ERNEST A. LACHNE

and MARTIN L. WILE

Populations of

the Polytypic Species

Nocomis leptocephalus

W (Girard) with a

Description of a

New Subspecies

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 92
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 92

Ernest A. Lachner Populations of

and Martin L. Wiley , i n i , o

7

 the rolytypic Species

Nocomis leptocephalus

(Girard) with a

Description of a

New Subspecies

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Lachner, Ernest A., and Martin L. Wiley. Populations of the Polytypic Species

Nocomis leptocephalus (Girard) with a Description of a New Subspecies. Smithsonian Contributions to Zoology, number 92, 34 pages, 1971.—Three subspecies

of the bluehead chub, Nocomis leptocephalus (Girard), are recognized. Our

conclusions are based on the examination of more than 20,000 specimens housed

in American institutions. The typical form, N. I. leptocephalus, ranges from the

Potomac drainage southward on the Atlantic slope to and including the Santee

drainage, and across the Appalachian Divide into the New River. The new subspecies, N. I. interocularis, occurs in the Savannah, Altamaha, and Chattahoochee

rivers and across the divide in a limited area of the Tennessee River in southeast Tennessee. The nominal form, N. I. bellicus, extends from the Alabama

River westward to streams draining into the Mississippi River from the east in

Louisiana and Mississippi, and in a limited area of the Tennessee River across

the divide in northern Alabama.

The diagnostic characters differentiating these subspecies are the number and

distribution of head tubercles. These three populations have diverged almost to

the species level. Intergrades of N. 1. interocularis X N. I. leptocephalus are

recognized from the Edisto River, South Carolina. Two basic color forms are

described, a blue and a brassy reddish, the evolutionary significance of which is

not understood. The distribution of these two color forms does not correspond

entirely to the distribution of the described morphological forms. The whorled

intestine, a specific character of N. leptocephalus, is often straight in specimens

from the Chattahoochee River.

We postulate that in Nocomis, decrease in tubercle number and increase in

tubercle size may represent a specialized condition that is especially demonstrated

among the subspecies of N. leptocephalus.

Official publication date is handstampcd in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

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

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Contents

Page

Introduction 1

Acknowledgments 1

Material Studied 2

Methods 2

Nocomis leptocephalus (Girard) 5

Nocomis leptocephalus interocularis, new subspecies 6

Nocomis leptocephalus leptocephalus (Girard) 10

Nocomis leptocephalus bellicus Girard 10

Nocomis leptocephalus intergrade interocularis X leptocephalus . . . 10

Coloration 10

Geographic Distribution and Ecology 12

Evolution and Dispersal 14

Plates 15

Literature Cited 19

Tables 21

Appendix: Specimens Studied 30

Nocomis leptocephalus interocularis, new subspecies 30

Nocomis leptocephalus leptocephalus (Girard) 34

Nocomis leptocephalus bellicus Girard 35

Nocomi s leptocephalu s intergrad e interoculari s x leptocephalu s .. . 3 5

Ernest A. Lachner

and Martin L. Wiley

Populations of

the Polytypic Species

Nocomis leptocephalus

(Girard) with a

Description of a

New Subspecies

Introduction

The status and relationships of the nominal species

Nocomis leptocephalus (Girard) and N. bellicus

Girard have received little study since they were

described (1856:213). They were included in the

synonymy of Hybopsis kentuckiensis (Rafinesque)

for many years until Hubbs (1926) resurrected N.

leptocephalus. Subsequent listings by Lachner

(1946), Moore (1968), and Bailey et al. (I960) have

recognized the two forms, but with little clarification of their morphological differences.

Preliminary studies indicated the existence of an

abundant, wide-ranging Piedmont species complex

whose range extended from the Potomac River

drainage southward to the Savannah River drainage and westward to tributaries of the Mississippi

River. First inspection of the data suggested that

the several river populations might be interrelated

under one species, with characters which formed a

cline from the northeast to the south and southwest portion of the range. Analysis of many

specimens, however, particularly the adult males,

Ernest A. Lachner, Division of Fishes, Department of Vertebrate Zoology, National Mtistum of Natural History, Smithsonian Institution, Washington, D. C. 20560. Martin L.

Wiley, Chesapeake Biological Laboratory, Solomons, Maryland 20688.

revealed characteristic abrupt breaks in character

values between certain adjacent populations, and

that three highly diverged populations were actually involved. The objectives of this paper are to

describe the morphological components of the

species complex of N. leptocephalus, including a

new subspecies, to discuss the interrelationships of

the several populations and to discuss the distribution, zoogeography, and evolution of the species

complex.

Ack now ledgmen ts

Hundreds of individuals have contributed to this

study by their collecting efforts in amassing over

1,300 collections and over 20,000 specimens from

southeastern United States. To these individuals,

we offer our great appreciation. Other individuals

have provided field observations, museum records,

have loaned collections, or made available specimens and facilities to the writers. We are especially

grateful for this cooperation, and we express our

thanks to the following individuals for their special

efforts; Joe Bailey, Duke University (DU); Reeve

M. Bailey and Robert Rush Miller, Museum of

Zoology, University of Michigan(UMMZ); Herbert

Boschung, University of Alabama (UAIC); J. H.

Elrod, Auburn University (API); Harry Freeman,

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

College of Charleston; Robert E. Jenkins, Roanoke

College; John Ramsey, Cooperative Fisheries Research Unit, Auburn University; Edward C. Raney,

Cornell University (CU); Robert D. Ross, Virginia

Polytechnic Institute; Frank J. Schwartz, Institute

of Marine Sciences, University of North Carolina;

Donald C. Scott, University of Georgia (UG);

William F. Smith-Vaniz, Institute of Marine and

Atmospheric Sciences, University of Miami; Royal

D. Suttkus, Tulane University (TU); Benjamin R.

Wall, Jr., University of Alabama; and Ralph Yerger, Florida State University (FSU).

Some specimens were also received from, or examined at, the University of Kansas (K.U); Academy of Natural Sciences, Philadelphia (ANSP); the

University of Minnesota; Oklahoma State University Museum of Zoology (OAM); and the Mississippi Game and Fish Commission (MGFC).

The staff of the National Museum of Natural

History (USNM) contributed the following: Dorothy Hubbs made X-ray photographs, summarized

various numerical data, and constructed the distribution map; Carolyn Bartlett Gast made a detailed

drawing of the new subspecies; A. M. Awl drew

the heads of three subspecies; and the Photographic Services Division made all of the photographs except the head view of N. I. leptocephalus

(Plate 6, upper) which was provided by Edward C.

Raney.

Robert E. Jenkins read the manuscript and offered valuable suggestions.

The first writer was partially supported by

Smithsonian Research Awards 3316 and 3347; the

second writer was assisted by both awards.

Material Studied

Most of the collections are housed at Cornell

University, University of Alabama Ichthyological

Collection, University of Georgia, University of

Michigan Museum of Zoology, National Museum

of Natural History, and Tulane University. The

number of collections and specimens studied, segregated by drainage, is summarized below. The total

number does not include many of the collections

containing only young specimens which were too

small to provide data on tubercle development, but

which were included on the distribution map.

Nocomis I. leptocephalus

Drainage

New

Potomac

Rappahannock

York

James

Chowan

Roanoke

T a r

Neuse

Cape Fear

Pee Dee

San tee

Savannah

Ogeechee

Altamaha

Chattahoochee

Collections

51

4

2

5

50

13

141

11

23

13

54

395

Subtotal 762

Nocomis I. interocularis

190

3

56

62

Subtotal 311

Specimens

783

69

2

159

1,471

334

4.746

230

211

70

1,133

5,343

14,551

2,275

8

1,011

789

4.083

Edisto

Nocomis leptocephalus intergrade

interocularis X leptocephalus

Nocomis I. bellicus

Mobile Bay

Pascagoula Bay

Pearl River

Mississippi River

tributaries

Tennessee River

Subtotal

TOTAL

Methods

147

12

48

33

4

244

1,322

153

1,111

185

248

433

23

2,000

20,789

The methods used in counting meristic characters

and measuring structures and body proportions

were similar to those discussed by Lachner and

Jenkins (1971). All references to body length are

to standard length (SL) in millimeters, unless

stated otherwise. The preliminary data of various

morphometric characters as well as meristic characters were segregated by size, sex, and river system.

The initial tabular data for each character were

taken from samples of from ten to twenty specimens when available, and the samples were selected

over the range of a population within a river system insofar as possible. When preliminary analysis
NUMBER 9 2

of the data revealed no major divergence within or

among certain river populations, collation was by

river drainage or by subspecies.

The cephalic tubercle counts were easily made,

almost always without the use of any great magnification, because the developed tubercles, spots,

buds, or scars are very large, comparatively few in

number, and are readily discernible. There is no

difficulty in obtaining accurate counts, unlike the

problem in counting the smaller, numerous tubercles of members of the Nocomis micropogon group.

The areas of the head referred to in tubercle distribution and location are given designations similar to those presented by Lachner and Jenkins

(1971), such as: transverse anterior internasal line

(AIN); posterior internasal (PIN); anterior interorbital (AIO); mid-interorbital (MIO); posterior

interorbital (PIO); anterior occipital (AOC);

mid-occipital (MOC); and posterior occipital

(POC). The posterior development of tubercles was

recorded as the line nearest the most posterior

tubercle (or tubercles). The supraorbital tubercles

were not considered in the above designations since

they have a comparatively similar development in

all three forms, and are independent of the progressive development of the principal head tubercles with increase in body length.

The distribution and number of head tubercles

are the two main characters involved in the separation of the subspecies of N. leptocephalus. These

two characters were also of primary importance in

the differentiation of the species of the micropogon

group. A detailed discussion of tuberculation in

Nocomis is presented by Lachner and Jenkins

(1971). The total number of head tubercles for

N. I. leptocephalus in the northern part of its range

is summarized by size groups in Table I. Because

significant differentiation in the number of head

tubercles occurred in each of the two southern

river populations of N. I. leptocephalus (Pee Dee

and Santee), these are summarized separately by

size groups in Tables 2 and 3. No notable differences were observed in total number of head

tubercles for the new subspecies in the Savannah,

Altamaha, and Chattahoochee (Apalachicola) river

drainages, and the data in Table 4 were combined

and summarized by size groups; the same was done

in Table 5 for similar tubercle data for N. I. bellicus from the Alabama River westward to tributaries of the Mississippi River. The allometric

relationship of increase in tubercle number with

increase in body size for these five diverged populations is shown in Figure 1. The curves were

drawn to best fit the means of the body-length

groups. The data for these curves are based on a

sample of 2,467 specimens, including 196 N. I. bellicus; 663 N. I. interocularis; and 843 Santee, 160

Pee Dee, and 605 northern specimens of N. I.

leptocephalus.

The total number of cephalic tubercles of specimens ranging in size from juveniles to adults and

segregated by subspecies, intergrades, and major

river drainages is summarized in Table 6. This

summary shows the difficulty in identifying juvenile specimens, particularly those involving the

subspecies N. 1. leptocephalus and N. I. interocularis. Because of the allometric growth relationship

of the number of tubercles with body length, specimens 90 mm SL and larger are compared by subspecies, intergrades, and river drainages in Table 7.

Most specimens of the bluehead chub are mature

at about 90 to 100 mm, and at this size the full

complement of tubercles is present in all populations except for N. I. leptocephalus in the SanteePee Dee drainages (see Figure 1). Figure 2 shows

the mean (Av.), range, standard deviation (SD),

and the mean ± two standard errors for the total

number of tubercles in the five populations. The

values are illustrated for two size groups, 70 to 79

mm SL and all specimens 90 mm SL or greater.

The statistical data for this graph are given in

Table 11. The differences are much less in the

smaller size group, and only the adult or subadult

specimens are of practical systematic importance in

this problem.

In order to demonstrate the sharp breaks in

tubercle numbers among the three subspecies, the

southern drainage populations of N. I. leptocephalus are compared with N. I. interocularis and N. I.

bellicus in specimens greater than 65 mm, but excluding the supraorbital tubercles (Table 8). Supraorbital tubercles vary somewhat by subspecies,

averaging notably fewer in N. I. bellicus.

Frequency distributions comparing the occurrence of tubercles posteriorly on the head, segregated by subspecies and river drainages, are given

in Table 9. The relationship between body length

and the extent of posterior development of head
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

S

CM

CM

CM

CM

CO

Ul

d8

UJ _

m 5

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tc

UJ

m *

2 -

z

CM

Z ""

UJ

s o

CD

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rt

CM

'A

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4

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/ X • ^ M **

/ / A .-

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/ .-;•'"" .

