130 mm; snout long; body not compressed and deep in the adults;

dark lateral band on body moderately or well developed in preserved specimens of juveniles

and adults; breast almost always well scaled; tubercles posterior to midinterorbital line

comparatively more widely spaced than those anterior to line; pharyngeal arch weaker, the

limbs longer and narrower Bigmouth chub, N. platyrhynchus, new species

The micropogon Group

DESCRIPTION.—Nuptial males develop a moderate

to large number of tubercles (Figure 4) on head,

dorsally, from tip of snout to midoccipital area;

tubercles erect or directed laterad, and moderate to

large in size; tubercles present over lachrymal in

one to three rows; a large more posterior nuptial

crest developed by older (larger) males; breeding

coloration of nuptial male rosy pink on lower head

and lower body, olive-greenish on upper body; a

wide, dark anterior and posterior scale border in

adults; dark caudal spot in juveniles small, indistinct; intestine almost always S-shaped; dentition

4-4; scale size intermediate, body circumferential

scales range from 28 to 39; body more elongate;

snout more pointed; posterior edge of preopercle

usually slanted anteriorly below; males attain

largest maximum sizes of all Nocomis; males of all
NUMBER 85 17

three species in group construct large mound-nests,

the largest in genus, using small to large size gravel.

The species include: N. micropogon, widely ranging

over much of northeastern United States, from New

York and the James River, Virginia, westward, not

reaching the Mississippi River; N. raneyi, of five

central Atlantic slope drainages, and N. platyrhynchus of the New River drainage.

Nocomis raneyi, new species

BULL CHUB

FlCURES 7, 9, 10-17

Hybopsis kentuckiensis —Jordan, 1889:123 (Roanoke dr. [in

part], 126 (Tar dr.), 129 (Neuse dr. [part]).

Hybopsis species.—Robins and Raney, 1956:31.

Hybopsis (Nocomis) species 2.—Reno, 1969b:739.

Nocomis micropogon.—Burton and Odum, 1945:186 [part].

Nocomis species.—Raney, 1950:184.

DIAGNOSIS.—Differs from the other micropogon

group members in: mouth small (Figure 5); snout

short (Figure 6, Tables 17, 20, 21) ; body deep, more

compressed laterally in adult males; head tubercles

in adults often more than 100 (Table 5) and densely

developed from tip of snout to interorbital area;

caudal fin more sharply forked; pharyngeal tooth

arch stout with short limbs (Figure 18).

DESCRIPTION.—Morphometry: Proportional measurements expressed in thousandths of the standard

length are given in Table 23 for the holotype, 5

male paratypes, 53 to 227 mm, and 5 female paratypes, 68 to 180 mm. Because of the allometric

growth, additional proportional data are given in

Table 25 for 13 characters of 24 mature males

having a mean length of 200 mm and ranging from

172 to 242 mm. N. raneyi is compared in these characters with the adults of the three other chub

species from the study area. Many proportions increase with increase in body length. The diameter

of the orbit is an exception, decreasing with increase in body length, a characteristic of all known

species of Nocomis. A number of morphometric

values may appear higher in males than in females,

but when plotted against body length there are no

appreciable differences. The higher values in the

males relate only to the larger body size attained by

them.

The head is pointy and the body slender in the

young and juveniles of N. raneyi, resembling N.

FIGURE 9.—Head of male Nocomis raneyi showing distribution

of tubercles, tubercle scars, and contour. Paratype, USNM

166404, 229 mm, captured in Roanoke River at Salem, Virginia

on 21 June 1946: a, lateral view; b, dorsal view. (Drawn by

A. M. Awl.)

platyrhynchus and unlike the more robust body and

deeper head of N. leptocephahts (Figure 17). As the

males approach maturity the facial features become

greatly modified, the forehead becomes deep and

rounded and the body somewhat slab sided (compare Figure 17, juveniles; Figure 16, tubercle-bud

stage in male; Plate 14, adult female in tuberclespot stage; Figures 10, 12, 15, nuptial tubercle stage

of breeding male). In the development of the facial

features at maturity N. raneyi resembles N. platyrhynchus more closely than N. micropogon or N.

leptocephalus.

Two morphometric characters, although showing

allometric growth, proved significant in segregating

a majority of specimens of N. raneyi from its two

species cognates in the micropogon group; these

were gape width (Figures 5, 7) and snout length

(Figure 6), when related to body length. The relationships of gape width plotted against standard
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

length (Figure 5) is based on 146 N. raneyi, 88 N.

platyrhynchus, and 151 N. micropogon from specimens of all sizes, sexes combined, and sampled over

FIGURE 10.—Lateral view of heads of two species of Nocomis.

Note uniform size of tubercles and their presence on snout

and subnasal areas. Nuptial crests are in different stages of

development, a, N. raneyi, USNM 194625, 232 mm, taken

23 May 1964, Craig Creek, James drainage, Virginia; b, N.

raneyi, USNM 197684, 198 mm, taken 28 June 1963, Roanoke

River, Virginia; c, N. platyrhynchus, TU 25902, 195 mm, taken

19 June 1962, South Fork New River, North Carolina.

the ranges of the species. The tests of difference

of slopes and heights (intercepts) of pairs of regression lines were performed by analysis of covariance (Snedecor, 1956:394-399). F values yielding

probabilities greater than .95 were considered significant. The slope of the regression of gape width

(as percent of SL) on standard length of N. raneyi

is significantly different from AT. platyrhynchus

(F=64.18) and N. micropogon (F=60.51). The

slope of the regression lines for N. micropogon and

N. platyrhynchus do not differ significantly

(F=2.97) but the heights are different (F=42.19).

The regression equations are N. micropogon,

y=.0294X+6.24, n=92; N. platyrhynchus, y=

.0247X4-6.07, n=103; N. raneyi, y=.0075X+6.68,

n=96. The same collections were used, as above in

plotting snout length against standard length for the

same three species (Figure 6). The consistently

shorter snout (average) is shown for any respective

body length in N. raneyi, although the differences

are not of the same magnitude as in the gape width.

The influence of increase in body length on the

gape width and snout length is shown in Tables 17,

20 and 21 for N. raneyi and N. platyrhynchus, the

FIGURE 11.—Variation in tubercle development in Nocomis

raneyi and N. platyrhynchus: a, N. raneyi, USNM 194625,

232 mm, taken 23 May 1964, Craig Creek, James drainage.

Tubercles widespread in interorbital and anterior occipital

areas and crowded on snout, b, N. raneyi, USNM 194606, 211

mm, taken 5 July 1961, Blackwater River, Roanoke drainage.

Slightly reduced tuberculation is shown in interorbital area

compared to specimen on left. Tubercle size smaller than in

most N. raneyi. c, N. platyrhynchus, CU 47590, 198 mm, taken

3 May 1964, Toms Creek, New drainage. Only a few tubercles

are present in the interorbital area and a hiatus is nearly

present on snout, d, N. platyrhynchus, USNM 194819, 200 mm,

taken 10 May 1964, New River, Virginia. A slightly greater

number of tubercles is shown in the interorbital and snout

areas compared to specimen on left.
NUMBER 85 19
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

specimens divided into three size groups, 50 to 99

mm SL, 100 to 149 mm SL and 150 mm SL and

larger. When gape width is plotted against snout

length for each specimen, only 60 percent separation

occurred in the smaller sized group and about 88

percent in the two larger groups.

The preopercle-opercle suture, as it approaches

the angle on lower head, is directed forward in almost all specimens of N. raneyi. In a sample of 400

specimens from three drainages, the James, Roanoke, and Chowan, the suture was considerably

directed forward (an angle of 10 to 20 degrees) in

75 percent of the sample; 20 percent with suture

slightly directly forward; and 5 percent with suture

perpendicular or posteriorly directed in respect to

the horizontal axis of body.

The fin shapes among the species of the micropogon group are relatively similar (Figures 12, 13,

14, 17, 25) except the caudal fin in N. raneyi, which

is more deeply forked, especially noticeable in the

subadults and adult females.

The intestine is simple, elongate and S-shaped,

frequently with an accessory loop (47 of 49 specimens from over its range). This organ differs greatly

from that of N. leptocephalus, in which it is

whorled and extends under the intestional bulb

(Figure 8).

The following are other general features of head

and body. The dorsal contour of the head on specimens without cephalic swelling is almost straight to

gently curved and slopes slightly to moderately

downward from the occiput to the internasal region,

more so in the anterior portion. A slight to moderate dip in contour, sometimes causing a small

hump, is usually present in the internasal region,

except on some of the large males. The snout anteriorly is rather short and subconical or moderately

rounded. On some specimens the tip of the snout

appears more elongate and/or acuminate. The head

ventrally is almost straight and slightly less oblique

to the horizontal axis than the dorsal contour of

the head.

The mouth is small in the young to moderate in

size in adult males; it is oblique and subterminal,

being slightly to moderately overhung by the snout

tip. The lower jaw is included. The jaw angle is

positioned at or slightly posterior to a line ventrad

from the posterior nostril. In lateral view the upper

lip is generally oblique, being slightly to moderately

slanted downward; however, in some specimens the

anterior half of the upper lip is directed slightly

above the horizontal, the posterior half directed

below. The lower lip is straight and directed somewhat downward. In ventral view the lips are semicircular or they are round anteriorly, the sides diverging, straight, or curving medially. The lateral

portion of the upper lip extends to the ventral head

margin except at the jaw angle where it is slightly

less than the head width at that point. The premaxilla is protractile and a deep groove is present

around the entire lips except along the anterior portion of the lower lip where it is absent. The lips

are fleshy, appearing more enlarged in nuptial

males. The single pair of terminal maxillary barbels

FIGURE 12.—Crested, nuptial male of Nocomis raneyi, USNM 194633, holotype, 242 mm, taken

29 May 1964 from Craig Creek, James River, Virginia.
NUMBER 85 21

are short and sometimes not excised from the deep

groove behind the jaw angle, and are abundantly

supplied with taste buds.

The eye is small, the fleshy orbit is round or

slightly obliquely or horizontally elongate. The

lachrymal bone is oblique to the horizontal, especially in adult males, and reaches the upper lip just

above which an obvious crease in the skin is present

along its anterior end. The branchiostegal membranes are united to the posterior isthmus, the distance between the points of insertion of the membrane are about half the gape width. The epidermis

in the interbranchial isthmus area is longitudinally

folded; below the mouth it is folded at right angles

to the longitudinal axis.

The body is elongate and subterete, slightly compressed in subadults to moderately compressed in

nuptial males, widest and deepest midway between

occiput and origin of dorsal fin. The back is slightly

to moderately elevated above the occiput level,

highest at the dorsal fin origin or just anterior to

this. The predorsal body contour is straight or

slightly convex, somewhat oblique to the horizontal

axis, and is continuous with the posterior head dorsum contour, or a slight abrupt rise to the body

level is sometimes present at the occiput. The postdorsal contour is straight and slopes slightly more

downwardly than the predorsal contour; it is upturned beginning just anterior to the procurrent

caudal fin rays. The dorsal fin base is straight but

usually more oblique to the horizontal axis than

the postdorsal contour. The ventral body contour

is gently rounded, generally not deep, from the

isthmus to the anal fin origin, straight and moderately directed upward at the anal fin base, and

then almost straight and sloping slightly upward to

a point just anterior to the procurrent caudal fin

rays where it is downwardly curved. The caudal

peduncle is moderate in length and depth, rather

narrow in width.

The lateral line is complete, slightly downwardly

curved anteriorly, and straight posteriorly. The line

begins on the body at the posterior operde insertion, slopes quite obliquely downward for a short

distance over the cleithrum to a point slightly above

the midlateral level, then slopes slightly downward

through the first seven to nine vertical scale rows to

the midlateral level or to a point slightly below

this; in the latter case it slopes gently back upwards

and attains die midlateral level at a point usually

just posteriad to a vertical line from the dorsal fin

origin. Upon attaining the midlateral level it is

rather straight and horizontal to the posteriormost

scale overlying the bases of the midcaudal fin rays.

The urogential papilla in both sexes is fleshy and

moderate to rather large in size, the length about

1 to 1.5 times the greatest width. It changes in

shape from bulbous to subconical with a small oriFICURE 13.—A postnuptial male Nocomis raneyi (Paratype, USNM 166404, 229 mm), with tubercles

and tubercle scars on head, captured in the Roanoke River at Salem, Virginia on 21 June 1946

[hypoid area somewhat downflexed due to specimen previously being on stringer]. (Drawn by

Carolyn Bartlett Cast.)
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—An adult female Nocomis raneyi, USNM 191036, 179 mm, captured on 18 September

1959 from Contentnea Creek, Neuse drainage. The tubercle "spots" are evident in specimens of

this size. The pointy snout is characteristic of the adult females and of both sexes when juveniles.

(Drawn by Carolyn Battled Gast.)

fice in the adults. Much of the papilla rests in a

depression before the origin of the anal fin, being

more excised from this in larger specimens.

The entire body, except the breast, is scaled.

Laterally, the exposed portion of the anterior scales

is higher than long, especially just behind the cleithrum where they appear diamond shaped, although

the posterior margin is somewhat rounded. The

exposed portion of the lateral caudal peduncle

scales is about equal in height and length, appearing square. The posterior margin of lateral line

scales is usually slightly to moderately indented.

The dorsal and ventral scales are quite variable in

size and shape; anteriorly they tend to be smaller

and imbedded.

Cutaneous sense organs are present over the entire

head, laterally and dorsally on the scales of the first

one or two vertical scale rows, ventrally back to

about the anal area, and on the upper and lower

surfaces of the interradial membranes of the pectoral fin. They are largest on the head ventrally,

especially on the lips.

The alimentary tract is a simple loop, the longitudinal lengths of the tract are straight or slightly

curved. The intestinal bulb extends posteriad on

the right side to just before the vent and, unless

full of food, is only slightly wider than the intestine.

The intestine then bends sharply mediad and anteriad to just behind the pericardium (the ascending intestine), and it sometimes bends sharply to

the left side and posteriad to the vent (the descending intestine). Usually, instead of the ascending portion bending directly into the descending portion,

an additional short loop, about 10 to 30 percent

(sometimes to 50 percent) the length of the descending portion, occurs on the left side at the end of the

ascending intestine and lateracf or dorsad to the

descending portion into which it leads. Sometimes

a short anterior length of the ascending and descending portions is "kinked" laterad to the right

side.

The fins are moderate in length except the slightly elongate caudal. The margin of the pectoral fin

is moderately angulate to round. The free margin

is angulate from the tip of rays 1 to 10 (the apex at

ray 4, the longest ray) or moderately rounded between these rays; it is then broadly rounded along

the tips of the much shorter posterior rays to the

posterior base. The margin of the pelvic fin is broadly rounded or almost straight with rounded corners,

ray 3 or 4 being the longest. When appressed, the

pelvic fin may reach or is quite short of the urogenital papilla base. The tip of the anal fin, anteriorly,

is slightly or well rounded to its longest ray (ray 2

or 3), then straight or slightly incurved to the last

ray. The origin of the dorsal fin is positioned well

anterior to the midbody point and usually over the

origin of the pelvic fin, sometimes slightly posterior

to the latter. The anterior tip is pointed or slightly

round; ray 1 or 2 is longest and falls short of the
NUMBER 85 23

posterior rays when appressed. The margin of the

dorsal fin is usually straight, sometimes slightly incurved. The caudal fin is well forked, the fork apex

is broadly rounded or very obtusely angulate. The

lobes are always subequal in length; the upper lobe

always longer and its tip usually narrower. The tip

of the upper lobe in young and juveniles is pointed

or round, in adults it is acuminate. The lower lobe

is broadly rounded in young and usually quite

pointed in adults. The well-developed procurrent

caudal rays form a distinct middorsal and midventral ridge on the posterior caudal peduncle.

Meristic characters: Values for squamation counts

for N. raneyi are close to N. platyrhynchus and they

are somewhat intermediate between the lower values

usually for N. micropogon and N. leptocephalus

and higher values of N. effusus (Lachner and Jenkins, 1967). The range of the mean values for N.

raneyi taken usually from five drainages, with the

count of the holotype in parentheses, follow: circumferential scale rows (Table 12), 32.9 to 33.4

(35); lateral line scale rows (Table 8), 41.0 to 41.6

(41); scale rows above lateral line (Table 11), 7.3 to

7.7 (8); scale rows below lateral line (Table 11), 6.1

to 6.4 (7); caudal peduncle scale rows (Table 14),

16.1 to 16.7 (17); pectoral fin rays (Table 16), 16.8

to 17.1; total vertebral numbers (Table 15), 40.9 to

41.6.

Assigning numerical values for the degree of

breast squamation (Table 18) was useful in demonstrating specific differences in the micropogon group

as well as showing certain racial differences. N.

raneyi almost always has a scaleless breast in the

James, Chowan, and Roanoke Rivers, whereas it is

about 50 percent scaled in the Tar and Neuse

Rivers.

Tuberculation: Data pertaining to tubercle development, distribution, and numbers is summarized

in Tables 1, 3, 4, 5. Early tubercle development is

shown in Figure 3c. The relationship between increase in tubercle numbers with increase in standard

length is shown in Figure 4. The distribution of

head tubercles is illustrated in Figures 9, 10, 11, 12

(holotype), and 16 for immature specimens and

nuptial adults.

The tubercle pattern and the relationship between increase in tubercle numbers with increase in

body length is similar in both sexes. At first inspection this sex similarity may not appear correct beFIGURE 15.—Dorsal profile of Nocomis raneyi, holotype, USNM

194633.
2

4 SMITHSONIA

N CONTRIBUTIONS TO ZOOLOGY

cause the tubercle spots are difficult to discern in the

females (at most, they develop only to a fine bud

stage), except in the largest specimens, and initial

trials at counting will give lower improper values.

Also, the males attain a larger size and a greater

average number of tubercles. The tubercles first

appear in N. raneyi as small, light spots, often 15 or

more, and extend from the anterior internasal line

to the midinterorbital line (Figure $c). They rarely

commence development in specimens under 60 mm

SL, and they are often not discernible at lengths of

60 to 85 mm SL. With increase in body length, the

tubercles develop posteriorly on the head, but almost

always stop development at the posterior interorbi-

tal line (Table 1). The snout and subnasal tubercles

are last to develop in N. raneyi. The full comple-

ment of tubercles, which may exceed 200 in nuptial

males, extends from the tip of snout (Figure 9) to

the posterior interorbital area, and includes a supra-

orbital ring and two or three irregular rows on the

lachrymal area. The tubercles are closely spaced

from the anterior interorbital to the tip of snout.

N. raneyi differs in the latter character from N.

platyrhynchus, which sometimes shows reduced tu-

berculation before the anterior internasal area, and

differs considerably from N. micropogon, which

usually has a hiatus of tubercles in this area. A

character exclusive to N. raneyi is the continued

increase in numbers with increase in body length

(Table 5; Figure 4). No other species of Nocomis

has this characteristic. The head tubercles of N.

raneyi are relatively uniform in size, but they are

somewhat larger in the area extending from the

internasal to the interorbital. The largest tubercles

are about 1.8 to 1.9 mm in diameter. The smallest

tubercles occur on the snout. Tubercles average

somewhat larger in N. platyrhynchus. They are

large in N. micropogon and very large in N. leptocephalus. All of our data on size of tuberculate or

scarred adult specimens (Table 3) and the body

FIGURE 16.—The spot or bud stage in tubercle development

of three species of Nocomis. Note the sharper snout configura-

tion of these non-nuptial males compared with the specimens

in Figure 10. a, N. raneyi, USNM 194627, 194 mm, taken

6 June 1963, Craig Creek, James drainage, Virginia; b, N.

platyrhynchus, USNM 194877, 183 mm, taken 6 July 1963,

Big Walker Creek, New drainage, Virginia; c, N. micropogon,

USNM 166482, 173 mm, taken 17 October 1947, Susquehanna

River drainage, Pennsylvania.
NUMBER 85 25

FIGURE 17.—Juvenile stages of three species of Nocomis: a, N. raneyi, 83 mm, James drainage;

b. N. micropogon, 87 mm, Potomac drainage; c, N. leptocephalus, 81 mm, James drainage.

length when tubercle buds first appear (Table 4)

indicates that N. raneyi matures at an appreciably

larger size than N. micropogon or N. platyrhynchus.

Observations of nuptial males of N. raneyi attending their mound-nests corroborates this point.

Small tubercles are present on the dorsal portion

of the rays of the pectoral fin, in a single row per

ray, and they occur modally on rays 2 to 5 or 6,

rarely on rays 2 to 7 and 8.

Nuptial crest: Because of the difficulty in capturing the adult males, which are secretive, solitary,

and prefer the deeper waters, complete sequential

development of the nuptial crest with maturity is

not determinable from our material. The swelling

first appears in the internasal to snout area and enlarges to a rounded "cocks comb" that extends to

the occiput (Figure 12). An immature male specimen, 194 mm SL, (Figure 16a) taken 6 June 1963

from Craig Creek, has the typical sharp snout of

smaller immature males or mature females but has

a very slight internasal-interorbital swelling. Three

later stages in the development of the nuptial crest

are seen on the following specimens of N. raneyi:

198 mm SL, captured 28 June 1963 with moderately

swollen area from snout to occiput (Figure 106);

232 mm SL, captured 23 May 1964 with large crest

(Figure 10a); and 242 mm SL, taken 29 May 1964

with very large crest (Figure 12). Many very large

nuptial N. raneyi have been observed during the

reproductive period in Craig and Catawba Creeks,

James drainage over the past several years by the

first author and all specimens had large crests. A

postnuptial male, 229 mm SL (Figure 13), taken 21

June 1946 from the Roanoke River, has a moderate

head swelling, but whether this was its maximum

development during spawning or it had since receded somewhat was not determined.

The crest in N. raneyi is shaped and located dif-
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ferently than in N. micropogon and N. leptocephalus. Both of the latter species develop higher, more

dome-like crests, that are more forward on the head,

particularly in N. leptocephalus.

Pharyngeal dentition: The pharyngeal tooth

count is 4-4 in 21 pairs of arches in specimens from

the James, Roanoke, Tar, and Neuse River drainages. The teeth are very stout and moderate in

length. The arch is stout, broad, with short upper

and lower limbs. In the large N. raneyi (Figure 18)

the arch is massive, and probably stoutest of all

Nocomis. The pitted surface is deeply sculptured,

but the pits are not usually as extensively developed

as in other species of Nocomis.

FIGURE 18.—The pharyngeal arches of four species of Nocomis:

a, N. raneyi, 186 mm SL; b, N. platyrhynchus, 203 mm; c, N.

micropogon, 219 mm; d, N. leptocephalus, Savannah River,

206 mm.

Coloration: Juveniles, adult females and subadult

males from the James and Roanoke River drainages

show no significant local or geographic color difference in life or in preservation, or among these size

stages.

The characteristic scale color pattern on the body,

dorsally and laterally, ventrad to 1-3 scale-rows

above the pelvic fin base, has two basic aspects, the

dark scale margins and the light median scale color.

In the lateral line area the exposed portions of the

scales are sharply bicolored. The dark anterior margin (the excised scale pocket membrane) and the

posterior margin meet at the dorsal and ventral

edge of the scales, thus surrounding the light median area. The inner edge of the anterior margin

is vertically straight or slightly curved except in the

lateral line scale-row where it is indented at the

level of the lateral line canal pores. The inner edge

of the posterior margin follows the curve of the

actual scale margin; its outer edge extends to and

is darkest at the scale edge. The anterior margin is

generally slightly wider than the posterior margin.

The scale margins are slightly wider on the anterior

body scales than those on the posterior body. A

reduction in scale bicoloration occurs ventrad and

dorsad; the margin decreases in width and intensity ventrad until absent, and increases in width

and and intensity dorsad, where, on the dorsum,

the dark margin nearly or entirely fills in the light

median area. Smaller juveniles generally have narrower margins than the larger adults; thus they

have average lighter colors. The colors of the margins are, on the dorsum, usually black-olive, some

brown-olive, and grade from dark olive dorsolaterally, olive or yellow-olive in the lateral line area

to pale olive or pale yellow-olive below. Some tints

of green iridescence also appear on the margins. In

some specimens there is a sharp body bicoloration,

effected by appreciable lighter colors in the lateral

line area.

The medians of the lateral body scales are iridescent silver with a cast of yellow to olive in most

specimens. Iridescent violet and green hues are also

present in the lateral region of many specimens,

and these colors are particularly evident in sunlight. The silver color is almost entirely replaced

dorsad by the dark scale margins. The belly is

pearly white whereas the breast, where scales are

absent, is less lustrous or gray-white. An iridescent
NUMBER 8 5 27

silver-yellow to silver-olive middorsal stripe about

one scale in width is developed from the occiput

to the dorsal fin and on the anterior half to the

entire postdorsum of specimens up to 120-160 mm

SL.

A horizontal lateral band occurs in most smaller

juveniles and in a few specimens up to about 130

mm SL, including some adult females. When fully

developed, it extends from the tip of the lachrymal

bone (interrupted by the orbit) through the lateral

line area to the area over the caudal base, and

ranges 1.5-2.5 scales in width, narrowing just before

the caudal base area and then widening slightly.

The intensity is variable, generally being darkest,

but rarely blackish, in smaller juveniles. It is generally darkest over the caudal peduncle and, when

faint, it is usually present only in this area. The

lateral band is effected by a relative darkening beneath the medians of the scales and lighter medians

1-2 scale rows above.

The basicaudal spot is small, generally amorphous, confluent or slightly separated from and

usually darker than the end of the lateral band.

It is almost always absent from specimens larger

than 120 mm SL, and often in many smaller ones.

A black or blue-black, slightly curving bar occurs

from the pectoral fin base to the dorsum behind

the opercular opening, entirely over the cleithrum

or extending slightly posterior to it.

The head, dorsally, to the snout tip is black,

brown, or dark olive. The occipital line is slightly

lighter than the head dorsum. In some large adult

males a dark blue cast occurs on the head and anterior dorsal part of the body. The head, laterally,

over the upper operculum, below the orbit and

over the lachrymal is largely yellow-olive to dark

olive with iridescent silver, gold and green sometimes masking the olive; the iridescence is most intense along the upper edge of the operculum. The

outer, upper lip is a lighter olive than the snout,

dorsally. The ventro-lateral head is silvery with

some golden reflections. The head ventrally, including the lower jaw, is white to gray-white. The

opercular membrane is olive laterally, and pale

olive ventrally. The inner iris in almost all specimens is narrowly bordered by iridescent yellow,

gold, orange, or combinations of these colors. This

iris ring is sometimes interrupted by the black to

light olive areas of the outer iris, or it is absent.

Fin coloration is variable in its intensity and

areas of development. The pale yellow to reddish

colors are almost entirely confined to the interradial membranes and the membranes between ray

branches; sometimes they shade into the rays of

juveniles. Membranes lacking these colors are hyaline to dusky gray, sometimes dusky olive in the

larger specimens. Portions of the fin rays appear

blackish since melanophores occur along the rays

and ray joints, generally being heaviest on the dorsal fin and lightest, or sometimes absent, on the

anal and pelvic fins; a greater deposition occurs in

larger specimens. Color is almost always lacking

along the fin margins, the margin being hyaline to

dull white, generally widest on the anal and narrowest on the caudal.

The typical caudal fin coloration is a light red in

the distal I/3-I/2 of the fin membrane, deepest in

the distal portion of the lobes and usually extending basally from these. In a number of specimens

the basal portion of the medial membranes is suffused with pale red. The reddish shade is generally

deeper in larger specimens. In some juveniles it is

orangish red; in an immature female 110 mm SL,

it was orange. In a few specimens the membrane

color is very weak or absent. A recently preserved,

large adult female from the Neuse River drainage

had a light red outer caudal. The rays are yellowish

olive to dark olive, tending to be colorless and clear

in small specimens. In larger specimens, the thick

skin over the outer edge of the most ventral ray is

light gray.

The distal half of the dorsal fin membrane is

usually light red to orangish red; sometimes this

color is suffused through the entire membrane. The

red is deepest in the distal portion of the anterior

membrane. The rays are yellow-olive to olive, or

colorless in smaller specimens.

The pectoral fin rays are dusky yellow to pale

olive, being the most colorful distally. The membrane is gray-white, often with an orange suffusion

distally. The thick skin over the first ray is very

dark except for the gray-white anterior edge.

The distal pelvic fin ray is light or moderate

yellow to yellow-olive, and grades basally to light

olive or colorless; the medial and distal rays often

have a tingle of light orange. Most of the first ray

and the distal portion of rays 2 and 3 are milky white

or yellow-white. The membrane is clear to light gray.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The anal fin is the least colorful. The rays are

pale yellow to pale olive over i/2-2,4 of their length.

The first ray is white or yellow-white. The membrane is clear to dull white.

The striking color features of the nuptial male

in life are a pink or rosy on the head, body and fins

and the accentuation of certain other body colors

over those of nonbreeding adult males. The most

highly colored male we examined (USNM 194625,

237 mm SL) had well-developed tubercles and a

large crest, captured in the James drainage, 23 May

1964, probably in the midst of breeding activities

for many fresh nests were about. The most intense

nuptial color was rosy, located from the posterior

end of the postopercular bar, in the level from one

scale-row below the lateral line to the upper pectoral fin base, posterior to the pelvic fin base. The

color was deepest in the anterior portion of this

area. It extended as light pink to the area over the

anterior anal fin base, where it ended. The color

graded to faint pink on the belly and was absent

from the scaleless breast area.

The lower head, laterally, below a line from

about the jaw angle obliquely to midheight of the

opercle, was pink to light pink with some portions

of this area being suffused with olive. The upper

opercular area was olive. The head, ventrally, including the lower jaw (but not the upper jaw) was

pink, deepest over the distal portion of the branchiostegal area.

A heavy silver-bronze iridescence colored the medians of the scales of the body, laterally, above the

lateral line and it was present in the medians of

only the lightest dorsal scales. A faint pink suffused

the medians of the third to fourth scale-rows above

the lateral line between the postopercular bar and

pelvic fin levels.

The dorsal fin membranes were light pink (with

some dusky olive in the basal third). In addition

to the yellow, olive, and orange colors of nonbreeding specimens, the pectoral and pelvic fins had a

suffuse pink, slightly lighter than that of the dorsal

fin. There was only a very slight pink suffusion in

the distal anterior membranes of the otherwise typically colored anal fin. The caudal fin lacked pink

coloration. A milky-white color occurred along the

tips of the dorsal, pectoral, pelvic, and anal fin rays.

Three highly tuberculate (one with many tubercle scars) males, 197, 212 and 242 mm (holotype) SL

with small to large crests, were captured on 28 June

1963, 15 May 1963, and 29 May 1964, respectively,

from the upper Roanoke drainage (first specimen)

and the James drainage. These and many observations by Lachner of spawning males provided additional data on life coloration. The significant feature of the breeding N. raneyi (and other Nocomis)

is to "turn on" his colors to high intensities just

prior to or during the spawning acts.

Only slight color differences were found among

these males and the male described (USNM 194625,

237 mm SL) but the intensities differed appreciably.

The intensity and area of development of the pinkrosy coloration was about the same as described

above but it could be more or less intense depending on the spawning "peak" of the male. Pink color

was usually absent from the fins. In some males in

the "peak" of spawning all fins had a lemon-yellow

margin. The lateral body medians of the scales

above the lateral line in the holotype were iridescent silver-bronze. In some, it was iridescent oliveyellow, a slightly brighter yellow than that seen in

any other specimens of N. raneyi.

The light red caudal fin coloration occupied the

distal half of all membranes in most of the males,

but it was confined to the distal half of the upper

lobe in the holotype. In some active, spawning

males the caudal fin is dusky-olive. Melanophore

deposition along the rays and ray joints was heaviest in the largest nuptial males, particularly in the

distal half of the dorsal fin, but in the smaller males

it was no heavier than that in large nonbreeding

adult males. The same is true of the dark scale

margins. No trace of a dark lateral band was

observed.

After chubs are placed in preservative (10%

formalin and alcohol), the iridescent colors are rapidly lost. A few days thereafter all that remain are

the dark colors of the scale margins, fin rays, lateral

band and caudal spot and various tones of blackgray to light gray.

Melanophores underlie the dark scale margin

and dark head coloration seen in life. Their patterns in alcohol are about the same as the overlying

life colors and are shown in the illustrations.

With the fading of the iridescent median scale

colors, certain patterns appear and some become

more pronounced on the body. These are produced

by the "subscale" melanophores which are located
NUMBER 85 29

on the undersides of scales and on the musculature.

The dark horizontal midlateral band on the body

seems to remain in well-preserved specimens that

possessed it in life. The band is produced largely

or entirely by a heavier concentration of subscale

melanophores. Other shades of lighter lateral and

blackish dorsal medians of scales are also due mainly to subscale melanophores. The light middorsal

stripe of living specimens turns darker than the

adjacent dorsum color in preservative. In many

specimens, slightly dark horizontal lines, running

nearly or entirely the length of the body, occur on

the body laterally, above the lateral line, and sometimes to 3 to 4 scale-rows below the lateral line.

The lines are produced by the slightly heavier deposition of subscale melanophores in the areas beneath the dorsal and ventral edges of the scales

rather than beneath the medians of the scales.

A slight average difference in coloration exists in

the lateral margins of the scales between the sexes,

this being noticed only in preserved specimens.

Many adult females have slightly wider anterior

and posterior scale margins than those of males of

similar size.

POPULATIONS.—The drainage populations of N.

raneyi are similar. Only two characters show distinctive interdrainage divergence, the degree of

breast squamation (Table 18) and the number of

vertebrae (Table 15). Both characters partially differentiate the populations in the James, Chowan

and Roanoke drainages from those in the Tar and

Neuse. Breast scales in the James, Chowan and

Roanoke drainages are almost always absent (93%

of 138 specimens) and the breast was never half

scaled. The Tar-Neuse populations had only one

of 41 specimens naked and about 80 percent had

the breast scaled about one third or more. The

mean values for total vertebral numbers were practically the same for the Tar and the Neuse drainages (40.9 and 41.0) and lower than in the three

more northern drainages (41.4, 41.6, and 41.4).