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• / >

/ ^ A

^ • ^^

A*'

4*"'

leptocephalus (PeeDse)

leptocephalus (Sontae)

~ i

* .^^^ ^

leptocepholus (Cape Fear to Potomac)

• •» * A

• •"" A

A A

•

interocularit •

bellicus o

•• TM" o

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30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 180 190 200

STANDARD LENGTH (mm)

FIGURE 1.—The relationship between body length and number of head tubercles in three subspecies of N. leptocephalus. Three populations of AT. /. Uptocephalus are also compared. The

curves were fitted to the mean values by inspection. Data for the sexes were combined. The

mean values were plotted from data in Tables 1 through 5.

tubercles is summarized by size groups for the

Santee River drainage in Table 10.

Vertebral numbers, pharyngeal dentition counts,

and circumferential and lateral-line scale numbers

are compared by subspecies and by various drainages in Table 12, 13, and 14.

Because of the large number of specimens examined and identified, only the river drainage and

the museum catalog number are given in the "Appendix: Specimens Studied" (pages 34-35) for N. I.

leptocephalus and N. I. bellicus. "Appendix: Specimens Studied" (pages 30-34) for N. I. interocularis

includes the collections for each river system

grouped by state and county. Also given are museum catalog number, date of collection, number

of specimens (in parentheses), tributary, and locality. Type material is designated only from the

Savannah River drainage. Collection data pertaining to the intergrades are listed separately on

page 35.
NUMBER 9 2

SPECIMENS 70-79m m S.L.

Itptocaphalus (Cap* Fear to Potomac)

l«ptoc«phalut

l«ptoc«phalut (Sant**)

inttrocularlt

44

16

104

99

31

btllicus

SPECIMENS 90mm S.L. and OVER

toptoctphalus (Cap* F*ar to Potomac)

242

17

ltptoc»phalu» (P«* D*t)

129

l*ptoc*phalu* (Santo*)

inttrocularis

219

43

beliicus

1 2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 16 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33

NUMBER OF TUBERCLES

FIGURE 2.—A comparison of the number of cephalic tubercles in two size groups of five populations of N. leptocephalus. The graph is based on data presented in Table 11. The following

statistics are illustrated: the small vertical line, mean; horizontal line, range; clear bar, mean HK

one standard deviation; dark bar, mean ± two standard errors (After Hubbs and Perlmutter,

1942, American Naturalist, volume 76, pages 582-592). The magnitude of difference among the

populations is not nearly as great in the 70-79 mm size group as in the size groups 90 mm SL

and over.

Nocomis leptocephalus (Girard)

The description, ecology, distribution, dispersal,

and evolution of N. leptocephalus were treated by

Lachner and Jenkins (1971) and compared with the

micropogon group, because the several species are

sympatric in the central Appalachians. Only the

descriptive data which show some measure of divergence and of possible intraspecific systematic

value or of evolutionary significance are presented

below.

In our preliminary analysis when we had but a
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

few mature specimens of the N. I. interocularisbellicus forms and when we had not yet examined

all museum collections of N. leptocephalus, we

presumed that the morphological differences were

of a clinal nature. The two major characters, number of tubercles and the extent of the posterior

distribution on the head, appeared to have higher

values and to extend farther back on the head in

the northern river populations, while there were

progressively lower numbers and a reduction in

extent of posterior distribution toward the south

and southwestern portion of the range. This concept was abandoned when collections from all the

major rivers became available and analysis of the

data was made. Two major breaks are discernible

in the data of Tables 6 through 9 and 11—one

between the Santee and Savannah drainages and

the second between the Chattahoochee and the

Alabama rivers systems. The three major populations separated by these drainages are within themselves homogeneous in respect to cephalic tubercle

numbers and distribution, especially the two southern forms, N. I. interocularis and N. I. bellicus. The

northern form, N. I. leptocephalus, diverged significantly in tubercle numbers in both the Pee Dee

and Santee drainages, when compared with drainages farther northward and across the Appalachian

Divide into the New River system. This divergence

was also another factor for dropping the clinal

theory, because the higher tubercle values occurred

in drainages where lower values would be expected

for proper clinal relationships.

The comparatively small population of N. leptocephalus in the Edisto River drainage shows intermediacy in characters between N. I. leptocephalus

and N. I. interocularis. This small river, located

below the fall line, at one time may have provided a more abundant habitat for the bluehead

chub. The number of tubercles of mature Edisto

specimens is nearly intermediate between that of

specimens of the adjacent Savannah and Santee

populations, but the posterior distribution on the

head is fairly extensive, approaching that of N. I.

leptocephalus.

The taxonomic rank that we propose for these

populations is influenced by the magnitude of

the two major character breaks as well as the intermediate nature of the Edisto population. We

recognize three highly diverged subspecies: N. I.

leptocephalus; N. I. interocularis, new subspecies;

and N. I. bellicus. The Edisto population is considered to be an intergrade of N. I. interocularis X

N. 1. leptocephalus. All nuptial males of N. I. bellicus are distinct from N. I. interocularis. Only a few

nuptial specimens of the latter form show overlap

with the typical form; however, since the distinguishing characters are so greatly influenced by

allometric growth, it is practically impossible to

demonstrate this divergence when attempting to

identify a few specimens. The "polytypic species"

concept associates evolution with differentiation of

these forms. The exact status of N. leptocephalus

will remain in doubt until we have experimental

data. The reproductive behavior involves elaborate

nest construction and requires a comparatively spacious reproductive area, thus indoor breeding and

rearing studies may be impossible. Since we now

have a great amount of information on reproduction, reproductive ecology, breeding behavior, and

hybridization in N. leptocephalus (as well as most

of the other species of Nocomis), it would be more

feasible to conduct breeding experiments in selected, small natural streams. It would be interesting and easy to place together stocks of the two

most divergent forms, N. /. bellicus and N. I.

leptocephalus.

A comparison of the major characters of the

three subspecies is given in Table 15.

Nocomis leptocephalus interocularis, new subspecies

PLATES 1, 2, 4, 5; FICURE 3

Ceratichthys biguttatus.—Jordan and Brayton, 1878: 38, 43.

Nocomis leptocephalus—Fowler, 1945: 26 [part], 168, 232

[parti.

ETYMOLOGY.—The subspecific name, interocularis, refers to the characteristic occurrence of tubercles posteriorly to the interorbital area of the

head.

HOLOTYPE— USNM 200757. North Carolina,

Transylvania Co., Toxaway R. just below Lake

Toxaway Dam. Caught on hook and line. 15 May

1966.

SPECIMENS STUDIED.—A total of 4,083 specimens

from 4 drainages, as shown in the tabulation on

page 2, were studied. For a listing by localities, see

Appendix, pages 30-34.
NUMBER 9 2

DIAGNOSIS.—Tuberculation intermediate between

N. I. leptocephalus and N. I. bellicus; total number

of head tubercles averages 8, ranges from 7 to 9 in

95 percent of adults and subadults, and does not

exceed 14 in study material; tubercle distribution

on head not extending posteriorly beyond anterior

occipital area, almost always before mid-interorbital area.

DESCRIPTION AND COMPARISON.—Tuberculation.

Body length at first tubercle appearance. The

light tubercle spots are first visible in juvenile specimens of varying sizes. Young-of-year specimens do

not show visible spots. A few specimens of N. I.

interocularis develop one or two spots at 30 to

39 mm. At 40 to 49 mm, 30 of 56 specimens had

developed from one to five spots. A few females,

80 to 109 mm, had no visible spots. In the northern

population of N. I. leptocephalus (Cape Fear River

and northward), only a few specimens in the 30 to

39 mm size group had one or two spots, but in the

40 to 49 mm group 67 percent had one to seven

spots; almost all specimens above 50 mm had spots.

The spots first appear in the Santee and Pee Dee

populations at sizes similar to the northern N. I.

leptocephalus. The spots appear in N. I. bellicus at

somewhat larger sizes; none were observed in the

30 to 39 mm and 40 to 49 mm size groups. At 50

to 59 mm, only 14 percent had one or two spots; at

60 to 69 mm, 76 percent had one to five spots,

nearly the full compliment. All but one specimen

examined above 70 mm had spots.

Development of tubercles. The anterior orbital

spots usually develop first at about 40 mm in N. I.

interocularis, followed by the internasal to postinternasal spots at about 50 to 60 mm; the two

postorbital spots appear last, at 70 to 80 mm. The

development is somewhat similar in the two other

subspecies, except that N. I. leptocephalus often

develops one or two more supraorbital tubercles on

each side, and N. I. bellicus may lack the posterior

supraorbital tubercle. A number of specimens in

the Pee Dee population at 70 to 80 mm lack supraorbital tubercles.

Time of budding. Small tubercles or buds may

appear in specimens captured in any month of the

year, as in the micropogon species group. Development is associated with maturity, and the variable

and more rapid rate of growth of some of the males

enhances tubercle growth. A few males may have

nearly completely developed tubercles in the

months of October and November, and slight head

swellings may also appear. A 143 mm male from

the Santee River drainage had small tubercles

when captured in mid-August. No differences in

time of budding among the three subspecies was

observed, but we lack comparative monthly samples.

Crowding of tubercles at the IN and PJN areas.

The tubercles develop very uniformly in the internasal and posterior-internasal area of N. I. interocularis (Plate 5, middle, and Figure 3); in N. I.

bellicus the development is the same except that in

almost all specimens the PIN tubercles are absent,

leaving only a pair of tubercles in the IN area

(Plate 5, lower). There is slight crowding of tubercles in the northern population of N. I. leptocephalus (Plate 6, upper), but there is a characteristic and considerable crowding in the Pee Dee and

Santee populations (Plate 5, upper, and Plate 6,

FIGURE 3.—Lateral and dorsal view of tuberculate head of

male X. I. interocularis, with small crest, 154 mm SL, captured 31 March 1949 from the Altamaha River drainage.

(USNM 161701. Drawn by A. M. Awl.)
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lower). The increased tuberculation of these populations is not spread uniformly over the head, but

occurs in the internasal-interocular area. The occipital tuberculation is less crowded, having only a

few tubercles more than the same area in specimens

of the northern population. Some tubercles occasionally appear just anterior to the nares in the

Pee Dee River population.

Supernumerary nape tubercles. Small spots and,

rarely, small buds occur on the nape just before the

scaled area; usually a pair is present (Figure 3,

bottom), but sometimes there is only one. They

occur most often in N. I. interocularis (Savannah

River, 5 specimens; Altamaha, 1 specimen) and the

Santee population (6 specimens) of N. I. leptocephalus. Only one specimen in over 9,000 of the

northern population of N. I. leptocephalus examined had such spots; none were observed in N. I.

bellicus. These spots may be the remnants of a

once fuller and more extensive tuberculation in

the precursors of the present N. leptocephalus. We

have not seen similar spots in other species of

Nocomis where they could be easily observed, as in

the micropogon group.

Rate of increase of tubercles. The general increase in tubercle numbers with increase in body

length for five populations of N. leptocephalus is

illustrated graphically in Figure 1. The slopes of

the curves level off at about 100 mm for N. I. bellicus, N. I. interocularis, and the northern population of N. I. leptocephalus. The curves for the

Santee-Pee Dee populations level off at about

120 mm. Neither maximum body size nor growth

rate is a factor in tubercle growth rate. The largest

bluehead chubs that have been collected are seven

male N. I. interocularis from the Savannah River

drainage (Toxaway River above Lake Toxaway),

all of which exceed 200 mm SL and have the normal number of tubercles for that subspecies. At

about 50 mm, when N. I. bellicus commences to

have 1 visible tubercle spot, N. I. interocularis

averages about 2, and the Pee Dee population of

N. I. leptocephalus averages about 6. At the 100-

mm size group, N. I. bellicus averages about 5

tubercles, N. I. interocularis about 8, the northern

population of N. I. leptocephalus about 17, the

Santee about 18, and the Pee Dee over 20. The rate

of tubercle increase with increase in body length is

given for N. 1. interocularis in Table 4; N. I. bellicus, Table 5; the northern race of N. I. leptocephalus in Table 1; and the Pee Dee-Santee race

in Tables 2 and 3.

Number of tubercles. The total number of

cephalic tubercles is a primary character in the

separation of the three subspecies. N. I. interocularis is the intermediate form, averaging about 8 in

the adult-subadult specimens, ranging from about

7 to 9 in 95 percent of the study material and never

exceeding 14; N. I. bellicus averages about 5, with

4 to 6 in 95 percent of the study sample, and the

maximum number does not exceed 7; N. I. leptocephalus averages 17 or more in the races, ranging

from 16 to 25, and the maximum number was 33.