REPRODUCTION AND GROWTH.—The male of N.

raneyi constructs a mound-nest (as in all species of

Nocomis) of gravel that he transports in his mouth.

The completed nest may exceed three to four feet

in diameter. (The reproductive behavior of the

species of Nocomis, including N. raneyi will be

treated in part 6, see p. 2). N. raneyi attains the

largest size of all the species in the genus, the largest

male examined was 266 mm SL and the largest

female, 180 mm SL, both from the Neuse drainage.

All nuptial males exceeded in length the largest

female captured. Several large males over 200 mm

SL aged by scale analysis were in their fourth and

fifth year of life.

ETYMOLOGY.—This species is named after Edward

C. Raney, scientist and teacher, Cornell University,

Ithaca, New York, whose enthusiasm and guidance

placed many American students on the professional

pathway to ichthyology.

The vernacular name, bull chub, was composed

by us because of the large size and aggressive habits

of the crested nuptial males during the spawning

period. The nuptial N. raneyi sometimes engages

in lengthy battles with other nuptial males of his

own kind when nesting territories overlap. The

native people in southwestern Virginia refer to

N. raneyi as a chub, pinkbelly chub, hornyhead or

nestmaker, but they generally do not differentiate

among the four species in the area.

MATERIALS EXAMINED.—A total of 106 collections

from 84 localities in five drainages were examined.

Holotype: USNM 194633, a nuptial male 242 mm

in standard length, captured in Craig Creek, Craig

County, Virginia, 2.2 road miles on Rt. 311 southwest (upstream) from Newcastle, at first bridge from

Newcastle, 29 May 1964 with shocker by Robert E.

Jenkins and Robert L. Miles. Five para types, USNM

194266, were taken with the holotype.

The Craig Creek system of James drainage and

the Roanoke drainage materials are designated as

paratypes.

The collections are listed below by drainages and

major tributaries in progressive upstream order;

following the collection date is the number of specimens captured (in parentheses). Designations given

as "same locality" or "same collection" refer to the

immediately preceding locality or collection.

James River Drainage, Virginia

Craig Creek, Botetourt County—USNM 194624: at 1 mi

above mouth, second bridge over Craig from Eagle Rock,

23 May 1964 (3). USNM 177397: at 2.5 mi above mouth,

3 airmi SW Gala, 3 July 1958 (26). USNM 194625; jet. Rtes.

615-683 (at suspension bridge) 3.3 rdmi NE (downstream)

Oriskany, 15.4 rdmi NE Newcastle on Rte. 615-42, 23 May

1964 (3).

Craig Creek, Craig County—USNM 194626: bridge on Rte.

606 leading to Camp Easter Seal (Rte. 606 turns off Rte. 615
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

at 9.2 rdmi NE Newcastle), 24 May 1964 (2). USNM 194627:

at 2.3 airmi NE Newcastle, 6 June 1963 (29). USNM 194628:

Pinetop, 2 airmi NE Newcastle, 11 May 1963 (15). USNM

194629: Rte. 616 ford, 1.2 airmi NE Newcastle, 16 August 1957

(133). USNM 194630: same locality, 12 October 1960 (71).

USNM 194631: same locality, 15 May 1963 (172). USNM

194632: holotype locality, 16 May 1957 (16). USNM 194634:

Rte. 311 bridge, 3 rdmi SW Newcastle, 19 August 1958 (4). UR

1538: Rte. 311 bridge, 5.6 rdmi SSW Newcastle, just N Abbott,

27 June 1963 (2).

Johns Creek, Craig County.—CU 20356: along Rte. 42 in

Newcastle, about 05 mi above mouth, 5 June 1951 (15).

CU 24859: same locality, 7 July 1952 (13). USNM 194635:

same locality, 3 October 1953 (12). USNM 194637: at 1.8 mi

W (upstream from) Newcastle in Nutter Mountain gorge,

16 August 1957 (4). USNM 194638: at 4.8 mi W Newcastle,

16 August 1957 (25). UMMZ 181830: at 7 mi above Newcastle, 27 August 1941 (1). CU 48564: at 4.35 rdmi W jet.

Rtes. 311-658 on Rte. 658, 7.75 airmi W Newcastle, 10 May

1965 (8). CU 48445: at 1.3 rdmi SW jet. Rtes. 632-635 on Rte.

632, 10.8 airmi WSW Newcastle, 27 March 1965 (2). USNM

194639: most downstream Rte. 632 bridge, 12 airmi WSW

Newcastle, 4.75 airmi NE (downstream) Maggie, 1 October

1957 (2). USNM 194640: second Rte. 632 bridge upstream

from Newcastle, 2.8 airmi NE Maggie, 30 May 1964 (7). UR

1796: first Rte. 632 bridge 2 rdmi NE jet. Rtes. 632-658 at

Maggie, 27 June 1963 (24). CU 47594: at 1 rdmi NE jet.

Rtes. 632-656 at Maggie, along field off Rte. 632, 29 May 1964

(27).

Chowan River Drainage, Virginia

Sappony Creek, Sussex County.—CU 16911: Rte. 40 bridge,

22 mi W town of Stony Creek, 27 March 1949 (3).

Stony Creek, Dinwiddie County—CU 47335: at 1.2 mi S

Dinwiddie on Rte. 1, 21 March 1948 (18).

Nottoway River, Brunswick-Dinwiddie County line.—USNM

177357: Rte. 1 bridge, 29 July 1958 (8).

Meherrin River, Lunenburg-Mecklenburg County line.—

USNM 194642 (2 collections lumped): (a) Rte. 138 bridge,

Union Mill, 5.1 airmi NNE South Hill, 29 July 1958; and

(b) bridge just N Ogburn, 7 airmi NW South Hill, 3 September 1958 (42). CU 43628: same collection as (b) (1). USNM

194258: Rte. 49 bridge, 31 July 1958 (2).

Roanoke River Drainage,

Virginia and North Carolina

DAN RIVER SYSTEM

North Carolina, Rockingham County—USNM 171727:

Leaksville, 6 April 1947 (10).

Virginia, Henry County—USNM 171593: Beaver Cr., 15

mi NW Martinsville, 22 June 1946 (1). USNM 194261:

Town Cr., trib. Smith R., Rte. 604 bridge, 1.75 mi SW Henry,

near railroad bridge, 6 August 1958 (1).

ROANOKE RIVER SYSTEM, VIRGINIA

Otter River, Bedford County.-USNM 171595: at 4 mi NNE

Bedford, 21 June 1946 (13). USNM 166406: at 4 mi NNW

Bedford on Rte. 43, 5 September 1947 (4). USNM 166405:

at 5 mi NW Bedford on Rte. 43, 9 June 1947 (11).

Otter River, Campbell County.-UMMZ 181434: Rte. 682,

S of Evington, 12 June 1956 (3).

Pigg River, Franklin County.-USNM 163436: Rte. 220

bridge, 1 mi S Rocky Mount, 3 April 1948 (106) . USNM

194601: same locality, 19 June 1962 (2).

Blackwater River, Franklin County.-USNM 194602: Rte.

616 low-water bridge, 3.7 airmi SE Scruggs, 7.8 airmi SSE

Haleford, 11 August 1958 (49). USNM 194605: same locality, 5 July 1961 (28). USNM 194606: at 0.75 mi downstream

from Rte. 670 bridge and gauging station, 3.1 airmi SSW

Scruggs, 4.6 airmi W Bedford-Franklin-Pittsylvania Co. line,

5 July 1961 (1). USNM 194603: Rte. 670 bridge by gauging

station, 2.7 rivermi upstream from mouth Gills Cr., 12 mi

S Haleford, 11 July 1958 (3). OSUMZ 5991: same locality,

5 July 1961 (68). USNM 194609: at 4.8 airmi SW Scruggs,

8.6 airmi SSW Haleford, 11 August 1958 (18). USNM 194610:

near Crafts Church, 7.8 airmi W Bedford-Franklin-Piltsylvania Co, line, 6 July 1961 (62). USNM 177395: at 3.8 airmi

NE Redwood, 8.2 airmi ENE Rocky Mount, 5 July 1956 (19).

USNM 177376: Rte. 122 bridge 4 airmi NE Rocky Mount, 5

July 1956 (6). USNM 194611: same locality, 7 July 1958 (7).

USNM 194612: at Blaine in rocket fuel plant area just E of

Rte. 220 bridge, 3 airmi NNE Rocky Mount, 22 August 1957

(11). USNM 194613: same locality, 4 July 1961 (109). USNM

194614: same locality, 6 April 1963 (69). USNM 194615: Rte.

220 bridge, 2.8 mi NNE Rocky Mount, 21 August 1957 (3).

CU 25032: Rte. 919 (old Rte. 220) bridge, 25 airmi NW Rocky

Mount, 22 June 1946 (1). CU 47595: same locality, 5 July

1956 (21). UMMZ 181433: same locality, 17 May 1931 (3).

USNM 194617: jet. Rtes. 643-732-737, 4*75 airmi WNW Rocky

Mount, 15 October 1961 (41). USNM 194618: Rte. 740 bridge,

1.8 airmi NE Calloway, 15 October 1961 (9). USNM 194619:

North Fork Blackwater along Rte. 641, 2 airmi NE Calloway,

14 October 1961 (14).

Blackwater System, Gills Creek, Franklin County.—USNM

194607: about 45 rivermi above mouth in Blackwater, 5 airmi

SE Hales Fort Church, 2.1 airmi SW Scruggs, 6 July 1961 (2).

USNM 194604: 25 mi SW Scruggs, 5.9 airmi SSW Haleford,

7 August 1958 (1).

Back Creek, Roanoke County.-USNM 194620: Rte. 657

bridge, 05 mi from Rte. 220 intersection, 45 airmi S Roanoke,

3 July 1961 (7).

Roanoke River, Roanoke County.—USNM 40240: at or near

Roanoke, 31 July-2 August 1888 (1). USNM 166404: Salem,

21 June 1946 (1). UMMZ 181431: at 2 mi above Salem, 13

June 1956 (3). CU 20769: Glenvar, 6 June 1951 (1). CU

24771: same locality, 30 June 1952 (2). UR 2025: same locality (at RR station just off Rte. 11), 11 July 1963. (11).

USNM 197684: first Rte. 639 bridge from Rte. 11, 15 airmi

ESE Riverside, 35 airmi ENE Lafayette, 28 June 1963 (19).

TU 25885: at 8 mi SW Salem on Rte. 11, 18 June 1962 (3).

USNM 194255: Roanoke-Montgomery Co. line, Rt. 11 bridge,

11 September 1956 (7).

South Fork, Roanoke River, Montgomery County.—USNM

194621: bridge off Rte. 11, 1 mi S Elliston, 2.3 mi N Shawsville, near Bench Mark 1282, 12 August 1958 (1). CU 24788:

Rte. 11 bridge, 2.4 mi S Elliston, 9 July 1952 (3). CU 24728:
NUMBER 85 31

about 1 airmi SE Shawsville along Rte. 609. [Figured in detail in Schwartz and Dutcher (1962: 370)], 9 July 1952 (2).

USNM 194265: same locality, 13 September 1954 (1). USNM

194254: same locality, 2 July 1956 (2). USNM 194622: same

locality, 12 August 1958 (1). USNM 177283: same locality,

31 August 1958 (2). USNM 171594: at 2 mi N Alleghany

Springs, 22 June 1946 (1). USNM 194623: at 0.75 mi N

Alleghany Springs, 2 mi S Shawsville, 16 October 1951 (2).

Tar River Drainage, North Carolina

Tar River or Fishing Creek, Edgecombe County.—USNM

191108: Rte. 1500 bridge about 4.5 mi SW Lawrence, 5.5 mi

NNE Tarboro, 19 September 1959 (10). TV 34466: Rte. 44

bridge, 1.9 mi NW Tarboro, 6 September 1964 (8). USNM

40324: at 2 mi below Rocky Mount, 29 August 1888 (2).

Tar River, Nash County—USNM 191049: railroad bridge

just N Rocky Mount, 19 September 1959 (10). TU 34508:

Rte. 64 bridge, 7.7 mi NE Pilot, 6 September 1964 (96).

Tar River, Granville County.—UMMZ 181832: Rte. 15

bridge, 7 mi SSW Oxford, 4.5 mi NNE Hester, 30 August

1946 (3). UMMZ 181432: 05 mi W Providence, 26 November 1953 (23). CU 19505: at 1 mi SW Providence (presumed

same locality as UMMZ 181432), 30 March 1951 (9).

Neuse River Drainage, North Carolina

Nahunta Swamp Creek, Wayne County.-CU 11892: at 1.2

mi N Pikeville, 27 June 1946 (1).

Contentnea Creek, Wilson County.-USNM 191036: about

3 mi SW Wilson, golf club below Rte. 301, 18 September

1959 (25). DU F-122: 4 August 1961 (1).

Falling Creek, Lenoir County—TU 34439: Rte. 70 bridge,

8.2 mi W Kinston, 5 September 1964 (1).

Little River, Wayne County.-USNM 40386: just W of

Goldsboro, 29 August 1888 (6).

Crabtree Creek, Wake County.-UMMZ 161963: Raleigh,

11 October 1929 (2). UMMZ 181430: same locality, 9 May

1930 (3).

Neuse River, Wake County.-USNM 194272: Raleigh, 1890

(2). USNM 40462: below dam at Milburnie, near Raleigh,

27 August 1888 (16). USNM 190975: same locality, 16 September 1959 (4). TU 34532: Rte. 64 bridge at Milburnie,

7 September 1964 (5). USNM 190997: Falls, below bridge,

17 September 1959 (8).

VO N platyrhynchus

A N micropogon

• N raneyi

FIGURE 19.—Distribution of Nocomis platyrhynchus, N. micropogon and N. raneyi in the central

eastern region. The Atlantic-Gulf drainage divide is indicated by the heavy, broken line; the

Fall Line by the North-South solid line. The Kanawha Falls, West Virginia, is marked by a heavy

bar just downstream from the Gauley River (the last river tributary to the New River downstream inhabited by iV. platyrhynchus). The areas west of letter A and B are enlarged in

Figures 20 and 29. The names of important rivers west of the divide are labeled in Figure 28.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Neuse River, Durham County.—UMMZ 181429: Rte. 15

bridge, 9 mi NE Durham, 29 August 1946 (23).

Eno River, Orange County .-USNM 188555: 1.6 mi NE Rte.

70 on Rte. 1567, and 1.1 mi NW on Rte. 1569, 10 May 1964

(30). USNM 188736: same locality, 25 May 1964 (12). DU

(unnumbered): below Hillsboro, 18 November 1941 (2).

DISTRIBUTION.—N. raneyi is a central Appalachian, Atlantic slope species with its center of distribution in Virginia. It occurs in the James, Chowan,

and Roanoke drainages of Virginia and in the Tar

and Neuse drainages of eastern North Carolina

(Figure 19). We examined 116 collections from 84

localities in the five drainages. Ecological distribution is discussed in the Ecology section.

It apparently is restricted in the James drainage

to a small portion of the upper basin, in Craig and

Botetourt counties (Figure 20), and was found downstream on only one occasion. We had extensive

collections (182) from the James drainage, in which

at least one species of Nocomis was taken from 141

localities scattered throughout the drainage, the majority from the upper half of the basin; more than

3400 Nocomis specimens are involved.

This species probably is continuously distributed,

or nearly so, in the 116 rivermile length of the

Craig-Johns Creek system (except for its headwaters)

since it was taken in 34 collections from 23 localities

and was represented in almost every collection made

within the middle and lower system; 742 specimens

were taken. One collection of four specimens (CU

48533) is from Catawba Creek at 0.4 rivermiles above

its mouth where breeding males have also been

regularly observed recently. One hybrid N. micropogon x N. raneyi was found in middle Catawba

Creek (UMMZ 174826) and another (USNM 194636)

in Johns Creek at Newcastle. One specimen (CU

48491) was collected in Pedlar River, Amherst

County, 4 rivermiles above its mouth, which is 52

rivermiles downstream from the Catawba Creek

mouth. (See discussion of atypical specimens of N.

raneyi, pp. 32-36).

The bull chub occurs in the Virginia portion of

the Chowan drainage (only six collections) in two

of its three main tributary systems, the Nottoway

and Meherrin Rivers.

It is widely distributed in the upper montane and

upper Piedmont portion of the middle Roanoke

drainage. Fifty-one collections from 37 localities

were examined. All localities in the upper Roanoke

are along the South Fork and upper main channel

except for one that is in Back Creek. In the middle

basin, populations are present in the Blackwater

(Figure 29), Pigg, and Otter Rivers. Its apparent

absence from other upper and middle Roanoke

tributaries, some rather well collected, is noted in

the Ecology section. It has been taken on only three

occasions in the Dan River system, the largest southern montane and Piedmont Roanoke tributary.

JV. raneyi is widespread in the Tar and Neuse

drainages; 24 collections from 21 localities were

examined.

Atypical Nocomis raneyi

Specimens that generally agree with our concept of

N. raneyi, but which are divergent in certain characters have been collected from the James drainage

and are referred to herein as atypical N. raneyi.

Atypical specimens were collected mainly between

the ranges of two apparently disjunct populations

of typical N. raneyi and partially within the interrupted range of N. micropogon in the James (Figure 20). The 13 collections of atypical specimens are

from the main James channel and lower portion of

all except one of its major tributaries from Craig

Creek to 47 miles downriver. The specific localities

are given at the end of this discussion. The most

upstream locality, in Craig Creek, is 17.5 rivermiles

above its mouth, where one atypical tuberculate

male, in spawning or just entering postnuptial condition, was captured in 1967 with a typical nuptial

male. Sixteen days earlier an apparently eggbound,

atypical female was picked up dead at the same

locality. Three collections, totaling 47 typical specimens only, were taken in a four year period earlier

at the same locality. Typical and atypical specimens

were also collected together at the mouth of Craig

Creek. In three collections from Craig Creek within

0.5 to 3 miles from its mouth, 32 typical specimens

only were taken. The two forms were collected together in one lower Catawba Creek collection and

the atypical form was collected alone on three occasions from this stretch of stream including James

River about 100 feet above the Catawba mouth.

Typical specimens captured outnumbered atypical

ones in lower Craig while the opposite was true in

lower Catawba. Recent breeding behavioral observations in lower Catawba, however, involve several
NUMBER 85

typical nuptial males and only one atypical (or hybrid?) specimen. The main James has yielded only

atypical specimens. The most downstream locality

for an atypical specimen is the mouth of Rocky Row

Run, which is 11 rivermiles above the mouth of

Pedlar River within which a typical specimen of

N. raneyi was found. N. micropogon is widespread

in the large Maury River system, which enters the

James four miles above the Rocky Row Run mouth;

N. raneyi is not known from this system.

There are 79 atypical specimens which range

from 42 to 229 mm SL, including several prenuptial

and postnuptial males and a tuberculate male. Two

or three small specimens could not be classified with

certainty as atypical or typical N. raneyi.

There are no apparent differences between atypical and typical N. raneyi in the following characters

in which typical N. raneyi differs significantly from

N. micropogon; circumferential and lateral line

scales, vertebral numbers (Table 27), breast squamation, all aspects of body morphometry, size attained,

coloration in life and in preservative, and the

pharyngeal arches.

Some features in the atypical series generally are

somewhat intermediate between typical N. raneyi

and N. micropogon. Other characters in which the

atypical form differs from N. raneyi are also unlike

those of N. micropogon and they are not intermediate between the two species. The main differences

between atypical and typical N. raneyi are in certain

aspects of tuberculation, involving tubercle numbers,

posterior distribution, early development, and size.

In larger juveniles and adults, the numbers are

generally intermediate between typical N. raneyi

and N. micropogon (Figure 21). The most posterior

distribution also is somewhat intermediate between

these two forms, ranging in adults from the PIO to

AOC lines, more often ending at the AIO line; when

0 mil.. 5

• N RANEYI

O N. MICROPOGON

O BOTH SPECIES

X HYBRIDS

FIGURE 20.—Distribution of Nocomis raneyi and N. micropogon in the upper James and Roanoke

drainages (see letter A of Figure 19). The common occurrence of N. leptocephalus in the area

was omitted to avoid confusion in symbols. Hybrid combinations are indicated: X1 = N. micropogon X N. raneyi; Xs

 = N. leptocephalus X N. micropogon, in Catawba River. Solid dot1 =

atypical and typical N. raneyi; solid dot1 = atypical N. raneyi only. Sinking Creek and other

headwater tributaries of the New drainage are shown in the upper left part of the map; certain

main mountain ridges are indicated by stippling.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

extending into and beyond the interorbital area, the

tubercles usually are more scattered than in typical

N. raneyi. The most posterior tubercles are on or

near the dorsal midline, as in typical N. raneyi,

rather than dorso-laterally as in N. micropogon.

The reduction and scattering of posterior tubercles

in the atypical form results in lower tubercle numbers compared with typical N. raneyi. Almost all

atypical specimens have tubercle spots and/or buds

that are larger than the maximum size of spots and

buds in typical N. raneyi and, in many of the former, these structures exceed even the maximum size

of those in N. micropogon of similar body length.

The spots first appear in smaller atypical specimens

than in any species of the micropogon group. The

spots of most of the smallest atypical specimens have

buds. All young or small juveniles, including females, from 42-62 mm SL, have 7-18 spots or buds.

Thus, in small atypical specimens, the pattern of

tubercle appearance and their increase is most simi100

• *-

O

0) 80

JQ

E

• _- _ Typical N. raneyi

o Atypical N. raneyi

A MMBam^BMM N . micropogo n

1

80

1

100

1

120

1

140

1

160

1

ISO

1

200

1

220

1

240

I

20

Standard Length in mm.

FIGURE 21.—The relationship between number of tubercles and standard length in typical N.

raneyi (all drainages; data from Table 5), atypical N. raneyi (James drainage) and N. micropogon

(James drainage; see Table 10). The curves for typical N. raneyi and N. micropogon were fitted to

the mean values of the 10 mm size groups. Data for atypical N. raneyi are plotted from individual

specimens.
NUMBER 85 35

lar to JV. micropogon, but at larger sizes it conforms

better to the JV. raneyi pattern. Although there is a

tubercle reduction in the atypical form, there is no

trend toward development of a hiatus between the

snout and internasal tubercles, which agrees with

JV. raneyi and not JV. micropogon.

Other characters in which the atypical specimens

differ from the species of the micropogon group are

discussed below. The snout profile of most young

and small juveniles is usually quite rounded, appearing somewhat swollen. The lower caudal fin

lobe is considerably shortened and more rounded.

The anterior rays of the anal fin are often reduced

in length, resulting in a smaller fin with a broadly

convex margin. The paired fins are frequently

shorter and more broadly rounded.

The following factors are considered in an attempt

to determine the origin and significance of the atypical specimens: (1) distribution and relative abundance with regard to opportunity for intergradation

or interspecific hybridization between JV. micropogon and JV. raneyi; (2) atypical JV. raneyi morphology in relation to that of Nocomis intergrades

and hybrids; and (3) possible influence(s) by environmental disturbance.

Almost all atypical specimens were taken from an

area connecting the two main populations of JV.

micropogon (one above Craig system, one from

Maury system downstream) and the main typical

JV. raneyi population (Craig Creek area). The range

of the atypical specimens may thus be one in which

the forms met and freely interbred in the past. However, the presence of typical JV. raneyi below the

atypical area (known from one specimen from Pedlar River), and die occurrence of JV. micropogon

with typical JV. raneyi in the Craig and Catawba

systems are difficult to explain by intergradation,

particularly since JV. raneyi probably entered the

James drainage in the Craig Creek area (see Dispersal section, p. 70). Relative rarity of one parental

species is frequently cited as an important factor in

production of hybrids. Although JV. micropogon is

unknown from the James River section with atypical JV. raneyi and the lower portion of most tributaries of this section, these larger waters have not

been well collected. Even if JV. micropogon is present but rare in this section, the abundance of atypical specimens and small numbers of typical JV. raneyi

(only in a small portion of the section) is not explained by hybridization within the section.

The atypical specimens conform to JV. raneyi in

most diagnostic characters; in the others they are

roughly intermediate between JV. micropogon and

typical JV. raneyi or differ from both species. The

atypical form does not agree with the concept of an

intergrade population, variably intermediate in

most or all diagnostic characters (as we have found

in subspecific intergrades of JV. leptocephalus and

populations of JV. micropogon that show effects of

apparent introgressive hybridization with JV. platyrhynchus). Interspecific hybrids are generally intermediate but may deviate to the extent that they are

extreme in certain characters, thus not excluding

atypical JV. raneyi from this category.

James River water is very darkly colored, due to

the effluent from a large pulp mill at Covington,

Virginia, downstream through the upper half or

more of the atypical JV. raneyi section. The peculiar

morphology of the atypical JV. raneyi may be induced by pollution. Indirect evidence from local

anglers indicates that JV. raneyi spawn in the main

channel of the James. It would then be in direct

contact with pollution beginning at early developmental stages. Lindsey (1956:770) gave evidence that

fin shape in certain Canadian populations of Couesius plumbeus may be influenced by environmental

conditions. De Sylva, Kalber, and Shuster (1962:29)

noted deformed fins in striped bass collected in

proximity to a chemical plant. Many cyprinid fishes

with malformed mouths and fins were found by

Larimore and Smith (1963:353) near a pollution

source in an Illinois stream. Beadles (1966) concluded that industrial and domestic effluents could

have produced variations in meristic and morphometric characteristics of subpopulations of three

cyprinid species in an Oklahoma stream. Additional

literature treating morphological variations caused

by environmental conditions in fishes was summarized by Barlow (1961).

It is more probable that the atypical condition of

JV. raneyi is due to its development in polluted

waters rather than to other alternatives treated.

Solution of this problem must include additional

collections, observations, and experimental work.

MATERIAL EXAMINED.—Atypical JV. raneyi from

the James drainage, Virginia. Included are museum

numbers and number of specimens of typical JV.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

raneyi taken with three of these series. Collections

are listed in progressive downstream order; following the collection date is the number of specimens

captured (in parentheses).

Botetourt County.—USNM 201895: Craig Cr. at jet. Rtes.

614-683, 7.5 airmi W. Eagle Rock, 20 May 1967 (1). USNM

201898: same locality at USNM 201895, 6 June 1967 (1);

taken with one typical specimen, USNM 201897. CU 50596:

Craig Cr. at mouth, James River, 30 and 31 March 1965 (3);

taken with two typical specimens, CU 48427. CU 52208:

James R. at Eagle Rock, about 05 mi downstream Craig Cr.

mouth, 1966 after 1 June (14). CU 47586: James R. about

100-200 ft. above mouth Catawba Cr., 24 October 1964 (15).

CU 47568: Catawba Cr. 0.2 mi above mouth, 24 October

1964 (3). CU 52597: Catawba Cr. 0.4 mi above mouth, 10

May 1965 (6) taken with four typical specimens, CU 48533.

CU 50820: same locality as CU 52597 (1). TU 25858: James

R. at Springwood, 18 June 1962 (10). CU 47587: same locality as TU 25858, 23 October 1964 (10). CU 48438: Looney

Cr. 0.3 mi above mouth, 30 March 1965 (3). CU 48426:

James R. at mouth Jennings Cr., 29 March 1965 (11) .

Amherst County—CU 48549: Rocky Row Run at mouth,

James R., at Snowden, 10 airmi S Buena Vista, 9 May 1965 (1).

PROBLEMATIC SPECIMENS FROM THE CRAIG CREEK

AREA.—Craig Creek, Catawba Creek, and the James

River near their mouths, all part of the James

drainage, is one of the most critical areas in this

study, for conclusions on the status of the forms of

the micropogon group have been based on specimens found in this drainage area. Since some of the

series from this area include small, poorly preserved,

and difficult to identify specimens of apparent hybrids, and typical and atypical specimens, a discussion of our methods (some subjective) used to

identify these and other series is warranted. These

collections also assume considerable importance due

to a recent impoundment of a section of the Johns

Creek system, the proposed impoundment of much

of Craig Creek (Mussey, 1948:20), and channelization of the lower 17 miles of Johns Creek. This is

the only area in which three species of Nocomis

have been collected together.

No adult N. micropogon were available from the

Craig area. However, juvenile N. micropogon were

taken from upper Johns Creek (TU 25469, 6 specimens), upper Craig Creek (UMMZ 135413, 2 specimens), and upper Catawba Creek (CU 50610, 1

specimen). These have the tubercle pattern of adult

N. micropogon well developed or incipient and their

meristic (Table 27) and morphometric values are

well within the ranges of N. micropogon. Also clearly determined are three specimens (UMMZ 135401)

taken with one N. raneyi (UMMZ 181830) from

middle Johns Creek. The three range from 60-80

mm SL and have the juvenile tubercle pattern, low

meristic values, long snout and larger mouth of N.

micropogon. The specimen identified as N. raneyi

lacks tubercle spots (as expected for this species at

60 mm SL but not for N. micropogon), has 35 circumferential scales, 41 vertebrae, and, in coloration

and morphometry, resembles N. raneyi.

The species were taken together on two other

occasions from the lower half of Johns Creek. The

most upstream collection, was 1.5 miles east of Craig

Healing Springs and consisted of two specimens

(CU 50340, 36 mm SL; and CU 52598, 42 mm SL).

Both are too small to be identified by tubercle

spots and morphometry, although CU 52598 has a

rather long snout, like N. micropogon. The siiQut

of the other specimen is more like N. raneyi. The

apparent N. micropogon specimen has 30 circumferential scales; no N. raneyi was found with 30

circumferential scales and a 31 count is rare. Its

vertebrae number 41, which is occasional for N.

micropogon but typical for N. raneyi; the lateral

line count of 40 applies to both species. Thus CU

52598 is identified as, probably, N. micropogon. CU

50340 is identified as, probably, N. raneyi from its

appearance and vertebral count of 42; we have not

found a vertebral count of 42 in N. micropogon

over its range. The circumferencial count of 32 and

lateral line of 40 are common to both species.

The other collection from Johns Creek, at 4.2

roadmiles on Rte. 311 from Newcastle, includes seven

N. raneyi (CU 50422), 37-116 mm SL, one N. micropogon (CU 50423), 39 mm SL, and several N. leptocephalus. The 39 mm specimen has more heavily

pigmented scale margins, a longer snout than the

other young specimens, 30 circumferential scales,

and 40 vertebrae, thus apparently representing N.

micropogon. One specimen with tubercle development, 116 mm SL, shows the typical juvenile pattern of N. raneyi. The other six specimens apparently

are young N. raneyi based on scale counts, and

vertebrae of 41 (2), 42 (3), and 43 (1).

All except one specimen from Craig Creek, near

its mouth, and the James River in the immediate

vicinity are typical or atypical N. raneyi. The exception, a specimen (CU 47584), 65 mm SL, lacks

tubercle spots, has 41 vertebrae, 40 lateral line scales,
NUMBER 85 37

and 30 circumferential scales. The circumferential

count falls out of the range of N. raneyi. The gape

width of 6.8 percent SL and snout length of 8.5 percent SL is intermediate between N. micropogon and

N. raneyi. It is tentatively identified as N. micropogon.

The nearest locality upriver (USNM 132064

taken around 1880) is in lower Mill Creek, a James

River tributary about five miles above the mouth

of Craig Creek. The 14 specimens are very poorly

preserved and small, 45-80 mm SL. The unsealed

breasts and the unwhorled intestines exclude these

being N. leptocephalus. They are identified as N.

micropogon mainly by the meristic values: circumferential scales 29 (1), 30 (5), 31 (5), 32 (2); lateral

line scales 38 (3), 39 (8), 40 (2); total vertebrae 39

(6), 40 (7), 41 (1). The tubercle spot pattern, typical

of juvenile N. micropogon, was discernible only in

the largest specimen.

The characters of atypical specimens, but basically N. raneyi, from lower Craig and Catawba Creeks

and the James River are given above. We found two

specimens from the Craig Creek area, one from

lower Johns Creek and one from middle Catawba

Creek, that do not fit our concept of the atypical

specimens, but rather, are intermediate between

N. micropogon and N. raneyi in most of the diagnostic characters. These have been determined as

hybrids between the two species. We have not found

atypical specimens from the vicinities in which these

specimens were taken. They will be described in

paper 5 on hybridization (p. 2).

Nocomis platyrhynchns, new species

BIGMOUTH CHUB

FIGURES 7, 10, 11, 16

Ceratichthys biguttatus.-Cope, 1868: 226,240 [(ANSP) Walker

Creek].

Hybopsis kentuckiensis.—Goldsborough and Clark, 1908:36

[New River drainage, West Virginia].

Hybopsis kentuckyensis.—Breder and Breder, 1923: 11 [South

Fork New River].

Hybopsis micropogon.—Raney, 1941:3 [New River].—Ross

and Perkins, 1959 [(part) New drainage records].

Hybopsis species.—Robins and Raney, 1956:31 [New River].

—Ross, 1959: 12, 16, 25 [New River above Kanawha Falls].

—Ross and Perkins, 1959:12, 19, 24, 27 [(part) New drainage records].

Hybopsis leptocephala.—Ross and Perkins, 1959 [ (part) New

drainage records].

Hybopsis (Nocomis) species 1.—Reno, 1969b, p. 739.

DIAGNOSIS.—A species resembling N. micropogon

in body features and approaching N. raneyi in certain aspects of head tuberculation and squamation;

mouth wide (Figure 5); snout long (Figure 6, Tables

17, 20, 21); body subterete; maximum number of

head tubercles in largest specimens approaching

average values of about 100 or less (Table 6); tubercles not densely developed on snout in adults;

tubercles characteristically present in the occipital

area in most adult specimens, sometimes extending

to the posterior occipital line (Table 2); breast

typically scaled, usually from 50 to 100 percent in

all river populations (Table 18); caudal fin oliveorange in subadults and moderately forked; pharyngeal arch of moderate proportions, the edentulous

processes longer than in N. raneyi (Figure 18).

DESCRIPTION.—Morphometry: Proportional measurements, expressed in thousandths of the standard

length, are given for the holotype, 5 male paratypes,

57 to 195 mm SL and 5 female paratypes, 65 to 142

mm SL in Table 24. Additional proportional data

are given in Table 25 for 13 characters of 31 adult

males having a mean length of 178 mm and ranging

from 159 to 214 mm SL. The general growth pattern

is similar to that described for N. raneyi. In the

young and smaller juveniles, all three species of the

micropogon group are very similar in their body

features. At about 70 mm SL (Figures 5, 6) the gape

width and snout length begin to show different

developments in N. platyrhynchus (and N. micropogon) compared with that of N. raneyi. The wide

mouth, longer snout, and more rounded, subterete,

body of the adults, particularly the nuptial males,

give the former two species a stouter, more robust

appearance. The systematic significance of the gape

width and snout length in the micropogon group

was treated under N. raneyi (p. 18).

The preopercle-opercle suture is most often perpendicular in relation to horizontal body axis, or

the lower portion is slightly posteriorly directed. In

a sample of 100 specimens, 5 had suture line directed forward at an angle of about 10° to 20°; 22

specimens had suture very slightly directed forward;

73 specimens had suture perpendicular or it was

slightly posteriorly directed, in relation to horizontal body axis.

The intestine is simple, elongate, and S-shaped

(84%) or with a very short anterior bend (16% of

44 specimens).
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Other general features of the head and body are

very close to those of N. micropogon: Head moderately elongate and wide, but not deep; head widest

at posterior part of opercle and deepest at the occiput. Dorsal and ventral body contours, excluding

nuptial males, as in N. raneyi, except snout only

moderately elongate and pointed to slightly or

moderately rounded as in N. micropogon; mouth

moderate in size in young to large in adult males;

mouth slightly oblique, subterminal and slightly to

moderately overhung by the tip of snout; angle of

jaw at or slightly posterior to vertical from posterior

nostril; shape and position of lips, barbel, eye and

lachrymal bone is in N. raneyi.

The body is elongate and subterete, only slightly

compressed in adult males compared to N. raneyi;

body deepest and widest midway between occiput

and origin of dorsal fin; predorsal body contour

slightly convex; postdorsal body contour is straight

and slopes downward; ventral body contour gently

rounded from head to origin of anal fin; lateral

line, urogenital papilla, scale pattern except breast,

and cutaneous sense organs as in N. raneyi; shape

and size of fins are much like those of N. raneyi,

except caudal fin is more shallowly forked, the

notch more broadly rounded, than in N. raneyi,

particularly in adult males (resembling caudal of N.

micropogon); lobes of caudal fin sometimes slightly

subequal, upper lobe a little longer in young and

juveniles, and the lower lobe slightly longer in

adult males.

Meristic characters: Squamation values are close

to those of N. raneyi. The range of mean values of

specimens from the main channel of the New River

and its major tributaries follow, with holotype data

in parentheses: Circumferential scale rows, 32.7 to

34.2 (34) (Table 12); lateral line scale rows, 39.6 to

40.3 (41) (Table 8); scale rows above lateral line,

6.6 (8) (Table 11); caudal peduncle scale rows, 16.8

(17) (Table 14); pectoral fin rays, 17.5 (Table 16);

total vertebral numbers, 40.5 to 40.6 (Table 15).

The breast (Table 18) is one half to fully scaled

in most specimens, the modal values for the main

New River and its major tributaries show about

a three fourths scaled condition. In breast squamation, N. platyrhynchus is unlike the James, Chowan,

and Roanoke River populations of N. raneyi, but it

is close to the Tar and Neuse populations of N.

raneyi, certain drainage populations of N. micropogon and to the consistently high values for N.

leptocephalus.

Tuberculation: Data pertaining to tubercle development, distribution and numbers are summarized in Tables 2, 3, 4, and 6. The relationship

between increase in tubercle numbers with increase

in standard length is shown in Figure 4. The distribution of head tubercles is illustrated in Figures

10, 11, and 16. Tubercle pattern and increase in

tubercle numbers with increase in body length is

fairly similar in the sexes. The males attain a larger

size than the females (Table 6) and thus a greater

average number of tubercles. N. platyrhynchus is

characterized as having fewer tubercles than N.

raneyi but more than N. micropogon; their distribution on the head extends, dorsally, over a greater

area than in both of the latter species; the increase

in tubercle numbers slopes off at about 150 mm SL

(Figure 4). The tubercles first appear as small, light

spots in the internasal area, intermediate in numbers between N. raneyi and N. micropogon (Figure

3). The tubercles rarely can be seen at body lengths

up to 60 mm SL. Both males and females may have

no visible tubercle spots at 70 mm SL in some specimens. With increase in body length the tubercles

develop posteriorly on the head, the complete

tubercle pattern being developed in some specimens about 100 mm SL. (Table 2). The snout and

subnasal tubercles develop more slowly than those

on the top of the head. Up to three rows of tubercles may be present in the lachrymal area of the

larger males. Tubercles on the snout of N. platyrhynchus are more abundant than in N. micropogon

and do not form a hiatus in the adults as in the

latter species, but some specimens may show a

tendency towards a hiatus in having reduced tuberculation between tip of snout and the internasal

area. Average tubercle size is approximately intermediate between that of N. micropogon and N.

raneyi; the largest tubercles occur in the internasal

area or just behind. The tubercle bud stage first

appears at about 80 mm SL (Table 4) at a size

similar to the first development in N. micropogon,

and appears in both species at a much smaller size

than in N. raneyi.

Small tubercles are present on the pectoral fin

rays as in N. raneyi, but they are most often present

on rays 2 to 6.

Nuptial crest: Small or moderate nuptial crests
NUMBER 85 39

are present on tuberculate specimens 195 to 200

mm SL. A large male, 195 mm SL, taken 19 June

1962 (Figure 10) has a moderate crest. A specimen

183 mm SL captured 6 July 1963 (Figure 16) has

no trace of head swelling and has the typical sharp,

pointy subadult snout. Two specimens, 198 mm SL

taken on 3 May 1964 and 200 mm taken on 10

May 1964, have moderate to large crests. The crest

is highest just posterior to the interorbital area. No

male N. platyrhynchus in the "peak" of crest development, which occurs in other Nocomis during

spawning or just before, has been captured or observed in life.

Pharyngeal dentition: The tooth count is 4-4

(12 pairs of arches), the teeth being moderate in

length and width. The arch is only moderately

broad, the edentulous processes comparatively long

and slender (Figure 18). The pitted surface is sculp

tured more extensively than in N. raneyi. The arch

in N. platyrhynchus is very similar to that of N.

micropogon.

Coloration: The coloration of JV. platyrhynchus is

similar to that of N. raneyi except that the former

retains the dark, lateral body stripe in the adults

of both sexes (as in N. micropogon) and the caudal

fin is more olive-orange (as in certain populations

of N. micropogon) rather than the reddish color of

N. raneyi. Some adult males of N. platyrhynchus

have lemon-yellow pelvic fins, not noted in any

other species of Nocomis.

The dark horizontal midlateral stripe is retained

in large adult females, nonbreeding and tuberculate adult males. In both sexes of N. raneyi, however, the stripe is lost before these stages of maturity.

In life, the holotype and topoparatypes of adult

females, subadults or nonbreeding adult males and

juveniles of N. platyrhynchus captured elsewhere

have the median scale color along the lateral band

area light to moderate pale green. The color appeared to be deep-lying. The scale edges were

rather sharply defined from the iridescent scale

medians dorsad and ventrad. Almost immediately

upon placing each specimen in formalin, the green

disappeared and was replaced by dark gray. The

occurrence, area of development, and intensity of

the lateral stripe are variable. The stripe is present

in most size classes of both sexes including the three

available tuberculate males. The stripe is sometimes absent from juveniles, subadults, and adults

in the same collections containing similar specimens with a stripe. It is absent most often from

poorly preserved or faded specimens. In most specimens with a stripe, it is developed mainly on the

posterior half of the body, but it is developed in

many specimens the entire body length. The dark

scale margins on the lateral body are slightly wider

in N. platyrhynchus than in N. raneyi and slightly

narrower than in N. micropogon.

The caudal fins of many juveniles and subadults

observed in life and fresh specimens captured by the

use of fish poisons were olive-orange. Some postnuptial and nonbreeding adult males had reddish colors

on the outer membranes of the caudal and dorsal

fins. The pelvic fins of recently preserved, large

specimens (USNM 194870) were yellow over most

of their length, but this color was absent in many

other specimens captured and observed before preservation.

We have not seen a living, nuptial male in closeup or a freshly captured specimen. A large, crested

tuberculate male, 200 mm SL (USNM 194816) captured and observed on 10 May 1964 by James E.

Carico, Lynchburg College, had the "ventral part of

the head and belly pink to the anus."

REPRODUCTION AND GROWTH.—Over the past several years large nests of the bigmouth chub were

observed in the Greenbrier River system during

May. The nests, often, were over three feet in diameter and always in the deeper, swifter channels of

the stream where it was particularly difficult to carry

on observations. The length-frequencies for 255

males and females is summarized in Table 6. The

largest specimen was a male, 222 mm SL. As in

N. raneyi, the males attain a much larger size than

the females, only a few females exceeding 120 mm.

ETYMOLOGY.—The specific name, platyrhynchus,

and the vernacular name, bigmouth chub, is in reference to the large gape width.

MATERIALS EXAMINED.—Holotype: USNM 194817,

a nontuberculate male 163 mm in standard length,

captured in Big Walker Creek, Bland County, Virginia, 0.25 roadmiles west of Bland-Giles county

line on Rte. 670 on 6 July 1963 with shocker by

Robert E. Jenkins, Robert L. Miles, and Fred D.

Hinson. Seven paratypes, USNM 194877, were taken

with the holotype.

Only materials from Virginia and North Carolina

are designated as paratypes.
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The collections are listed below by drainages and

major tributaries in progressive upstream order;

following the collection date is the number of specimens captured (in parentheses).

New River Drainage, Virginia and North Carolina

VIRGINIA COLLECTIONS

Main Channel, New River, Montgomery-Pulaski line.—

USNM 194870: about 05 mi downstream McCoy, 3-5 July

1964 (13).

Main Channel, New River, Grayson County.—USNM 194819:

at 1 mi upstream Rte. 58-221 bridge, 4 mi W Galax, 10 May

1964 (1).

East River, Giles County.—UMMZ 95253: Glenlyn, 11 September 1928 (1).

Wolf Creek, Bland County.-USNM 56871: Rocky Gap,

23 July 1900 (4). USNM 177380: at 15 rdmi E. Grapefield,

5 August 1957 (21). USNM 194812: Clear Fork, at 15 airmi

SW Rocky Gap, 7 August 1957 (3). USNM 194815: at Camp

Roland, 3.1 airmi W Bastian, 6.75 airmi SW Rocky Gap,

6 August 1957 (8). UMMZ 139446: between Bastian and

Norris, 22 August 1936 (3).

Big Walker Creek, Bland County.-CU 47593: at 1 mi SE

Crandon, 11 July 1957 (23). USNM 177353: same locality,

12 July 1957 (4).

Little Walker Creek, Pulaski County.-USNM 194822: at

6.8 rdmi above mouth, 29 August 1957 (2). USNM 194257:

on Rte. 601, 35 mi W jet. Rtes. 100-601, 7.1 airmi NW

Dublin, 1 July 1957 (10).

No Business Creek, Bland County.—USNM 194805: first Rte.

608 bridge, about 1 mi. N. of its mouth, 3 July 1957 (6).

Toms Creek, Montgomery County.—CU 47590: bridge just

downstream from mouth Poverty Creek, 65 mi W Blacksburg,

3 May 1964 (1).

Little River, Floyd County.—CU 24833: Rte. 8 bridge, 4.6

mi S Riner, 2 July 1952 (14). UR 887: same locality, 2 July

1958 (13). CU 41815: Rte. 221 bridge, 62 mi. NE Floyd,

31 March 1962 (6). USNM 194810: Graysontown, Rte. 699

bridge, 7 airmi S Radford, 23 August 1957 (8).

West Fork, Little River, Floyd County.-CU 24801: Rte. 8

bridge, 25 mi ENE Floyd, 2 July 1952 (5). CU 47592: (no

specific data) 28 April 1962 (6).

Big Reed Island Creek, Carroll County.-USNM 194808:

Rte. 654 bridge, 2 mi S. jet. Rte. 654-58, 9.7 airmi ESE Hillsville, 15 July 1957 (3). USNM 177400, New R. drainage (no

specific data) (29).

Snake Creek, Carroll County.-USNM 177381: Rte. 58

bridge, 6 airmi ESE Hillsville, 16 July 1957 (27).

Little Reed Island Creek, Carroll County.-USNM 194813:

at 05 mi N jet. Rtes. 52-769 on 769, 20 August 1957 (7).

USNM 194816: bridge at jet. Rtes. 769-771, 20 August 1957

(9).

Reed Creek, Wythe County.-USNM 89439: about 4 mi E

Wytheville, 6 August 1888 (1). USNM 194251: at 1.4 rivermi

from mouth, 6.8 airmi ESE Max Meadows, 12 August 1957

(5). USNM 194807: Max Meadows, 0.4 mi SW of town,

12 August 1957 (6). USNM 177403: at 1 mi above Rte. 11

bridge, 35 airmi W jet. Rtes. 11-52 in Wytheville, 13 August

1957 (34).

Cripple Creek, Wythe County.-USNM 194809: at 3.8 rivermi above mouth, 1.8 mi NW Ivanhoe, 13 August 1957 (7).

Elk Creek, Grayson County.—USNM 194260: Comers Rock,

10.8 airmi NNW Independence, 25 July 1957 (35). CU 25812:

mouth 7 mi W Galax on Rte. 58, 28 August 1953 (140).

Fox Creek, Grayson County.—USNM 194814: mouth Middle

Fox Creek, jet. Rtes. 678-711, 95 airmi NW Independence,

13 July 1963 (2). USNM 194821: just below mouth Little

Fox Creek, \2 mi NW on Rte. 711 from jet. Rtes. 58-711,

8.75 airmi W. Independence, 23 July 1957 (1). USNM 194811:

(no specific date) May 1964 (2).

NORTH CAROLINA COLLECTIONS

Little River, Allegheny County.-UMMZ 147552: Rte. 21

bridge, 1 mi SE Sparta, 27 August 1946 (11). UMMZ 165439:

at 7 mi NE Sparta, 23 July 1953 (34).

North Fork, New River, Ashe County.-USNM 171757: at

1 mi NNE Warrenville, 1 April 1940 (25). DU (unnumbered): near Bina, 100 yds above mouth Big Horse Creek,

25 July 1941 (2). DU (unnumbered): 25 mi above mouth,

26 July 1941 (2).

South Fork, New River, Ashe County.—DU (unnumbered):

at 3.5 mi NW Scottsville, 28 June 1949 (5). TU 25902: at

4 mi SW Scottsville, 19 June 1962 (17).

South Fork, New River, Watauga County.—CU 47591: Rte.

221 bridge, 1.6 mi E Boone, 8 September 1947 (33). USNM

166408: at 05 mi below Todd, 24 August 1941 (4). CU

21865: Rte. 221 bridge between Boone and Deep Gap, 22 July

1952 (43). DU (unnumbered: mouth Fighting Creek, 27

August 1949 (4).

Material not designated as paratypes.

NEW RIVER TRIBUTARIES, VIRGINIA

West Fork, Little River, Floyd County.-USNM 194252: Rte.

720 bridge 05 mi from jet. Rtes. 720-729, 30 July 1957 (1).

Dismal Creek, Bland-Giles County line.-USNM 194820: the

"Cascades," about 6.8 airmi NW White Gate, 10 October

1956 (2).

Wolf Creek, Bland County.—USNM 56666: Rocky Gap, 23

July 1900 (1).

NEW RIVER AND TRIBUTARIES, WEST VIRGINIA

Main channel, New River, Raleigh County.—USNM 203916:

8.2 mi below Hinton, 1 September 1966 (87).

Main channel, New River, Fayette County.—USNM 204451:

at Power Dam pool upstream from Rte. 21; just below Hawks

Nest State Park above Kanawha Falls, 18 September 1968 (1).

Indian Creek, Monroe County.—CU 25772: at 1.8 mi SE

Monroe line on Rte. 12, 25 August 1953 (5). UMMZ 118782:

midway between Red Sulphur Spring and Greenville, 15 July

1933 (3). UMMZ 118793: Greenville, 15 July 1933 (22).

Turkey Creek of Indian Creek, Monroe County.—UMMZ

119163: 28 August 1933 (8).

Blucstone River.-USNM 56828: Abbs Valley W. P. Hay,

collector (4). USNM 56982: Abbs Valley, 29 July 1900 (7).
NUMBER 85 41

Camp Creek, Mercer County.—UMMZ 95272: 11 September

1928 (19).

Rock Creek, Mercer County.-UMMZ 118737: 9 July 1933

(7). UMMZ 95261: along hwy. N of Princeton, 11 September

1928 (10).

Rich Creek, Mercer County—UMMZ 118745: 2 mi above

Spanishburg, 9 August 1933 (3).

Second Creek, Greenbrier River, Greenbrier-Monroe County

line.-UMMZ 95230: S of Ronceverte, 10 September 1928 (3).

UMMZ 118855: at Nickells Mill, 5 mi from mouth, 18 July

1933 (2).

Greenbrier River, Greenbrier County.—USNM 203917: first

riffle area below bridge at Anthony downstream to just below

mouth of Anthony Creek, 31 August 1966 (220).

Muddy Creek, Greenbrier River, Greenbrier County.—

USNM 203915: 1 mi above Alderson, 11 August 1966 (18).

Second Creek, Greenbrier River, Greenbrier County.—USNM

171697: 30 July 1931 (5). UMMZ 118842: near GreenbrierMonroe Co. line, 17 July 1933 (11).

Ozle Creek, Greenbrier River, Greenbrier County.—USNM

194824: on Rte. 60, 3 mi E White Sulphur Springs, 28 June

1955 (1).

Howards Creek, Greenbrier River, Greenbrier County.—

USNM 194825: W of White Sulphur Springs, 13 October 1956

(1). UMMZ 95222: south edge of White Sulphur Springs and

2 mi below, 10 September 1928 (27).

Greenbrier River, Pocahontas County—USNM 56560: Durbin, 21 July 1899 (2). CU 20673: Durbin, 5 June 1951 (9).

USNM 171610: near Bartow, 27 June 1935 (4). CU 23859:

just E. Bartow, 5 June 1951 (9).

East Fork, Greenbrier River, Pocahontas County.—USNM

194826: at 1 mi E Bartow, jet. Rtes. 28-250, 14 October 1956

(11).

Deer Creek, Greenbrier River, Pocahontas County.—USNM

194823: on Rte. 28, 4 mi E Cass near jet. Rtes. 8-28, 1 mi SW

Greenbank (5). UMMZ 118886: Boyer, 21 July 1933 (1).

North Fork, Greenbrier River, and Deer Creek.—UMMZ

118899: Greenbank, 18 July 1933 (1).

Gauley River, Nicholas County.—CU 20866: Rte. 41, 3 mi

S Summersville, 4 October 1951 (47).

Twenty-mile Creek, Gauley River, Nicholas County.—UM

MZ 95281: mouth about 6 mi above Gauley Bridge, 12 September 1928 (7). UMMZ 119065: Dixie, 15 August 1933 (5).

Peters Creek, Gauley River, Nicholas County.-UMMZ 119029:

at 3 mi W Summersville, 7 July 1933 (4).

Muddlety Creek, Gauley River, Nicholas County.—UMMZ

108168: 1 May 1932 (5).

Gauley River, Webster County.-UMMZ 119008: above Bolair, 7 August 1933 (1). UMMZ 165755: at 5 mi above Bolair,

8 August 1953 (28).

Cranberry River, Gauley River, Webster County.—UMMZ

165699: at mouth of Dogway, 4 August 1953 (5).

Williams River, Gauley River, Webster County.—UMMZ

165706: at mouth of Three Forks, 6 August 1953 (17). UMMZ

118951: by a beaver dammed trib. to Gauley, 3 August 1933

(25).

DISTRIBUTION.—N. platyrhynchus is a western Appalachian slope species (Figure 19), confined to, and

widely distributed in, the New River drainage of

West Virginia, Virginia, and North Carolina. Kanawha Falls, located at the town of Glen Ferris,

Fayette County, West Virginia, is herein regarded

as the geologic and faunistic divide of the NewKanawha River system into two major portions.

The New River drainage is that portion above this

barrier (including Sandstone Falls) to its headwaters, the North and South Forks in northwestern

North Carolina. The Kanawha River drainage is

the shorter portion below the Falls to its mouth in

the Ohio River. The division between the upper

and lower New drainage is recognized as the boundary of the Ridge and Valley province and the interior Allegheny Plateau. The New River leaves the

Ridge and Valley province at approximately the

Virginia-West Virginia state line (Dietrich, 1959:

12).

N. platyrhynchus occurs throughout the upper

New drainage. Fifty records from 48 localities are

from almost all major tributaries and the main

channel. Within the lower New, this species is

known from four major tributaries. Four records

are from Indian Creek, which enters the New just

downstream from the upper-lower New drainage

boundary, and ten records are from Gauley River,

which flows into the New about one mile above

Kanawha Falls. Entering the New between these

tributaries are the Greenbrier and Bluestone rivers,

from which we have 23 collections. One collection

of 87 specimens was taken in the lower main channel of the New River, (USNM 203916) 8.2 miles

below Hinton, resulting from a poison station to

survey the large river fauna by the West Virginia

Department of Natural Resources. The bigmouth

chub was the most common fish species encountered.

Many juveniles were not picked up. N. platyrhynchus is also abundant in the Greenbrier River. The

collection of 220 specimens (USNM 203917), taken

just below Anthony, of juveniles to subadults represent only a sample from a poison station also by

the West Virginia Department of Natural Resources.

The bigmouth chub was also the most common

species observed at this locality.
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Nocomis tnicropogon (Cope)

RIVER CHUB

FIGURES 16, 17, 22-25

Ceratichthys tnicropogon Cope, 1864: 277 [type, ANSP 5061,

Conestoga River, Pennsylvania]; 1869: 366, pi. 12: fig. 2

[redescription].—Fowler, 1909: 550, pi. 27.

Hybopsis kentuckiensis.—Jordan, 1889a: 110 [part]; 1889b: 9,

pi. 3: figs. 10, ll.-Goldsborough and Clark, 1908: 36 [part].

NOMENCLATURE.—N. micropogon was briefly described by Cope (1864:277) from a single specimen

from the Conestoga River, a tributary of the Susquehanna River, Pennsylvania, collected by a member of the Linnaean Society of Lancaster probably

around 1863. Cope (1869:366), when redescribing

the specimen, stated that it might be a hybrid

Ceratichthys [= Nocomis] X Hypsilepis [= Notropis

(Luxilus)] cornutus. Thereafter specific status of

micropogon was questioned or it was synonymized

with other species of Nocomis, until Hubbs (1926:

27-29) used micropogon as the name for a valid

species of Nocomis. Hubbs apparently based his decision on earlier descriptions of it by Cope (1864;

1869:366, pi. 12: fig. 2) and Fowler (1909:550-552)

and because only one species was known to occur in

the Susquehanna drainage.

Examination of the type (ANSP 5061, 67 mm SL)

shows that it apparently is a hybrid Nocomis micropogon x Notropis cornutus. Cope (1869) correctly

recognized that, in particular, its head resembles

that of Notropis cornutus. Its anterior lateral scales

are more elevated and the lateral line is more downcurved than in Nocomis and thus indicates Notropis

(Luxilus) parentage. Mouth position and angle, very

small size of the two barbels, and moderately large

eye size are approximately intermediate between

these two stocks. The breast is considerably more

scaled than the Susquehanna population of N. micropogon. The holotype resembles other hybrid

specimens of N. micropogon X N. cornutus, which

is a common combination in our hybrid studies.

The presence of a tooth in the minor row of the

type would also tend to confirm its hybrid origin.

The pharyngeal teeth in N. micropogon are always

4-4, in N. cornutus 2,4-4,2. Cope (1869) reported

the type to have 4-4 teeth but Fowler (1909:550)

gave 1?,4-4,1?. It is not clear, but probably the

arches have at least one minor tooth row; the numbers could be interpreted as 4-4 to 1,4-4,2 depending upon whether the cavities anterior to the major

tooth row are regarded as tooth scars or foramina.

Based upon our examination of many other Nocomis arches, at least one tooth in the minor row

was present but lost when dissected by Cope. The

parentage of N. cornutus is also indicated by the

long and well-hooked lowermost tooth of each arch;

the lowermost one in Nocomis is relatively short

with a conic, lesser hook on the head.

Other characters of the type are lateral line scales

about 39, caudal peduncle scales 16, anal rays 7,

pectoral rays 18. Other scale counts could not be

made since some scales are missing. Color pattern is

not obvious since the specimen is faded.

Article 17(2) of the International Code of Zoological Nomenclature (1961:17) states that a name is or

remains available even though "in the case of a

species-group name, it is found that the original

description relates to .. . an animal or animals later

found to be hybrid; . . ." In accordance with the

Code, we restrict the name Ceratichthys micropogon

Cope to that presumed parent of the type specimen.

Only a few particular references pertaining to the

central Appalachian area are considered in the synonymy of this wide-ranging, frequently encountered species.

DIAGNOSIS.—A species most closely related to N.

platyrhynchus but differs in having fewer tubercles,

reduced tubercle pattern, and larger scales. The

head tubercles in adults are grouped from the snout

to the interorbital area and do not extend posteriorly on the midline beyond the anterior interorbital

area, forming a V-shaped pattern. Snout tubercles

almost always not forming a confluent pattern with

those at anterior internasal area thus causing a hiatus

in tuberculation. Tubercles number fewer than 60

in almost all adults. The means for total tubercle

numbers for the various river populations range

from about 30 to 65. Scales larger, the mean values

for circumferential scale rows and lateral line scales

(Tables 8, 12) are all lower in N. micropogon than

in N. platyrhynchus.

DESCRIPTION.—Morphometry: Proportional data

is given in Table 25 for 51 specimens, 29 of which

were grouped from the Potomac, Rappahannock,

and James River drainages, with a mean length of

165 mm SL and ranging from 142 to 189 mm and

22 specimens from the Tennessee River drainage

with a mean length of 161 mm and ranging from
NUMBER 85 43

141 to 188 mm. Although the Tennessee population

shows the greatest reduction in tubercle numbers,

the morphometric data agrees with that of other

N. micropogon drainage populations. The head

length of the Tennessee population may be somewhat shorter (mean, 27.2% of body length) than the

Potomac-James group (mean, 28.4%).

The proportional features of N. micropogon are

generally similar to those of AT. platyrhynchus. There

is some difference in the gape width between these

species (Figure 5; see N. raneyi, p. 18), the heights

of the regression lines are statistically different.

This difference is of a subtle nature (Figure 7) and

often when disassociated with body size, it was of

no systematic use. A juvenile N. micropogon is

shown on Figure 17.

The slope of the preopercle-opercle suture is similar to that of N. platyrhynchus; the lower portion

is most often perpendicular to the horizontal body

axis or it is slightly posteriorly directed (70% of 200

specimens), 24 percent with suture line very slightly

directed forward, and 6 percent with suture line

directed forward at an angle of about 10° to 20°.

The intestine is simple, elongate and S-shaped

(39% in a sample of 75 specimens), or with a very

short kink (45%) or moderate kink (16%, see Figure

8a for N. raneyi).

Meristic characters: Many of the meristic characters were segregated by drainages in order to compare populations of this wide-ranging species, and

possibly to associate related populations with early

dispersal and stream capture. Most of the mean

values for the meristic characters in N. micropogon

are lowest for its species group, but are higher than

those for N. leptocephalus. The range and mean

values are given for as many as 19 drainages or

regions for circumferential scale rows (Table 12),

lateral line scales (Table 8), scale rows above and

below the lateral line (Table 11), rows of scales

around the caudal peduncle (Table 14), number of

pectoral fin rays (Table 16) and vertebral numbers

(Table 15). The higher means for the circumferential scale rows in the Monongahela, Youghiogheny,

and Potomac drainages may have been influenced

by relatively recent gene interchange. The higher

values in the Tennessee and Cumberland drainages

contrast with the low values in nearby Kentucky,

Big Sandy, and Kanawha populations. These differences also may be linked with recent drainage exchanges of N. micropogon stocks.

The breast squamation (Table 18) differs among

populations from those having scaleless or only

slightly scaled breasts to those completely scaled or

nearly so. There is also great variation in this character within some drainages. For example, the Tennessee and Cumberland drainages contain specimens

with breasts ranging from scaleless to fully scaled.

The Atlantic slope drainages from the Susquehanna

southward to the James consistently have few or no

scales on the breast. The major drainages west of

the divide, such as the Tennessee, Cumberland,

Kentucky, Kanawha tributaries, Monongahela and

Allegheny, show consistently higher values for the

degree of breast squamation.

Tuberculation: The development, distributional

pattern, and numbers of tubercles for N. microFICURE 22.—Head of nuptial male Nocomis micropogon,

USNM 166416, 165 mm, captured in Stone Creek, Juniata

River, Susquehanna drainage, Pennsylvania, 3 June 1948:

a, lateral view; b, dorsal view. (Drawn by A. M. Awl.)
4

4 SMITHSONIA

N CONTRIBUTIONS TO ZOOLOGY

A

a

a

/ ¥ ' ,

pogon is summarized in Tables 3, 4, 7, 9,10, and 13.

The relationship between increase in tubercle num-

bers with increase in body length for three impor-

tant drainage populations of AT. micropogon, and

compared with N. leptocephalus, N. platyrhynchus,

and N. raneyi, is shown in Figure 4. Early develop-

ment and pattern of tubercle spots is illustrated in

Figure 3. The distribution of head tubercles in non-

breeding males and nuptial males is shown in Fig-

ures 16,23-25. The tuberculation in N. micropogon

is characterized as having fewer tubercle numbers

than N. platyrhynchus (and much fewer than N.

raneyi) but appreciably more than N. leptocephalus

(Figure 4). The tubercles average slightly larger than

those of N. platyrhynchus but much smaller than in

N. leptocephalus (Figure 24). The tubercles are dis-

tributed from the snout to the anterior interorbital

area, with a hiatus almost always present between

the snout and the anterior internasal area.

The tubercles first appear as light spots in the

internasal area (Figure 3). They first appear in speci-

mens 50 to 60 mm SL of both sexes, but they are

sometimes not developed at sizes ranging from 60

to 75 mm SL. With increase in body length the

tubercles develop on the tip of the snout and also

spread posteriorly to the anterior interorbital area,

where a V-pattern is formed by the absence of tuber-

cles along the midline. The increment in tubercle

numbers with increase in body length is considerably

reduced at about 120 mm SL for all the river popu-

lations except the Potomac, in which reduction oc-

curs at about 150 mm SL. The lachrymal tubercles

are slowest in developing. There is usually only one

or two rows of lachrymal tubercles in JV. micropogon.

Tubercles are present on pectoral fin rays 2 to 4

(2 specimens); 2 to 5 (5); 2 to 6 (37), and 2 to 7 (10).

Significant aspects of the tuberculation in N. micropogon in the Potomac and Monongahela drainFIGURE 23—The distribution and number of head tubercles

and crests of nuptial males of Nocomis micropogon and

Nocomis leptocephalus. Observe absence of snout and sub-

nasal tubercles in N. leptocephalus: a, N. micropogon, USNM

166485, 166 mm, collected 3 June 1948, Juniata River, Sus-

quehanna drainage; b, N. micropogon, CU 41642, 168 mm,

collected 24 May 1961, Tioughnioga River, Susquehanna

drainage; c, N. leptocephalus, Duke University Collection,

145 mm, Eno River, Neuse drainage; d, N. leptocephalus,

USNM 194715, 157 mm, collected in May 1964, Roanoke

drainage.
NUMBER 85 45

ages are the higher numbers of tubercles and a more

posterior distribution, sometimes extending to the

posterior interorbital area. In tuberculation, and in

FIGURE 24.—Dorsal views of heads of Nocomis micropogon

and N. leptocephalus showing size and distribution of tubercle

development, a, N. micropogon, USNM 166485, 168 mm, taken

3 June 1948, Juniata River, Susquehanna drainage, b, N.

micropogon, CU 41642, 168 mm, taken 24 May 1961, Tioughnioga River, Susquehanna drainage. A hiatus, typical for

AT. micropogon, is present between tubercles on snout and

internasal areas. The hiatus is absent in the specimen above.

c, N. leptocephahts, USNM 194715, 159 mm, taken May 1964,

Roanoke drainage. Virginia. Large tubercles develop in the

internasal-interorbital area and never on the snout in this

species.

some meristic characters, N. micropogon of these

drainages closely approaches character values of N.

platyrhynchus. Possible introgressive hybridization

of these two species resulting from exchanges of

stocks via the Greenbrier-Monongahela-Potomac

drainages is discussed in the section on dispersal

(p. 66). The Potomac and Monongahela populations

are very similar in respect to total number of tubercles, increase in numbers with increase in body length

and the relative variation of tubercle numbers by

size groups (Table 7). The James drainage population (Table 10), which is to the south of the Potomac, and the Susquehanna population, which is to

the north (Table 13), have similar numbers of tubercles and appreciably fewer than the Potomac population. N. micropogon from the Tennessee drainage

has the lowest number of tubercles (Table 9) for the

species. The average values for tubercle numbers of

mature males of varying body lengths were: Potomac-Monongahela population, about 50 to 65;

James, Susquehanna, and west of Allegheny Mountains, about 40 to 50; Tennessee drainage, about 30.