The comparative uniformity in the total number

of tubercles present in N. I. interocularis and N. I.

bellicus by river drainages is shown in Tables 6, 7,

and 8. The higher maximum numbers in the Pee

Dee-Santee race compared to the northern race of

N. I. leptocephalus is also shown in these Tables.

The three subspecies show a significant differentiation in the total tubercle count at the 70 to

79 mm size group and more so at 90 mm and over

(see Figure 2 and Table 11). The differences in the

means of the Pee Dee and Santee populations may

be due to the small Pee Dee sample and, also, that

the lower values of the Congaree and Saluda rivers,

Santee drainage, were lumped with the CatawbaBroad rivers specimens (Table 6). The data in

Table 8 show the high number of tubercles in the

internasal-posterior internasal area for the Santee

and Pee Dee drainages compared with the lower

values for N. 1. interocularis and N. I. bellicus. The

crowding is somewhat greater in the Catawba

River, Santee drainage, than in the Pee Dee drainage, and more so in the Catawba than the BroadCongaree-Saluda rivers.

The largest N. I. interocularis females were in

the 140 to 149 mm and 150 to 159 mm size groups

and these three specimens had an average tubercle

complement of eight. Some specimens, much

smaller than these, in the 70 to 79 mm and 80 to

89 mm groups had more tubercle spots (Table 4).

There is considerable variation in tubercle numbers within size groups, the greatest occurring

among the juvenile sizes (Tables 1 to 5). Both sexes

showed this variation. There were only small tubercle differences between the sexes in most size

groups among the three subspecies, the males aver-
NUMBER 9 2

aging slightly higher numbers. Greater tubercle

differences occurred between the sexes in the Santee population (Table 3). The males in all subspecies attain the larger size, as shown in Tables 1

to 5. This is also true for all other species of

Nocomis.

Occurrence of head tubercles. The progressive

development of tubercles posteriorly on the head

is shown in Table 10 for N. I. leptocephalus from

the Santee River drainage. While the general trend

is an increase in posterior occurrence of tubercles

with an increase in body size, specimens of both

sexes in the juvenile or subadult size group of 70

to 79 mm have the definitive tubercle distribution

in the posterior occipital area. All drainage populations of N. I. leptocephalus over 60 mm (Table 9)

have the majority of specimens with tubercles

(buds or spots) already present in the mid-occipital

or post-occipital areas. The occurrence of head

tubercles in particular areas is one of the best

characters for separation of the subspecies (Tables

9 and 15). All nuptial males of the three subspecies

can be distinguished by this character, no overlap

in tubercle distribution occurring. Specific rank for

these populations was not accorded because of the

intergrading Edisto population. Table 9 shows that

a small juvenile of N. I. leptocephalus just showing

tubercle spots has a tubercle pattern similar to an

adult N. I. bellicus. Tubercle distribution, as in

tubercle numbers, is comparatively homogeneous

within each subspecies. The evolution of the tuberculation has resulted in major character breaks in

which three highly differentiated subspecies can be

recognized.

Nuptial crests. A discussion of the nuptial

crests in Nocomis is given by Lachner and Jenkins

(1971). The three subspecies of N. leptocephalus

appear to develop different sized crests (Table 15);

N. I. inter ocular is (Plate 2) has a crest intermediate in size between the smaller one of N. I. bellicus

(Plate 7) and the larger one of N. I. leptocephalus

(Plate 3). We lack sufficient comparative material

to accurately determine this possible differential

development.

Vertebral Numbers. The total number of vertebrae ranged from 38 to 41 in the three subspecies

(Table 12). N. I. interocularis had 40 vertebrae in

86 percent of counts and N. I. bellicus had 39 or

fewer vertebrae in 88 percent of counts. The northern river drainages of N. I. leptocephalus had

higher values, approaching 40, compared to those

south of the James which had modal values of 39.

The Edisto intergrades showed a good intermediate

trend (39.4) between those of N. I. interocularis

(about 40) and those of the southern rivers of N. I.

leptocephalus (about 39).

Dentition. The number of pharyngeal teeth in

all three subspecies (Table 13) is almost always 4-4

(83 specimens); five specimens had 4-3 or 3-4;

three had 3-3 ;and one had 2-4. Most of the tooth

reduction occurred in N. I. leptocephalus. (See

Lachner and Jenkins, 1971, for discussion of evolutionary significance of tooth reduction in Nocomis).

Squamation. Initial studies suggested possible

scale differences among the three subspecies. When

samples were increased and selected from many

localities within and among river drainages, the

results showed only small differences which were

of no use in identifying subspecies. Mainly, the

mode shifted among the drainages involving one or

sometimes two scales in the circumferential count

and less in the lateral-line scale count. The difference in the circumferential count is a reflection of

the degree of crowding of the belly scales. Lateralline and circumferential scale counts are summarized in Table 14. In N. I. interocularis the Chattahoochee drainage specimens have somewhat lower

values than those of the Savannah-Altamaha, and

they are very similar to counts for N. I. bellicus.

The scale counts for N. I. leptocephalus were summarized chiefly from data used by Lachner and

Jenkins (1971) in comparing scale differences in the

four central Appalachian species of Nocomis.

Intestine. The whorled intestine characteristic

of N. I. leptocephalus (for description and illustration, see Lachner and Jenkins 1971) does not hold

up as a systematic character, mainly in N. I. interocularis of the Chattahoochee River drainage. A

highly or moderately whorled intestine is almost

always found in N. leptocephalus over its range.

A slightly whorled intestine is sometimes found in

specimens from the Cape Fear, Saluda, and Broad

rivers. The typical whorled condition is also found

in the Savannah and Altamaha specimens of N. I.

interocularis and over the range of N. I. bellicus,

except for one collection. Over 200 specimens of

N. I. interocularis in eight collections from the

Chattahoochee River had straight intestines; other
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

specimens from these and other collections in the

Chattahoochee had whorled intestines. The straight

intestine condition resembles that of the micropogon group. The specimens are normal otherwise.

One specimen of N. I. bellicus from a tributary of

the Tallapoosa River, Alabama drainage, had a

straight intestine. We associate no systematic trend

to the occurrence of the straight intestine among

these collections. It is a primitive condition in

Nocomis, and the Chattahoochee specimens may

represent merely reversions.

Nocomis leptocephalus leptocephalus (Girard)

PLATES 3-6

Ceratichthys leptocephalus Girard, 1856: 213 [type locality,

Salem, North Carolina, probably Winston-Salem, from a

tributary of Yadkin River, Pee Dec River drainage].

Nocomis leptoccpha.lvs.—Lachner and Jenkins, 1971: 50.

SPECIMENS STUDIED.—A total of 14,551 specimens

from twelve drainages, as shown in the tabulation

on page 2, were studied. For a listing by localities,

see Appendix, pages 34-35.

Nocomis leptocephalus bellicus Girard

PLATES 4, 5, 7

Nocomis bellicus Girard, 1856, p. 213, [type locality, Black

Warrior River, Alabama, Mobile Bay drainage].

Ceratichthys biguttatus.—Hay, 1881:512.

Hybopsis kentuckiensis.—Gi\ben, 1891:154.

Hybopsis bellica.—Cook, 1959:132.

Nocomis leptocephalus.—Fowler, 1945:26 [part].

Nocomis leptocephalus bellicus.—Smith-Vaniz, 1968:42.

SPECIMENS STUDIED.—A total of 2,000 specimens

from five drainages, as shown in the tabulation on

page 2, were studied. For a listing by localities, see

localities, see Appendix, page 35.

Nocomis leptocephalus intergrade

interocularis x leptocephalus

SPECIMENS STUDIED.—A total of 155 specimens

from the Edisto drainage, as shown in the tabulation on page 2, were studied. For a listing by

localities, see Appendix, page 35.

Comparison. The development and occurrence

of head tubercles of the Edisto River population is

intermediate between N. I. interocularis and N. I.

leptocephalus. The four adult specimens (Table

7) had 16 tubercles, a value higher than the mean

(8) and maximum number (14) for N. I. interocularis but lower than the mean (17 in northern race,

18 in Santee, 23 in Pee Dee) and much lower than

the maximum number (33) in N. I. leptocephalus.

Further, intermediacy in tubercle numbers of the

Edisto population compared with the above two

subspecies is shown in a summary of all material

that could be counted (Table 6), and in specimens

over 65 mm SL, excluding the supraorbital tubercles, in Table 8. The intermediacy of the occurrence of tubercles posteriorly on the head in juveniles and adults (specimens over 60 mm SL) of

the Edisto population is compared with the above

two subspecies in Table 9. The mode for the

Edisto population occurs on the AOC line, a fairly

intermediate position between the AIO area, usual

for N. I. interocularis, and the MOC-POC area,

usual for N. I. leptocephalus.

The vertebral numbers of 39 (17 specimens), 40

(10), and 41 (1) of the Edisto population are additional intermediate values between the adjacent,

southern drainage populations of N. I. leptocephalus (about 39 vertebrae) and N. I. interocularis (about 40 vertebrae).

The adult and subadult specimens from the

Edisto drainage are insufficient to compare this

population more precisely with the adjacent populations. The above comparisons of three characters show intermediate values for the Edisto

population that are not found among the three

drainages (Savannah, Altamaha, and Chattahoochee) in which N. I. interocularis occurs nor in

drainages from the Santee northward inhabited by

JV. /. leptocephalus. We presume that the Edisto

population has received N. leptocephalus stocks

from both north and south, after these two stocks

had diverged into N. I. leptocephalus and N. I.

interocularis. Thus, the last entry of a N. leptocephalus stock into the Edisto was probably comparatively recent.

Coloration

The general coloration of N. I. leptocephalus is discussed and compared with the other species of

Nocomis inhabiting the central Appalachian area
NUMBER 9 2 11

by Lachner and Jenkins (1971). Also described are

two different primary color forms of N. I. leptocephalus among the living nuptial males. A form

with a brassy orange-olive coloration on the body

laterally and sometimes forming a stripe occurs in

the northern race from the Cape Fear drainage

northward. Another form with a bluish body

occurs in the Pee Dee-Santee drainages.

The blue form also occurs in N. I. interocularis

of the Savannah River drainage. Following is a description of a prenuptial male, 187 mm SL, collected on 15 May 1966 in Mill Creek, a tributary

of Lake Toxaway, Savannah River drainage

(USNM 200757). This coloration is characteristic

of the Savannah drainage population. The specimen was captured with three females from beneath

a cut bank near a nest. Its nuchal crest was not

fully developed and the head tubercles were normally distributed, but were not enlarged as in some

males. The nuchal crest was brown, the rest of the

head slate-blue; eye, yellow-brown; opercular membrane, dark gray; dorsum of the body, brown with

a few golden highlights on the postdorsal area; posterior margins of the scales dorsally on the sides

above the lateral line, black; lateral band, diffuse,

a greenish gray or olivaceous color; belly, bluish

white; dorsal fin, slate-blue base with dark gray

rays, the posterior having a yellow tinge, and the

fin membrane, smoky; pectoral fins, smoky yellow,

the rays grayish with the first ray darker; pelvics

and anal fins, yellow with a gray cast; caudal fin,

a burnt orange, the median rays darkest. A large

gravid female in the same collection had the

nuchal area and dorsum, brown, with a faint,

light, predorsal stripe; lateral band, black with a

narrow, diffuse, light brown stripe above and extended anteriorly and prominently onto the postocular area with a faint bar continuing to below

the nostrils; opercular membrane, dark gray; body

below the lateral band, yellowish; belly, white;

dorsal and caudal fins, rust-orange; the other fins,

a brownish yellow.

The following coloration was recorded from a

living tuberculate, prenuptial male, 144 mm SL,

captured in Amy's Creek, Habersham County,

Georgia, Chattahoochee River drainage, on 2 April

1970 by Lachner and Wiley: dorsal fin, light

pinkish orange, melanophores along base of dorsal

fin; caudal fin, olive-orange, lighter near the base

with melanophores along basal portion of rays;

pectoral fin, deep pink-orange; pelvic fin, deep

pink-orange with some white on first ray; anal fin,

pink-orange, the outer edge lighter; iris, red; body

above lateral line grayish with slight bronze; ventral part of head, light reddish; dorsal part of head,

slightly tan to pale olive; belly white to milky. The

head tubercles were moderately developed, but the

head was only slightly swollen. A gravid female,

119 mm SL, captured with the above male, had the

same general color pattern in life except that the

fins were very light, showing only traces of color,

and the body was pale.

Other color data on adult N. I. interocularis are

from a color slide of a male, 164 mm SL, collected

by E. C. Raney in the Chattahoochee River drainage, Georgia. This tuberculate, prenuptial male

had light orange fins, the rays of the dorsal and

caudal having a darker burnt orange color. Good

comparative color data of nuptial males from the

Altamaha drainage are not available.

Observations by the authors and others indicate

that the breeding coloration of N. I. bellicus in the

Alabama River system of the Mobile Bay drainage,

Alabama, is essentially the same as the northern

color form of N. I. leptocephalus, e.g., with the

blue head and the lateral orange stripe anteriorly

on the body changing to yellow caudally. A color

description of a large, prenuptial male N. I. belliens with large tubercles, was recorded from a fresh

specimen captured on 16 March 1957 in the Alabama River system, Elmore County, Alabama, by

R. D. Suttkus (RDS 2592). The sides of the head

were bluish gray; head, dorsally, and entire nape

posteriorly to origin of dorsal fin and down on the

sides to lateral line, very brassy orange; rest of

back and upper sides, bright olive; iris, bright

orange-red; caudal fin, olive; leading edge of anal

and pelvics, milky; remainder of anal, pelvics and

pectorals, olive; dark bar on cleithrum immediately

behind opercle. Smaller specimens in same collection have bright orange caudal fin, orangy dorsal

fin, and bright orange iris.

A color description of a tuberculate postnuptial

male, captured over a nest on 1 July 1965 by R. D.

Suttkus (RDS 3717) in the Alabama River system,

Clarke County, was recorded in the field as follows:

head below lower margin of tubercles, below eye,

and below top of opercular opening, pale blue; top
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of head, golden brown; iris, bright red-orange; back

and upper sides, brown with golden olive overlay;

center of scales, golden, especially anteriorly on

body and more yellow posteriorly on body; predorsal stripe, golden and broad, one scale row in

width; postdorsal stripe tapering posteriorly and

more yellowish than predorsal stripe; lower sides

of body and belly, pale; pectoral fin, yellowish, the

anterior one fourth bright yellow, the color pronounced on anterior rays; pelvic fins, yellowish,

more so on anterior rays; dorsal fin, mostly olive,

the base bluish, and some yellow on leading edge;

caudal fin mostly yellowish olive; and anal fin,

yellowish anteriorly, with branching areas of rays

also yellowish as in pelvic fins.

A subadult N. I. bellicus, 107 mm SL, and a

young specimen, 36 mm SL, taken by H. T. Boschung in a tributary of the Cahaba River system,

Bibb County, 22 August 1967 (UAIC 2629-1), preserved in 10 percent formalin and ional solution,

still retained lifelike colors in January 1968 (H. T.

Boschung, personal communication). The larger

specimen was colored as follows: body dorsally,

dark olive above lateral line, and light olive from

lateral line to about two scale rows below; lower

body, light to pale; head, dark olive dorsally, light

olive laterally, and light blue ventrally; body with

no evident lateral band; dorsal fin, bright orange

with some olive near base; caudal fin, bright

orange; pectoral fins, light olive; pelvic fins, very

light olive, with light posterior margins; and anal

fin, light olive, with touches of orange along

middle portion of rays, and the outer margin, light.

The young specimen had the body light brown to

dusky above the pronounced dark lateral band and

light to dusky below the lateral band; dorsal fin,

light orange, the outer margin light to slightly

dusky; caudal fin, medium orange, the outer rays

light to slightly dusky; and pectorals, pelvics, and

anal fins, light to only slightly dusky.

Throughout its range, N. leptocephalus is commonly observed to have variable amounts of orange

or reddish coloration in the dorsal and caudal fins

(and, frequently, in the other fins) in young, juvenile, subadult male and female, and mature female stages.

There is great difficulty in evaluating color differences (aside from those associated with individual variations and populational differences),

because of the gradual color changes related to

growth and maturity. Also, among the nuptial

males, the changes occur throughout the reproductive season and "peak" colors develop immediately

at the time of spawning. The characteristic coloration of the nuptial males compared to that of nonnuptial males and all females is discussed by

Lachner and Jenkins (1971). The elaborately developed nuptial coloration common to Nocomis

biguttatus, N. effusus, N. micropogon, N. raneyi,

and N. platyrhynchus is found in N. leptocephalus,

except that the problem of interpreting the systematic significance of the coloration is compounded

in the latter species by the presence of two basically

different nuptial color patterns. The blue body

form of N. I. leptocephalus of the Pee Dee and

Santee river drainages is trenchantly different from

the brassy reddish colored northern race of N. I.

leptocephalus (in all drainages northward from the

Cape Fear). These two color forms within N. I.

leptocephalus correlate with the tubercle differences noted above, i.e., the northern race has fewer

tubercles and is brassy reddish while the southern

race has more tubercles and is blue. A similar blue

color form appears to be typical for N. I. interocularis. We are not certain if the nuptial males of the

Altamaha drainage develop the blue coloration

typical of N. I. interocu laris of the Savannah drainage. The orange-olive color form reappears in the

Alabama drainage and westward in N. I. bellicus.

In this subspecies, the nuptial males appear to have

brighter orange-red colors in the fins than the

northern race of N. I. leptocephalus. Most significant is the fact that great differentiation in nuptial

coloration has occurred within N. I. leptocephalus,

and probably N. I. interocularis and N. I. bellicus

are also completely different in nuptial coloration.

Geographic Distribution and Ecology

The extensive distribution of N. leptocephalus

along the Atlantic and Gulf Coast drainages is

shown in Figure 4. Collections of the three subspecies and the Edisto intergrades are individually

plotted and their distributions are shown in respect

to the fall line and the Appalachian Divide. The

distribution of N. I. leptocephalus is discussed by

Lachner and Jenkins (1971) in detail where the

northern race occurs sympatrically with three other
NUMBER 9 2 13

FIGURE 4.— The distribution of N. leptocephalus. The three subspecies are plotted with separate

symbols. The intergrades, N. 1. interocularis x N. I. leptocephalus, in the Edisto drainage are

indicated by a single star, but several collections are actually represented by this plot. When two

or more collections were made at (or near) a locality, only one was plotted. The single plot in

northern Alabama of N. I. bellicus in the Tennessee drainage also represents several collections.

The two squares representing N. leptocephalus in southeast Tennessee, Tennessee drainage, are

based on small specimens and could not be identified to subspecies with assurance. Data for plots

arc listed in the text, and those that merit special attention are discussed. The Appalachian

Divide and the fall line are indicated.

species of Nocomis in certain river drainages. N. I.

leptocephalus is widely distributed in the James

drainage, and in limited headwater localities of the

Rappahannock and Potomac drainages, where it

occurs sympatrically with N. micropogon. Across

the divide in the middle and upper New River

system, its distribution overlaps considerably with

N. platyrhynchus. In portions of die James,

Chowan, Roanoke, Tar, and Neuse drainages, it is

more or less commonly associated with N. raneyi.

N. I. interocularis is widely distributed in the

Savannah and Altamaha drainages, occurring far

below the fall line. All of our records occur above

the fall line in the Chattahoochee system. N. mi-
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cropogon is associated with N. I. interocularis in the

upper Savannah drainage, where the former was

probably derived by stream capture from the Little

Tennessee River, Tennessee drainage. The presence

of AT. leptocephalus in a restricted area of the

Tennessee drainage of North Carolina (probably

N. I. interocularis) may be associated with stream

capture with the Savannah drainage (see discussion

by Lachner and Jenkins, 1971). We have examined

two collections from this area, totaling eight specimens, all of which were young or juveniles (TU

29663, six specimens, Cedar Creek, tributary to Lake

Thorpe, 4.4 mi north of Cashiers. Jackson County;

TU 28001, two specimens. Little Tennessee River,

Cullasaja River, above Cullasaja Falls, 4.3 mi NW

of Highlands, Macon County). Here, N. leptocephalus is again sympatric with the widely distributed N. micropogon of the Tennessee drainage.

The one plot of AT. /. interocularis in the upper

Coosa River system is based on a typical male

specimen, 118 mm SL, captured by R. D. Suttkus

in Cartecay River, 7.8 mi SE of Ellijay, Gilmer

County, Georgia, on 18 June 1965 (TU 38213).

William F. Smith-Vaniz (personal communication)

informs us that there is considerable bait fishing in

the impounded waters of the area. Populations of

N. I. interocularis in the nearby Chattahoochee

drainage are a ready source of bait. Several species

of fishes, including Nocotnis (Figure 4), are apparently prevented from entry into the Coosa

River system by the falls at the Fall Line (SmithVaniz, 1968:124, 130).

The southwestern subspecies, N. I. bellicus, finding suitable habitat below the fall line, occurs

from the Alabama River system westward to certain tributaries flowing into the Mississippi River

from the east. It also is common in a restricted area

of the Tennessee drainage, where B. R. Wall, Jr.,

(personal communication) has obtained collections

in more than twenty localities of the Bear Creek

system (indicated in Figure 4 by one plot mark).

Smith-Vaniz (1968:124) cites headwater stream capture in the area involving the Tennessee and the

Tombigbee rivers systems and lists four species other

than Ar. /. bellicus that also made entry into the

Tennessee River from the Tombigbee River system. The presence of AT

. wicropogon as well as

many other species, in the Coosawatee system,

Coosa-AIabama drainage, is probably the result of

a stream capture with the Hiwassee system of the

Tennessee drainage.

The ecology of N. leptocephalus is reviewed by

Lachner and Jenkins (1971). Over its large range,

the bluehead chub is one of the most abundant

species, preferring small stream habitats of about

10 to 50 feet in width and avoiding the larger

streams and rivers as well as the extreme headwaters. Clear streams of gravel-rubble bottom composition, particularly current-swept areas, are preferred and necessary for these gravel-mound nestbuilding chubs (Plate 8). Slack waters are avoided,

although the species thrives and is abundant in

typical Piedmont streams where lower gradients,

sandy bottoms, shifting sands, and some turbidity

are characteristic of the habitat, but in these areas

some gravel-rubble riffles also occur. The rarity of

the bluehead chub along the middle-Atlantic

Coastal Plain is probably related to the general

absence of riffles or exposed gravel and rubble;

however, these ecological conditions also prevail

below the fall line of the southern Gulf Coastal

area. The widespread distribution of the chub is

undoubtedly related to its ability to survive Piedmont and certain Coastal Plain conditions. It probably was able to enter new drainages by way of the

mouths of adjacent streams during intermittent

periods of sea-level recessions, as well as to enter

new drainages by stream capture, because of its

ability to survive in relatively small, headwater

streams.

Evolution and Dispersal

Lachner and Jenkins (1971) present a detailed account of routes of dispersal in the central Appalachian area.

The enlargement and loss of tubercles probably

represents an advanced condition in Nocomis. The

adaptive significance of large, stout, nuptial tubercles is not known. They would serve as effective

weapons in agonistic behavior but nuptial males of

N. leptocephalus, unlike some other species of

Nocomis that have smaller, more numerous tubercles on the head, have not been observed to engage

in either prolonged or violent fighting or both

during the nest-building activities of reproduction.

It may be that the large, easily visible tubercles
NUMBER 9 2 15

PLATE 1.—Lateral view of tuberculate male N. I. inter ocular is, 154 mm SL, from the Altamaha

drainage. (Specimen same as in Figure 3, USNM 161701. Drawn by Carolyn Bartlett Gast.)

PLATE 2.—Lateral view of nuptial male N. I. inter ocular is, showing cephalic tuberculation and

anterior position of moderately developed crest. Photograph of a paratype, USNM 200176,

156 mm SL from the Savannah River drainage, captured 8 July 1957.

PLATE 3.—Lateral view of a tuberculate, highly crested male N. I. leptocephalus, 196 mm SL,

captured 16 May 1966, from the Santee River drainage. This specimen (USNM 200758) illustrates

the crowding of tubercles (numbering 29) in the Santee-Pee Dee populations.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 4.—Lateral head profiles comparing the tuberculate

males of three subspecies of N. leptocephalus: upper, N. I.

leptocephalus, USNM 200758, Santee drainage, 196 mm SL,

captured 16 May 1966 (29 tubercles); middle, N. I. interocularis, USNM 200176, Savannah drainage, 156 mm SL, captured 8 July 1957 (8 tubercles); and lower, N. I. bellicus,

USNM 200756, Mobile Bay drainage, 145 mm SL, captured

3 May 1966 (5 tubercles).

PLATE 5.—Dorsal head profiles comparing the tuberculate

males of three subspecies of N. leptocephalus (data as in

Plate 4): upper, N. I. leptocephalus; middle, N. I. interocularis; and lower, N. I. bellicus.
NUMBER 9 2 17

PLATE 6.—Dorso-lateral and dorsal profiles of heads of male

N. I. leptocephalus representing typical tuberculation patterns of the northern (upper photograph) and southern

(lower) races. Upper specimen is from Brush Creek, New

River system, Montgomery Co., Virginia, captured June 1951,

140 mm SL with 17 tubercles ((photograph courtesy of Edward

C. Raney, field number 2027). Note large tubercles on anterior head and very little crowding compared with specimen

shown below. Lower specimen is from a tributary of

the Catawba River, Santee drainage, McDowell Co., North

Carolina, captured 1 May 1966, 157 mm SL with 31 tubercles.