Nuptial crest: Nuptial crests or swellings were observed on specimens of various sizes. Some swelling

and moderate to large crests were found on specimens in the following standard length size-groups:

100-119, 2 specimens; 120-139, 2 (4 with crests);

140-159, 5 (19); 160-179, 6 (10); 180-199, 3 (5);

200-219, 1 (1). These data, as with that pertaining

to the development of tubercle buds (Table 4) and

the presence of tubercles or scars (Table 3), suggest

that different males mature at different sizes or ages,

or that they mature and spawn more than once.

Since tubercle scars are present on nonbreeding

males that are developing tubercle buds, it is assumed that some males spawn more than one season.

Pharyngeal dentition: The tooth count is 4-4

(Figure 18) in 34 pairs of arches in which the specimens were sampled over the range of the species.

The teeth and arch are like that of N. platyrhynchus.

Coloration: Only particular aspects of the coloration of N. micropogon will be discussed because

much of the salient color and pattern has been

treated above under the two other group members.

Lachner (1952: 437-439) described the coloration of

N. micropogon, chiefly from specimens from the

northeastern portion of its range, the Allegheny,

upper and middle Susquehanna and Lake Ontario

drainages. It was noted that a dark horizontal mid-
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lateral stripe is prominent in the young and juveniles of N. micropogon while only faintly visible in

formalin preserved specimens of adult males. In the

large number of preserved adults now available, we

find that a majority of specimens have a dark lateral

stripe, but it may be faded or absent in preservation. The lateral stripe is present in specimens over

the geographic range of the species. Its relative frequency of occurrence, portions of the body on which

it is developed and the intensity in adult males and

females agree with that of N. platyrhynchus.

Reighard (1943:400) observed that living young

males of N. micropogon with tuberculate heads have

a well-marked lateral stripe. He did not state this

for the large nuptial males engaged in reproductive

activities. Lachner (1952:438) stated that nuptial

male N. micropogon do not show a conspicuous

dark lateral stripe in life as was observed in N.

biguttatus. Our recent observations on nest-building

males from the upper Susquehanna drainage show

that a light stripe is effected by light green medians

of the scales, or a moderate to heavy dark stripe by

the blackish scale margins on the midlateral level;

the intensity of the stripe can change rapidly. The

light green medians of the scales were also seen in

living juvenile and adult males and females from

the James, Potomac, and upper Tennessee drainages. Upon preservation, the medians of the scales

turn light to dark slate color. The lateral stripe was

similar to that of living N. platyrhynchus.

The pink-rosy nuptial coloration of the male is

developed in the same area of the body as in N.

raneyi and N. platyrhynchus. It has been observed

in males from Lake Ontario, upper and middle

Susquehanna, Rappahannock, James, upper Tennessee, and Allegheny drainages. Reighard (1943:

400) reported it from Michigan. The most intensely

colored male we have seen, 160 mm SL (CU 48579),

was captured in the upper Susquehanna on 30 May

1965 during the spawning period. Its rosy color was

developed dorsad to three scale-rows above the lateral line. The medians of the scales in the lateral

stripe area were very light pink, lighter than those

above and below, with the light green color barely

present.

The olive-yellow middorsal stripe described for

juveniles of both sexes and smaller adult females of

N. raneyi is generally present in the same life stages

and sexes of N. micropogon as well as in N. platyrhynchus. It was also present, but faint, on the

largely blackish dorsum of a captured living nuptial

male N. micropogon of 150 mm SL.

The coloration of the paired and anal fins of N.

micropogon is similar to that of both N. raneyi and

N. platyrhynchus. The yellow pelvic fin coloration

of a nuptial male after one week in preservative

approached that of the specimens of N. platyrhynchus (USNM 194870). Lachner (1952:438) noted

white coloration on the tips of the rays of the paired

and anal fins of nuptial males and we have observed

it on the tips of the anterior dorsal fin rays of a

nuptial male.

Caudal and dorsal fin coloration may show some

geographic differentiation. There appear to be two

types of caudal color pattern, an orange-reddish one

in populations from the southern part of the range,

and an olive, olive-orange, or olive-yellow one in

northern populations. The distinctiveness of these

color differences is not clear and little data are available on variation in shades. The caudal fin in juveniles to adults of the southern populations is fairly

reddish in color, approaching that of N. raneyi, and

much like that of N. platyrhynchus. The distal

caudal and dorsal membranes were light orange-red,

the basal membranes being oHve in several postnuptial males and females and' nonbreeding adult

males collected in the upper James drainage during

late July 1963. A highly tuberculate crested male

with nuptial color from the Rappahannock drainage had an orange-reddish outer caudal. The most

striking caudal coloration yet seen in N. micropogon

was that of five postnuptial males (USNM 194689)

having tubercles or tubercle scars, crests, and pink

lower coloration, taken in the upper North Fork of

Holston River, Tennessee drainage on 2 July 1963.

The caudal fin color, a bright orange-red, was developed over most of the fin which was largely olive

basally.

The orange-red caudal fin (color over half of

entire fin and, in some, over anterior distal dorsal

fin) has been observed by us in living or freshly

preserved specimens of the following growth stages:

lower Potomac, living juveniles; Shenandoah of Potomac, preserved adult males; upper Rappahannock,

preserved juveniles and an adult male; upper James,

living juveniles and adults; Powell, Clinch, and

North Fork Holston of upper Tennessee, living

juveniles and adults. Carter R. Gilbert (personal
NUMBER 85 47

communication) has observed N. micropogon with

reddish caudals in the upper Cumberland drainage.

The "olive" caudal fin appears to be characteristic

of northern populations. In Catatonk Creek, upper

Susquehanna drainage, juvenile and adult females

and nonbreeding males generally have yellow-olive

rays, sometimes appearing amber. Distally the membranes are translucent grayish olive; basally they

are more grayish, tending to be clear. All specimens

from Catatonk, except one nuptial male captured

in May 1965 lacked reddish or orangish coloration.

The exception, a highly rosy-colored male N. micropogon had a faint yellow-orange in the distal one

half to two thirds of the membranes of the caudal

lobes and a tinge of this color in the distal portion

of the medial membranes. Lachner (1952:438) stated

that the dorsal and caudal fins of N. micropogon

were yellowish with some red near the tips of the

rays; his specimens were from Lake Ontario tributaries, upper and middle Susquehanna, and the

Allegheny drainages. Hubbs and Lagler (1958:69)

state that the caudal fin of N. micropogon is not

red, in contrast to the red caudal fin of young

N. biguttatus in life. Trautman (1957:86, 295) stated

that N. micropogon in Ohio (Great Lakes and Ohio

River basins) has a slate-colored caudal that is never

flushed with red, that it may be a faint orange in

some young and that it may have a reddish or orange

tinge in specimens only from polluted waters. Information is lacking on details of caudal fin coloration

in the Ohio basin of Indiana and much of Kentucky

and West Virginia. Most of the freshly preserved

juveniles and a ripe female in two series from Deer

Creek, Susquehanna River, near its mouth, taken

during the 1965 spawning season, had faint tinges

of reddish orange on the distal membranes of the

caudal.

In summary, there appears to be some evidence

that two caudal fin color forms exist. The difficulty

in evaluating the significance of these color forms is

that we lack comparative data. We have observed

the two forms from different geographical areas, but

we know little of the exact areas or drainages each

occupies, and of the intermediate areas or regions

in which both forms may occur. We know that some

colors can be intensely developed, or "turned on,"

in Nocomis within a few seconds, but this was not

noted for caudal color. We have resorted to color

guides only in the laboratory; none were used in

the field other than, occasionally, colored objects,

such as differently colored plastic chips. The shades

of red, reddish orange or olive-orange, and the reflections from these colors, in free-living and freshly

captured specimens may be variously interpreted by

different individuals at different times. Specimens

were seen, or available for observation, one at a

time, spread out over years of observations by any

one individual, and thus comparative, objective data

were not obtained.

POPULATIONS.—The northern drainage populaFIGURE 25.—Crested, nuptial male Nocomis micropogon, USNM 166416, 165 ram, captured in

Stone Creek, Juniata River, Susquehanna drainage, Pennsylvania, 3 June 1948. (Drawn by Carolyn

Bartlett Cast.)
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tions of the river chub, from Michigan to New York,

are very much alike. The principle area of differentiation, excluding the caudal fin color forms discussed above, is in the drainages of the central

Appalachian region. Here the interesting divergence

is in the Potomac and upper Monongahela drainage

populations, both having finer scales, higher tubercle numbers, and more extensive distribution of

tubercles than in river chubs of adjacent drainages.

The Tennessee drainage population has a notable

reduction in numbers of head tubercles, as well as

having shorter heads. The most southern population of the Atlantic slope, in the James drainage,

resembles the northern populations in respect to

tubercle numbers and distribution. Populations in

the Kentucky drainage and in tributaries of the

Kanawha drainage are fine-scaled compared to Cumberland and Tennessee drainage stocks. N. micropogon of the Atlantic slope streams typically have

reduced squamation on the breast or have the naked

condition.

We find no significant differentiation over the

range of the river chub to merit recognition of subspecific populations. The similarities in some of the

drainage populations may be related to stream capture and early dispersal of stocks.

REPRODUCTION AND GROWTH.—The nest-building,

reproduction, age and growth of N. micropogon is

discussed by Lachner (1952) and compared with N.

biguttatus and N. leptocephalus. Of the thousands

of specimens recently examined, one specimen of

N. micropogon (Table 7) from the Potomac drainage just exceeded 200 mm SL and another, by far

the largest specimen known, measured about 270

mm SL. This large crested nuptial male (TU 8928)

is recorded from Lake Pontchartrain, Mandeville,

Louisiana, May 1877, "collection of G. Kohn, New

Orleans, La." A few tubercles are intact and these

plus the tubercle scars total 60, a typical number

for the species. The tubercle pattern is also normal

for the species. The locality is completely out of the

known range for N. micropogon. The specimen consists of the head and entire skin of the body attached

to the head, including the fins. It probably was an

item of curiosity and could have been hand-transported down the Mississippi by the river traffic of

the time.

MATERIALS EXAMINED.—Data were taken from 395

collections. The total number of collections studied

from each of 18 drainages is given and these are

further subdivided by the institution and number

of collections housed.

Susquehanna 24: CU 8, UMMZ 2, USNM 14. Chesapeake

Bay tribs. 10: CU 3, UMMZ 7. Potomac 30: CU 8, UMMZ 2,

USNM 20. Rappahannock 19: CU 8, UMMZ 3, UR 1, USNM

7. York 3: UR 1, USNM 2. James 54: ANSP 1, CU 11,

UMMZ 7, USNM 34. Savannah 4: CU 1, TU 2, USNM 1.

Tennessee 77: CU 3, DU 1, UMMZ 39, USNM 34. Cumberland 27: CNHM 1, CU 5, Ind. U. 1, SU 1, TU 2, UMMZ 13,

USNM 4. Kentucky 21: CNHM 1, KFW 2, UMMZ 5, USNM

13. Big Sandy 8: CU 1, UMMZ 1, USNM 6. Guyandot 5:

CU 1, UMMZ 1, USNM 3. Coal, Kanawha 10: CU 1, UMMZ

1, USNM 8. Elk, Kanawha 10: CU 5, UMMZ 5. Ohio tribs.,

W. Va. 5: CU 3, USNM 2. Monongahela 52: CU 6, USNM

46. Allegheny 20: CU 1, UMMZ 4, USNM 15. Lake ErieOntario, N.Y. 23: CU 7, USNM 16.

DISTRIBUTION.—N. micropogon is native to a large

portion of northeastern United States and does not

occur west of the Mississippi River. Its range on the

Atlantic slope is southward from the Susquehanna

drainage in New York to the James drainage in

Virginia, absent southward except for a population

in the upper Savannah drainage. The only known

population on the Gulf of Mexico slope occurs in

one river system of the upper Mobile Bay drainage.

West of the Appalachian divide it occurs from the

Tennessee drainage tributaries of northern Alabama

and southwestern Tennessee, and from die Wabash

drainage of Illinois (P. W. Smith, 1965:7), up (eastward) throughout most of the Ohio River basin. It

is found in the Great Lakes drainage of the Lower

Peninsula of Michigan eastward to Lake Ontario

tributaries in New York and Ontario. The river

chub is apparently absent from significant portions

of some drainages, and the entire length of other

drainages, mainly within the southwestern lowland

portion of this general range.

N. micropogon occurs in all major drainages tributary to the western shore of Chesapeake Bay, from

the Susquehanna to the James (Figure 19). Records

are available from throughout most of the Susquehanna drainage of New York and Pennsylvania (except from some large, poorly collected, and polluted

areas), to near its mouth in Maryland. It occurs in.

all minor Chesapeake Bay drainages of sufficient

size from Swan Creek to Patuxent River, Maryland.

Most Potomac drainage records are from throughout

the upper and lower main basin in Maryland and

Virginia including its large tributary, the Shenan-
NUMBER 85 49

doah River system of Virginia. This species may be

rare in the North and South Forks and North and

South Branches of the Potomac River, largely in

West Virginia and western Maryland. Relatively

few specimens were seen from this montane region,

but it has been poorly collected. The river chub

occurs in all major upper Rappahannock drainage

tributaries, Virginia, and has been taken from its

lower freshwater portion just above the Fall Zone.

Six locality records of N. micropogon from the York

drainage, Virginia, indicate that it is widespread

therein.

N. micropogon occurs throughout most of the

James drainage of Virginia and West Virginia. The

river chub was taken in 68 collections from 63 localities of the 182 James drainage collections that include Nocomis. The majority of the collections are

from major upper tributaries, such as the Jackson

River, Back, Dunlap, and Potts Creeks and, in the

middle portion, the James, Maury (North) River

and its tributaries. Three collections are from the

Piedmont-based Appomattox River, the longest

James tributary; two are from its headwaters, the

other in Swift Creek, a tributary entering the Appomattox near its mouth in the James River estuary.

The river chub occurs commonly above and below

the restricted range of N. raneyi in the James but is

rare within the range of N. raneyi (Figure 20). Only

13 or 14 specimens of the river chub were captured

in the Craig Creek system compared to 742 specimens of N. raneyi. Six specimens of N. micropogon

(TU 25469) were taken with one specimen of N.

leptocephalus and three hybrid N. leptocephalus x

N. micropogon from upper Johns Creek, near Maggie. N. micropogon and N. raneyi are known to have

been collected together only three times, all from

the lower half of Johns Creek. The upstream and

downstream localities are believed to be accurate

but the location of the middle station is approximate. Specimens UMMZ 135401 and 181830 were

from the latter station, which was reported by G. W.

Burton to be "seven miles above Newcastle." It was

plotted by measuring seven miles above (upstream

from) Newcastle along State Route 311 and County

Route 658. Two specimens of N. micropogon

(UMMZ 135413) are from Craig Creek, and were

collected by Burton, supposedly 25 miles west of

Newcastle. No point on Craig Creek is 25 air- or

roadmiles west of Newcastle. The collection is located as 25 roadmiles on Route 311 and County

Route 621 southwest (upstream from) Newcastle.

This probably is still erroneous since the locality

would be in the extreme headwaters. Craig Creek,

from this locality to four miles downstream, is quite

small, about 4-10 feet in width, smaller than streams

typically inhabited by the river chub. Presumably,

the specimens are from Craig Creek, although probably from somewhat below the given locality. From

correspondence of Burton it is indicated that he

collected only in the Craig system, James drainage.

Our numerous attempts to find a river chub in the

Craig system were unsuccessful although one juvenile specimen (CU 47584), tentatively identified as

this species, was collected at the mouth of Craig

Creek.

Only one specimen of N. micropogon (CU 50610)

from Catawba Creek was found, this being from the

headwaters. Hybrid N. leptocephalus x N. micropogon (USNM 171685) and N. micropogon x N.

raneyi are known from the middle Catawba. N. leptocephalus is common throughout Catawba Creek.

JV. micropogon apparently is common in sizable

streams surrounding the Craig and Catawba Creek

areas. Upriver, nearest to the range of N. raneyi, the

river chub (USNM 132064) was collected from Mill

Creek in 1885, near its mouth in the James River

at Gala, approximately five rivermiles upstream

from the mouth of Craig Creek, and in other streams

shown in Figure 19. It is widely distributed in the

Maury River whose mouth is 16 rivermiles above

that of the Pedlar River system, from which a specimen of N. raneyi was taken.

The five specimens upon which Fowler (1945:77)

based his record of N. micropogon from the Roanoke drainage were found to be Ar

. leptocephalus.

The ten records (CU, TU, USNM) of N. micropogon from the Chatooga and Keowee River systems, upper Savannah drainage, South Carolina and

Georgia, indicate widespread occurrence therein.

Another locality is in the Savannah River, Abbeville

County, South Carolina. Most of these localities

were plotted by Lachner and Jenkins (1967:fig. 7).

N. leptocephalus was taken in only one of the eleven

collections.

Our only records of N. micropogon on the Gulf

slope (TU, 5 collections) are from the Coosawattee

system (upper Coosa) of the Mobile Bay drainage in

Gilmer County, northwestern Georgia. These col-
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lections were noted by Suttkus and Ramsey (1967:

139-140).

Within the New-Kanawha system, N. micropogon

is known only below Kanawha Falls, from Coal and

Elk Rivers, the two largest tributaries of the Kanawha drainage. Specimens were taken from upper

and lower Coal River; those from Elk River are all

from the upper portion. No specimens of chubs

were available from other Kanawha tributaries. The

fauna here and elsewhere in West Virginia has been

largely extirpated by mining (Goldsborough and

Clark, 1908: 31-32; Kinney, 1964: 22-24) and industrial pollution.

The wide occurrence of N. micropogon in the

southwestern Ohio River basin was discussed and

illustrated by Lachner and Jenkins (1967). Its distribution north of the Ohio River, in Indiana and

Ohio, was analyzed and mapped respectively by

Gerking (1945:48-49, map 25) and Trautman (1957:

296, map 61). The river chub is also widely distributed in most of the remainder of the Ohio basin of

West Virginia, Pennsylvania, and New York.

Within the Great Lakes drainage, N. micropogon

occupies streams of the entire lower Peninsula of

Michigan (Hubbs and Lagler, 1958:78), thence eastward, within Ontario Province streams of the Lake

Huron basin south of Georgian Bay and the northern Lake Erie basin, to approximately the midlength

of the northern Lake Ontario basin (Radforth, 1944:

59-60, fig. 22). It ranges eastward, from the southeastern corner of Michigan, in streams of southern

Lake Erie, including the Maumee River drainage

of Ohio (Trautman, 1957:296-297, map 61), and

Lake Ontario basins to the Finger Lakes drainage of

New York. However, within the southern Lake Ontario basin, this species is apparently absent from

the entire Genesee River drainage, since it was not

reported by Greeley (1927) and later collectors. It is

common in Salmon Creek which enters Lake Ontario

six miles west of the Genesee mouth. East of the

Salmon Creek mouth there is a 52 mile gap in its

distribution, ending at Blind Sodus Creek, whose

mouth is 16 miles west of the mouth of Oswego

River, the Finger Lakes outlet. Within the Finger

Lakes drainage, N. micropogon is known from Catherine Creek, which enters the south end of Seneca

Lake, the Cayuga Lake basin and one record is from

a Seneca River tributary downstream from Cayuga

Lake.

The leptocephalus Group

DESCRIPTION.—Nuptial males develop few tubercles (Figure 4) on head, numbering fewer than 30

and often less than 20 (Table 19). In populations

of the south Atlantic and Gulf slope drainages the

nuptial males have even fewer tubercles, in some,

averaging about 6. Tubercles extend from internasal

area to occiput, depending on subspecies. Tubercles

absent on snout and lachrymal areas. A large, nuptial crest developed by larger males positioned more

anteriorly than in the micropogon group. Breeding

coloration of nuptial male bluish on head and body

or, body with orange, brassy or tan lateral stripe.

Dark marking on anterior portion of scale in adults

narrower than micropogon group. Caudal spot dark,

small, indistinct in juveniles (compared to large,

distinct spot in biguttatus group). Intestine almost

always with a ventral whorl (except in some collections from southeastern United States). Dentition

4-4 to 3-3. Scales large, body circumferential scales

range from 26 to 33. Body comparatively short and

stocky; snout blunt; posterior edge of preopercle

usually vertical or sloped posteriorly below. Males

attain an intermediate size for genus except one

southeastern population which is larger. Males of

the three subspecies construct .small to moderately

sized moundnests of fine to small gravel. The three

subspecies include Nocomis I. leptocephalus of the

Atlantic slope from the Potomac and New drainages southward to the Savannah River; a new subspecies (see paper 2, p. 2) of the southeast Appalachian slope in the Savannah, Altamaha and Apalachicola drainages; and Nocomis I. bellicus of the

southern Gulf slope in the Mobile drainage westward to certain eastern tributaries of the Mississippi

River.

Nocomis leptocephalus (Girard)

BLUEHEAD CHUB

FIGURES 17, 23, 24, 26, 27

Ceratichthys leptocephalus Girard, 1856: 213 [type locality,

Salem, North Carolina; Yakin River system].

Hybopsis kentuckiensis.—Jordan, 1889a: 123 [ (part) Roanoke

drainage].

Hybopsis leptocephala.—Ross, 1959: 12, 16, 25 [New drainage].—Ross and Perkins, 1959 [(part) New drainage].

Hybopsis micropogon.—Ross, 1959: 12-13 [(part) Potomac

drainage] .—Ross and Perkins, 1959: 16 [ (part) New drainage]-
NUMBER 8 5 51

Hybopsis species.—Ross and Perkins, 1959: 12, 19, 24, 27

[ (part) New drainage].

Nocomis leptocephalus.—Jackson and Henderson, 1942: 95

[ (part) Roanoke River].—Burton and Odum, 1945 [ (part)

James drainage],

Nocomis micropogon.—Fowler, 1945: 77 (Roanoke drainage).—

Burton and Odum, 1945 [(part) New drainage].

DIAGNOSIS.—A stout bodied species differing from

all other Nocomis in the reduced numbers of head

tubercles, almost always fewer than 25 in the typical

subspecies of the central Atlantic slope and reduced

to 6 for nuptial adults in the southern, Gulf slope

form; tubercles absent on snout, subnasal, and lachrymal areas; intestine whorled in almost all drainage populations, in all from the central Atlantic

slope; scales large, mean values for circumferential

scales for drainage populations less than 30; values

for most meristic characters are lower than in all

other species of Nocomis.

DESCRIPTION.—Morphometry: Table 25 contains

proportional data for 13 characters of 47 specimens

from the James, Roanoke, and New drainages. The

body features are fairly constant within size-groups

among the drainage populations of the Atlantic

slope as well as among the three subspecies over the

entire range of the species. Most of the variation in

characters is associated with changes related to allometric growth.

Compared to other species of Nocomis, the body

is short, stout, and comparatively deep. The snout

is short (averages 11.6% SL), the interorbital wide

(9.3% SL), and the lachrymal short (6.9% SL). The

dorsal and anal fins in the adults have a rounded

contour; the pectoral and pelvic fins are short and

rounded. Compare the deeper and more blunt head

and deeper body of the juvenile with N. micropogon

and N. raneyi in Figure 17.

The intestine is unlike any other Nocomis, having

a pronounced whorl or anterior loop extending to

the right side and covering part of the anterior alimentary tract (Figure 8), except in certain collections of the southeastern Atlantic slope subspecies.

All populations of the typical subspecies ranging

north of the Savannah drainage show constant and

relatively uniform development of the intestine. It

is typically coiled in small juveniles at 25 to 30 mm

SL. Its dark, coiled outline may be seen through

the pale belly tissue in these small sizes without

cutting open the abdominal cavity.

The slope of the preopercle-opercle suture is directed forward at the angle only slightly in 43 specimens measured and either perpendicular or posteriorly directed in 157 specimens.

Meristic characters: Summaries of meristic characters, usually for five drainage populations of the

central Atlantic area, including the range for mean

values, are as follows: circumferential scale-rows,

28.1 to 29.4 (Table 12); scale-rows along the lateral

line, 38.1 to 39.9 (Table 8); scale-rows above lateral

line, 6.1 to 6.8, and scale-rows below lateral line,

4.8 to 5.4 (Table 11); caudal peduncle scale-rows,

16.0 to 16.3 (Table 14); number of pectoral fin rays,

16.5 to 16.9 (Table 16); total number of vertebrae,

38.9 to 39.8 (Table 15). Most of the meristic characters have lower values than for all other species of

Nocomis, but they were homogeneous, and showed

little divergence and were of little value in solving

the systematic boundaries of the infraspecific populations. Specimens from certain tributaries of the

New drainage, such as Fox Creek, had higher circumferential and lateral line counts compared with

those in other New drainage localities or those of

the James and Roanoke drainages. The same characters for specimens in the Tar and Neuse Rivers had

somewhat lower values compared with those of the

James and Roanoke. We cannot attach any particular significance to these differences.

The breast is almost always fully scaled (Table

18). In a sample of 107 specimens from several drainages, the breast was about one half scaled in 4, three

quarters scaled in 69, and fully scaled or nearly so

in 34 specimens. N. leptocephalus differs from the

micropogon group in the constantly high degree of

breast squamation.

Tuberculation: The number of head tubercles in

specimens from the James, Roanoke, and upper New

drainages, segregated by sex and size groups is summarized in Table 19. A comparison of tubercle

numbers by size-groups with the species of the micropogon group is shown in Figure 4. The typical, early

tubercle development and pattern is illustrated in

Figure 3. The distribution of head tubercles in nuptial males is shown in Figures 23, 24, 26, 27. Tuberculation in N. leptocephalus is characterized by a

great reduction in numbers, the lowest for any species of Nocomis; by an enlargement of the tubercle

size, the largest for any species of Nocomis; and by

the absence of tubercles from the snout, subnasal,
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 26.—Head of male Nocomis leptocephalus, CU 10542,

153 mm, showing nuptial crest and tubercle development,

taken 23 June 1946 from a tributary of the Ararat River,

Yadkin River system: a, lateral view; b, dorsal view. (Drawn

by A. M. Awl.)

and lachrymal areas. The tubercles first appear as

large, light spots in the internasal-anterior interorbital area, at about 30 to 40 mm SL. With increase

in body length the tubercles increase in numbers

and spread to the posterior occipital line in the

adults. Some specimens of both sexes may not have

visible spots at body lengths of 35 to 45 mm SL.

The distribution and numbers of tubercles is the

primary character in the separation of the three

subspecies of N. leptocephalus. The data on tubercle numbers and distribution above (Table 19) pertains to the typical subspecies, in which the numbers

are highest and the adult pattern most extensively

developed. Analysis of the subspecific populations

of N. leptocephalus will be treated in a separate

paper. Tubercle numbers for N. I. leptocephalus are

almost always fewer than 25. N. I. bellicus has the

lowest number, 7 or fewer in the adults. The early

tubercle pattern for N. I. leptocephalus, as shown

for small juveniles in Figure 3, is actually the definitive adult pattern in N. I. bellicus.

N. leptocephalus has more tubercles on the pectoral fin rays than species of the micropogon group.

Tubercles are present on pectoral fin rays 2 to 6 (8

specimens), 2 to 7 (18), 2 to 8 (23), 2 to 9 (1), and

2 to 10 (2).

Nuptial crests: The development of the crest in

N. leptocephalus is similar to its development in the

micropogon group. In N. leptocephalus large crests

develop on smaller size specimens than in the micropogon group. It's development is undoubtedly associated with age and maturity. The crests of N.

leptocephalus become more greatly developed at a

position somewhat more forward on the head than

in the micropogon group, the swelling is often very

large just posterior to the internasal area. Crests,

moderate to large, were present on the following

size-groups (in mm): 110-119 (1 specimen), 120-129

(4), 130-139 (5), 140-149 (10), 150-159 (5), 160-169

(4)-

Pharyngeal dentition: Tooth numbers were 4-4

in 90 percent of 92 specimens sampled over the

range of all three subspecies. Counts of less than

4-4 comprised the other 10 percent of the sample,

these being 3-4 or 4-3, 3-3, and 2-4. The structure

of the arch and the pitted surface is similar to the

arch of N. micropogon.

Coloration: Two different primary color forms

exist in living nuptial males of N. leptocephalus.

N. I. leptocephalus has both forms as described by

Lachner (1952:439). Those in the Roanoke and

Neuse drainages have blue on the head laterally, an

orangish lateral stripe, and variable amounts of

shades of orange in the fins while in the Pee Dee

drainage the body laterally is bluish instead of

orange. Further observations by the authors and

others have shown that the orange-sided form is

also in the New and James drainages in the northern part of the range while the blue-sided form also

exists in the Santee drainage south of the Pee Dee

drainage. Additionally, males of the northern color

form, at the height of nuptial coloration, have the

lateral stripe abruptly changing from orange to

olive-yellow, caudally, from about the vertical from

the middle of the dorsal fin. Both forms at the peak

of color intensity have a tan to brown nuchal crest

and the dorsum above the lateral stripe is colored

brown to bluish black.

Nuptial males develop a light to moderate blue

on the lateral and ventral head, deepest on the opercle and on the cheek, faintest on the lower jaw and

branchiostegal regions. The tubercle region is dark
NUMBER 85 53

tan to brown in living, large males; the tan turns

to dark olive after several hours in formalin. Of

ten tuberculate males captured in the upper James

drainage during 10-30 May 1963-1965, the deepest

shade developed on the lateral head was moderate

blue in one specimen, a light blue in five specimens

(faint blue-gray or blue color absent on the ventral

head) and the blue was not detectable in four specimens. In one specimen the light blue color turned

noticeably darker while being examined out of

water. The color also turned darker in all specimens with blue upon being in formalin for about

fifteen minutes; those without the color developed

a light or moderate blue. A postnuptial male with

Saprolegnia over its tubercle scars was observed in

the stream to have a fairly dark blue head. The

blue head color of living and preserved N. leptocephalus is lighter than the bluish black color that

sometimes develops on the head of recently preserved nuptial male specimens of N. raneyi and N.

jnicropogon. The coloration of the head of the latter species is unlike the nuptial head color of N.

leptocephalus in life.

The other nuptial color component in the northern male N. leptocephalus; radically different from

the pink-rosy lower sides of the micropogon group,

is the horizontal stripe of faint to dark, coppery to

rusty-orange. This is developed in the scale medians

from the postopercular bar to about the dorsal fin

base level, within 3.5 scale-rows above to 1 scale-row

below the lateral line at its widest point (anteriorly)

and tapering slightly to the dorsal fin level. The

edge of this stripe is fairly sharply delimited below

by the whitish lower sides, somewhat less above

since the dorso-lateral scale medians have brassy

iridescence. The stripe then grades posteriad to the

level above the anal fin base into a bright olivish

yellow, this shade being more yellowish than the

dorso-lateral region just above it, and then grades

over the caudal peduncle to a less bright, light

yellow-olive, about the shade of the dorso-lateral

scale medians.

A deviation from the typical orange stripe was

seen in nuptial males from Craig Creek, captured

on 15 May 1963 where many fresh nests were present. In three of the five nuptial males taken, the

scale medians were pink in the anterior lateral stripe

area from the postopercular bar posteriad to the

level of the anterior base of the dorsal fin; the most

ventral row of scale medians in that area had a

suffuse yellow added. The posterior portion of the

midlateral stripe was typical olive-yellow. Many

nuptial males more recently taken or observed in

the upper James, several of which were from Craig

Creek, had an orange area or stripe anteriorly on

the side of the body.

The head coloration of all sizes except the nuptial males and some postnuptial males is very similar to that of the members of the micropogon group;

it differs in that the dorsum is slightly lighter, ranging from tan-olive to dark olive. The inner "color

ring" of the iris is wide, often occupying half of its

FIGURE 27.—Crested, tuberculate male of Nocomis leptocephalus, USNM 166392, 165 mm, from

the Dan River, Roanoke drainage, collected 8 June 1947. (Drawn by Carolyn Bartlett Cast.)
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

width, and is colored bright orange, red-orange, or

red. It is very rarely interrupted by the blackish

outer iris color. Most specimens of N. leptocephalus

can be separated in the field from the micropogon

group by this character alone.

The body color grades from dark above to white

on the belly. The color of the median of the scales,

laterally on the body, tends to be lighter in some

specimens, being more of an iridescent yellow-green

than yellow-olive and often with brassy iridescence.

The dark midlateral stripe and basicaudal spot is

often absent but prominent in some living and preserved juvenile and adult females and juvenile males.

Preserved adult males rarely have a well-developed

stripe and, when present, it is usually only on the

caudal peduncle. An iridescent yellow-olive, middorsal stripe is generally present in all but the larger

adult males; this soon turns darker than the adjacent dorsal area in formalin. A moderately dark

lateral stripe and light middorsal stripe were seen

in a highly tuberculate male of about 130 mm SL

that was building a nest in the upper James drainage. No specimens were observed in life with a light

greenish lateral band as seen in N. micropogon and

N. platyrhynchus.

The fins are colored yellow-olive, usually with

small to large areas of orange or, infrequently,

slightly reddish orange. These colors are generally

restricted to the rays but in some specimens they

extend slightly into the membranes. The membranes

are clear to clouded with white and some olive. An

accentuation of the fin colors has not been noted in

nuptial males. One nuptial male with intense body

color had pale dorsal and caudal fins, lacking the

yellow and orange, and it had only slight shades of

these colors in the other fins. The pelvic fin rays

were milky white along the branches. The fins of

N. I. leptocephalus have a heavier deposition of

melanophores along the rays and ray joints than in

the micropogon group; it is heaviest in the dorsal

fin, the rays appearing almost entirely black in

larger nuptial males.