PLATE 7.—Lateral view of a typical tuberculate male N. I. bellicus, with a small crest, 145 mm

SL, captured 3 May 1966, from a Tallapoosa River tributary, Mobile Bay drainage (USNM

200756).
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATE 8.—Gravel mound-nests of N. I. leptocephalus: upper, large nest, over three feet in width,

observed in a tributary of the Saluda River, Santee drainage, on 16 May 1966. A large tuberculate

male, 196 mm SL (USNM 200758) was captured nearby; and lower, typical, small nest of the

bluehead chub in Lapsley Run, James drainage, observed during May and early June 1968 and

1969. The larger males usually build the larger nests.
NUMBER 9 2 19

serve as recognition signals to the female (or they

may have some other behavioral significance).

The evolution of the leptocephalus complex is

not clear with respect to existing data on morphology and distribution. This species complex, although

distinct from the micropogon and biguttatus

groups, is most closely related to a N. micropogon

stock; however, its evolution and dispersal cannot

be related to that of the micropogon group. The

basic N. leptocephalus stock became isolated on the

present Atlantic-Gulf slope probably during the

Teays Period. It appears most likely that a N.

leptocephalus stock existed, at first, in the upper

Santee-Pee Dee drainage area. This stock could

have evolved from a N. platyrhynchus type from

the Old Teays River (now the upper New River).

The high numbers of tubercles, their crowding in

the forepart of the head before the eyes, and their

occurrence from the internasal to the occipital area

are characters of N. I. leptocephalus of the SanteePee Dee that are also common to N. platyrhynchus.

This southern N. I. leptocephalus stock found dispersal routes to the northeast on the Atlantic slope

and westward on the gulf slope at different times

and rates via stream capture and by the confluence

of fresh waters at the mouths of rivers during

Pleistocene sea-level fluctuations.

The westward extension of the N. leptocephalus

stock ended abruptly at the Mississippi River. It is

not known to occur farther westward. Following

the dispersal and establishment of N. leptocephalus

stock on the gulf slope, a considerable period of

isolation of the Savannah-Chattahoochee stock from

that of the Alabama and more western rivers must

have been necessary in order for the divergence of

N. 1. bellicus from N. I. interocularis to have

reached the present level of differentiation. The

distribution of N. I. bellicus could have been

variously effected during geological times, but its

morphological homogeneity suggests considerable

gene exchange among the several drainages it now

occupies or a recent entrance into the western areas

via the upper Mobile drainage.

The old escarpment of the Coosa at about the

fall line prevented its entrance into this system.

Its limited distribution in the Tennessee River in

northern Alabama is a well-documented, recent

stream capture involving several Mobile drainage

forms. The speciation of other fishes on the gulf

slope was reviewed by Smith-Vaniz (1968).

The homogeneity of the characters of Ar. /. interocularis suggests a past history in the SavannahChattahoochee area similar in time to that of N. I.

bellicus, mentioned above. The recent entrance of

N. micropogon into the upper Savannah via the

Tennessee River is also discussed under geographic

distribution.

The northward dispersal of N. leptocephalus

stock from the Santee-Pee Dee area must have

occurred during at least two major periods of

exchange. The early dispersal of this multituberculate, blue-bodied form could have been northward to the Roanoke-James drainages, and subsequent isolation could have permitted divergence of

the northern race toward reduced tuberculation

and a brassy orange coloration. The more recent

entry of the northern race of N. I. leptocephalus

across the Appalachian Divide into the New River

system and its capture in the James, Rappahannock, and Potomac drainages are discussed by

Lachner and Jenkins (1971).

The intermediate population in the Edisto River

suggests very recent exchanges of N. I. interocularis

from the Savannah drainage and N. I. leptocephalus from the Santee drainage. The factors

related to the development of the blue body coloration in the southern form of N. I. leptocephalus

and probably all N. I. interocularis are not understood. It is of interest that both forms of the

leptocephalus complex at opposite ends of the geographic distribution diverged similarly in respect

to the basic body coloration of the nuptial male.

Literature Cited

Bailey, R. M., E. A. Lachner, C. C. Lindsey, C. R. Robins,

P. M. Roedel, W. B. Scott, and L. P. Woods

1960. A List of Common and Scientific Names of Fishes

from the United States and Canada. American Fisheries Society, special publication number 2, 102

pages. (Second edition).

Cook, F. A.

1959. Freshwater Fishes in Mississippi. 239 pages. Mississippi Game and Fish Commission.

Fowler, H. W.

1945. A Study of the Fishes of the Southern Piedmont

and Coastal Plain. The Academy of Natural Sciences

of Philadelphia. Monograph 7, 408 pages.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Gilbert, C. H.

1891. Report of Explorations Made in Alabama during

1889, with Notes on the Fishes of the Tennessee,

Alabama, and Escambia Rivers. Bulletin of the

United States Fish Commission, number 9 (1889),

pages 143-159.

Girard, C.

1856. Researches upon the Cyprinoid Fishes Inhabiting

the Fresh Waters of the United States of America,

West of the Mississippi Valley, from Specimens in

the Museum of the Smithsonian Institution. Proceedings of the Academy of Natural Sciences of

Philadelphia, 8:165-213.

Hay, O. P.

1881. On a Collection of Fishes from Eastern Mississippi.

Proceedings of the United States National Museum,

3 (179): 488-515.

Hubbs, C. L.

1926. A Check-list of the Fishes of the Great Lakes and

Tributary Waters, with Nomenclatorial Notes and

Analytical Keys. Miscellaneous Publications, Museum of Zoology, University of Michigan, number

15, 77 pages.

Jordan, D. S., and A. W. Bray ton

1878. On the Distribution of the Fishes of the Alleghany

Region of South Carolina, Georgia, and Tennessee,

with Descriptions of New or Little Known Species.

Pages 7-95 in Contributions to North American

Ichthyology . . . HI. United States National Museum

Bulletin 12.

Lachner, E. A.

1946. Studies of the Biology of the Chubs (Genus Nocomis, Family Cyprinidae) of Northeastern United

States. 78 pages. Unpublished Cornell Universitythesis.

Lachner, E. A., and R. E. Jenkins

1971. Systematics, Distribution, and Evolution of the Chub

Genus Nocomis Girard (Pisces, Cyprinidae) of Eastern United States, with Descriptions of New Species.

Smithsonian Contributions to Zoology, (85:97).

Moore, G. A.

1968. Fishes. Part two, pages 22-165, in Vertebrates of the

United States. Second edition. New York: McGrawHill Book Company, Inc.

Smith-Vaniz, W. F.

1968. Freshwater Fishes of Alabama. 211 pages. Auburn

University (Alabama), Agriculture Experimental

Station.
NUMBE

R 9 221

It§

1

8i

t8?i l

It

<•© .JJ

5 23

zzzjii

oX 25 26 -*COsaCMO»17 18 <oaasCM

t-l-oo00-MS«CMO

eSL/oi

O -1

M> OKCM- •aX X

r» .» CM

00 r^ co

m M» o>CMMj

** in

• <o • CO e* r*

.

. CO

— a ^ *» IA m w> r* 2«CM

X XX

20-2

30-3

40-450-560-6

oCO

M)

CM• ^

>

-3

-3

fl CM

-c?

1-1 l-l

t-< CM

• J^(3)

i • • ^ "b o70-7

o

00

-1 1 «2

-*cCO CO •» 1 CM CO "2CM t-l(1) (Z)

C M.

1 1 • ^ ••b o- ON80-8

I1«

~

CO ^ "3

ft en

~.

2

CM*(4) (X)

ON90-9

ms-eoCOCO5 4

(3) (2) <X) 3(X)

• bo

109 100-

00

M)

sN3CM , CO , "> 1 ^3CM ^(X) "b o119 110-

oCO•-11-m•n•- 129 120-

oCOCOs»CMCM•n•b139 130-

.8.4 -tCO-CMCMto149 140-

co-1•- •- 1

r*% 159 150-

o

-a•••i;-• 169 160-
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

2 E

N

 3 2

OJ * ff> *

III I

2

°> f> o-

" S . «>••!. •sS " 3

S 'S -S -3

"S '3 S

C p

•H W

illS
NUMBER 9 2 23

1

m r^. eo en o

d ao oo H d

<3

E

1

2 t:

•S 1

I t>J I «M en

^f PIHHWrt

« C M <n in co in r*. I-I en >-* <f i CM<M I-I

3

8

r » H HN

S

a

c

i en CM

X X X X X "b CH- X '. • O- X

• I I I I

o o o o o

— -< i— —• CM r-i

l l • • I l

o o o o o o

o ^ ©
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 5.—The number of head tubercles in N. 1. bellicus from the drainages of Mobile Bay,

Pascagoula Bay, Pearl River, and eastern tributaries of the Mississippi River, segregated by

sex (X = not sexed; females in parentheses; mean values with sexes combined).

SL/mm

30-39

40-49

50-59

60-69

70-79

80-89

90-99

100-109

110-119

120-129

130-139

140-149

150-O59.

170-179

X

X

X

X

cf

9

X

cf

9

X

<f

9

X

cf

9

cf

9

cf

9

cf

9

cf

cf

cf

0

11

23

18

8

( 1 )

1

2

2

(2)

2

1

4

3

(2)

(4)

1

(1)

NUMBER

3

3

3

3

(1)

2

1

(1)

OF

4

15

2

(2)

10

4

(3)

11

1

(3)

3

(1

2

(2)

TUBERCLES

5

2

1

-

1

-

1

1

6

3

2

3

1

2

2

2

4

(1)

1

(1)

1

(1)

1

1

7

1

1

(1)

2

2

1

N

X

11

23

21

34

12

14

6

N

cf

13

10

5

3

6

3

4

2

1

1

N

9

7

8

4

2

4

1

1

0.0

0.0

0.2

2.6

3.7

3.8

4.3

5.0

5.3

6.5

6.2

5.5

6.0

7.0
NUMBER 9 2 25

IN — CO — 0 0

— <-< <-> F~ O <

'22'

1 S 5 2

'SS2

S 11 §8 U s

L I

III ll .28

I (I <9 V

. Z U O- C

i l : s

• 3 *- « xi

556 5 3<
TABLE 8.—The high number of internasal and posterior-internasal tubercles in

N. 1. leptocephalus from the Pee Dee and Santee drainages compared with the

lower number in N. 1. interocularis and N. 1. bellicus for specimens over 65 mm.

(Supraorbital tubercles not counted.)

SUBSPECIES

Drain-age

leptocephalus

Pee Dee

Santee, Catawba

Broad, Congaree

Saluda

Intergrades - Edisto

interocularis

Savannah

Altamha

Chattahoochee

bellicus

racca-oula Bay

Pearl

Mississippi tributaries

0

1

2

-28

5

9

r*

1

1

1

2

6

1

17

3

5

':

2

2

1

5

12

8

63

11

5

V

J •

12

10

25

3

4

6

10

7

44

8

7

1

1

NUMBER OF

4

7

32

40

32

2

113

15

17

5

12

48

27

ZL

10

6

3

7

TUBERCLES

6

6

41

15

21

3

1

1

3

7

4

18

1

2

3

8

5

11

1

3

9

-

5

-

10

2

5

2

11

1

1

1

TABLE 9.—The extent of tubercle development posteriorly on the head in subspecific populations of N. leptocephalus larger than 60 mm.

SUBSPECIES

Drainage IN PIN

AREA OF HSAD

AIO MIO PIO AOC POC

leptocephnlus

NevJ Janes,

York, Roaioke

Tar, Neuse,

Cape Fear

Pee Doe

Sar.tee, Catawba

Broad, Conjarea

Saltda

intergrades - EHisto

Interocularis

Savannah V

Chattahoocbee

2

3

10

8

15

3

15

3

4

10

14

11

8

7.

71

13

8

10

5

9

-

3

12

8

12

3

32

3

12

35

21

1?