Some slight but rather consistent differences from

the micropogon group are seen in the melanophore

pigmentation of alcoholic specimens. The anterior

margins of the scales are narrower, sometimes absent (when the scale just anterior entirely covers the

scale pocket below), while the posterior margins of

the scales are wider and do not have a sharply defined inner edge. Frequently the posterior margins

do not extend to the actual posterior edge of the

scales. Adult females have wider scale margins than

adult males.

REPRODUCTION AND GROWTH.—The nest building,

spawning, and reproductive behavior of this species

is now well known, based on recent studies by Lachner, which will appear in the sixth paper of this

series (p. 2), in which the reproductive behavior

of all species of Nocomis will be treated. The age

and growth of N. leptocephalus was treated by

Lachner (1952); the largest specimen studied was a

male in the 115-120 mm SL size class. In Table 19

we report on one male specimen in the 170-179 mm

SL size class. After having examined about 30,000

specimens from over its range we encountered one

collection of 11 tuberculate males that dwarfed all

others known. This collection (TU 29534) was taken

in Toxaway River, above Lake Toxaway, Transylvania County, North Carolina, on 14 July 1962. Ten

of the specimens ranged from 192 to 214 mm SL,

and the smallest was 172 mm SL.

MATERIALS EXAMINED.—The 124 collections from

five drainages are given followed by the institution

and number of collections housed.

James 25: CU 11, UMMZ 5, USNM 9. Roanoke 45: CU 6,

UMMZ 1, USNM 38. Tar 10: CU 9, USNM 1. Neuse 12:

CU 4, USNM 8. New 32: CU 11, UMMZ 1, USNM 17, DU 3.

DISTRIBUTION.—Nocomis I. leptocephalus is distributed on the Atlantic slope from the Potomac

drainage in Virginia southward to the Santee drainage and west of the Appalachian Divide in the

upper New drainage (partly shown in Figure 28).

The only known collections of N. leptocephalus

from the Potomac drainage were taken in 1956 and

1958 by R. D. Ross and his associates from Back

Creek and Middle River, tributaries of the upper

South Fork of Shenandoah River, Augusta County,

Virginia (all USNM collections). The single specimen known from the Rappahannock drainage was

captured in South River, a tributary of upper Rapidan River, Greene County, Virginia, in 1951 by

E. C. Raney and C. R. Robins (CU 46283). The

York drainage was first included within the range

of N. leptocephalus by Raney (1950:161). Six collections were available, all from tributaries of the Pamunkey River, one of the two major tributaries of

the York estuary.
NUMBER 85 55

eoi

.771

! • N.leptocephalu

CHOWRN

ROANOKE

39

37

801

FIGURE 28.—Distribution of Nocomis leptocephalus in the northern portion of its range, on the

Atlantic slope from the Potomac drainage to the Roanoke drainage and west of the divide

in the New drainage. Its distribution within the Blackwater River system (directly west of

Letter B) is shown in detail in Figure 29. The horizontal bars represent the drainage divide line

and the solid line the Fall Line.

N. leptocephalus occurs widely within the James,

Chowan, Roanoke, Tar, and Neuse drainages.

This species also has a wide distribution within

the upper New drainage of Virginia but, apparently, it does not occur in its headwaters or the lower

New. The farthest downstream record is one specimen taken in 1885 from Wolf Creek, Giles County,

which enters the New just upstream from the

Virginia-West Virginia state line. As known from

collections and angling, this species is common in

much of Big Stony Creek, the first major tributary

just upstream from the mouth of Wolf Creek. Fox

Creek is the uppermost tributary known to be inhabited, whose mouth is just downstream from the

Virginia-North Carolina state line, where it was

taken commonly in 8 collections from 6 well-spaced

localities.

Ecology, Association, and Frequency of

Hybridization

REGIONAL ECOLOGY.—The habitats of the four

species of chubs are basically similar although there

is some ecological variation within their geographic

ranges. Generally, the species of Nocomis prefer

clear, moderate to warm water streams of intermediate gradients. They are carnivorous fishes common

over bottoms composed of gravel, rubble, and boulders, and usually with scant or no higher aquatic

vegetation.

The primary ecological distinction between the

species groups is their preference for different size

streams. The three species of the micropogon group

generally inhabit larger streams, becoming the dominant chub in streams averaging wider than 40-60
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

feet. This preference has been noted throughout

most of the range of N. micropogon (Lachner, 1952:

434-435 and included references; Gerking, 1945:15,

48; Trautman, 1957:297) and is likewise true for

N. raneyi and N. platyrhynchus.

There may be limited upstream movement correlated with the breeding season. Miller (1964:316)

noted that sometimes mature N. micropogon did

not appear in certain breeding areas until early May

in Catatonk Creek, Susquehanna drainage, New

York. Raney (1950:161) mentioned an upstream

movement of this species to spawning areas in April

and May. We note that nuptial males, ripe females,

and juveniles of N. raneyi were seen or commonly

captured on 29-30 May 1964 in an upstream portion

of Johns Creek averaging 15-25 feet in width.

Whether the adults remain upstream after spawning in this relatively narrow section is unknown. A

number of older adult males die soon after spawning.

Less information is available concerning the distribution and abundance of river chubs in the large

river channels. They occur locally in certain rivers

where adequate sampling was made. The James

River where (atypical) N. raneyi was taken fairly

commonly is 60 to 150 yards in width. N. raneyi was

also captured in the larger sections of the Tar and

Neuse rivers. Several adult N. platyrhynchus were

caught by sport fishermen at localities in the main

channel of the New River in Virginia where large

nests, obviously of this species, were seen. Large

nests have also been observed in the deeper waters

of the Greenbrier River and the Knapp Creek tributary, West Virginia, over the past several years.

Several records are available of N. micropogon from

the lower Potomac River down to Great Falls just

above tide level. Lachner (1952:434) mentioned that

larger adults of this species are taken from deep

river holes by fishermen; these and more recent observations were made in western Pennsylvania. H. M.

Smith (1907:104) stated that in North Carolina N.

kentuckiensis (Rafinesque) shows a preference for

larger streams. His records are probably based on

specimens of N. raneyi in the east and N. micropogon from the Tennessee drainage.

N. leptocephalus generally is most abundant in

small streams, although there is usually much overlap with species of the micropogon group in intermediate stretches. This species is common to abundant in streams averaging from 10 to 50 feet in

width, but it rarely occurs in extreme headwaters.

The juvenile and adult chubs commonly occupy

both the riffle and pool habitats. These life stages

were collected in about equal numbers in moderate

to rapid riffles and short to moderate length pools

of the upper James, Roanoke, and New drainages.

They seldom occur in the long stretches of slack

water. N. leptocephalus is frequently common in

rapid, small mountain streams that are stocked with

trout.

A stream bottom composition partly of gravel and

rubble is an important factor in the maintenance

of a sizeable population of Nocomis, especially since

these materials are essential for nest building. Other

bottom types in various combinations are occupied.

The largest populations of the four species in Virginia and of N. micropogon in Ohio (Trautman,

1957:297), Indiana (Gerking, 1945:15, 48), and elsewhere are most commonly found where currentswept gravel, rubble, and boulders predominate.

The decline or disappearance of populations of the

widely distributed N. micropogon with increased

turbidity and siltation is generally observed. N.

raneyi populations apparently have also declined in

more silted sections of the Roanoke drainage.

The well-established populations of N. raneyi,

N. micropogon, and N. leptocephalus in typical

Piedmont streams indicate their adaptiveness to this

type of habitat. Lower gradients, sandy bottoms and

somewhat greater turbidity generally characterize

streams of this province but gravel, rubble, and

sometimes boulder riffles occur. N. raneyi and N.

micropogon do not tolerate sandy bottom conditions

as extreme as those where N. leptocephalus is sometimes common; this may account for the apparent

absence of N. raneyi from several Piedmont tributaries of the Roanoke and its rarity in others such

as the middle Dan River.

The rarity of chubs in streams of the Coastal

Plain province is probably related to the general

absence of riffles, exposed gravel and rubble. The

courses of the eastern rivers through the Fall Zone,

the transitional belt of varying width (whose eastern

limit is the Fall Line) between the Piedmont and

Coastal Plain, are generally marked by a series of

rapids and cascades which provide some habitat.

From the James northward the Coastal Plain rivers

and most of the generally small tributaries are tidal

or brackish up to the Fall Line. However, the rivers
NUMBER 85 57

south of the James do not drop to sea level so quickly, and their channels are on crystalline rock with

rapids occurring for about 20 miles after entering

the Coastal Plain (Fenneman, 1938:13, 129). The

presence of riffles in some upper Coastal Plain tributaries may be accredited to drainage of higher terraces during which time some have cut deposits to

the crystalline rocks of the Piedmont (Clark, et al.,

1912:26). Headward extension with exposure of

gravel and rock by erosion, beginning with the postPleistocene elevation of the Coastal Plain (Clark

and Miller, 1912:200), seems to be still in effect.

N. rarteyi and N. micropogon occur in the Fall

Zone and sparingly in the upper Coastal Plain province. Jordan (1889a: 109-110) noted that a rapid and

rocky portion of Swift Creek, James drainage, just

below the Fall Zone has a fauna "essentially that of

the upland streams." The southeastern limit of N.

micropogon (excluding the upper Savannah drainage population) is the James drainage, and for reasons stated above, its distribution terminates at or

near the Fall Zone. N. raneyi has available a rather

wide lowland region in southeastern Virginia and

northeastern North Carolina but few streams are

probably suitable. It is not known from the Coastal

Plain portion of the Chowan and Roanoke drainages although one collection from the Chowan is in

or near the Fall Zone. Of the 24 collections of N.

raneyi from the Tar and Neuse drainages seven

were taken from the Fall Zone and nine below. Of

the latter, some were from moderate to rather large

streams with riffles, bordered at least in part by

swampland, and with lightly brown-colored water.

One stream, Nahunta Swamp Creek, Neuse drainage, averages only 10 feet in width. No species of

Nocomis is known to inhabit the predominantly

slow blackwater type of stream or inner portions of

swamps.

N. leptocephalns was taken in the Fall Zone from

the York and the Chowan drainage southward to

the Neuse and in most cases with N. raneyi. The

former species apparently avoids Coastal Plain

streams in the central Atlantic region, for we know

of only one collection from this area in a Roanoke

tributary.

N. micropogon, the only member of the species

group inhabiting central United States, shows a distinct preference for recently glaciated regions within

the northern and middle, lowland portions of its

range in the Ohio River drainage basin. It is absent

from the region covered by the Illinoian glacier

south of the Wisconsin glacier limit in Indiana

(largely the lower Wabash drainage) except for a

borderline record (Gerking, 1945:15, 48-49, map

25). The habitat here is generally unsuitable, prevailing conditions being similar to or more limiting

than those on the Atlantic Coastal Plain. Jordan

(1889a: 160) stated that the lower Wabash tributaries

"are mostly sluggish and yellow with clay and mud".

Gerking (1945:17) indicated the same for the lower

Wabash flood plain. Trautman (1957:11) noted that

most unglaciated and hence undisturbed and almost

base-level streams of southeastern Ohio have no

well-developed riffles while the glacial streams in

this region have relatively high gradients. Some of

the latter streams are inhabited by the river chub

(Trautman, 1957:296-297, map 61). Its absence from

other Ohio River tributaries in Kentucky may be

related to similar sluggish conditions.

Higher aquatic vegetation is not common in most

areas inhabited by the micropogon group and N.

leptocephalus. Generally, habitats suitable for good

plant growth are borderline situations for these species. An exception is the well-established growth of

water willow, Justicia, in portions of the upper

James and New drainages supporting a thriving

chub population, and similarly reported in Ohio by

Trautman (1957:297). This stout emergent herb

often grows in dense stands along the shore and in

shallow pools and riffles. Shockley (1949:259-261)

found stream areas supporting a growth of Justicia

to be more productive than those where it was absent. Submerged portions of water willow usually

show retention of much organic detritus and an

abundance of aquatic insects. Chubs also are found

in streams with good growths of the algae Cladophora and the vascular riverweed, Podostemum.

Lachner (1950:230; 1952:434) found a large population of N. micropogon in streams with an abundance of algae, chiefly Cladophora, and vascular

plants (several species of Potamogeton), in Lake

Ontario tributaries of New York. Smith and Bean

(1899:183) reported this species as common over

grassy bottoms of the Potomac River.

Lachner (1950) found the food of N. micropogon

in western New York to be largely aquatic insect

larvae with lesser percentages of Crustacea and Mollusca. In many instances, algae, chiefly Cladophora,
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and vascular plants were taken in large quantities.

Much of the plant material is probably ingested

incidentally with animal food since it appears to be

undigested. Fishes are rarely included in the diet of

chubs. Differences in food ingested by N. micropogon

were found among young, juveniles, and adults and,

within the juvenile stage, among certain seasons

(Lachner, 1950). Comparative food studies of the

species of chubs occurring sympatrically might reveal differences, especially since N. leptocephalus

has a long, whorled intestine, differing from all

other Nocomis. Flemer and Woolcott (1966:83-84)

analyzed food intake of 249 specimens of N. leptocephalus from a typical, lower piedmont tributary

of the James River just above the Fall Line. They

found a great preference for plant food by this species, forming a high percent of total items eaten.

They concluded that the plant material was specifically selected by N. leptocephalus.

ASSOCIATION AND FREQUENCY OF HYBRIDIZATION.—

The following discussions refer to the extent that

the species of chubs occur syntopically (occupy the

same macrohabitat) or together (Rivas, 1964). All

species occurring in the same drainage have been

collected together. However, differences among the

species of the micropogon group in their frequency

of occurrence with N. leptocephalus are revealed

and the apparent noncompatibility between N.

raneyi and N. micropogon is discussed. The data

support our preceding statements concerning stream

size preferences. The relative frequencies of hybridization between certain of the species are reported

below. Factors involved in hybrid formation will be

discussed in papers 5 and 6 on the study of Nocomis

hybrids and reproduction.

The data summarized in Table 26 are derived

largely from intensively sampled tributary systems

of the James, Roanoke, and New drainages. In order

to eliminate bias from small collections and those

taken by selective angling, the data indicating syntopy or allotopy (Rivas, 1964) are from collections

in which a minimum of ten specimens were taken.

The sampling intensity for the four species in intermediate size streams was about the same.

In the James drainage, N. leptocephalus was taken

with N. raneyi in 89 percent of the collections

(Table 26) from the Craig Creek system (Figure 20).

In approximately the center of this system, in five

collections from three localities in lower Craig Creek

just below Newcastle, N. raneyi was always found

to be the more abundant species, appreciably outnumbering N. leptocephalus (429 to 239 specimens).

In upper Craig Creek within a radius of about 3.5

miles above Newcastle, N. raneyi was taken in three

collections from two localities, but in less numbers

than N. leptocephalus (26 to 52 specimens). Two

juvenile N. raneyi were taken about 5.5 miles above

Newcastle, but none have been found above this

area. N. raneyi was found to extend well up Johns

Creek to within 0.75 mile of Maggie, the latter being

where N. micropogon and hybrids of N. leptocephalus X N. micropogon were taken. In three large

collections, two in spring and one in summer, from

within 3 miles of Maggie, 58 N. raneyi and 92 N.

leptocephalus were taken, the latter species being

more numerous in each. The stream in this area

averages about 15 to 25 feet wide. N. leptocephalus

apparently becomes progressively less numerous below the Newcastle area. A sparse spawning population occurs down into the middle section of lower

Craig Creek. Only N. raneyi was taken in four collections from the lower 3 miles of Craig Creek.

The Blackwater River of the Roanoke drainage

in Franklin County, Virginia, is a well-surveyed

stream in which similar distributional relationships

were found between N. raneyi ahd N. leptocephalus

as observed in Craig Creek (Figure 29). The Blackwater is a typical Piedmont river; its lower portion

is about equal in size to lower Craig Creek. Of the

22 collections (15 localities) in which N. raneyi was

taken, all but two are in the main channel from

lower North Fork near its mouth in Roanoke River.

N. leptocephalus was taken at all but one location.

A progressive upstream decrease in the relative

numbers of N. raneyi is indicated when the main

channel is divided into three sections of approximately equal lengths and by calculating the percentage composition of each species per section. N. raneyi

comprised 93 percent of the 246 chubs taken in the

downstream section; in the middle section, 76 percent of 296 specimens; and upstream, 21 percent of

505 specimens. The progressive change in relative

numbers is also indicated in almost all individual

collections. In the two collections from the lower

portion of the most downstream, and major, tributary of the Blackwater, N. raneyi comprised only

three of the 49 chubs taken. N. leptocephalus is
NUMBER 85 59

^-""^o ) BLACKW&TER

y FRANKUN CQ

RIVER, ROANOKE DRAINAGE

O N . RANEYI

O N . LEPTOCEPHALUS

• BOTH SPECIES

2pJ ( ^ nuMUNox^

/'

FIGURE 29.—Distributional relationships to stream size of Nocomis raneyi and N. leptocephalus

within the Blackwater River system of the Roanoke drainage. The most important ecological

difference between these species over the ranges was the preference by N. raneyi for the larger

and deeper streams and a partial preference for the smaller tributaries by N. leptocephalus;

the latter habitat was almost entirely avoided by N. raneyi. The map is an enlarged portion

of the Roanoke drainage directly west of the letter B in Figures 19 and 28.

common to abundant in the remainder of the tributary system in which N. raneyi is absent or rare.

The distributional relationships in the intensively

collected lower South Fork and upper main channel

of the Roanoke River, Montgomery and Roanoke

counties (Figure 20) are somewhat different. Although within the Appalachian province, the bottom of these streams is generally much more silted

than that of Craig Creek to the north, even though

riffles are more frequent in the Roanoke River than

in Craig Creek. The Roanoke River at the city of

Salem, Virginia, averages about the same in width

as lowermost Craig Creek. N. leptocephalus was captured in all nine lower South Fork collections (five

localities) in which N. raneyi was taken. The uppermost locality is near the town of Alleghany Springs,

where the stream averages about 30 to 40 feet in

width. AT. raneyi comprised only 3 percent of 510

chubs captured. Only N. leptocephalus was taken

in numerous additional collections from lower South

Fork and farther upstream. Although N. raneyi

showed an expected increase in relative numbers in

the downstream direction, it was taken from the

upper main channel only at seven localities in ten

collections, in which it accounted for 29 percent of

the 228 total chub catch. The large majority of the

recent collections (through 1970) from the main

Roanoke from Salem upstream did not include N.

raneyi, although AT. leptocephalus was, at least, common. The only collection (USNM 197684) in which

AT. raneyi was common was from a bottom area with

less silt than at all other main Roanoke localities.

The upper Roanoke record from below Salem was

made in 1888. The Roanoke River from the city of

Roanoke downstream is now badly polluted.

The known case of hybridization between these

species occurred in the upper Roanoke. Ten specimens of AT. leptocephalus, one of N. raneyi, and two

hybrids were taken.

Elsewhere in the Roanoke and in the Chowan,

Tar, and Neuse drainages about the same distributional relations prevail between AT. raneyi and N.
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

leptocephalus as that in the James and Roanoke

drainages.

In contrast to N. raneyi, N. micropogon was

found less frequently with N. leptocephalus in the

James drainage (61% of the comparable collections,

Table 26). Included in this percentage are six collections with hybrids of N. leptocephalus X N. micropogon taken with nonhybrid specimens of only

one or the other species. The occurrence of one or

more hybrid specimens was relatively high, hybrids

being taken in 60 percent of 25 collections. Only

three of the 15 hybrid collections contained less

than ten specimens; thus it appears that this hybrid

combination may occur fairly frequently (87 specimens examined). Hybridization occurred in almost

all tributaries in which the two species were taken

together and, within each tributary, in a frequency

ratio about equal to the relative number of collections from that tributary.

Ecological conditions in the upper New River

basin of the Blue Ridge and Appalachian provinces

are varied but generally montane. Individual tributary systems in Virginia are described by Ross and

Perkins (1959), Shoup (1948), and Burton and Odum

(1945). In the Virginia portion of the upper New

the relations between N. platyrhynchus and N. leptocephalus are rather similar to those between the

latter species and N. micropogon in the James drainage. N. leptocephalus was absent from the West

Virginia portion of the New River drainage; collections of N. platyrhynchus from this area are not included in the following comparison. The two species

were syntopic in 63 percent of 16 comparable collections. Hybrids were taken in three (20% of 15

samples), each from a different tributary system.

The distributional relationships between N. raneyi and N. micropogon in the James drainage have

ecological and zoogeographical significance. N. micropogan has been taken about as commonly and in

equal numbers throughout the James, exclusive of

the Craig Creek system, as N. raneyi has been taken

in the Craig system (excluding the stream section a

short distance below Newcastle that is eutrophically

enriched). N. micropogon, however, is rare within

the Craig system, being taken in only four or five

collections and in small numbers (see p. 49). Two

of the collections are from the headwaters, a habitat

marginal for the micropogon group because of the

small stream size. N. raneyi is generally distributed

downstream and one of the most dominant members

of the fauna. As might be expected to occur between

closely related ecological homologues, this pattern

of distribution and relative abundance suggests that

N. micropogon is being largely displaced from the

Craig Creek system (as well as the lower Catawba)

by competition with the apparently better adapted

N. raneyi. N. micropogon may be the more successful of the two species in tributaries upriver from

Craig Creek and downriver in the Maury River system. Perhaps here the period of their contact, if

any, has been of such short duration that changes

in distribution have not occurred. One hybrid N.

micropogon X N. raneyi was taken in lower Johns

Creek and one was taken in lower Catawba Creek

(see p. 49), where observations in the last several

years have revealed only high breeding populations

of N. leptocephalus and constant breeding numbers

of N. raneyi.

SUMMARY.—On the basis of similar ecological

preferences, and apparent competition in the Craig

Creek system, the three species of the micropogon

group are regarded as ecological homologues. N.

raneyi almost always coexists with N. leptocephalus

in moderate-size streams. There is a definite tendency for N. leptocephalus to occur less frequently

with N. micropogon and N. platyrhynchus. The

greater frequency of occurrence of N. raneyi with

N. leptocephalus suggests that competition is somewhat more intense between the latter species and

N. micropogon and N. platyrhynchus.

There are differences among the species of the

micropogon group in their frequency of hybridization with JV. leptocephalus, this being inversely proportional to the frequency of syntopic occurrence.

Hybridization between N. raneyi and N. leptocephalus is rare, only two specimens are known from a

habitat more disturbed than that of the other

streams which were intensively sampled. N. micropogon and N. leptocephalus hybridize comparatively commonly (87 specimens known). Although the

number of collections in which N. platyrhynchus

and N. leptocephalus were taken together, and the

number of specimens involved, is less than those of

the other two combinations of chubs, the frequency

of hybridization is appreciably (5 specimens are

known to us) less than that of the N. leptocephalus

X N. micropogon combination.
NUMBER 85 61

Dispersal: Biological and Geological Evidence

The ichthyofauna of central eastern United States

presents many stimulating zoogeographical problems, such as the dispersal of the species of the

micropogon group, the leptocephalus group and of

several of their associates. Possible routes of entry

into various drainages have been recognized. These

are often indicated by biological evidence of present distributional patterns and relationships and

past geological events. In some cases, however, routes

have been conjectured only from biological evidence

since complementary geological information is meager or nonexistent. While this procedure may not

be justifiable since distribution patterns are known

to change, the fact remains that populations have

crossed drainage divides in some manner. Therefore,

attempts are made herein to offer explanations of

how dispersal may have occurred. Past and present

ecological conditions are considered.

Three kinds of geological events have operated in

the dispersal of chubs. These are stream captures,

eustatic changes of the Atlantic Coastal Plain, and

Pleistocene drainage modifications. An obvious

fourth means of dispersal is movement from one

drainage system to another through past and existing interconnecting main rivers.

STREAM CAPTURE.—The geological literature on

the occurrence of and evidence for captures in western Virginia and adjacent portions of West Virginia

is extensive. It appears that numerous captures have

occurred which made it possible for stocks of fishes

to spread among several drainages. Major papers

treating this subject are by Wright (1931, 1934,

1936), Thompson (1939), and Dietrich (1959), and

numerous additional references are in their bibliographies. It is interesting to note that Dietrich

(1959:30), a geologist, invoked biological evidence

from fishes in support of a geological hypothesis. A

century ago, Cope (1869) concluded that the faunal

similarities among opposing drainages in western

Virginia may have resulted in part from headwater

transfers. Recently Ross and Carico (1963:7-12)

documented two stream captures to support hypotheses on the dispersal of certain fishes between

the New and Tennessee drainages. Other authors

(Robins and Raney, 1956:31; C. R. Gilbert, 1961:

456; 1964:106) mention possible use of stream cap

tures by fishes in western Virginia. Other instances

of isolation, subsequent differentiation, and present

distributional patterns of fishes in this region can be

accounted for primarily by these events.

When invoking stream capture as an agent of

dispersal, the biology of the fishes, ecology of the

region, and nature of stream capture are factors

that should be considered. Some factors are species

abundance, interspecific competition, habitual movement and migration of species, size and flow of

streams, and other ecological expedients or barriers.

Some important geological factors are the magnitude of the theater of stream capture, the duration

of the water connection between opposing drainages

and the numbers and sequence of captures that occurred in a region. The absence of certain species

today from a drainage from which its associated

species have successfully entered is perplexing and

may be explained by subsequent change in stream

conditions, competition, and extirpation.

Another problem in correlating dispersal widi

stream capture is the uncertainty or lack of information concerning the dates of occurrence. Thompson's statement (1939:1353) that "captures that can

now be definitely recognized belong to comparatively recent geologic times" provides a positive

argument for adopting the concept of capture. Many

captures described can probably be assigned to postHarrisburg time, a period of stream rejuvenation

which began in the late Tertiary (Wright, 1934:38).

In addition, since Wright (1931:246) pointed out

that evidence of capture soon disappears with active

dissection of the capture area, we may safely assume

that many more captures occurred dian are presently detectable [see also Ross (1969) and Jenkins,

Lachner, and Schwartz (ms)].

EUSTATIC CHANGES OF THE ATLANTIC COASTAL

PLAIN.—Dispersal can be equated with eustatic

changes of the Atlantic Coastal Plain during or

somewhat after Pleistocene time. During such times

the several drainages now entering the western shore

of Chesapeake Bay, from south of the present Susquehanna River mouth to the James drainage, were

sometimes tributary to the extended freshwater

Susquehanna, hereafter called the Greater Susquehanna River (Shattuck, 1906:134, pi. 31). This resulted from a general increase in elevation of the

central Atlantic Coastal Plain and/or a lowering of

sea level (Flint, 1957:270). Lougee (1953:264-265)

stated that sea level was at or near its lowest level
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

during the last glacial climax. It is also possible that

a drainage similar to the Greater Susquehanna was

in existence during earlier periods of Pleistocene

glaciation (Flint, 1957:270).

At the same time it is likely that similar events

occurred in North Carolina, in present Albemarle

Sound where the Chowan drainage would have been

a tributary of the Greater Roanoke River, and in

Pamlico Sound where the Tar and Neuse drainages

would have been conjoined to form the Greater

Pamlico River. Some authors consider the Chowan

to be a present Roanoke tributary, but this is not

7 7

•//////// .

GROUP OR SPECIES BASIC STOCK

N . BIGUTTATU S

OZAR K CHU B

N . EFfUSU S

MICROPOGON GROUP

N. MICROPOGON

IEPTOCEPHAIUS GROUP

FIGURE 30.—The present distribution of certain species and species groups of Nocomis superimposed over stages in drainage development. The probable principal dispersal routes of basic

Nocomis stocks are indicated by wide and narrow arrows. Stages of formation of the Mississippi

and Great Lakes drainage basins are only approximations, probably present during the late

pre-Pleistocene and/or Pleistocene periods. The Laurentian River is shown to occupy the present

Great Lakes basin. The distributions of the species or species groups, in shaded areas or outlined

by dotted lines, represent total ranges; some areas within these ranges may be uninhabited.

Drainage patterns adapted from Leverett (1902), Tight (1903), Fenneman (1938), and Janssen (1953).
NUMBER 85 63

likely since the large lower channels of these drainages are separated by tidal flats (Hubbs and Raney,

1944:5), thus largely or entirely preventing dispersal

of most freshwater species between them. Opportunity for dispersal of species considered herein

through extended channels is thought not to have

occurred earlier than the Pleistocene since there was

a Pliocene submergence of the Coastal Plain (Clark

and Miller, 1912:215).

PLEISTOCENE DRAINAGE MODIFICATIONS NORTHWARD.—Several means of late Pleistocene dispersal

in the Great Lakes and adjacent regions are discussed under the following species accounts.

The vast preglacial Teays River system (Figure

30) served as a reservoir for a large portion of the

North American ichthyofauna and provided an

early means of dispersal. Its upper and middle portions are of much importance in this study. The

upper Teays, approximately, is regarded as the NewKanawha River system after the occlusion of the

middle Teays and formation of the present Ohio

River during Pleistocene glaciation; however, some

drainage relationships of the Teays with the earlier

Ohio, Tennessee and Cumberland drainages are uncertain. Detailed discussion of the history of the

Teays system is presented by Leverett (1902), Tight

(1903), and others and, of a more general nature,

Janssen (1953) and Flint (1957:170-171).

Nocomis platyrhynchus: The basic stock of the

micropogon group probably occupied the middle

and upper Teays River system (Figure 30), perhaps

during late Pliocene and certainly part of Pleistocene times.

The bigmouth chub may represent the form

closest to the basic stock of this group and probably evolved in the New River drainage (upper

Teays) where it is presently confined. Kanawha Falls,

approximately 24 feet in height, and Sandstone Falls

just upstream, apparently served as an effective

barrier for isolation of the segment of its ancestral

stock from that below. The period of inception of

Kanawha Falls is not dated, however; it may have

been during pre-Pleistocene times. If the ancestor

of N. platyrhynchus was below the Falls and was

prevented from directly entering the New, it may

have made its entry through stream capture between

Elk River of the Kanawha drainage and Gauley

River of the New, an event for which there is geological evidence (Campbell, 1896:669-670).

The New drainage has a unique fauna, partly

characterized by several endemic forms and further

marked by the absence of a large fraction of the

fauna of the Ohio basin. Ross (1959) listed and discussed the members of this fauna. Forms, in addition to N. platyrhynchus that most likely evolved

in the New are the cyprinids Notropis scabriceps

and Phenacobius teretulus, and the percids Etheostoma osburni and Etheostoma kanawhae. Most of

these species occur widely within the New, from its

headwaters to Gauley River, but none are known

below Kanawha Falls. It therefore seems that Kanawha Falls has been the major impediment to their

establishment in the Kanawha drainage. The several tumultuous rapids, cascades, and low falls in

the New River gorge of West Virginia (Campbell

and Mendenhall, 1896; Reger, 1926:3, 90) also may

limit downstream dispersal.

Despite its endemics, the New drainage has a depauperate fauna in relation to the size of the basin.

There is evidence to suspect that the earlier New

drainage fauna was similar to that of the Ohio basin

and that its depauperate nature probably resulted

from extirpation. The Kanawha drainage fauna is

essentially that of the middle Ohio basin. Particular

examples of probable extirpated forms are species

of redhorse suckers (genus Moxostoma, subgenus

Moxostoma). The occurrence today of some species

of this group in Atlantic slope drainages from the

James southward may be best explained as their

having crossed the Appalachian divide from the New

drainage. Ross and Perkins (1959) discuss factors

that possibly combined to limit this fauna. Some of

the ecological factors mentioned (ibid.: 10) were "the

general absence of aquatic plant life, the narrow

poorly developed flood plains, the hard bottom,

steep gradient and high velocity which by abrasion

kept the stream bottom well scoured." Speciation in

the New also may have been in response to some

of these conditions.

N. micropogon: Certain stages in the dispersal of

N. micropogon can be readily postulated whereas

others are not clear. The present range of AT. micropogon south of the Great Lakes drainage may reflect much of its early distribution. A population

of the ancestral stock of the micropogon group remaining below Kanawha Falls in the Teays system

probably speciated to N. micropogon. Subsequently

this species may have occupied the middle Teays
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

drainage of West Virginia, Kentucky, Ohio, and

Indiana including the future upper Wabash drainage (Figure 30). Alternatives available for its ingress

to the Tennessee and Cumberland drainages are

discussed on page 50. N. micropogon probably never

occurred west of the Mississippi, even if it once did

reach the extreme lower Ohio basin. This would

attend the possibility that the lower Teays (early

Mississippi) and Ohio were separate systems (Fenneman, 1938:89-90). On the other hand, these large

rivers conjoined in the extensive lowland region

would still have been more of a barrier than an

agent of dispersal.

More than one region may have served as a refugium from which the river chub spread northward

during inter- and postglacial times, to repopulate

northern Ohio River tributaries and enter the Great

Lakes drainage. Evidence does not favor the lower

Wabash drainage as having been a refugium for a

Teays stock. This drainage was almost entirely overwhelmed by the Illinoian glacier (Gerking, 1945).

During glaciation the nonglaciated lower Wabash,

although certainly of a character different from that

today, probably was adversely affected. Seasonal and

diurnal variations of proglacial discharge, sometimes

perhaps catastrophic in proportion, combined with

sediment and ice loads (Dyson, 1962:77-80; Flint,

1957:174-175) would have greatly restricted fish

inhabitants.

Ancestors of populations now occupying drainages of unglaciated Kentucky and West Virginia are

likely to have moved northward after glacial recession into northern Ohio River tributaries.