13

86

18

16

98

60

*9

8

94

16

11

31

15

13

1

136 197 41

Alabama to !*5.ssis8ippi 4?
NUMBER 9 2 27

TABLE 10.—The relationship between standard length and the extent

of posterior development of cephalic tubercles in N. 1. leptocephalus

from the Santee River drainage (size groups marked with an X were

not sexed).

Zl/wr.

30-39

40-40

50-59

60-69

70-79

8C-°o

90-99

lCO-109

110-119

120-129

130-139

140-149

I5O-I59

I5SS

X 2

x 5

x 5

cf

9

x 3

cf

9»

X

cf

9

X

cf

9

X

cf

9

cf

9

cf

9

cf

cf

cf

a

cf

pn

13

8?

54

18

(7)

(1)

ATO

5

18

50

(1)

??.

r

CO

3

(2)

(1)

(1)

1

WTO

6

35

1

17

3

1 .

(5)

-

(2)

(2)

-

(1)

PIO

2

6

1

16

2

(8)

-

(1)

1

(1)

(1)

-

AOC

-

14

1

2?

4

(13)

T

r%

CO

1

(i)

1

1

1

MOC

1

10

1

37

3

(2)

6

21

on

(i'-O

2

(5)

1?

10

(2)

9

2

3

2

POC

1

5

1

5

(3)

(?)

32

(1)

5

2

2

1

TABLE 11.—Statistics comparing the total number of cephalic tubercles of two

size groups of five populations of N. leptocephalus. These data are shown

graphically in Figure 2.

SUBSPECIES

leptocsphalus

Cape Fear, northward

Pee Dec

Santee

lnterocularis

bellicus

90

No.

242

17

129

219

.3

Tin fltncl

X

17.1

22.8

18.4

8.2

5.2

over size

SD

3.^2

5.03

5.33

1.80

1.37

group

SE

0.22

1.23

0.47

0.32

0.21

X + 2 SE

16.6-17.5

20.4-25.3

17.^19.3

7.9- 8.4

4.8- 5.6

2p_

No.

'44

16

104

99

31

to 79

X

11.6

13.9

11.9

5.2

3.8

mm siz e

SD

4.42

5.61

3.98

2.31

1.52

Kroup

SE

0.67

1.40

0.39

0.23

0.27

X + 2 SE

10.3-13.0

11.1-16.7

11.1-12.6

4.7- 5.6
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 12.—The number of vertebrae in three subspecies of

Nocomis leptocephalus segregated by major river drainages.

SUHSPECISS - Sive r

1

4

1

2

1

VERTEPRAL XWBKRS

39

24

14

9

12

U

37

2

1*

3

3

1*

1?

2

2

40

22

10

12

21

4

11

1

1

2

10

11

12

34

41

2

2

3

1

1

1J. 1 .

Potomac

ToHc

Janes

Chomn

Noose

Cipe Fe«r

- Wlsto

I* i*

Altanahn

Chatt«hooch«e

1». 1. helllcus

!'6bil« Bay

Riscagoula Bay

Pearl

Mississippi tributaries

2

13

13

9

10

TABLE 13.—The number of pharyngeal teeth in Nocomis

leptocephalus separated by drainages.

5t!33FECIES SUI-SER PEtt BIGHT-LEFT ARCH

4 - 4 4- 3 3- 4 3- 3 2- 4

5

6

1 1

3

3

1 4

17

5
NUMBER 9 2 29

TABLE 14.—The number of circumferential and lateral-line scales in N. leptocephalus. Data for N. 1. leptocephalus from the New and James rivers southward

to the Santee River were grouped; data for N. 1. bellicus from the Alabama River

westward were grouped.

Drainage 25

leptocephalus

intergrados - SHisto

interocularis

Savannah

Altamaha

Chattahooehe<% 1

bellies

26

4

1

1

CIRCUMFERENTIAL SCALES

27

27

3

1

4

5

28

293

10

22

15

17

13

29

206

12

17

10

9

15

30

131

13

13

9

3

7

31

57

5

8

3

5

32

19

1

4

2

33

4

3

36

3

37

32

l

l

1

LATERAL-LINE SCALES

38

173

5

6

1

4

5

39

227

21

15

8

20

17

40

180

10

33

19

11

15

41

62

2

14

8

2

7

42

24

3

2

1

1

43

3

TABLE 15.—Comparison of the major characters of the subspecies

of N. leptocephalus.

CHARACTER

Head tubercles

Total mirfoer

I7u?>iher, 95?- of subadults, adults

Mean n'OTbor

1'jaxi.mar. nuinber

Head tubercle distribution

Posterior occurrence

T-'axiwun posterior

occurrence \n all

specimens examined

Nuptial crost

Vertebral number

Coloration

leptocephalus

Ikny

16-25

Greater than 17

33

Sictenciire

To mid- or postoccix>ital areas in

almost all sobadults

and adults.

POC

Very large

T'odally J) cr *»C

Two nupti-il rale

color patterns;

blue and brassyorange

SUBSPECIES

interccilaris

Intermediate

7-9

o

14

T'oderato

Almost alvays to

anterior or vAAinterorbital

areas.

AOC

Large

liO

Cne| blue

bellicus

Pew

4-6

5

7

Restricted

In internasal

or posterior

irteenasal

areas.

AI0

J'oderate

39

One, orange
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Appendix: Specimens Studied

Nocomis leptocephalus interocularis, new subspecies

(paratypes)

SAVANNAH RIVER SYSTEM (GEORGIA)

Burke County.—USNM 161675: Stream from Utley's Cave.

12 October 1951 (14 specimens).

Jefferson County.—USNM 166504: Brushy Cr., tributary of

Brier Cr. about 0.4 mi S of Wrens, US 1, 18 September 1947

(2). CU 17377: Brushy Cr., tributary of Brier Cr., 0.9 mi S

Wrens. US 1, 24 March 1950 (1). USNM 166503: Reedy Cr.,

3.9 mi NE of Wrens, US 1, 18 September 1947 (7).

Richmond County.—TU 38597: Rays Cr., at crossing of

Walton Way extension in Augusta, 5 July 1965 (3). TU

38605: Rays Cr., at crossing of Berkman Road in Augusta,

6 July 1965 (2). CU 11980: Rays Cr., 2 mi W of Augusta,

23 March 1948 (8). UMMZ 132758: Small brook in pine

woods 2 mi W of Augusta, tributary of Rays Cr., 27 December 1940 (5).

Columbia County.—TU 38588: Little Kiokee Cr., 103 mi

N of Evans, 27 June 1965 (9). TU 39486: Little Kiokee Cr.,

103 mi N of Evans on dirt road, 26 October 1965 (2).

USNM 161684: Lake Spring Cr., i/2 mi from Savannah R.,

24 July 1948 (9). USNM 161683: East Fork Cliatt Cr., 24

July 1948 (19). USNM 161682: West Fork Cliatt Cr., 24 July

1948 (13).

McDuffie County.—USNM 161692: Big Cr., 22 July 1948

(22). USNM 161694: Madison Cr., 22 July 1948 (15). USNM

161693: Harts Cr., 20 July 1948 (3).

Columbia-Lincoln County.—USNM 161685: Little R., 23

July 1948 (4).

Lincoln County.—USNM 161690: Cherokee Cr., 23 July

1948 (2). USNM 161689: Gray's Cr., 23 July 1948 (7).

USNM 161691: Lloyd's Cr., 23 July 1948 (7).

Oglethorpe County.—UG 283: Buffalo Cr., 6 mi E of Lexington, 29 April 1953 (3). UAIC 1463: Buffalo Cr., about 3

mi E [W?] of Wilkes-Oglethorpe County line, 5 September

1964 (10). USNM 161699: South Fork Broad R., 16 May

1948 (4).

Elbert County.—CU 19719: Morea Cr., 1.3 mi S of Nuberg

on Rte. 77, 27 March 1951 (6). UG 636: Little Dove Cr.,

0.8 mi E of Deep Creek Baptist Church, 11 April 1959 (20).

USNM 161686: Small Cr., 23 mi E of Broad R., Georgia

Hwy. 72, 15 April 1950 (12).

Madison County.—USNM 161697: Small Cr., 03 mi E of

Carlton, Georgia Hwy. 72, 15 April 1950 (8). UAIC 1613:

Fork Cr., E side of Carlton, Georgia Hwy. 72, 4 June 1965

(5). UG 308: Fork Cr., 2 mi NE Comer, 19 April 1954 (12).

UG 22 E: South Fork Broad R., 2 mi S of Carlton, 11 April

1959 (16). TU 26181: South Fork Broad R., 2 mi S of Carlton, 21 April 1962 (19). UG 22 D: South Fork Broad R., 1

mi S of Carlton, 19 April 1954 (7). UG 269: South Fork

Broad R., 2 mi W of Comer, 1 November 1952 (3). CU

41277: Stream [South Broad R.?] crossing Rte. 72, 1.8 mi W

of Comer, 5 June 1961 (1). TU 26178: South Fork Broad R.,

I mi W of Comer, 21 April 1962 (11). UAIC 1612: South

Fork Broad R., about 2 mi E [W?] of Comer, Georgia Hwy.

72, 4 June 1965 (4). USNM 161696: 23 mi SE Danielsville.

Dewey Powers Farm, 19 June 1950 (1). CU 51112: Tributary

of Broad R., 5 mi NE Danielsville on Rte. 29, 27 March 1952

(27). UG 173 A: Small Cr., 3 mi SE Danielsville, 19 April

1954 (4). USNM 166502: Broad R. tributary, 0.7 mi SW of

Danielsville, US 29, 10 September 1947 (22). UG 309: Holly

Cr., 3 mi N of Carlton, 19 April 1954 (2). USNM 161695:

Mason's Cr., 11 mi W of Royston, US 29, 15 April 1950 (16).

USNM 165720: Broad R. trib., 10 mi SW of Royston, US 29

(below dam), 5 April 1952 (18). CU 22689: Tributary of the

South Fork [of the Broad R.], 2.4 mi NE of Madison County

line on Rte. 29, 27 March 1952 (15).

Banks County.—TU 37510: Hudson R., about 2 mi N of

Homer, on Hwy. 441, 24 April 1965 (65). CU 21433: Hudson

R.. 17.6 mi S of Cornelia on Rte. 441, 9 June 1952 (2). CU

21591: Tributary of Hudson R., 1.7 mi N of Homer on Rte.

441, 9 June 1952 (9).

Franklin County.—UG 696: Middle Fork Broad R. NVV

[SW?] of Carnesville, 23 September 1959 (3). CU 19620:

Tributary of Nail Cr., 0.7 mi SW of Ashland, 27 March 1951

(24).

Stephens County.—USNM 161700: Tributary of Broad R.,

29 August 1948 (6). UG 637: Carnes Cr., 2 mi S of Toccon,

II April 1959 (58). USNM 171671: North Fork Broad R.,

3.7 mi W of Toccoa, 9 April 1947 (33). CU 17445: Toccoa

Falls, about 2.5 mi from Toccoa, 1 April 1950 (32). CU

21593: Tributary of North Fork of Broad R., 5 mi W of

Toccoa on Rte. 123, 8 June 1952 (31). UG 638: Toccoa Cr.,

just upstream from Georgia Hwy. 17, 11 April 1959 (36).

Hart County.—USNM 161688: Beaver Dam Cr., 3 mi E of

Royston at US 29, 15 April 1950 (10). CU 19724: Tributary

of Savannah R., 3.3 mi S of Hartwell on Rte. 77, 27 March

1951 (15). UMMZ 94568: Creek S of Hartwell, 12 November

1931 (10). UG 687: Savannah R., 1 mi below Hwy. 29

bridge. 1 September 1959 (37).

Habenham County.—UG 310: Panther Cr., 3 mi S of Tallulah Falls, 23 April 1954 (1). FSU 7418: Panther Cr., 102

mi NE of ClarkesvUle, US Hwy. 441, 16 July 1961 (5). TU

29322: Panther Cr.f 3.9 mi SW of Tallulah Falls, US Hwy.

441, 19 June 1962 (9).

Rabun County—FSU 7422: Tallulah R., 2.2 mi SE of

Wiley (about 10 mi S of Clayton), US Hwy. 441, 16 July 1961

(13). TU 29329: Tallulah R., 13 mi S of Wiley, US Hwys.

441 and 23, 19 June 1962 (24). UG 492: Bowen Branch

(=Boen Creek, Tiger Creek-Tallulah R. tributary just N of

Wiley), 14 May 1957 (6). TU 12231: Timpson Cr., 5.4 mi W

of Clayton, US Hwy. 76, 8 October 1955 (6). FSU 4078:

Timpson Cr., 5.4 mi W of Clayton, Rte. 76, 8 October 1955

(2).. UG 317: Persimmon Cr., tributary, 6 mi WNW of

Clayton, 24 April 1954 (9). CU 25174: Tributary of Tallulah R., 6.1 mi W of Clayton on Rte. 76, 3 September 1953

(11). CU 37919: Stream 6 mi N of Wiley on US Hwy. 441

(detour to Clayton), 6 June 1961 (1). UG 711: Persimmon

Cr., 2 September 1959 (10). UG 600: StonewaU Cr., on Harley McCall's farm, 16 October 1958 (9). UG 491: Worley

Cr., 15 May 1957 (14).
NUMBER 9 2 31

SAVANNAH RIVER SYSTEM (GEORGIA-SOUTH CAROLINA

STATE LINE)

Rabun-Oconee County.—TU 26979: Chattooga R., at

Georgia-South Carolina state line on Hwy. 28, just above

junction with West Fork, 13 July 1962 (1, cataloged with

A\ micropogon specimens).

SAVANNAH RIVER SYSTEM (SOUTH CAROLINA)

Allendale County.—USNM 162567: Savannah R., at mi

134.5, just above Ring Jaw Point, 15 May 1952 (1). USNM

200157: East Mill Cr., a branch of Tower Runs [Lower

Three Runs?] Cr., 3.0 mi E of Martin, South Carolina Hwy.

37, 12 June 1957 (1). CU 15152: Tributary of Lower Three

Runs Creek, 3.6 mi NW of Appleton on Rte. 28, 7 June

1949 (24).

Barnwell County.—USNM 200164: Branch of Three Runs

Cr., 1.5 mi S of Donora, 2.5 mi E of Dunbarton, 20 August

1951 (1). USNM 200186: Branch of Lower Three Runs Cr.,

slightly NE of South Carolina Hwy. 40, SE of Donora, 2-3

mi E of Dunbarton, 8 October 1951 (1). USNM 200151:

Branch of Lower Three Runs Cr., 3.5 mi S of Donora, 1 mi

N of South Carolina 40, 1 October 1951 (2). USNM 200176:

Branch of Lower Three Runs Cr., 1-1.5 mi S of Donora, 2-3

mi E of Dunbarton, 8 July 1957 (1). USNM 192858: Branch

of Lower Three Runs Cr., 3-4 mi S of Donora, 8 July 1957

(6). USNM 200144: Poplar Br. of Lower Three Runs Cr.,

0.75 mi W of South Carolina Hwy. 39, 0.6 mi N of R.R.

crossing, 1.5-2 mi N of Dunbarton, 12 June 1957 (1). USNM

200171: Fork of Lower Three Runs Cr., 5-6 mi NW Snelling,

on South Carolina Hwy. 53, 1-1.5 mi E of South Carolina

Hwy. 39 just E of Joyce Branch, 11 June 1957 (1). USNM

200179: Joyce Branch of Lower Three Runs Cr., 4-5 mi N

of Dunbarton, 1 mi W of South Carolina Hwy. 39, 11 June

1957 (3). USNM 200187: Joyce Branch, 1 mi W of Hwy. 39,

4.5 mi N of Dunbarton, 9 June 1952 (4). USNM 200145:

Hattie Cr., 3.5 mi S of Meyer's Mill, at South Carolina Hwy.

28, 13 August 1951 (19). USNM 200159: Hattie Cr., 3.5-4

mi S of Meyer's Mill, 1.5 mi NE Hattieville, South Carolina

Hwy. 28, 13 August 1952 (2). USNM 200172: Same locality

as USNM 200159, 12 August 1958 (2). USNM 200182: Steel

Cr., 3.5 mi W of Dunbarton, Hwy. 64, 8 August 1951 (8).

USNM 200158: Branch of Steel Cr., 1.5 mi NE of Meyer's

Mill, South Carolina 122, 8 August 1951 (3). USNM 200181:

Branch of Steel Cr., 1 mi E of Meyer's Mill, South Carolina

55, 8 August 1951 (5). USNM 200184: Steel Cr., 2 mi W of

Meyers Mill, South Carolina 56, 10 August 1951 (9). USNM

192856: Steel Cr., 2.5 mi W of Meyer's Mill, South Carolina

Hwy. 56, 7 July 1952 (4). USNM 200141: Same locality as

USNM 192856, 5 August 1958 (7). USNM 200178: Spring

Branch, 0.5 mi W of Meyer's Mill, South Carolina 56, 13

August 1951 (1). USNM 200166: Upper East Branch of Steel

Cr. about 0.5 mi W of Meyer's Mill, South Carolina 56, 7

July 1952 (1). USNM 200183: Same locality as USNM

200166: 5 August 1958 (4). USNM 200175: Branch of Steel

Cr., just S of Meyer's Mill, South Carolina 28, 13 August

1951 (1). USNM 200142: Same locality as USNM 200175,

5 August 1958 (4). USNM 200156: Branch of Steel Cr.. 05

mi SW of South Carolina Hwy. 28, 3 mi SSW of Meyer's

Mill, 17 September 1951 (4). USNM 200168: Same locality

as USNM 200156, 12 August 1958 (3). USNM 200177: Pen

Creek, 5 mi SE of Ellen ton at South Carolina Hwy. 28, 26

July 1951 (1). USNM 200146: Pen Cr., at intersection with

C. & W.C. R.R., 5 mi SE of Ellenton, 6 August 1951 (2).

USNM 200174: Pen Cr., about 6.5 mi SE of Ellenton, 2 mi

SW of South Carolina 28, 3.5 mi NE of Savannah R., 14 July

1952 (4). USNM 200160: Indian Grave Cr., 3.5 mi SE of

Ellenton, South Carolina Hwy. 64, 3 August 1951 (4).

USNM 200173: Pen Cr., 5 mi SE of Ellenton, South Carolina

64, 3 August 1951 (4). USNM 200185: Pen Cr., 5 mi W of

Dunbarton, about 15 mi N of South Carolina Hwy. 64, 12

August 1958 (1). USNM 161674: Pen Cr., 10 October 1951

(3). USNM 200165: Upper Four Mile Cr., 2 mi E of Ellenton, at South Carolina 48, 19 May 1952 (1). USNM 161673:

Four Mile Cr., 10 October 1951 (1).

Aiken County.—USNM 200154: Upper Four Mile Cr., 5.4

mi E of Ellenton, 0.5 mi NE South Carolina 149, 25 July

1951 (6). USNM 162551: Upper Three Runs Cr., at South

Carolina Hwy. 28, 3 May 1952 (4). USNM 192854: Upper

Three Runs Cr., Tims Branch, 1 July 1958 (19). USNM

200167: Tims Branch, 2 mi E South Carolina Hwy. 19, on

South Carolina 148, 25 July 1951 (1). USNM 200161: Branch

of Tinker Cr., 1 mi W of Kennedy's Lake, 1 October 1951 (2).

USNM 200180: In Pond at grist mill on Reedy Branch of

Tinker Cr., a branch of Upper Three Runs Cr., 8 mi SW of

White Pond, 13 June 1957 (3). USNM 200150: Mossy

Branch of Tinker Cr., 0.1 mi SW of Mossy School, 10 mi

NNW of Dunbarton, South Carolina Hwy. 124, 7 July 1952

(1). USNM 200143: Same locality as 200150, 28 June 1957

(4). USNM 200149: Tinker Cr., a branch of Upper Three

Runs Cr., 2.5 to 3 mi downstream from Kennedy's Lake, 26

June 1957 (2). USNM 200162: Upper Three Runs Cr., 9 mi

W of White Pond, 4 mi E of Talatha, South Carolina Hwy.

781. 27 June 1952 (1). USNM 200163: Henry's Pond and

stream behind pond, on Beulah Fork of Upper Three Runs

Cr., 0.5 mi N of South Carolina Hwy. 781, 7 November 1952

(1). USNM 200148: Same locality as 200163. 14 June 1957

(1). USNM 200169: Turner Branch, 15 mi SE of Hawthorne, 13 July 1951 (1). USNM 200202: Same locality as

200169, 16 July 1951 (4). USNM 200152: Same locality as

200169, 30 June 1952 (2). USNM 200147: Same locality as

200169, 27 July 1958 (8). CU 24384: Turner Branch of

Upper Three Runs Cr., 15 mi SE of Hawthorne, 30 June

1952 (4). USNM 200153: Turner's Pond, below the dam at

headwater of Turner Branch of Upper Three Runs Cr., 1

to 15 mi SE of Hawthorne, 15 mi E of South Carolina Hwy.

19, 27 June 1958 (5). USNM 200155: Beulah Branch of

Upper Three Runs Cr., 0.8 mi W of South Carolina Hwy.

781. 14 June 1957 (4). USNM 200170: Upper Three Runs

Cr., 3.8 mi E of South Carolina Hwy. 19 on road 10A to

Aiken, 25 July 1958 (1). UG 364: Pond, AEC Savannah

River Plant, 15 June 1954 (22).

Edgefield County.—USNM 86190: Sweetwater Cr., 24 August 1922 (5). TU 38623: Stevens Cr. tributary, 4.6 mi E of
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Parksville, county Rte. 533-138, 7 July 1965 (4). TU 39477:

Log Cr., at US 25 crossing 4 mi N of 25-25A intersection just

N of Edgefield, 25 October 1965 (9).

Edgefield-McCormick County line.—TU 38613: Stevens

Cr., 1.4 mi N of Parksville, county Rte. 533-21, 7 July 1965

(15).

Abbeville County.—CU 19593: Calhoun Cr., 7.6 mi E of

Calhoun Falls on Rte. 72, 27 March 1951 (25). CU 19648:

Little R-, 5.6 mi E of Calhoun Falls, 27 March 1951 (4).

Anderson County.—CU 19668: Beaver Dam Cr., 25 mi E

of Anderson on Rte. 29, 25 March 1951 (1). TU 26190.

Tributary of Savannah R., [Little Generostee Cr.?] 5 mi W

of Iva, South Carolina Hwy. 184, 21 April 1962 (30). USNM

162959: Tributary 6 mi SW of Anderson, US 29, 4 April

1952 (27). CU 11446: Tributary of Generostee Cr., 5 mi W

of Anderson, 9 April 1947 (31). CU 19602: Twenty-three

Mile Cr., 0.9 mi N\V of Sandy Springs, 11.1 mi NW of An

derson on Rte. 76, 25 March 1951 (1). CU 19749: Eighteen

Mile Cr., 1 mi NVV of Pendleton on Rte. 76, 25 March 1951

(5). USNM 166501: Seneca R. tributary, 0.9 mi NW of

Pendleton, 10 September 1947 (66). CU 37916: Hurricane

Cr., Rte. 28 bridge, 0.9 mi NW of junction of Rte. 28 and

bypass 28, 3 June 1961 (3).

Oconee County.—CU 19752: Cane Cr., 3 mi SW [SE?] of

Walhalla on [off?] South Carolina Rte. 28, 25 March 1951

(21). USNM 94577: Little Cane Cr., Walhalla, 28 March

1930 (39). USNM 94578: Cane Cr., Walhalla, 28 March 1930

(6). USNM 94579: Pond, 8 mi NE of Walhalla, 31 March

1930 (15). TU 26955: Little R., 8 mi NE of Walhalla,

South Carolina Rte. 11, 5 July 1962 (12). TU 26971:

Thompson R., Whitewater R. tributary, 1.4 mi S of North

Carolina-South Carolina state line in Sumter National

Forest, 10 July 1962 (5). TU 29461: Bearcamp Cr., near

junction with Horsepasture R., 05 mi SE of North Carolina

state line, 10 July 1962 (2). USNM 168102: Chauga Cr.. 3

mi NE of South Carolina-Georgia state line, US 123, 12 September 1954 (2). USNM 171628: Coneross Cr. tributary, 5.3

mi S of Seneca, 9 April 1947 (4). CU 11448: Tributary of

Coneross Cr., 5.3 mi S of Seneca, 9 April 1947 (4). CU

17365: Tributary of Tugaloo R., 4.7 mi N of Fair Play on

Rte. 59, 1 April 1950 (6). CU 17307: Tributary of Tugaloo

R., 2.4 mi W of Fair Play on Rte. 59, 0.9 mi E of Tugaloo

R, 1 April 1950 (1). USNM 171690: East Village Cr., Chat

tooga R. tributary just S of Mountain Rest on Rte. 28, 30

August 1946 (13). TU 26974: Chauga R., 15.9 mi S of

Georgia-South Carolina state line on Hwy. 28, 13 July 1962

(5). UMMZ 165635: Cedar Cr., Chauga R. tributary 9 mi

NW of Walhalla, 31 October 1940 (11). UMMZ 165544:

Bonecamp Cr., 10 mi NW of Walhalla, 31 October 1940 (77).