Perhaps the most important center of dispersal

for river chub populations now occupying the Ohio,

Great Lakes, and Atlantic slope drainages was a

southern portion of the Old Allegheny River system, probably the Old Lower Allegheny. The three

separate preglacial component drainages of the Old

Allegheny system flowed northward to the present

Lake Erie basin (Leverett, 1902:129-138). N. micropogon presumably entered the Old Allegheny

through stream capture with the Teays. Following

interconnection of the Allegheny components with

the Ohio River due to glacial advance, the river

chub may have spread downriver into Ohio drainage tributaries when uncovered from the ice sheets.

Deglaciation of the northern portion of the Allegheny basin would have permitted influx to glacial

Lake Maumee (Lake Erie basin), through Conneaut

Creek, a present Lake Erie tributary in Pennsylvania whose upper portion was involved in a drainage reversal with the Allegheny in an area of the

Old Middle Allegheny (Leverett, 1902:214). Its presence in the lower Allegheny during glacial times

would enhance the probability of its reaching the

vicinity of Conneaut Creek before its diversion.

Dispersal to the upper Allegheny probably postdated the Cuba Outlet connection between the

Allegheny and Genesee drainages since N. micropogon is not known from the latter. Ross (1958a:

17-18) discusses a complex distribution problem in

this region involving subspecies of Campostoma

anomalum.

Once N. micropogon entered the developing Great

Lakes basin in the Lake Maumee area, it could have

entered future Lake Erie tributaries in Ohio and

Ontario and the Maumee drainage of northeastern

Indiana. Passage to the Wabash drainage from the

Maumee through the Fort Wayne Outlet would follow if this outlet remained open sufficiently long

after the Conneaut Creek diversion and/or if ample

water connection through the low divide in the Fort

Wayne region existed during more recent periods

of high water (Gerking, 1945:8).

Dispersal northward could have ensued into the

developing Lake Huron basin including its tributaries in Ontario south of Georgian Bay and those

of the eastern shore of the Lower Peninsula of

Michigan. Some movement to the western shore of

the Lower Peninsula may have been around its

northern end, but this is unlikely since the river

chub apparently did not reach (or has not persisted

in) streams of the Upper Peninsula. The eastern

shore of Lake Michigan was probably populated

after movement across the Lower Peninsula through

a later stage of the Grand River connection. Since

N. micropogon has not appeared in the Illinois

drainage, it probably did not reach the region just

north of the upper Illinois until after the Chicago

outlet was closed.

Dispersal northeastward from Conneaut Creek

could have followed the development of Lake Erie

and thence entry of streams of the present western

Lake Ontario basin, probably during the Lake

Lundy stage. Entrance to the Lake Ontario basin

at a later stage would apparently necessitate passage
NUMBER 85 65

over and establishment below newly created Niagara Falls (Fenneman, 1938:495, 499).

Natural occupation of Lake Ontario streams may

have progressed slowly. A notable (52 mile) break

occurs in the river chub's distribution in the southern Lake Ontario drainage, from just west of the

mouth of the Genesee River to 16 miles west of the

mouth of Oswego River. Two records break the

distributional gap, both from the same area and

constitute the only occurrence in a tributary of the

eastern lakeshore (excepting those from the Finger

Lakes drainage). Dispersal along the lakeshore may

involve a relatively long period of time for the river

chub, especially since for long distances only small

streams enter the lake and unsuitable habitats are

present (these conditions not permitting establishment of large populations for continuing outward

dispersal). Since the New York lakeshore tributaries

have been, comparatively, thoroughly collected

(Greeley, 1940:42), the easternmost records may indicate local extirpation or unsuitable habitat in the

gap area, bait-minnow introduction, or dispersal by

natural means. All four alternatives are feasible.

Some of the streams do not provide the larger water

type of habitat for the river chub; however, this

species is common in a relatively short, western Lake

Ontario streams and in Salmon Creek, its easternmost stream before the gap. The lower Genesee

drainage, whose mouth is just east of Salmon Creek,

would provide a large area of apparently suitable

habitat. The area is heavily fished by sportsmen.

The hardy chubs, frequently used as bait, could

have been discharged from a bait pail. Natural

means of dispersal would entail the river chub moving northward from the Susquehanna drainage, via

the Horseheads Outlet during the Lake Newberry

stage (Fairchild, 1934:1097, fig. 8, 1099), through the

Finger Lakes drainage to the lakeshore. A problem

with the latter hypothesis is that N. micropogon is

relatively rare, and the recent records are widely

scattered in the Finger Lakes basin, while it is common throughout the adjacent upper Susquehanna.

Greeley (1928:84, 97) stated that the Finger Lakes

drainage was extensively collected, and reported

only a single collection of the river chub from

Catharine Creek, the Seneca Lake inlet. He attributed its presence there to a canal connection with

the Susquehanna drainage. Although this is possible, it is now known to have a wider, although localized, distribution in the Finger Lakes drainage whidi

may have resulted from the same factors considered

for the Lake Ontario shore area.

The river chub probably occupied at least some

of the Atlantic slope drainages during glacial times

and it had a history somewhat different from that

which occurred northward and westward. Evidence

for a relatively long eastern occurrence is that its

spread throughout all the drainages of its central,

eastern range probably required considerable time,

and its movement throughout most of the James

should predate the entry of N. raneyi into this

drainage. Populations south of the Susquehanna

may be derived from stocks older than the northern

forms. C. R. Gilbert (1964:109) has given biological

support in suggesting that Notropis cornutus entered this area at an early time.

N. micropogon possibly entered the Susquehanna

by stream capture from the Allegheny drainage.

This view conflicts with Bailey's statement (1945:

125) that "some Great Lakes forms found in the

Susquehanna system are absent from the upper Ohio

basin; thus, stream capture from that [latter] source

fails to supply a tenable explanation" for dispersal

to the Susquehanna.

N. micropogon probably first gained the central

Atlantic slope by entering the Potomac drainage

through stream captures from either or both Youghiogheny and Cheat River tributaries, Monongahela drainage (once part of Old Allegheny system).

Biological and/or geological evidence for these captures is cited by Ross (1958b:5-6), Gibbs (1957:205),

and Schwartz (1965). Dispersal to the James and

Rappahannock was possible during a stage in the

development of the Shenandoah River of the Potomac, during the period the Shenandoah attained

its drainage by a series of piracies (Stose, 1922:7;

Watson and Cline, 1913).

The Greater Susquehanna River may have been

an ample means of faunal exchange among all

Chesapeake Bay tributary drainages, thus accounting^ for the presence of the river chub in the York

drainage, the minor Chesapeake drainages, and the

Appomatox River which now enters the James

River estuary. Certain factors, however, militate

against assumption that the Greater Susquehanna

was the major means of dispersal among its tributaries. This obviously was a large river, certainly

so during late Pleistocene periods of precipitation,
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in which smaller stream and some large stream

fishes would not have found suitable conditions.

Lower Potomac species such as Ericymba buccata,

Percopsis omiscomaycus, and Percina caprodes semifasciata might be expected to have spread more

widely through the Greater Susquehanna but are

not known south of the Potomac on the Atlantic

slope. The presence of Notropis s. spilopterus in

this region, only in the Susquehanna and Potomac

including their lower portions, is good reason to

believe it entered the latter by headland stream capture (Gibbs, 1957:194-205, fig. 3) and not through

the Greater Susquehanna. Other factors may confuse

this issue. Some species may have entered the upper

portions of larger Chesapeake drainages at a relatively recent period and sufficient time for spread

through the Greater Susquehanna may not have

been available.

Our interpretation of the relationships of the

species of river chubs and their populations in the

lower New, upper Monongahela, and Potomac

drainages is given further credence by consulting

the stream history of the area. We have discussed

certain morphological similarities that suggest an

exchange of river chub forms occurred between the

New and Monongahela. A likely avenue of exchange

would be a capture that occurred between the upper

Greenbrier River of the New and upper Cheat River

of the Monongahela. This is supported from geological evidence by Fridley (1933) and Wright (1934:

55). Additional biological evidence for a capture in

this area is the distribution of Percina oxyrhyncha.

This species is widespread in the New-Kanawha

system but the only known population in the Ohio

system above the Kanawha River mouth occurs in

the upper Monongahela (Hubbs and Raney, 1939:

1-3). (Additional possible agencies of ingress of N.

micropogon into the New drainage are the piracies

of Greenbrier River tributaries by three tributaries

of Back Creek of the upper James drainage (Thompson, 1939:1352-1353) and the two captures documented by Ross and Carico (1963:7-12) involving

Holston River tributaries of the upper Tennessee

drainage, and tributaries of the upper New drainage). We have also indicated a relationship between

the Potomac and Monongahela populations of N.

micropogon. A stock of the Monongahela form of

N. micropogon could possibly have entered the Potomac drainage through the capture discussed by

Schwartz (1965) for Etheostoma blennioides, and

subsequently spread throughout the latter drainage.

The entrance of N. micropogon into, and spread

within, the southwestern Ohio basin includes several

complex facets. (See Figure 7 in Lachner and Jenkins, 1967, for the distribution of N. micropogon

and other Nocomis in this region.) The middle

Teays is shown (Figure 30) as it flowed in pre-Pleistocene times (through Ohio, Indiana, and Illinois);

this portion may have reformed during Pleistocene

interglacial periods (Janssen, 1953). The latter possibility is discussed below since N. micropogon may

have evolved during the Pleistocene. It is not known

whether the Kentucky and Licking drainages emptied directly into the middle Teays or into the early

Ohio River.

We presume that N. micropogon evolved from a

stock of the bigmouth chub, N. platyrhynchus of

the New River, the latter believed to be the most

primitive form in the micropogon group. The early

range of N. micropogon within the region of the

present southwestern Ohio basin possibly included

only the Big Sandy drainage, shown to be a Teays

tributary (Leverett, 1902), and perhaps the Licking

and Kentucky drainages. Another alternative, its

having evolved in the most southwestern parts of

the Ohio basin (Cumberland and Tennessee drainages), is unlikely since it would have needed considerable time to attain its wide range in nonglaciated regions east of the Ohio basin, and since much

of its range in the southwetsern Ohio basin appears

to be recently acquired.

The two alternatives available for the spread of

N. micropogon deep into the present southwestern

Ohio basin are through possible main river connections with the lower Teays or lower Ohio River,

and/or through stream captures with upper or middle Teays tributaries (present middle Ohio basin)

in the region about southwestern Virginia and eastern Tennessee and Kentucky. Some of the evidence

against the first alternative is the lowland and large

river conditions the species would have to traverse

in the long bend of the middle and lower Teays. It

could have used the more recent and shorter Ohio

River after the present course of the Ohio was determined by glacial advance but this route is also

unlikely due to river conditions similar to the Teays.

If N. micropogon did use a large river pathway

downstream to the Green, Cumberland, and Ten-
NUMBER 85 67

nessee, upon entering these drainages it could have

encountered competition from N. effusus, a species

more adapted to the lower gradients. Their distributional relationships in the Cumberland (Lachner

and Jenkins, 1967) support the latter hypothesis.

These species are allopatric except for being collected together on one possible occasion near a fringe

of their ranges. Even if the latter record is valid,

this does not mean that close competition is lacking.

The absence of N. micropogon from the Green

drainage is evidence that it did not use a downriver

course to the Cumberland and Tennessee or, if it

did, that this species and N. effusus were incompatible in the Green, the latter being dominant.

Ingresses of N. micropogon into the Tennessee

and Cumberland drainages were most likely through

stream capture since there is considerable biological

evidence for this, some geologic evidence, and a bulk

of evidence against lower main river entry. The

favored sequence of entrance may have been from

Big Sandy to Tennessee, then into the Cumberland,

and then the first (or another) entrance into the

Kentucky drainage. It could have been attained the

Kentucky and Licking drainages if these were middle Teays tributaries and probably entered them

once they were connected to the Ohio River.

Evidence is given above that the early range of

N. micropogon included the Big Sandy drainage, a

tributary of the middle Teays and presently a middle Ohio basin tributary. Much of the upper Tennessee drainage of Virginia is adjacent to the upper

Big Sandy. An unknown capture(s) is invoked to

account for transferal of N. micropogon from the

Big Sandy to the Tennessee. This species would not

have entered the Tennessee from the adjacent portion of the upper Teays (present New drainage)

since its cognate, N. platyrhynchus, occurs therein.

N. micropogon has spread throughout most of the

middle and upper Tennessee, but is apparently absent from most of its lower portion. Since the habitat

in some lower Tennessee tributaries appears quite

suitable, its absence may be related to a somewhat

slow downriver dispersal rate, large stream size and/

or lower stream gradients. The many uninhabited

streams are additional evidence that N. micropogon

entered the Tennessee from the northeast and not

through the lower Teays or Ohio Rivers.

Stream capture involving the Tennessee and upper

Cumberland is most likely. Even if N. micropogon

had dispersed upriver in the Cumberland through

the large area occupied by N. effusus, the former

species would have had to surmount Cumberland

River Falls—an impossibility—in order to attain its

distribution throughout the upper Cumberland.

The upper Cumberland has a depauperate fauna

(Kuehne and Bailey, 1961) compared to those of the

middle Cumberland and upper Tennessee. N. micropogon was either one of the relatively few species

to make the crossing, or to survive thereafter.

The well-established population of N. micropogon

in the middle Cumberland was probably derived

from washing over the falls and/or from stream

capture between Big South Fork of the Cumberland

and the Tennessee or between Rockcastle River, or

surrounding tributaries of the Cumberland and the

Kentucky drainage. Explanation of the presence of

the population in the Caney Fork system, middle

Cumberland, presents greater difficulties. N. micropogon may have entered Caney Fork directly through

stream capture with the middle Tennessee drainage.

Possibly, it is a relict population predating farther

upriver movement of N. effusus, and now cut off

from the main population by N. effusus (and impoundments). It probably is not the result of human

introduction since one specimen was collected in

1917, earlier than times of considerable trucking

about of bait minnows. The use of minnows as bait

in the recently created southwestern Ohio basin

impoundments is now heavy.

The Cumberland-Kentucky drainage divide was

very likely crossed through stream capture, with the

direction of movement largely into the Kentucky.

Kuehne and Bailey (1961) give good geological evidence for capture between these drainages and biological evidence for movement in the same direction.

The differentiation of N. micropogon observed in

eastern Kentucky has zoogeographic implications.

The largest differences in mean values of body circumferential scales are among adjacent drainage

populations sampled (Table 12) in the Cumberland

(grand mean 32.1), Kentucky (29.9), and Big Sandy

(31.9). The differences between the Kentucky and

the former and latter are, respectively, 2.2 and 2.0.

The next greatest differences from Atlantic slope

drainage populations are 1.3 and 1.2; all others are

less, usually considerably less than 1.0. N. micropogon is thus relatively consistent in circumferential

scale numbers over most of its range, except in
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

eastern Kentucky. A distinct bimodality exists in

the frequency distribution of the scale counts from

the Kentucky drainage. We have not found a bimodal distribution of meristic characters from any

other drainage population of Nocomis. Populational differences in other characters are present but less

marked. Lateral line differences approach those of

the circumferential scales. The Tennessee drainage

population has the most reduced tuberculation and

smallest head parts within the species. Too few

large specimens are available from eastern Kentucky

to determine relationships by these morphometric

characters.

Additional differences became evident after segregating the eastern Kentucky circumferential scale

counts by tributary systems or areas within the Cumberland and Kentucky drainages (Lachner and Jenkins, 1967, Table 3). The difference (1.3) between

the means in scales between the middle and upper

Cumberland populations, separated by the Falls,

indicates their partial isolation; some individuals

may still go over the Falls.

Within the Kentucky drainage, 86 specimens from

the South and Middle Fork systems (the two more

westerly of the three Forks of the upper Kentucky

drainage) have a mean of 30.2. There are no significant differences or trends among the individual samples from the two systems. Thus the western upper

Kentucky drainage mean of 30.2 and upper Cumberland mean of 32.7 are appreciably different and

represent the greatest scale difference among adjacent populations of N. micropogon and within all

other species of Nocomis. The upper Cumberland

and Kentucky are in the same or similar physiographic areas; thus these differences probably are

genetically based. The same was concluded by

Kuehne and Bailey (1961:4) for a species of darter

(Percidae) with two subspecies that have similar

differences and distributions. It would be difficult

to think, because of this difference, that the South

and Middle Fork populations are derived from the

upper Cumberland population through relatively

recent stream capture, were it not for the fact that

the populations of the North Fork and Red River

systems, Kentucky drainage, also are apparently

different from the South and Middle Fork populations. The mean circumferential scale values from

the Red River (29.3, 20 specimens), a middrainage

tributary, and the North Fork tributaries (29.0, 26

specimens) are lower than those from the South and

Middle Forks, and also lower than in any other

population of AT. micropogon (Table 12). The strongly assymetrical frequency distributions (Lachner and

Jenkins, 1967, table 3) for both the North Fork and

Red River samples suggest that selection is operating strongly against individuals with scales larger

than those found in these populations.

Several implications are derived from the Kentucky drainage data. It appears that a trend toward

a large-scaled form began in the Kentucky, probably

prior to the entrance into the upper Kentucky of

a finer scaled population from the upper Cumberland. The stream capture described by Kuehne and

Bailey (1961) occurred between the upper Cumberland and a tributary of South Fork of the Kentucky.

Such a capture would interject the finer scaled Cumberland form within the range of a larger scaled

differentiate of the Kentucky. Downstream movement within the South Fork and then through the

main Kentucky River would lead a fine-scaled form

first to the Middle Fork. Two aspects of the number

of circumferential scales for the South Fork-Middle

Fork sample are notable: (1) The mean is somewhat

intermediate between those for the upper Cumberland drainage and the North Fork-Red River samples although much closer to the latter. (2) A distinct

bimodality exists for the South Fork-Middle Fork

sample. An intermediate mean may be expected

with either occurrence or absence of interbreeding

between a fine-scaled upper Cumberland form and

a large-scaled North Fork-Red River form. The bimodality may indicate divergence of these forms to

a level higher than that of a race. The Nocomis of

the Kentucky, upper Cumberland, and adjacent

drainages warrant further study, particularly to determine if possible differences in fin coloration (see

p. 46) correlate with scale differences.

The question of dispersal from the Kentucky River

of the large-scaled form cannot be adequately appraised at present since we have little data on adjacent populations. Drainages of nonglaciated Kentucky and West Virginia are believed to have served

as refugia for N. micropogon during glacial times.

Study of Ohio and Indiana populations may yield

clues to the determination of which refugia held

their ancestors.

The 15 specimens of N. micropogon from the

Licking drainage, between the Kentucky and Big
NUMBER 85 69

Sandy drainages, have a mean value of 30.7 circumferential scales, distributed as 28 (in 1 specimen),

30(3), 31(10), 32(1). Four specimens (CU 25595)

from the Little Sandy River, northeastern Kentucky,

have counts of 29(1 specimen), 30(1), 31 (2). The

mean for 56 specimens from the Big Sandy drainage

is 31.9. The means are similar or somewhat lower

from southern Ohio River populations farther upriver from the Big Sandy, in West Virginia. These

data indicate that the large-scale Kentucky drainage

population may not have spread widely south of

the Ohio River.

Why N. biguttatus is not at least moderately distributed in the southwestern Ohio basin is a perplexing question. N. biguttatus generally inhabits

streams of lower gradient than does N. micropogon

(Lachner and Jenkins, 1967:573). The latter apparently is absent from most of northern Kentucky, a

region partly suitable for occupancy by N. biguttatus, and thus it would not be a competitor. One

reason may be that the eastward extension of N.

biguttatus is probably largely or solely related to

postglacial drainage events north of the Ohio River

and its habit of keeping within glaciated regions in

the East. Its entire range north of the Ohio River

is Recent and is almost entirely within or peripheral

to regions of Illinoian glaciation; it is thus absent

from the major portion of southern Ohio and Indiana (and Illinois). With its current absence from

these areas, it is somewhat unlikely that it crossed

the Ohio River even though it occurs in the northern Ohio basin. The single southeastern population,

known in Elkhorn Creek, lower Kentucky drainage,

in all probability stems from bait bucket escapees.

The Elkhorn has been a popular smallmouth bass

fishing stream for the last century (Towles and McClane, 1965:446), and chubs are favored bait.

The upper Savannah drainage population of N.

micropogon was probably derived from the Little

Tennessee River of Tennessee drainage by stream

capture. Capture here is well documented by geological and additional biological evidence (Ramsey

and Woolcott, ms).

The presence of N. micropogon in the Coosawattee system of the Mobile drainage may be attributed

to stream capture with the Hiwassee system of the

Tennessee. We have not further examined this or

considered other possibilities.

N. raneyi: This species probably evolved in the

Roanoke drainage from an ancestral stock of N.

platyrhynchus after its transfer by stream capture

from the New drainage. Ample opportunity for

passage to the Roanoke seems to have been available. Wright (1934:61-72), Thompson (1939), and

Dietrich (1959) argued that drainage loss by Little

River and other adjacent New River tributaries was

effected by headward erosion of the lower elevated

upper Roanoke in the region of present North

and South Forks of the Roanoke River. Wright

(1934:61) believed the Roanoke captured an area of

approximately 200 square miles formerly drained by

the New since the close of the Harrisburg cycle (late

Tertiary). Dietrich (1959:29-32) also supported a

hypothesis that the headwaters of the Dan River,

Roanoke drainage, have captured portions of Big

Reed Island Creek of New drainage. N. raneyi is a

member of the rich and distinctive Roanoke fauna

composed of several endemic and semi-endemic

forms, many of which have their heritage to the west.

With the Roanoke drainage as its center of dispersal, N. raneyi spread northward and southward.

Ingress to the Chowan drainage may, in part, have

been during the Pleistocene when the latter was

tributary to the Greater Roanoke. Some events have

certainly facilitated dispersal between these drainages since recent collecting has shown the Chowan

fauna to be very similar to that of the Roanoke. A

major problem lies in how N. raneyi and several

other species, now common to the Roanoke, Tar,

and Neuse, entered one of the latter two from the

Roanoke.

Although specific geological evidence is lacking

for stream captures between the Chowan, Roanoke,

Tar, and Neuse drainages, apparently because the

evidence would be obliterated on the easily eroded

Piedmont and Coastal Plain cover, captures may be

inferred to have occurred during late Pliocene to

Recent times. Clark and Miller (1912:215) stated

that coincident with the Pliocene submergence of

the Coastal Plain there seems to have been a slight

elevation and tilting of the Piedmont west of the

shoreline, and that (op. cit.:200) successive uplifts

of the Coastal Plain occurred during the late Pleistocene period of shore terracing. Accompanying rejuvenation and subsequent headward erosion, with

lack of structural control in these provinces, may

well have resulted in captures. The lower Roanoke

presently has no large tributaries entering from the
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

north or south; these were probably lost by capture

to the Chowan, Tar, and Neuse. Further, as the

Coastal Plain emerged from inundations, swampy

conditions must have once been more continuous

than at present. Some swamps are not yet discontinuous, but are now becoming so (R. D. Ross,

personal communication).

Biological evidence confirms these hypotheses. A

large number of species is shared by the four drainages, some of which are small stream forms that are

less likely to have utilized river-mouth connections.

In addition to transfer with Piedmont captures, the

occurrence of common upland species in Coastal

Plain streams, such as N. raneyi, suggests the possibility that they may at times have gotten through

some swamp connections to adjacent drainages.

Members of the more typical Coastal Plain fauna

probably attained part of their wide ranges in similar manner.

The entry of N. raneyi into the James drainage

from the upper Roanoke is an interesting problem

with several aspects, best treated after establishing

some pertinent distributional facts. Its limited range

in the James is roughly paralleled by that of three

additional species of the upper Roanoke drainage

that are not known from north of the James drainage or the New drainage. These are a sucker Moxostoma cervinum, a madtom catfish Noturus gilberti,

and a darter Percina crassa roanoka. Of the three,

M. cervinum appears to have spread farthest within

the James, perhaps due to greater vagility. This

species is known from the middle and lower Craig

Creek system and only downriver, in lower Catawba

and Jennings Creeks, and in lower Buffalo Creek of

the lower Maury River system. The known range

within the James of N. gilberti and P. crassa roanoka

is entirely within the Craig Creek system. These

three species are taken in about equal abundance

in the Craig Creek system and the upper Roanoke

drainage. These peculiar distribution patterns stand

out after consideration of the entire James and

Roanoke fauna. These two drainages share numerous other species, some of which range widely or are

more or le&s restricted to the central Appalachians;

all, however, range throughout most of the James

and Roanoke drainages or generally occur where

their preferred conditions exist. (The single exception is Pimephales notatus, a popular bait and farm

pond culture species, widely distributed in the New

drainage, whose small range within the upper Roanoke appears to have resulted from introductions.)

In order to have achieved their wider distributions

these species probably were exchanged at a date

earlier than that of the first-mentioned four species

and/or at more than one locality. In addition, isolation of some geminate pairs has been sufficiently

long for differentiation to the specific or subspecific

level (see C. R. Gilbert, 1961:455-456; 1964:107, for

discussion of these cases).

The similar and limited ranges of the four species

strongly suggest that they entered the Craig Creek

area of the James from the upper Roanoke through

a relatively recent connection. They do not appear

to be restricted by lack of suitable stream conditions

elsewhere in the James since their preferred habitat,

although unoccupied, is readily accessible. No distinct ecological differences are apparent between the

Craig Creek system and other upper and many middle James tributaries. Some evidence (p. 49) indicates that N. micropogon probably occupied the

Craig Creek system prior to the entry of N. raneyi

into the James. Competition with N. micropogon

may be one factor limiting the further dispersal of

N. raneyi within the James. As evidenced by the

limited distributions of the three other species, however, which do not appear to be restricted by competition, their entrance to the James was probably

of such recent date that sufficient time to achieve

wide distributions has not elapsed.

Robins and Raney (1956:31) suggested that the

populations of M. cervinum and N. gilberti in the

Craig Creek system were derived from the Roanoke

drainage through stream capture, but that Hybopsis

"sp" (= N. raneyi) could have come from the Roanoke or New drainages. Present morphological

knowledge of the chubs excludes the latter alternative.

Connections between the Roanoke and James

drainages (Figure 20) were probably available. There

appears to have been a complex of recent captures

in the present valleys of North Fork of Roanoke

River and Catawba Creek, involving these streams,

a Craig Creek and a New drainage tributary (Wright,

1934:68). Trout Creek, a small upper Craig Creek

tributary that heads against upper North Fork and

flows through a gap in the Craig Creek-North Fork

divide, probably captured a portion of North Fork.

This seems to be the only possible route through
NUMBER 85 71

which fishes could have entered Craig Creek directly from the Roanoke since they are separated elsewhere by relatively unbroken mountain ridges. Another possible capture is one between North Fork

and Catawba Creek. Although fishes might cross the

existing low divide as it is flooded during a period

of excessive rain, the divide may not have shifted

much at the expense of either stream (R. D. Ross,

personal communication). A third alternative may

be connection in Fincastle Valley between Catawba

Creek and upper Tinker Creek, the major Roanoke

River tributary at Roanoke. Evidence for this capture is given by Wright (1934:62-63).

The distributions of these four species have some

significance, bearing on which of these possible

routes may have been used. Although N. raneyi is

not known from North Fork of Roanake (not taken

in five collections containing N. leptocephalus), it

may be in its lower portion. At least two of the three

other species were taken from North Fork. All four

probably occurred in lower Tinker Creek which

now suffers from pollution. N. raneyi and M. cervinum occur in lower Catawba Creek, but the two

other species are absent or rare therein since they

were not taken in the eight known collections from

lower and middle Catawba. Thus, although N. gilberti and P. crassa roanoka may have entered the

James from either the North Fork or Tinker Creek,

their apparent restriction to the Craig Creek system

suggests that they entered via a North Fork-Trout

Creek capture. This cannot be said for N. raneyi

and M. cervinum since they range more widely.

Further complexity results from the existence of the

atypical population of N. raneyi in the main James,

between the Craig and Catawba Creek populations

and the Pedlar River population. Whatever factors

have affected this population, they probably operated subsequent to the spread of the typical form

from Craig or Catawba Creeks to Pedlar River. The

bull chub and sucker apparently have not dispersed

to James tributaries upriver from Craig Creek. The

main channel and tributaries between Craig and

(upstream to) Potts Creek, however, are poorly sampled or they have not been collected.

We have considered that perhaps the four species

were introduced by man into the James from the

Roanoke. This may be suspected owing to the limited ranges and only recent records of the species.

Our records of N. raneyi in the James date from

1941, when G. W. Burton (Burton and Odum, 1945)

collected a single specimen. The three other species

were not reported from the Craig Creek system by

Burton and Odum (1945) and our records of these

date from 1951. Apparently the only fish collection

made in this system before Burton's was by Cope in

1867 (Cope, 1868). None of Cope's material has been

labeled as from Craig Creek, and none of it represents any of the four species. Very few specimens

from Burton's work are at the University of Michigan Museum of Zoology, probably since only the

"necessary specimens [were] preserved" (Burton and

Odum, 1945:183). Recent collecting in the Craig

system above Newcastle yielded ten native species,

in addition to those discussed above, that are relatively common but were not listed by Burton and

Odum. Other species taken at only a few stations by

Burton were found to be comparatively widespread.

The lack of early records of the four species is

probably related to insufficient collecting and loss of

collections.

There is evidence that the four species were not

introduced by man. Since N. gilberti is generally an

uncommon species in the Roanoke there is slight

chance of it being captured and introduced by

fishermen. The chances are also slight that the common but relatively delicate darter would survive an

extended period in a bait bucket. Few bait collectors really get into riffles, which are the preferred

habitat of these two species. The hardy N. raneyi

are more easily taken by bait collectors, but this

species is generally uncommon in the upper Roanoke. The establishment of the Craig Creek and

atypical populations of N. raneyi downstream almost

certainly predates any recent introduction. The five

spillway-type dams across the James River between

Jennings Creek and Pedlar River have probably

been in effect since the latter part of the last century. Their height, from 8-25 feet, would impede

movement.

A final and important point is to reexamine the

historical and problematical status of M. cervinum

in the James drainage in light of recent data. The

probable absence of M. cervinum in the James

drainage and its early confusion with Moxostoma

(Thoburnia) rhothoecum, widely distributed in the

James, was discussed by Hubbs (1930:43-44), Raney

and Lachner (1946b:218), and Raney (1950:159).

Soon afterward, Robins and Raney (1956:31) report-
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ed the capture of M. cervinum from Johns Creek

at Newcastle and later it was found to range an

appreciable distance downstream. The two suckers

occur syntopically in the Craig Creek system. Cope

(1868:236) described and gave the type locality of

M. cervinum as the headwaters of the Roanoke and

James Rivers, Virginia. From Cope's paper it is

clear that he collected in the Craig Creek system

and he mentioned no other upper James tributary.

M. rhothoecum was not recognized by Cope. Robins

and Raney (1956:25-26) found that Cope's James

drainage specimens, designated by Fowler as cotypes

of M. cervinum, were M. rhothoecum. It appears

then, that Cope's color plate clearly depicting the

diagnostic coloration of M. cervinum, was based on

specimens from the Roanoke drainage where he also

collected, and where this species is common. The

possibility remains, though, that Cope saw M. cervinum from the Craig Creek system but, as was sometimes the case, the specimens were not saved or were

subsequently lost.

It should be noted that most of the more downstream records of Thoburnia rhothoeca listed by

Burton and Odum (1945:186-187, tables 4 and 5)

probably were, in part or entirely, M. cervinum since

this species replaces the former in larger streams and

is the more abundant of the two in the Newcastle

area today. Jordan (1889a: 109), prior to the description of a Thoburnia, reported M. cervinum from

Elk Creek, a short, direct tributary of the James

River about eight river miles upstream from the

mouth of Maury River, and from lower Buffalo

Creek, apparently just upstream from the locality

at which M. cervinum was recently taken. For several reasons, one being Jordan's color description

of his specimens, Raney and Lachner (1946b: 218)

concluded that "they were largely Thoburnia" (M.

rhothoecum). Hubbs (1930:43) found at the Museum of Comparative Zoology one of Jordan's specimens of "cervinum" from Buffalo Creek to be M.

rhothoecum and we located three of the same among

the old Indiana University collection now at the

Museum of Zoology, University of Michigan and six

at the National Museum of Natural History. It still

appears, however, that characters of both suckers

were included in Jordan's description and that at

least some of his "older" specimens were M. cervinum. After leaving the James drainage, Jordan collected in the upper Roanoke where he may have

taken M. cervinum and then compounded the colors

of at least two species of suckers under his account

of M. cervinum from the James. It is strongly indicated that M. cervinum and the other three species

were in the James drainage long before fishermen

with minnow buckets became common and transportation easy.

N. leptocephalus: The origin of the leptocephalus

group is not clear, but after differentiation of the

more northern form, N. I. leptocephalus, considerable dispersal occurred that included entry into

several drainages. Its further advance northward

along the Atlantic slope and westward may have

been through some of the same connections utilized

by N. raneyi. N. leptocephalus typically inhabits

smaller streams than the members of the micropogon group. It would thus be suspected, more than

the latter group, to cross drainage divides with

greater frequency through small headwater captures.

The dispersal of N. leptocephalus raises the possibility that there may have been many undetected or

geologically unconfirmed captures in the central Appalachians. Once N. leptocephalus entered a drainage, a wide distribution within the drainage would

probably have taken longer than for a large stream

inhabitant.

We have no data that indicate whether N. leptocephalus occupied the Neuse, Tar, Roanoke, and

Chowan before N. raneyi. Its entry into the James

probably predates that of N. raneyi because it is

widespread in the James and it occurs farther northward. The bluehead chub may have entered the

James drainage by captures along the broad front

of adjacency of the Roanoke and Chowan to the

James. The Greater Susquehanna River is invoked

to account for the presence of N. leptocephalus in

the York drainage.

The occurrence of N. leptocephalus within single

stream systems of both the Rappahannock and Potomac drainages suggests it got into these by stream

captures from the adjacent James drainage. There

is no evidence for or against the possibility that

these may be due to introductions. A third alternative, through the Greater Susquehanna, is quite unlikely, particularly in the Potomac case, since the

streams where N. leptocephalus is known in the two

drainages are headwaters, appreciable distances from

the drainage mouths. Inspection of topographic

maps of the upper Piedmont in the region of tribu-
NUMBER 85 73

taries of the Rapidan River, Rappahannock drainage, and those of the Rivanna River of the James,

show that headwaters of these streams are opposed

to each other although no definite elbow of capture

is evident. Clear geological evidence is given by

Thompson (1939:1350-1351) for a capture of the

Potomac by a Rivanna River tributary in the Blue

Ridge near the Albemarle-Augusta County line. If

N. leptocephalus used this connection it would have

entered South River of the Potomac at a point approximately midway between the two nearby Potomac streams in which it was taken.

Entry of N. leptocephalus into the New drainage

was probably through one or more of the recent

captures by the Roanoke. This species is known

only from Virginia tributaries of the New. Three

additional species, Chrosomus oreas, Notropis albeolus, and Notropis cerasinus, are also known in the

New almost exclusively from the Virginia portion

and may also be recent arrivals. The most upstream

tributaries where these four species were taken are

Fox and Wilson Creeks, Grayson County, which

enter the New just downstream from the VirginiaNorth Carolina state line. Geological barriers to

further dispersal up the New are not known to have

existed in the past.

Downstream in Virginia near the Virginia-West

Virginia line, C. oreas is known from Big Stony

Creek, Giles County (Ross and Perkins, 1959:27).

Ross and Perkins (1959:27) and C. R. Gilbert (1964:

139, map 3) gave records of N. cerasinus from Big

Stony. The range of N. albeolus extends farthest

downstream of the four species. C. R. Gilbert (1964:

156) reported a collection from the lower Greenbrier River and believed its presence there was due

to very recent dispersal or introduction, as it was

not collected earlier from this stream and in streams

somewhat farther upriver in West Virginia and Virginia just across the state line. The Bluestone Dam,

completed in 1949, on the New River one mile

above the mouth of the Greenbrier, and its impounded waters extending up to the state line,

obstructed recent dispersal. The presence of N.

albeolus in the Greenbrier may be related to earlier

dispersal or introduction.

Evolution of Nocomis

Our interpretation of die phylogeny of the species

of Nocomis is based on morphology, coloration,

zoogeography, and life history. The most primitive

Nocomis is probably the biguttatus group (Lachner

and Jenkins, 1967). The tubercles in the species of

this group are rather evenly spaced dorsally over the

head. They are absent on the snout. The tubercles

are small to moderate in size (but large compared to

almost all American cyprinids). The general pattern

appears to be the basic one that has become modified in various manners within the micropogon and

leptocephalus groups. Generally, the cyprinids with

unspecialized breeding habits tend to have small

tubercles distributed over the head in no outstanding pattern, although some differences in the patterns usually exist among the species. Development

of particular tubercle patterns and larger tubercle

size correlate in many of the North American cyprinids known to have specialized reproductive behaviors, such as nest building, special mode of spawning and highly developed territorialities. Examples

of these specialized forms, in addition to Nocomis,

include Campostoma, Semotilus, Pimephales, and

certain Notropis. The two related, nest-building

species of Exoglossum are notable exceptions.

The tubercles are slightly to considerably larger

in all members of the micropogon and leptocephalus groups compared with those of the biguttatus

group. A probable specialization of tubercle distribution, shown only by the micropogon group, is

their development on the anterior snout and lachrymal areas. A trend toward this condition is actually

seen in the biguttatus group since the dorsal tubercles are developed slightly anterior to the anterior

internasal line and, in adults, there are 1-3 tubercles located in the subnasal area but not over the

lachrymal bone.

Large body size would be of selective advantage

for nest construction and defense. N. biguttatus is,

over its range, the smallest species and it may follow

that the larger size of all of the other species is the

derived condition.

Another important phylogenetic character is the

pharyngeal dentition. The general trend in North

American cyprinids is probably toward loss of teeth

in the major and minor rows and a subsequent loss

of the minor row. The dentition of N. biguttatus

and the Redspot chub, an undescribed species (part

3) in die Arkansas and Red River drainages, is

almost always 1,4—4,1, sometimes 1,4-4,0; loss of the

minor tooth row would result in the dental formula
74

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of an early stock of the micropogon and leptocephalus groups. N. effusus, considered an advanced form

in the biguttatus group (Lachner and Jenkins, 1967)

also lost the minor tooth row.

Linear (rowed) arrangement of head tubercles

does not appear to be a primitive condition in Nocomis. The head tubercles in the biguttatus group

do not align in definite rows when the adult complement is approached or fully developed. Occasionally several tubercles form a longitudinal line over

a portion of the head but this appears to be the

result of chance. We do not know how Branson

(1962:536) construed that most head tubercles occur

in rows on the nuptial male of N. biguttatus. The

pattern of early tubercle development in juveniles

of the biguttatus group also does not indicate a

rowed condition since the first several tubercle spots

that appear in the many juveniles examined are

widely scattered. Only occasionally in small juveniles of N. platyrhynchus and N. raneyi do some of

the tubercle spots tend to align into longitudinal

rows through the internasal and anterior interorbital areas; in such cases several randomly located

tubercle spots are also present. With growth in all

of these species, rows are not evident.

In apparently advanced species of Nocomis in

which a moderate to low number of tubercles occur,

there is a trend for individual tubercles to develop

consistently in specific locations and, in some cases,

they appear to be arranged linearly. In N. micropogon the tubercles in the anterior middorsal region

often develop in four rows, two curving posterolaterad on each side from the anterior internasal

area to the anterior edge of the orbits. This trend

is most evident in the Tennessee drainage population of N. micropogon which averages the lowest

total tubercle numbers within the species. The first

tubercle spots that develop in juveniles of this species are included in the internasal rows.

A similar situation but more extreme than in

N. micropogon occurs in the leptocephalus group.

These forms have the largest tubercles (the largest

of North American cyprinids) and the lowest number in Nocomis, and the tubercles generally develop

more constantly in specific areas of the head than in

N. micropogon. This is particularly shown by the

southern form, N. leptocephalus bellicus, adult

males of which almost always have only six tubercles

and which develop in nearly the same places on all

specimens. The posterior internasal and anterior

interorbital tubercles of the leptocephalus group are

those that particularly develop in the same location

of the head. A row typically curves just above the

dorsal half of the orbits.

In the micropogon and leptocephalus groups the

tubercles are always largest where tubercle spots first

develop, in the internasal and interorbital areas.

The tubercles in the biguttatus group are largest in

the occipital areas; spots first develop in the internasal and interorbital areas, but before many appear in these areas they also develop in the occipital

region.

Developmental patterns parallel to those in Nocomis occur in other species of North American

cyprinids. The large majority of those species with

high tubercle numbers on the head dorsally do not

have rowed tubercles. When rows are present, they

often occur over rather narrow bones and other

supporting structures. An example is the distinct

tubercle rows over the mandible in species of the

subgenus Pteronotropis, genus Notropis (Bailey and

Suttkus, 1952, pi. 2). The rows over the lachrymal

bone in all species of the micropogon group is another example. When present on the fins, tubercles

occur on the rays. Tubercles often tend to have a

linear development near edges or folds of skin such

as around the orbits and above the lips. When

present on scales, the tubercles are usually at the

free, posterior margin, forming a row. All species

of Pimephales have a low number of tubercles and

they are large in size, comparable to the leptocephalus group. Also in Pimephales, the tubercles develop

in rather definite locations and form rows (Hubbs

and Black, 1947:13-17). From developmental patterns of tubercles, other morphological aspects and

zoogeography, Gibbs (1957:187, 189-192) postulated

that the occurrence in rows of head tubercles is

primitive within the subgenus Cyprinella, genus

Notropis, and that the scattered tubercle condition

is derived. Results of a detailed study of tubercle

development (Koehn, 1965) in one of the presumably derived species of Cyprinella indicate that its

early tubercle pattern is similar to the adult pattern

of primitive species. Examination of many nuptial

males of Cyprinella indicates that dorsal head tubercles tend to be more linearly arranged in the species

with fewer and larger tubercles, which may also be

the primitive condition within Cyprinella.
NUMBER 85 75

Synthesis of these comments on tubercle development is difficult since there appear to be several

factors involved. Limited area for basal attachment

would be a simple reason for linear development

of tubercles over narrow supporting structures.

Tubercle rows about the orbit and other structures

could be a protective mechanism during agonistic

behavior and nest-building activities. Tubercles developed along the edge of scales would come into

more use than if they were more basal, near the

imbricated scale base. Genetic fixity for confinement

of few tubercles to discrete areas may be associated

with specific behavior patterns. Tubercles that develop in rows, such as on the snout of Pimephales

and the dorsal part of the head of species of Cyprinella, may serve more efficiently in special breeding

behaviors.

The biguttatus group is most closely related to

the micropogon group; neither is particularly closely related to the leptocephalus group. The nuptial

male coloration, consisting in the biguttatus and

micropogon groups of pink and reddish over the

lower head and body, is quite unlike the blues and

oranges in the leptocephalus group. The whorled

intestine of the latter group, readily derived from

the simple S-shape intestine of the two other groups,

is also unique within Nocomis.

The nuptial crest appears to be a functional and

adaptive character. It is prominently developed in

the micropogon and leptocephalus groups. Some

nuptial male specimens of N. effusus have a very

slight head swelling which has not been observed

in N. biguttatus; this may be a lesser development

of the crest. Possibly the crest arose independently

within the micropogon and leptocephalus groups.

The fact that the crest is alike in development and

general form in these two groups, and that it is unknown in other cyprinid fishes, suggests common

ancestry among the two groups and that it was not

derived independently.

Crest development, lateral curving of the tips of

tubercles and loss of the minor tooth rows relate

early stocks of the micropogon and leptocephalus

groups. The leptocephalus group would be regarded as tfi<? most divergent from a common biguttatus

stock.

The crest is perhaps an adaptation for lessening

the effects of head impact from certain agonistic

behavior. The moderate enlargement of tubercles in

two of the three species of the micropogon group

and the considerable enlargement of tubercles in the

leptocephalus group may correlate with the development of large crests in these groups. A large, hard

(cornified) tubercle pressing into the cephalic region

upon impact would give more severe shock than

smaller tubercles. N. I. bellicus and an undescribed

subspecies (part 2) are apparently derived from N. I.

leptocephalus by loss of the occipital and medial

interorbital tubercles and show a trend toward reduction of the crest posteriorly. However, N. micropogon lacks tubercles in the interorbital and occipital areas where the crest is often well formed.

The direction in which the tips of tubercles point

is different in the groups. The head tubercles of

most cyprinid species have erect or antrorse tips.

The tubercles or their tips in N. biguttatus are

mainly antrorse, but a few are more or less erect.

The tips of most tubercles in the other two groups

of Nocomis are erect or curved laterally.

The development of a hiatus (absence of tubercles) between the anterior snout and internasal

tubercles occurs in the micropogon group. The absence of tubercles may have a functional basis, for

the development of tubercles, especially large ones,

would inhibit the protractility of the upper jaw. A

complete absence, or but a few tubercles, occurs between the anterior snout and internasal areas in N.

micropogon; tubercles are usually present in this

area in N. platyrhynchus and are always present in

N. raneyi. The absence of tubercles at the hiatus is

correlated with tubercle size in these three species,

N. micropogon developing the largest tubercles and

N. raneyi the smallest. Within the subgenus Cyprinella, 14 of 16 included species also develop a hiatus

in tubercle distribution (Gibbs, 1957:187), in the

same region as in the micropogon group. The hiatus

is absent in two species that have small tubercles

and, for additional reasons, are regarded as a distinct species group within Cyprinella (Gibbs, 1961).

The hiatus occurs over the areas where the vertex

of an angle is produced when the mouth is opened

and the upper jaw is protracted, or the skin in this

area during protraction is stretched to a greater

extent than in adjacent areas. Some specimens have

wrinkles or folds of skin in the hiatus area, apparently the result of upper jaw movement. The bases

of large Nocomis tubercles developed in the hiatus

area would tend to impede jaw action or be eroded
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

off from such action. This is particularly apt to

occur during nest building in Nocomis, as the males

very often open the mouth to about its full extent

when picking up large gravel. Small-based tubercles

in the hiatus area probably do not produce such an

effect and/or may not be as readily shed from the

skin stretching beneath them.

A red postocular spot occurs only in the biguttatus

group. It probably arose in the biguttatus group

after the origin of the other groups. The postocular

area in all species of Nocomis is usually lighter in

shade than the immediate surrounding area. Genetic

fixity for retention of a red blush color by the nuptial male may have occurred since this could be an

effective species-recognition mark. Its variable development among the species of the biguttatus group

is discussed by Lachner and Jenkins (1967).

SUMMARY.—The basic stock of the micropogon

group, after some differentiation from its precursor

(biguttatus line), may be characterized as having

tubercles completely distributed over the head and

lachrymal bone; tubercles small to moderate in size

with many tips directed laterad; rosy nuptial body

color; nuptial crest present, perhaps small, in breeding males; pharyngeal tooth arch structurally moderate to heavy; teeth 4-4; body size moderate to

large. The form presently closest to this prototype

is N. platyrhynchus. The head tubercles in N. platyrhynchus are widely distributed, often posterior

to the midoccipital or posterior occipital lines as in

the biguttatus group. They are more widespread

than in N. raneyi, and tubercles never occur in the

occipital, and rarely in the interorbital areas in N.

micropogon. Adults of N. biguttatus generally have

a fairly sharp downward break of the contour of the

nape at the occiput, down to a somewhat flat head

dorsum. Such contours are best developed in N.

platyrhynchus, less often and not as well developed

in N. micropogon and N. raneyi.

The area of origin of N. platyrhynchus was probably the upper Teays River system. Its prototype,

derived from a stock of the biguttatus group, moved

into the lower and/or middle Teays. Kanawha Falls

on the New-Kanawha River provided opportunity

for differentiation of N. platyrhynchus stock in the

New drainage above. Geographically, the upper

Teays is central to the present ranges of the two

species derived from N. platyrhynchus stock, N.

micropogon and N. raneyi.

Stream capture between the Roanoke and New

River drainages permitted transferal of a N. platyrhynchus stock into the Roanoke drainage and

could have lead to the origin of N. raneyi in the

Roanoke through subsequent isolation and differentiation.

N. raneyi is the most divergent species in the

micropogon group and relatively unique within

Nocomis in some of its diagnostic characters. The

head and its parts have small features, particularly

the mouth. The body is moderately deep and compressed in adult males. Juveniles have the most

streamlined body form in Nocomis. The caudal fin

area is large, the lobes varying in larger specimens

from subequal to considerably unequal, the upper

lobe always longer. This unequal condition of the

caudal fin lobes was thought by Uyeno and Miller

(1965:38) to be unique among North American

cyprinids to Pogonichthys, a monotypic western

genus. N. raneyi is the largest species in the genus

and in this respect rivals the fallfish, Semotilus

corporalis, the largest cyprinid in eastern United

States.

The tubercle distribution in N. raneyi approaches

the primitive condition and it is not greatly different from N. platyrhynchus (or the basic group

stock). The most posterior tubercles in adult males

generally occur in the PIO and AOC areas. The small

tubercles are rather closely spaced dorsally over

most of the head, extremely so on the snout and

internasal areas. Two other probable primitive conditions in Nocomis are seen in characters of N.

raneyi. The pharyngeal arches are heavy and their

anterior edentulous processes are short, as in the

biguttatus group. The scales are relatively small in

size, though somewhat larger than in N. biguttatus.

N. micropogon probably arose from N. platyrhynchus or a stock common to both in a portion of the

middle Teays system, below Kanawha Falls, or in a

drainage that was involved in stream capture with

the upper Teays. N. micropogon has the most reduced tuberculation of the micropogon group, having the least number and most restricted distribution, but has the largest tubercles.

N. micropogon and N. platyrhynchus differ from

N. raneyi (as reviewed above) and, in some characters, from all other Nocomis in the development of

the tubercle pattern, tubercle numbers, broad head,

slender pharyngeal arch, dark lateral body stripe,
NUMBER 8 5 77

size of intestinal loop, and nature of caudal fin.

In the two characters which best separate N. micropogon from N. platyrhynchus, the latter is more

similar to but still different from N. raneyi. One is

the occurrence of tubercles in the interorbital and

occipital areas. This is probably the primitive condition within the group, best retained in N. platyrhynchus. The small scales in the circumferential

series of N. platyrhynchus may indicate inheritance

from the primitive micropogon group stock, or

adaptation to ecological conditions in the somewhat

higher altitudes of the New River drainage as

compared to most or all other regions inhabited by

group members.

Several conditions indicate that introgressive hybridization may have occurred in the past between

N. micropogon and N. platyrhynchus. The overall

similarity of these species is part of the evidence.

Tuberculation in N. platyrhynchus is extreme for

the micropogon group in their posterior distribution, since these sometimes occur in the POC area

(probably a primitive condition); however, it is also

the most variable species in this respect. A good

proportion of large juveniles and adults have tubercles only as far posterior as the AIO and MIO lines.

In addition, all specimens with tubercles developed

into the more posterior head areas have them widely

scattered, much more so than in N. raneyi. Thus,

although in some specimens tubercles occur in the

occipital area, the total tubercle number is not

increased (Table 2). The expression of other characters in N. platyrhynchus may also be due to hybridization with N. micropogon. The similarity of

populations of N. micropogon in the Monongahela

and Potomac River drainages, described above, sup

port a hybridization hypothesis. From the Monongahela-Potomac populations we obtain the highest

tubercle counts. This is effected by an extreme

crowding of internasal tubercles and increased posterior distribution. Stream capture apparently has

occurred between the New and Monongahela and

between the Monongahela and the Potomac drainages (see Dispersal, p. 66).

An interesting aspect of hybridization is the relatively common occurrence of the hybrid combination N. leptocephalus X N. micropogon in the James

River drainage where these species are sympatric

over a great area of the river system and are often

captured together. Eighty-seven specimens have been

examined of this hybrid combination. The hybrid

specimens have been taken in many tributaries and

their occurrence does not reflect any particular geographical rea. The James River drainage is the only

one in which these two species are known to occur

together abundantly. N. micropogon's close relative

in the New River system, N. platyrhynchus, is sympatric also with N. leptocephalus over a large area

of the system; however, we have found only five

specimens of N. leptocephalus x N. platyrhynchus.

The relatively uncommon production of hybrids

may be associated with different specific reproductive ecologies, or it may be related to genetic incompatibility. We know that N. leptocephalus prefers

smaller streams and occupies the swifter waters

(sometimes riffles) of these streams for its nest sites.

N. platyrhynchus prefers larger streams and nests in

the deeper pool areas, out of the riffles. N. micropogon and N. platyrhynchus are not known to be

sympatric in any portion of their range and thus we

cannot evaluate their genetic compatibility. We also

have the following interspecific hybrid combinations

of Nocomis: N. micropogon x N. raneyi, 2 specimens; N. leptocephalus X N. raneyi, 2 specimens;

N. biguttatus X N. micropogon, 1 specimen. Incompatibility between the latter two species is suggested

for they are sympathetic over most of northeastern

United States, they have been collected together

abundantly (N. micropogon and N. raneyi rarely),

in many intermediate streams they occur together

commonly and in approximately equal numbers,

both species build nests and have similar spawning habits, both forms have spawning periods that

overlap, and both forms hybridize fairly commonly

with members of other cyprinid genera. We have

placed these species in separate groups on the basis

of morphological characters, and we have done likewise with the parental stocks of the common hybrid

N. leptocephalus X N. micropogon. Intergeneric

hybrids involving Nocomis and other cyprinids most

frequently involve N. biguttatus, N. micropogon,

and N. leptocephalus. This hybridization may be

most facilitated by the great geographical distributions and abundance of these species and in their

choice of wider reproductive ecological sites.

The unknown precursor of Nocomis was probably

not an elaborate nest builder and probably had

fewer sexually dimorphic features. The trend for

selection to act more on sexually dimorphic charac-
78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ters is characteristic of Nocomis, probably related

to the functional aspects of nest construction, nest

guarding and territorial maintenance by the male,

and the need, particularly by the females, to recognize the males. We thus find in the evolution of the

chubs, elaboration or adaptive reduction of important diagnostic characters as the extent of tuberculation, development of nuptial crests and develop

ment of particular reproductive colorations among

the three recognized groups and among the species

or forms within the groups. The immediate precursor of N. biguttatus probably was highly tuberculate. The dorsal head tuberculation is well developed and similar in the three living species of the

biguttatus group. Evolution in the group involved

the tuberculation of the lateral body scales, degree

of squamation, reduction in dentition and, to a

lesser degree, the development of the postocular

spot.

The primitive stock of the micropogon group,

best exemplified by N. platyrhynchus, shows relationships with biguttatus stock in the widely distributed, cephalic tuberculation, smaller tubercles, finer

squamation, and stouter pharyngeal arch. Evolution

within the micropogon group involved, mainly: the

reduction in tubercle numbers; reduction in cephalic

tubercle distribution; increase in tubercle size; crest

development; and in the differentiation of the pharyngeal arch, size of scales, special features of the

head (as mouth size) and the caudal fin. The leptocephalus group shows a greater reduction in tubercle numbers compared widi the micropogon group,

a reduction in cephalic tubercle distribution, great

increase in tubercle size, specific reproductive coloration, stouter body, and coarser scales. Evolution

within the forms of the leptocephalus group also involved further reduction of cephalic tubercle numbrs and distribution, and in the differentiation of

special reproductive color patterns.

Summary

The chub genus Nocomis Girard of North America

has been characterized and relationships among

three species groups in Nocomis were discussed. Two

new species were described from the central Appalachian region, both in the micropogon species

group. Characters separating Nocomis from other

barbeled cyprinids are that the nuptial males have

large breeding tubercles on the head that show a

phylogeny among the species in their number, distribution, and size; have expansive crests or swellings

on the head in two species groups that develop coincidentally with the spawning period; develop elaborate breeding coloration; construct large moundnests of gravel which they transport in their mouths.

The genus has large scales, the body circumferential

scales are fewer than 37; the pharyngeal teeth number 1,4-4,1 to 3-3; the scale radii are present only

in the posterior field and the total number of radii

in the adults ranges from 23 to 54; the total number

of vertebrae ranges from 38 to 43; and a single,

terminal maxillary barbel.

The three species groups were defined and their

characters summarized, namely, the biguttatus group

with three species, the micropogon group with three

species, and the leptocephalus group with three subspecies. The central Appalachian region is inhabited

by four species, N. micropogon (river chub), N.

platyrhynchus, new species (bigmouth chub), and

N. raneyi, new species (bull chub), of the micropogon group, all of which are sympatric with N. leptocephalus (bluehead chub) in one or more river

drainages. A discussion of the nomenclatural history

of the genus and nominal species was given and important diagnostic characters Useful in differentiating among the specific and several infraspecific

populations were discussed, evaluated, and summarized. The critical specific and subspecific characters,

such as cephalic tuberculation involving the distribution, number, size and developmental patterns

of tubercles, cephalic crests or swellings and nuptial

body coloration are sexually dimorphic, developing

elaborately in the males. Many morphometric characters, as well as tubercle numbers and distribution,

show an allometric relationship with body size. An

account of each species has included a synonymy,

diagnosis, description and comparison of meristic

and morphometric characters, coloration in life and

in preservation, populational differentiation, reproduction, growth and size attained, materials studied

and geographic distribution. The species are illustrated, special body features are drawn, distributions are plotted, and important character data are

shown in text figures.

N. micropogon and N. leptocephalus are widely

distributed, the former occurring over most of northeastern United States and the latter is found chiefly
NUMBER 85 79

on the Atlantic and Gulf slopes. In the study area,

N. leptocephalus is found east of the Atlantic drainage divide north to the Potomac River and west of

the divide in the upper New River (New-Kanawha

River system). N. micropogon occurs east of the

Appalachian divide from the James drainage north

ward and throughout the western slope including

the Kanawha River system (below the Falls). Both

of these species are common and are widely distributed in the James River drainage. N. raneyi is an

Atlantic slope form, occurring in the James, Chowan,

Roanoke, Tar and Neuse drainages, where it is

found commonly and is widely sympatric with N.

leptocephalus. These two species and N. micropogon

occur together in limited areas of the James drainage. N. platyrhynchus is found throughout the New

River system, where part of its range overlaps that

of N. leptocephalus.

The ecological requirements of the chubs were

reviewed and species preferences discussed. The primary ecological distinction among the species is

stream size. Regional ecology was reviewed in respect to occurrence and abundance of the species.

Generally, the species of Nocomis prefer clear, moderate to warm water streams of intermediate gradients. They are chiefly carnivorous, common over

stream beds of gravel, rubble, and boulders. Wellestablished populations of three species in typical

Piedmont streams indicate an adaptiveness to this

shifting, sandy habitat. All species of chubs are rare

below the Fall Line in the upper Coastal Plain

province.

Associations, interrelationships and frequency of

hybridization were compared among the four species. The three species of the micropogon group

were regarded as ecological homologues. N. leptocephalus almost always coexists with N. raneyi in

moderate-sized streams, where they occur sympatrically, and it occurs less frequently with N. micropogon and N. platyrhynchus. Differences are observed

among the species of the micropogon group in their

frequency of hybridization with N. leptocephalus.

Hybridization is somewhat inversely proportional

to the frequency of syntopic occurrence. Only two

natural hybrids N. raneyi X N. leptocephalus were

found, whereas N. micropogon x N. leptocephalus

hybrids are comparatively common (87 specimens

known). A population of N. raneyi in a limited area

of the James River is considered atypical. The

meristic and morphometrical data of this atypical

population were compared with typical N. raneyi

and N. micropogon from the James drainage. Introgressive hybridization may have occurred between

N. micropogon and N. platyrhynchus because it best

explains the similarity of populations of the former

in the Monongahela and Potomac River drainages

with the latter species in the New River system.

The biological and geological evidence providing

an explanation of the dispersal of the chubs and

other species of fishes in various interdrainage exchanges in the central Appalachian region was comprehensively reviewed. Possible routes of entry into

the several drainages were proposed. Present, detailed distributional patterns of the chubs and other

species were related with geologic events, and present and past ecological conditions, in an effort to

understand the conditions, barriers, and routes affecting dispersal. Three kinds of geological events

that operated in the dispersal of chubs are considered: stream capture, eustatic changes of the Atlantic

Coastal Plain, and Pleistocene drainage modifications.

The interpretation of evolution in Nocomis is

based on morphology, coloration, zoogeography, and

life history. The unknown precursor of Nocomis was

probably not an elaborate nest builder and probably had fewer sexually dimorphic features. The

trend for selection to act more on sexually dimorphic features is characteristic of Nocomis, probably

related mainly to the functional aspects of nest construction, nest guarding and territorial maintenance

by the male. Nocomis shows an elaboration or adaptive reduction of important diagnostic characters as

the extent of tuberculation, development of nuptial

crests, and development of particular reproductive

colorations. The most primitive Nocomis are probably of the biguttatus group, based on, among other

characters, the general pattern, small size and relatively large numbers of cephalic tubercles. This

general tubercle pattern appears to be the basic one

that has become modified in various manners within

the micropogon and leptocephalus groups. The

primitive stock of the micropogon group, best exemplified by N. platyrhynchus, shows relationships with

biguttatus stock in the widely distributed cephalic

tuberculation, small tubercles, fine squamation, and

stout pharyngeal arch. Evolution within the micropogon group involved, mainly: the reduction in
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tubercle numbers; reduction in cephalic tubercle

distribution; increase in tubercle size; development

of the nuptial crest; and in the differentiation of

the pharyngeal arch, size of scales, and certain head

and body features. The leptocephalus group, the

most advanced, shows a greater reduction in tubercle numbers compared widi the micropogon group,

a reduction in cephalic tubercle distribution, great

increase in tubercle size, development of specific

reproductive coloration, stouter body and coarser

scales. The forms of the leptocephalus group evolved

further in the greater reduction of cephalic tubercle

numbers and distribution, and in the differentiation

of special reproductive body colorations.

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84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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Tables

TABLE I.—The progressive development of tubercles posteriorly on the head with increase in body length in Nocomis

raneyi taken from Atlantic drainages. (Data for sexes combined; see text for explanation of symbols.)

Standard

length

(in mm)

60-69

70-79

80-89

90-99

100-109

110-119

120-129

130-139

140-149

150-159

160-169

170-179

180-189

190-199

200-209

210-219

220-229

230-239

240-249

PIN

1

3

6

5

3

2

.

1

AIO

1

5

4

12

17

18

13

4

1

2

1

-

-

1

MIO

1

2

9

18

24

21

10

5

3

4

-

1

1

PIO

5

14

17

13

16

4

7

3

2

5

2

2

2

_

1

AOC

1

1

-

2

2

1

-

1

.

1

.

1

1

1

MOC POC

1

1

-

2

1

1

TABLE 2.—The progressive development of tubercles posteriorly on the head with increase in body length in Nocomis

platyrhynchus taken from the upper New drainage. (Data for

sexes combined; see text for explanation of symbols.)

Standard

length

(ln nm)

50-59

60-69

70-79

80-89

90-99

100-109

110-119

120-129

130-139

140-149

150-159

160-169

170-179

180-189

190-199

200-209

210-219

220-229

PIN

1

8

9

6

2

1

.

1

AIO

3

9

17

20

14

7

3

1

1

1

1

-

1

1

MIO

1

2

7

10

6

7

4

3

4

1

4

1

2

-

1

1

PIO

_

1

5

5

4

3

3

6

2

2

2

2

1

AOC

1

.

2

1

2

2

1

-

1

1

MOC

1

1

.

1

1

1

-

2

1

1

1

POC

1

1

3

-

-

5

3

1

TABLE i.—The presence of tubercles or scars on mature male specimens, by size categories, of four species of Nocomis.

N.

N. z N.

raneyi

pla tyrhynchus

micropogon

lepCocephalus

Drainage

Susquehanna

Potomac

Rappahannock

James

Tennessee

Allegheny

Other northern

drainages

100-

109

1

1

1

110-

119

-

2

-

7

120-

129

1

1

-

2

3

14

130-

139

1

2

2

1

-

1

3

14

140-

149

-

8

2

-

-

4

3

7

18

150-

159

1

8

3

1

-

7

4

3

11

Standard

160-

169

1

7

4

1

1

6

2

2

4

170-

179

1

3

-

4

1

-

1

length

180-

189

1

3

1

4

3

1

190-

199

2

3

-

200-

209

2

2

-

210-

219

2

1

1

220-

229

1

1

230-

239

1

240-

249

1

TABLE A—The relative size at which tubercle buds develop in several species of chubs (Nocomis) segregated by sex. (Specimens of

N. micropogon from several northern, eastern, and central drainages.)

N.

N.

N.

Species

raneyi

platyrhynchus

micropoRon

d"

a

9

cC

9

7 0 -

79

3

1

8 0 -

89

2

6

4

9 0 -

99

4

4

9

4

100-

109

6

2

13

5

110-

119

4

3

9

12

120-

129

9

1

24

8

Standard

130-

139

6

-

28

4

140-

149

4

1

19

2

length

150-

159

1

6

16

160-

169

3

12

13

170-

179

2

10

1

180-

189

1

4

1

190-

199

2

2

200-

209

-

210-

219

1
86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 5.—The number of head tubercles in Nocomis raneyi, segregated by sex, taken from Atlantic drainages. (Females in

parentheses; mean values with sexes combined.)

Standard Number of tubercles on head

(la am) °"

 10" 20

~

 30" *°~ 50

~

 60

~

 70

~

 80" '°~ 10 ° " 110" 120* 13 ° " 140' 15 ° " 160' 17 ° " 18 ° " l90~ 20 ° " 9 19 29 39 49 59 69 79 89 99 109 119 129 139 149 159 169 179 189 199 209

240-249

230-239

220-229

210-219

200-209

190-199

180-189

170-179

160-169

150-159

140-149

130-139

120-129

110-119

100-109

90-99

80-89

70-79

60-69

50-59

40-49

1

<D

2

(3)

2

(7)

16

(11)

16

(*>

1

(1)

(2)

1

(7)

3

(13)

3

(3)

5

(2)

1

(1)

1

(2)

(5)

1

(9)

9

(7)

1

(2)

1

(1)

(2)

-

(4)

4

(11)

4

(8)

4

(5)

1

(2)

-

(2)

4

(1)

3

(7)

9

(3)

1

(2)

(1)

1 1 .. .

(1)

114---12 -

(1) - - - - (1) ' <D

1----221--1 1

(1) - (1)

1 1 2 12-331 2

(1) - - (1) - (2) (1)

1 1 2 4- 2

(2) - (5) (1) (3) - (1)

232225- 1

(2) (8) (5) (7) (3) (2)

4-232- - 1 1

(5) (5) (2) (4) (1)

4 1111 1

(3) (2) (2)

1

1

3

2

6

18

34

42

35

31

14

13

4

164.5

204.5

157.8

204.5

137.7

133.1

132.5

149.5

133.5

130.0

106.3

81.3

70.9

54.1

40.0

25.3

14.5

7.1

4.5

4.5

14.5

* Sexes combined.

TABLE 6.—The number of head tubercles in Nocomis platyrhynchus, segregated by sex, taken from the upper New drainage.

(Females in parentheses; mean values with sexes combined.)

Standard Number of tubercles on head

length In — — ^ Jot Jot SQI 60^ 70^ 80^ 90^ 100^ 110^ 120^ 130^ 5 ii i~

89 99 109 119 129 139

220-229

210-219

200-209

190-199

180-189

170-179

160-169

150-159 1

140-149

(1) (1)

130-139 3

120-129 2

(1)

110-119

100-109

90-99

80-89

70-79

60-69

50-59

(2)

2

(4)

7

(10)

3

(1)

(4)

4

( I D

9

(5)

2

(7)

1

(9)

3

(4)

(1)

(5)

3

(4)

3

(5)

3

(1)

1

(3)

(2)

8

(1)

2

(1)

(1)

(1)

5

(1)

3

-

4

1

2

2

6

8

12

14

9

8

10

15

15

18

12

10

12

9

3

2

6

8

13

19

19

9

10

84.5

109.5

84.5

91.2

87.0

89.5

88.1

77.8

79.5

69.5

65.0

57.1

42.2

34.5

20.0

12.6

5.6

4.5
NUMBER 8 5 87

TABLE 7.—The number of head tubercles, sexes combined, in

Nocomis micropogon from the Potomac drainage and the

Cheat and Tygart rivers of the upper Monongahela drainage.

(Data for the Monongahela specimens in parentheses.)

Standard

length

In trm

0- 10- 20- 30- 40- 50- 60- 70- 80- 90-

210-219

200-209

190-199

180-189

170-179

160-169

150-159

140-149

130-139

120-129

110-119

100-109

90-99

80-89

70-79

60-69

50-59

(2)

8

(1)

(1)

2

(2)

(2)

(2)

10

(3)

3

(2)

(3)

(2)

(1)

(6) (1)

(4)

N, - Potomac drainage

N- - Cheat and Tygart

54.5

79.5

69.5

65.8

67.8

54.5

49.2

39.9

39.5

34.5

TABLE 8.—The number of rows of scales along the lateral line

in four species of Nocomis.

Susquehanna

Chesapeake trlbs., Md.

Potomac

Rappahannock

York

Savannah

Cumberland

Kentucky

Big Sandy

Guyandot

Coal, Kanawha

Elk, Kanawha

Ohio trlbs., W. Va.

Monongahela

Allegheny

Lake Erie, Ontario, N.Y.

38 39

3

2

1

23

40

31

16

10

15

61

41

45

15

38

23

33

44

42

47

16

38

8

14

9

43

13

3

6

5

2

?

4 27

1 9

6 41

5 20

0

1 64

2 19

8 40

1 22

9 59

6 16

1 9

5 15

2 9

3 10

4 20

3 37

6 18

6 50

6 14

1 6

3 27

20

4

31

6

4

46

11

47

31

24

19

4

9

16

9

23

26

18

42

4

3

31

3

5

1

9

1

18

9

2

3

1

6

1

7

5

7

10

1

16

TABLE 9.—The number of head tubercles, segregated by sex,

in Nocomis micropogon taken from the upper Tennessee

drainage. (Females in parentheses.)

Standard

length in

Number of tubercles on head

10-

19

20-

29

30-

39

40-

49

50-

59

180-189

170-179

160-169

150-159

140-149

130-139

120-129

110-119

100-109

90-99

80-89

70-79

60-69

(1)

(1)

(4)

1

(6)

11

(6)

25

(3)

1

(2)

2

(4)

4

(10)

5

(6)

6

(1)

6

(1)

16

10

(1)

13

(3)

5

(1)

7

(6)

5

1

10

(1)

2

(2)

4

1

9

6

14

18

27

13

17

8

11

10

8

17

25

1

3

5

6

17

10

7

7

3

31.6

31.2

31.6

31.7

29.1

26.4

25.4

19.5

18.8

15.0

10.5

7.4

4.5

x Sexes combined.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 10.—The number of head tubercles, segregated by sex, in Nocomis micropogon taken from

the James drainage. (Females in parentheses.)

Standard

length In

mm

180-189

170-179

160-169

150-159

140-149

130-139

120-129

110-119

100-109

90-99

80-89

70-79

60-69

50-59

0-

9

(2)

(1)

3

(13)

24

(14)

3

(2)

10-

19

(1)

(4)

1

(6)

2

(12)

6

(8)

2

20-

29

(1)

(1)

(4)

(8)

5

(6)

5

(2)

2

30-

39

3

1

-

2

4

(2)

1

(8)

3

(2)

10

(2)

3

Number

40-

49

2

1

2

1

4

(2)

3

3

(1)

1

2

1

of tub<

50-

59

2

-

3

2

2

2

(1)

1

1

60-

69

1

1

2

N

<3

5

1

6

5

8

7

9

3

6

17

10

11

26

3

N

?

3

2

5

13

14

16

15

21

14

2

52.5

44.5

42.8

48.5

48.1

44.5

40.9

32.6

28.5

25.4

19.3

20.1

5.0

4.5

x Sexes combined.

TABLE 11.—The number of rows of scales above and below the lateral line in four species of

Nocomis.

Species

raneyl

James

Chowan

Roanoke

Tar

Neuse

pla t yrhynchus

New, Va.-N. C.

alcropogon

Potomac

Rappahannock

York

James

Tennessee

Cumberland

Kentucky

Guyandot

Kanavha

Motion gahela

Lake Ontario

leptocephalus

James

Roanoke

Xar

Neuse

New

6

1

1

2

1

1

5

4

79

12

26

26

Scale

7

28

7

37

4

7

25

39

17

3

57

35

12

24

3

2

12

20

14

54

1

5

34

rows

8

54

8

24

2

13

42

7

5

1

3

7

19

1

1

1

above

9

1

1

1

lateral

N

83

15

63

6

20

67

46

22

5

62

43

32

25

4

3

12

25

18

133

13

31

61

line

X

7.7

7.5

7.4

7.3

7.7

7.6

7.2

7.2

7.0

7.0

7.1

7.7

7.0

7.3

6.7

7.0

6.8

6.8

6.4

6.1

6.2

6.6

4

5

3

2

5

2

Scale

5

1

2

1

3

8

1

4

12

1

1

4

7

7

114

11

26

34

rows

6

45

15

49

5

16

26

44

16

3

59

38

32

12

3

2

8

18

6

16

25

below

7

33

1

12

1

3

38

2

4

5

4

4

1

lateral line

8 N

79

16

63

6

19

2 66

46

21

6

72

43

31

24

4

3

12

25

18

133

13

31

62

X

6.4

6.1

6.2

6.2

6.2

6.6

6.0

6.1

5.5

6.0

6.1

6.0

5.5

5.8

5.7

5.7

5.7

5.1

5.1

4.9

4.8

5.4
NUMBER 85

TABLE 12.—The circumferential scale rows in four species of Nocomis.

26 27 28 29 30 31 32 33 34 35 36 37 38 39

Greenbrle

Gauley

Cheaspeake trlbs., Hd.

Savannah

Cumberland

Kentucky

Big Sandy

Guyandot

Coal-Kanawha

Elk-Kanawha

Ohio trlbs. , W. Va

Youghlogheny

Allegheny

Lake Erie, Ontario

61 57 30

32 34 34

58 54 28 28

18 19 16

14 39 63 49

16 32 33 14

51 65 41

14 43 29 23 13

6 14 23 22 6

10 13 18

14 29 24

12 14 10

33.3

33.4

33.1

32.9

33.0

34.2

33.1

32.7

33.2

33.6

30.5

31.8

32.4

31.2

31.5

31.0

31.1

31.8

32.1

29.9

31.9

30.9

30.3

31.0

31.0

31.6

32.2

31.5

31.3

29.2

28.8

28.1

28.1

29.4

TABLE 13.—The number of head tubercles in Nocomis micropogon taken from the Susquehanna drainage and localities

over most of the range of the species west of the Allegheny

Mountains.

S. L.

mm

180-189

170-179

160-169

150-159

1A0-U9

130-139

120-129

110-119

100-109

90-99

80-89

70-79

60-69

50-59

(J-

?

1

1

1

1

1

Number

10-

19

2

1

1

3

4

1

of

45-

29

1

1

3

3

7

5

k

3

-

-

tuber

30-

3?

I

-

3

10

11

8

7

4

k

1

3

1

.lea o

40-

4?

2

8

5

12

10

6

5

3

k

a head

50- 60-

59 69

3

3 1

4 1

4

2

3

2 1

N

1

2

15

20

31

25

ih

18

15

12

8

3

1

1

TABLE 14.—The number of rows of scales around the caudal

peduncle in four species of Nocomis.

Caudal peduncle

Specie

14 IS 16 17

Cumberland

Kentucky

Kanavha

Honongahela

Allegheny

Lake Erie , Ontario

H. T.

36 28 16
90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 15.—The number of vertebrae in four species of Nocomis.

Species

Drainage

ranevi

James

Chowan

Roanoke

Tar

Neuse

platvrhvnchus

New, Va.-N.C.

Greenbrier

Bluestone

mlcropogon

Susquehanna

Potomac

Rappahannock

York

James

Tennessee

Cumberland

Kentucky

Big Sandy

West Virginia

Monongahela

Allegheny

Lake Ontario

Lake Michigan

leptocephalus

Potomac

York

James

Chowan

Roanoke

Neuse

Nev

Pre

20 21

16

1 7

2

2

5

23

1 15

1

4

1

1

1

1

1

caudal

22

3

1

9

7

14

93

21

5

22

38

34

7

59

19

28

42

18

8

46

4

25

7

24

7

14

6

40

9

17

71

15

16

76

18

35

19

6

1

1

3

1

12

8

1

9

3

2

17

4

1

1

11

2

6

1

13

24

4

2

5

16

2

1

3

2

1

1

1

1

7

1

3

Ci

17

1

6

4

3

19

12

30

4

29

7

2

15

4

2

22

9

13

3

20

7

33

10

22

udal

18

11

11

53

16

31

47

13

4

4

35

4

3

33

29

27

43

10

7

32

4

20

7

12

5

5

7

6

19

8

8

34

7

19

69

11

2

1

5

9

21

7

6

1

7

3

20

2

3

1

3

8

1

2

37 38

1

4

1

1

4

2

39

1

1

19

16

53

6

35

3

8

26

4

8

8

1

14

9

12

11

37

16

24

1

40

10

6

11

55

10

3

6

38

7

3

46

30

34

35

18

13

49

4

21

21

10

12

21

4

11

1

22

otat

41

66

11

60

30

37

79

17

3

3

9

22

10

6

10

1

10

3

2

2

3

2

42 43 44

50 4 1

16

31 4

3

13

7

N

131

27

95

39

61

142

28

6

25

58

64

9

90

55

52

67

32

14

67

4

33

24

26

21

36

16

52

19

48

41.4

41.6

41.4

40.9

41.0

40.6

40.6

40. S

39.2

39.7

39.0

39.3

39.7

40.4

40.0

39.7

40.2

40.1

40.0

40.0

39.8

40.0

39.5

39.6

39.8

39.8

39.8

38.9

39.5

TABLE 16.—Number of pectoral fin rays in species of Nocomis

segregated by river systems.

Species

raneyi

James

Chowan

Roanoke

platvrhvnchus

James

Tennessee

leptocephalus

James

Roanoke

New

15

1

1

3

3

16

14

8

2

4

4

7

29

11

17

29

8

48

4

14

26

17

58

13

18

5

12

14

35

18

2

9

2

19

1

2

2

1

N

50

8

68

23

56

50

27

99

29

T

16.8

17.0

17.1

17.5

17.6

17.4

16.9

16.7

16.5

TABLE 17.—The relationship between snout length and gape

width in 85 juveniles of two species of Nocomis ranging in

length from 50 to 99 mm. Data in parentheses refer to N.

platyrhynchus, the remainder, to N. raneyi. About 60 percent

of individuals of the two species do not overlap.

Snout length in per cent of S.L.

S.5 9.0 9.5 10.0 10.5 11.0 11.5

-j 9.5

•g 9.0

t 8.5

7-0

( 1 )

3

6

6

(1)

2

1

(2)

5

(O

5

2

(2)

2

(1)

1

(3)

_

(4)

1

(5)

(1)

(4)

U)1

(3)

(1)

( 1 )

( 1 )
NUMBER 8 5 91

TABLE 18.—The degree of breast squamation in four species of

Nocomis.

Species

Squamation ratings

raneyl

James

Chowan

Roanoke

Tar

Neuse

platyrhynchua

58

20

50

1

6

3

3

New, Va.-N. C.

Indian

Bluestone

Greenbrler

Gauley

mlcropogon

Susquehanna

Chesapeake trlbs., Md.

Potomac

Rappahannock

York

James

Savannah

Tennessee

Cumberland

Kentucky

Big Sandy

Guyandot

Coal, Kanawha

Elk, Kanawha

Ohio trlbs., W. Va.

Honongahela

Allegheny

Lake Erie, OntarloN. Y.

leptocephalus

2

1

20

18

31

20

3

30

17

3

6

4

1

16

1

3

16

1

-

18

8

25

12

5

41

7

9

18

9

5

4

3

18

6

22

8

5

26

1

-

3

5

1

2

1

2

10

6

12

2

2

1

5

3

8

8

9

47

4

1

2

13

1

1

15

9

14

4

3

10

8

3

2

6

45

10

4

20

17

26

6

37

2

1

1

4

2

3

6

TABLE 19.—The number of head tubercles, segregated by sex,

in Nocomis leptocephalus taken from the James, Roanoke,

and upper New drainages. (Females in parentheses.)

length In

170-179

160-169

150-159

140-149

130-139

120-129

110-119

100-109

90-99

80-89

70-79

60-69

50-59

40-49

30-39

(5)

(3)

1

(6)

9

(34)

22

(17)

11

(10)

(1)

Number of tubercles on head

10-

19

18

24

(5)

18

(8)

13

(28)

23

(13)

8

(18)

16

(10)

3

(1)

1

4

11

22

29

22

32

28

13

25

10

25

25

11

_

5

8

33

17

24

44

17

10

24.5

17.0

17.2

17.2

16.6

16.3

16.7

17.3

13.4

14.5

12.7

8.3

5.2

4.5

4.5

es, to Neuse and X - Sexes combined.

TABLE 20.—The relationship between snout length and gape

width in 121 juvenile and subadult specimens of two species

of Nocomis ranging in size from 100 to 149 mm in standard

length. The higher values, in parentheses, refer to N. platyrhynchus, the others to N. raneyi of the Atlantic slope. An

overlap of about 12 percent occurred in this sample.

Snout length in per cent of S. L.

8.5 9.0 9.5 10.0 10.5 11.0 11.5 12.0 12.5

11.0

10.5

10.0

9.5

9.0

8.5

8.0

7.5

7.0

6.5

6.0

(1)

1

2

7

u

2

1

7

5

13

3

3

1

(1)

5

6

7

2

2

(1)

(4)

1

(3)

1

2

3

(2)

(3)

(5)

(2)

(2)

1

(1)

(1)

(1)

(2)

(2)

(2)

(2)

TABLE 21.—The relationship between snout length and gape

width in 70 adults, chiefly males, of two species of Nocomis

ranging in size above 149 mm in standard length. The higher

values in parentheses refer to N. platyrhynchus, the lower

values to N. raneyi. An overlap of 12.9 percent occurred in

this sample.

Snout length in per cent of S. L.

10.0 10.5 11.0 11.5 12.0 12.5 13.0 13-5

ui

•s

1per

5width

s.

3

11.5

11.0

10.5

10.0

9.5

9.0

8.5

8.0

7.5

7.0

2

2

(3)

(2)

(1)

1

(2)

(2)

(4)

(2)

1

(3)

1

(2)

(1)

(2) (1)

(2)
92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 22.—Comparison of the three species groups of Nocomis.

Species-groups

leptocephalus

Nuptial tubercles

Size, number

Distribution,

head dorsum

lachrymal

body

pectoral fin rays

direction

Nuptial crest

Coloration

nuptial

caudal spot, yg.& Juv.

anal fin (life)

upper body (life)

scale (dark anterior)

Intestine

Pharyngeal teeth

Scales, size

body, circumferential

Vertebrae

Body form

Mouth

Snout, shape

lntcrnasal hump

Preop«rcle, posterior

edge

Caudal fin lobes

Lateral line curvature

Size attained

Distribution

Spec

moderate, many

mid-snout to occiput

absent

present or absent

2 to 5 or 6

almost always antrorse

absent or slight swelling

pink-rosy, lower head and

body

large, round, distinct

orangish, brightest fin

olive-greenish

wider

present

S-shape

1,4-4,1 to 4-4

smaller

31 to 38

38 to 42

stocky to somewhat elongate

more terminal, oblique

moderate to large, many

tip of snout to mid-occiput

present

absent

2 to 5 or 6

erect, some laterad

large, more posterior

pink-rosy, lower head and

body

small, indistinct

light orange to pale

olive-greenish

wider

absent

very large, few

Internasal to occiput

absent

absent

2 to 8 or 9

erect and laterad

large to moderate, more anterior

head and body bluish or body

1th orange stripe and pink belly

small, Indistinct

pale or light olive

olive-tannlsh

narrower

absent

S-shape or small ace

loop on left side

4-4

Intermediate

28 to 39

38 to 44

more elongate

ory

4-4 to 3-3

larger

26 to 33

38 to 41

more stocky

inferior, horizontal

rounded to pointed

usually absent

intermediate

usually equal

stralghter, passing

through lateral stripe

smaller to moderate

Mississippi, southern

Great Lakes, upper Red

River of North basins

more pointed

usually present

more blunt, decliv

usually absent

effusus

Ozark chub, n. sp.

ally slanted anteriad,

below, except large males

upper lobe usually longer

straighter, passing

through lateral stripe

larger

Ohio R. and southern

Great Lakes basins,

entral Atlantic slope

platyrhvnchus. n. sp.

raneyl, n. sp.

below

lower lobe often slightly longer

more down-curved passing

through lower lateral stripe

or below

moderate

central and south Atlantic

slope, west to southeast

Mississippi R. tributaries

1. leptocephalus

T. belllcus

T. n. subsp. (southeast)

TAB(-E 23.—Proportional measurements of Nocomis raneyi expressed in thousandths of standard

length.

Measurements

USNM catalogue number

Standard length, mm

Head length

Snout length

Fostorbltal length

Lachrymal,greatest

length

Orbit length

Interorbltal, bony width

Cape width

tipper ]•» length

Predorsal length

Caudal peduncle length

Caudal peduncle depth

Body, greatest depth

Body, greatest width

Holotype

194633

242

260

126

112

81

46

92

89

89

562

209

116

267

172

194626

227

258

119

118

81

46

84

83

87

530

225

110

259

153

Mai*

194684

198

261

121

119

82

47

90

92

87

552

215

116

280

176

194617

150

252

109

103

80

55

80

84

81

541

212

111

244

165

194603

96

254

97

109

70

64

74

68

72

548

226

118

259

161

194614

53

253

95

95

61

72

72

76

72

548

215

110

232

147

191036

180

244

109

107

73

51

84

74

77

523

198

119

264

169

Fe

194631

142

246

103

106

68

54

73

72

75

532

217

110

247

174

males

194602

121

245

99

104

66

58

74

74

70

546

211

11J

276

168

194614

93

254

95

99

66

58

75

78

72

530

217

109

241

165

194614

68

260

105

99

65

74

78

80

75

555

212

111

238

155
NUMBER 8 5 93

TABLE 24—Proportional measurements of Nocomis platyrhynchus expressed in thousandths of

standard length.

Measurements

Catalogue number

Standard length,

Head length

Snout length

gth

Holotype

USNM

194817

163

TU

25902

195

Mai

USNM

194804

159

es

USNM

194812

133

USNM

194817

82

USNM

194804

57

USNM

194257

142

USNM

194811

123

Females

TO

25902

109

USNM

194816

86

USNM

194257

65

Lachrymal, greatest

length

Orbit length

Interorbttal, bony width

Gaps width

Upper Jav length

Fredorsal length

Caudal peduncle length

Caudal peduncle depth

Body, greatest depth

Body, greatest width

285

125

126

89

54

83

102

92

551

230

114

252

172

292

135

137

96

100

98

517

234

115

256

151

289

128

131

87

50

84

106

99

550

225

116

241

177

274

123

114

77

93

90

538

229

115

226

159

268

108

104

69

67

82

85

520

235

113

224

150

283

108

115

66

84

75

84

87

581

213

119

234

154

279 281 275 284 268

117 122 117 112 107

121 120 117 119 109

79

57

70

80

92

543

212

110

230

151

77

58

72

88

91

559

208

113

274

191

77

61

78

87

90

561

219

111

271

180

76

69

68

78

93

549

210

114

248

159

71

65

71

84

76

546

219

116

223

140

TABLE 25.—Comparison of morphometric characters in adult males of four species of Nocomis.

Measurements are expressed in thousandths of standard length.

Species

Total sat

Drainage

Standard

iple

and subsample

length, mm : mean

range

24

James 12

Roanoke 11

Neuse 1

200

172-242

platyrhynchus

31

New

178

159-214

ml(

Potomac

Rappahannock

James

165

142-189

:rop<

~i13n

2

14

>gon

Tennessee

161

141-188

leptocephalus

47

James 32

22 Roanoke 9

New 6

138

121-173

Head length

Snout length

Postorbltal length

Lachrymal,greatest length

Orbit, bony length

Interorbltal, bony width

Gape width

Upper Jav length

Predorsal length

Caudal peduncle length

Caudal peduncle depth

Body, greatest depth

Body, greatest width

260

244-275

116

104-126

117

110-131

81

71-88

49

45-54

86

78-95

83

71-103

85

79-98

542

517-567

211

193-225

115

106-123

263

226-287

158

131-189

278

260-292

126

119-137

123

112-137

87

79-96

50

45-56

85

75-103

104

91-115

98

89-106

542

517-576

226

210-243

118

108-126

249

224-278

165

134-190

284

272-300

131

121-148

126

114-136

87

80-99

53

49-58

88

77-94

108

88-123

102

91-113

546

529-565

215

201-231

116

106-130

252

220-288

169

149-203

272

262-285

125

114-135

121

111-130

82

75-91

52

47-59

82

66-92

104

94-130

100

92-115

533

516-558

226

211-240

116

108-125

238

218-263

168

151-190

275

258-309

116

102-137

122

105-139

69

60-81

53

47-58

93

81-114

106

88-123

98

82-117

560

537-597

210

192-224

120

100-134

258

228-315

178

148-205
9 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 26.—Association and frequency of hybridization of Nocomis leptocephalus with three

related species of river chubs.

Number of Number of collections 1'

1/ In which with hybrids "eluding In columns

leptocephalus leptocephalus hybrids IV and V

occurred

James If 18 16 89 - 16 940

Chowan 4 4 100- 4 189

Roanoke 41 39 95 1 43 2098

Tar 4 3 75 - 3 64

Neuse 7 7 100 - 8 205

TOTAL - - .. . 3496

platyrhynchus

New, Va. 16 10 63 3 12 328

James 23 14 61 15 ^ 10 478

_2 / Includes only the col lections front Che Cra ig-Johns Creek system,

4/ Includes three collections lacking typical N. mlcropogon and three lacking typical £. leptocephalus.

TABLE 27' —Comparison of certain meristic characters of

Nocomis raneyi, atypical N. raneyi, and N. micropogon from

the James drainage.

CIRCUMFERENTIAL SCALES

28 29 30 31 32 33 34 35 36

raneyi

Craig

Cataw

TOTAL

atypical r

micropogon

Above

Craig

Below

ba

1

an

C

a

C

eyl

re.8*

raig

6

1

2 2

37

15

8

28

1

1

1

28

10

27

57

58

11

16

4

21

LATERAL LINE

38 39 40

53

1

54

17

5

2

4

27

1

28

20

3

1

SCALES

41 42

27

1

28

13

1

43

7

1

8

9

172 33.3

4 34.5

177 33.3

71 33.8

31.2

30.9

30.9

3 31 43 44 13

4

5

22

39

3

25

14

25

31

8

25

6

15

45

40

5

4

47

19

2

TOTAL VERTEBRAE

40

10

10

3

41

64

2

66

23

42

47

2

50

22

43

4

4

2

13

3

44

1

1

1

134

4

139

70

61

25

70

41.2

41.8

41.3

41.2

39.6

39.0

38.9

126

4

131

51

51

20

19

41.4

41.5

41.4

41.5

39.7

39.7

39.7

1 - Includes one specimen fron Pedlar

2 - Hill, Craig, Johns and Catawba Cr
Inde

x

albeolus, Notropis, 73

nmblops winchelli, Hybopsis, 14

amnis, Notropis, 14

anomalum, Campostoma, 64

Association, 58-60

Atypical Nocomis raneyi, 32-36, 56

Bigmouth chub, 16, 37

biguttatus, Ceratichthys, 37

Nocomis, 15, 69, 78

Semotilus, 13,15

blennioides, Etheostoma, 66

Bluehead chub, 16, 50

boucardi, Notropis, 14

Breeding tubercles, 7, 8

Brest squamation, 12

buccata, Ericymba, 66

Bull chub, 16, 17

Campostoma, 73

anomalum, 64

caprodes semifasciata, Percina, 66

Catostomus melanotus, 15

cerasinus, Notropis, 73

Ceratichthys, 14-15

biguttatus, 37

cyclotis, 15

leptocephalus, 50

micropogon, 15, 42

nebrascensis, 15

stigmaticus, 15

vigilax, 14

Ceratichthys x Hypsilepis, 42

cervinum, Moxostoma, 70

Chrosomus oreas, 73

Cladophora, 57

Coastal Plain province, 56, 57

Eustatic changes, 61-63

Pliocene submergence, 69

Coloration, 12

Nocomis leptocephalus, 51

Nocomis micropogon, 45-57

Nocomis platyrhynchus, 39

Nocomis raneyi, 26-29

Color forms, 47

cornu/tw, Notropis, 65

corporalis, Semotilus, 76

Coueiitw, 15

plumbeus, 35

crassa roanoka, Percina, 70

Crests (See Nuptial crests)

Cuba Outlet, 64

Cumberland River Falls, 67

cyclotis, Ceratichthys, 15

Cyprinella, 74, 75

darter, 70

Dentition (Sec Pharyngeal dentition)

Diagnostic characters, 7-13

Dispersal, 61-73

Kanawha Falls, 63

Nocomis leptocephalus, 72-73

Nocomis micropogon, 63

Nocomis platyrhynchus, 63

Nocomis raneyi, 69-72

Distribution

Nocomis leptocephalus, 54-55

Nocomis micropogon, 48-50

Nocomis platyrhynchus, 41

Nocomis raneyi, 32

Ecology (See Regional ecology)

effusus, Nocomis, 67, 74

Ericymba buccata, 66

Erimystax, 14

Etheostoma blennioides, 66

kanawhae, 63

osburni, 63

Etymology

Nocomis platyrhynchus, 37

Nocomis raneyi, 29

Evolution, 73-78

Summary, 76-78

Exoglossum, 73

Extrarius, 14

Fall Line (See Fall Zone)

Fall Zone, 56-57

Falls

Cumberland River, 67

Kanawha, 63

Geological events, 61

Eustatic changes, 61-63

Pleistocene drainage modifications, 63

Stream capture, 61

gilberti, Noturus, 70

glacier, Illinoian, 64

Greater Susquehanna River, 61, 65

harperi, Hybopsis, 14

Head crests, 7 (See ako Nuptial crests)

Head tuberculation, 7 (See abo Tuberculation)

hiatus, 75

Hybopsis, 14-15

amblops winchelli, 14

harperi, 14

fcentudfcienws, 17, 37 42 50

kentuckyensis, 37
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Hybopsis—Cont.

leptocephala, 37, 50

tnicropogon, 37, 50

Hybridization

Frequency, 58-60

Introgressive, 35, 45

Summary, 60, 77, 79

Hybrids

Ceratichthys x Hypsilepis, 42

Nocomis biguttatus x Nocomis tnicropogon, 77

Nocomis leptocephalus x Nocomis micropogon, 49, 58,

60,77

Nocomis leptocephalus x Nocomis platyrhynchus, 60

Nocomis leptocephalus x Nocomis raneyi, 59

Nocomis micropogon x Nocomis raneyi, 49, 60

Nocomis micropogon x Notropis cornutus, 42

Illinoian glacier, 64

Intestine, 11

Introgressive hybridization, 35, 45

Justicia, 57

kanawhae, Etheostoma, 63

kentuckiensis, Luxilus, 15

Hybopsis, 17, 37, 50

kentuckyensis, Hybopsis, 37

Key, 16

Lake Maumcc, 64

leptocephala, Hybopsis, 37, 50

leptocephalus, Ceratichthys, 50

leptocephalus group, 50

leptocephalus, Nocomis, 50

Coloration, 51

Description, 50-54

Diagnosis, 51

Dispersal, 72-73

Distribution, 54-55

Growth, 54

Key, 16

Materials, 54

Nuptial crest, 51

Reproduction, 54

Tuberculation, 51

Leuciscus, 14

Leucosomus, 14

Luxilus kentuckiensis, 15

Macrhybopsis, 14

Madtom catfish, 70

Materials, 6

Nocomis leptocephalus, 54

Nocomis tnicropogon, 48

Nocomis platyrhynchus, 39-41

Nocomis raneyi, 29-32

melanotus, Catostomus, 15

Meristic characters, 8

Nocomis leptocephalus, 51

Nocomis micropogon, 43

Nocomis platyrhynchus, 38

Nocomis raneyi. 23

Methods, 3

micropogon, Nocomis

Coloration, 45-47

Color forms, 47

Description, 42-47

Diagnosis, 42

Dispersal, 63-69

Distribution, 48-50

Evolution, 76-78

Growth, 48

Key, 16

Materials, 48

Nomenclature, 42

Nuptial crest, 45

Populations, 47-48

Reproduction, 48

Tuberculation, 43-45

tnicropogon, Ceratichthys, 15, 42

group, 17

Hybopsis, 37, 50

Morphometric characters, 9, 17-23, 37-38, 42-43, 51

Moxostoma, 63

cervinum, 70

rhothoecum, 71

nebrascensis, Ceratichthys, 15

Nocomis, 13

Nocomis

biguttatus, 15, 69, 78

Description, 13

Diagnosis, 13

Dispersal, 61-73

Evolution, 73-78

Key, 16

leptocephalus, 15, 37, 50-55, 72-73

leptocephalus bellicus, 50

micropogon 15, 16, 42-50, 63-69, 76-78

nebrascensis, 13

Nomenclatural history, 14

platyrhynchus (new species), 16, 37-41, 63, 76-79

raneyi (new species), 16, 17-37, 69-72, 76-78

Nocomis biguttatus x Nocomis micropogon, 77

effusus, 67, 74

leptocephalus x Nocomis micropogon, 49, 58, 60, 77

leptocephalus x Nocomis platyrhynchus, 60

leptocephalus x Nocomis raneyi, 59, 60, 77

micropogon x Nocomis raneyi, 49, 60

micropogon x Notropis cornutus, 42

notatus, Pimephales, 70

Notropis, 73

albeolus, 73

amnis, 14

boucardi, 14

<rerai«nu5, 73

foiTinftw, 65

scabriceps, 63

spilopterus, 66

Noturus gilberti, 70

Nuptial coloration, 12, 26-29, 39, 45-47, 51

Nuptial crests, 5, 25-26, 38-39, 45, 51

Old Allegheny River, 64
NUMBER 85 97

omiscomayctis, Percopsis, 66

oreas, Chrosomus, 73

Oregonichthys, 14

osburni, Etheostoma, 63

oxyrhyncha, Percina, 66

Percina caprodes semifasciata, 66

crassa roanoka, 70

oxyrhyncha, 66

Percopsis omiscomaycus, 66

Pharyngeal dentition, 26, 39, 45, 51

Phenacobius teretulus, 63

Piedmont, 69

streams, 56

Pimephales, 14, 73, 74, 75

Pimephales notatus, 70

Platygobio, 14

platyrhynchus, Nocomis (new species), 37

Coloration, 39

Description, 37-39

Diagnosis, 37

Dispersal, 63

Distribution, 41

Etymology, 37

Evolution, 76-78

Growth, 39

Key, 16

Materials, 39-41

Nuptial crest, 38-39

Reproduction, 39

Tuberculation, 38

Pleistocene

Drainage modifications, 62, 63

Glaciation, 62

plumbeus, Couesius, 35

Podostemum, 57

Pogonichthys, 76

Populations

micropogon, 47-48

raneyi, 29

Potamogeton, 57

Problematic specimens, 36-37

Pteronotropis, 74

raneyi, Nocomis (new species), 17

Atypical, 32-36

Coloration, 26-29

Description, 17-29

Diagnosis, 17

Dispersal, 69-72

Distribution, 32

Etymology, 29

Evolution, 76-78

Growth, 29

Key, 16

Materials, 29-32

Nuptial crest, 25-26

Populations, 29

Problematical specimens, 36-37

Reproduction, 29

Tuberculation, 23-25

Redspot chub, 73

Regional ecology, 55-58

Reproduction, 29, 39, 48, 54

rhothoecum, Moxostoma, 71

River chub, 16, 42

Saprolegnia, 8

scabriceps, Notropis, 63

Semotilus, 14, 73

Semotilus biguttatus, 13, 15

Semotilus corporalis, 76

spiloplerus, Notropis, 66

stigmaticus, Ceratichthys, 15

Stream capture, 61

Sucker, 70

Swellings, 5, 7

Teays River, 62, 63, 64, 67, 76

teretulus, Phenacobius, 63

Thoburnia (See rhothoecum)

Tubercle, 5, 7

anlagen, 3, 8

bud, 3, 7

evolution, 74, 75

function, 8

hiatus, 75

scar, 3, 8

spot, 3, 7

Tuberculation

evolution, 74, 75

Nocomis leptocephalus, 51

Nocomis micropogon, 43-45

Nocomis platyrhynchus, 38

Nocomis raneyi, 23-25

vigilax, Ceratichthys, 14

Yuriria, 14

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