UMMZ 165610: Chauga R, 2 mi SE of Mountain Rest, 4

November 1940 (57).

Pickens County.—CU 19795: Tributary of Twelve Mile

Cr., 3 mi NNW of Pickens on Rte. 178, 24 March 1951 (25).

CU 38134: Tributary of Keowee R., 8.4 mi E of OconeePickens County line on Rte. 11, 3 June 1961 (6). USNM

171733: Keowee R. tributary, just E of Keowee R.R. crossing

about 10 mi S of North Carolina state line, Rte. 288, 30

August 1946 (14).

Oconee-Pickens County line.—TU 26996: Confluence of

Horsepasture and Toxaway rivers, 1.5 mi S of North Carolina-South Carolina state line, 1 July 1962 (6). TU 29585:

Toxaway R., 0.25 mi S of junction with North Carolina state

line, elev. 1,060 ft, 23 July 1962 (13).

NORTH CAROLINA

Transylvania County.—TU 29420: Bearcamp Cr., Horsepasture R. tributary, 0.2 mi N of South Carolina state line,

28 June 1962 (8). TU 29391: Bearcamp Cr., 4 mi S of Oak

land, 25 June 1962 (12). TU 27001: Same locality as TU

29391, 24 July 1962 (34). TU 29784: Same locality as TU

29391, 27 August 1962 (12). TU 28023: Horsepasture R., 0.5

mi N of North Carolina-South Carolina state line, 6 June

1962 (15). TU 29319: Horsepasture R., at bridge 3 mi S

Oakland on gravel road, 18 June 1962 (7). TU 29314: Horsepasture R., 2.5 mi S of Sapphire-Oakland on Whitewater

road, 18 June 1962 (10). TU 29303: Horsepasture R., 1.1

mi S of Oakland on Sapphire road, 14 June 1962 (5). TU

29436: Horsepasture R., between Drift Fall and Rainbow

Falls, 2.6 mi SW of Oakland, 3 July 1962 (3). TU 29599:

Toxaway Cr., Toxaway R. tributary, 3 mi SW of Rosman,

31 July 1962 (27). TU 29422: Bearwallow Cr., about 1.7 mi

N of North Carolina-South Carolina state line, ?9 June 1962

(2). TU 29261: Bearwallow Cr., 1.9 mi due N of South Carolina state line, elev. 1,410 ft, 7 June 1962 (5). TU 29288:

Bearwallow Cr., at junction with Toxaway R. 2.0 mi N of

South Carolina state line, 10 June 1962 (11). TU 29449:

Toxaway R., just below Toxaway Falls, 2 July 1962 (344).

USNM 200757: Mill Cr. where it begins to back up at entrance into Lake Toxaway, about 50 yds below bridge. Also

in Toxaway R. just below Lake Toxaway Dam. 15 May

1966 (6). TU 29309: Toxaway R., above Lake Toxaway, 2.6

mi W of town of Lake Toxaway, 15 June 1962 (21). TU

29450: Same locality as TU 29309, [probably 2 July 1962] (1).

TU 29534: Same locality as TU 29309, 14 July 1962 (11).

Jackson County.—TU 28017: Chattooga R. tributary [Fowler Cr.?], 1.7 mi S of Cashiers, South Carolina Hwy. 107, 4

June 1962 (2). TU 29652: Chattooga R., 27 mi S of

Cashiers, 7 August 1962 (5). UMMZ 156134: Horsepasture

R., S of Lake Fairfield, 2 July 1947 (32).

Nocomis leptocephalus interocularis, new subspecies

(non-paratypes)

OGEECHEE RIVER SYSTEM (GEORGIA)

Warren-Hancock County line.—TU 39498: Ogeechee R.,

at Jewell, 26 October 1965 (6). OAM [no number]: Ogee

chee R., below dam at Mayfield, 30 August 1964.

Taliaferro County.—UMMZ 158016: Ogeechee R., 0.75 mi

S of Robinson, 27 August 1939 (2).

ALTAMAHA RIVER SYSTEM (GEORGIA)

Wilcox County.—USNM 166508: Ocmulgee R. tributary, at

Oscewichee Springs, 14 mi N of Fitzgerald, 17 September
NUMBER 9 2 33

1947 (1).

Monroe County.—UMMZ 88309: Tobesofkee Cr. tributary,

0.5 mi W of Forayth, 3 September 1929 (4).

Lamar County.—UMMZ 88302: Little Towaliga Cr., near

Barnesville, 3 September 1929 (86).

Henry County.—CU 17524: Walnut Cr., 1.5 mi N of MeDonough, 29 March 1950 (9).

Dekalb County.—CU 51015: South R., 4.9 mi SE of junction of Rtes. 12-78 on Rte. 12, SE of Atlanta, 7 September

1953 (3). UAIC 1447: Ocmulgee R. tributary, about 2 mi

W of Georgia R.R., SE of Stone Mountain, 30 August 1964

(39).

Gwinnett County.—USNM 165731: Ocmulgee R. tributary,

5.5 mi W of Loganville, 5 April 1952 (10). UAIC 1449: Ocmulgee R. tributary, 4 mi W Loganville, about 5 mi W of

Gwinnett-Walton County line, 30 August 1964 (5). USNM

165736: Tributary, 9 mi WSW of Snellville, 5 April 1952 (6).

UAIC 1448: Yellow R., about 2 mi E of Dekalb-Gwinnett

County line near Snellville, 30 August 1964 (2). TU 26140:

Yellow R., 7 mi E of Duluth, 20 April 1962 (13).

Laurens County.—UMMZ 88372: Rocky Cr., near Dudley

on Georgia state road 26, 2 mi from Beckley County line, 5

September 1929 (10). UMMZ 88388: Hunger and Hardship

Cr., 5 mi SW of Dublin, 5 September 1929 (II).

Jones County.—TU 12050: Commissioners Cr., 2.4 mi E

of James, 16 September 1955 (4).

Jasper County.—TU 37507: Little R. tributary. 12.1 mi

NE of Monticello, 24 April 1965 (20).

Newton County.—TU 26161: Little R. tributary, 2 mi N

of Newborn, 20 April 1962 (47).

Greene County.—UAIC 1620: Richland Cr., about 2 mi

W of Greensboro, 5 June 1965 (1). UMMZ 158005: Richland

Cr., 1.5 mi W of Greensboro, 27 August 1939 (14).

Walton County.—USNM 161703: Jack's Cr., at bridge at

High Shoals, on Goodhope Rd., 31 March 1949 (2). USNM

161702: Jack's Cr., below Mill dam, 1 mi E of Goodhope,

31 March 1949 (11). USNM 165730: Tributary. 1 mi NE of

of Monroe, 5 April 1952 (21). USNM 161701: Apalachee R.,

Mill Race at High Shoals, 31 March 1949 (2).

Oconee County.—CU 37846: West Branch Oconee R. [Apalachee R.?], at Morgan-Oconee County line on Rte. 129, 4

June 1961 (4). USNM 161698: Apalachee R., at High Shoals.

9 August 1950 (4). CU 51049: Barbers Cr.. Hope Mill near

Princeton, 3 September 1950 (8).

Oglethorpe County.—UAIC 1465: Creek on US 78 about 9

mi E of Athens airport, 5 September 1964 (7).

Clarke County.—UG 3: South Prong, 3 mi SE of Athens,

19 October 1947 (3). USNM 161678: McNutt's Cr.. 14 May

1948 (8). USNM 161676: McNutt's Cr.. at YWCA camp. 4

November 1947 (62). USNM 161677: McNutt's Cr., at

YWCA camp, 18 February 1950 (7). USNM 161680: Middle

Oconee R., at Princeton, 14 April 1950 (1). UG 340: Middle

Oconee R., 05 mi below dam at Princeton, 30 April 1954

(1) UG 265: Oconee R., 3 mi W of Athens, 2 October 1952

(2). USNM 161679: Middle Oconee R., 5 October 1947 (3).

UG 420: Middle Fork Oconee R., 26 October 1954 (4).

USNM 161671: Small tributary, Middle Oconee R., 29 April

1951 (7). USNM 161670: Oconee R., at Athens, 3 March

1951 (2). UMMZ 87059: Athens. 1853 (5). UMMZ 94578:

Biggers Cr., NW Athens, 12 November 1931 (3). USNM

161681: Bobbin Mill Cr., 20 September 1947 (34).

Barrow County.—USNM 161669: Beechwood Cr., 22 April

1951 (1). CU 21438: Tributary of Mulberry Fork. 1.2 mi

NW of junction of Rte. 124 and Rte. 211. 9 June 1952 (22).

USNM 166507: Tributary of North [Middle?] Oconee R., 2.5

mi N of Winder, 11 September 1947 (238).

Jackson County.—UMMZ 94529: Aliens Cr., N of Talmo,

12 November 1931 (103). CU 21444: Curry Cr., just inside

Jefferson city limit, 9 June 1952 (27). CU 21397: Tributary

of Oconee R., 3.3 mi SW of Commerce, 9 June 1952 (2).

Hall County.—CU 17341: Mulberry Fork, tributary of

Middle Fork of Oconee R., 13.8 mi S of Gainesville, 31

March 1950 (24). CU 21437: Tributary of Mulberry Fork,

4.1 mi NW of junction of Rte. 124 and Rte. 211, 9 June

1952 (7). CU 11203: Tributary of Oconee R.. 2 mi NW of

Gillsville, 9 April 1947 (2). CU 17427: Same locality as CU

11203, 1 April 1950 (9). CU 19809: Same locality as CU

11203, 26 March 1951 (28). CU 43581: East Branch [North

Fork] of Oconee R., W of Gillsville, 11 September 1962 (30).

TU 30496: East Fork [tributary of North Fork] of Oconee R..

about 2 mi N of Gillsville, 22 September 1963 (3). USNM

171734: [tributary of] Oconee R.. 2 mi NW of Gillsville, 9

April 1947 (20). TU 30484: North Oconee R., 5.9 mi NE

of Gainesville, 22 September 1963 (5).

CHATTAHOOCHEE RIVER SYSTEM (ALABAMA)

Lee County.—UAIC 1233: Wacoochee Cr T 19 N. R 29 E,

Sec. 23, 4 April 1964 (4).

CHATTAHOOCHEE RIVER SYSTEM (GEORGIA)

Upson County.—TU 27525: Potato Cr., Flint R. tributary,

5 mi SW of Thomaston. 23 April 1962 (5).

Harris County.—FSU 6688: Mulberry Cr. tributary, 7.9 mi

E of Hamilton, 17 August 1960 (11). CU 17514: Tributary

of Mulberry Cr., 7.7 mi E of Hamilton, 29 March 1950 (5).

UMMZ 157884: Mountain Cr.. 25 mi SE of Chipley, 23

August 1939 (3).

Troup County.—UMMZ 157892: Flat Shoal Cr., 72 mi

NW of Chipley, 23 August 1939 (2). TU 7527: Tributary of

Chattahoochee R., 5.8 mi SW of La Grange, 8 October 1953

(21).

Meriweather County.—FSU 7040: Sulpher Cr. tributary, 2

mi NE of Pine Mountain, 16 August 1960 (40).

Carroll County.—UAIC 1095: Wolf Cr., 5 mi NE of

Whitesburg, 8 December 1963 (1).

Douglas County.—UAIC 1093: 0.9 mi S of McWhorter,

7 December 1963 (31).

Cobb County.—USNM 161672: Nickajack Cr., at Camp

Highlands, 4, 11 March 1951 (14).

Fulton County.—UMMZ 88296: Nancy Cr., about 10 mi

N of Atlanta, 2 August 1929 (11). CU 51023: Nancy Cr.. 9.7

mi S of Roswell, 6 September 1953 (5). CU 17136: Vickery

Cr., at junction with Chattahoochee R. at limits of Roswell,

30 March 1950 (1). TU 12131: Vickery Cr.. 0.6 mi E of Ros-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

well, 6 October 1955 (1). TU 26128: Vickery Cr. tributary,

4 mi NE of Roswell, 20 April 1962 (20).

Gwinnett County.—UG 819: Richland Cr., 1 mi from

Chattahoochee R., 20 October 1962 (13).

Hall County.—USNM 93201: 7 mi NW of Gainesville, 1933

(4). CU 21445: Tributary of Chattahoochee R. 1 mi N of

junction of Rte. 52 and Rte. 129, 10 June 1952 (5). USNM

166505: Chestatee R. tributary, 12.4 mi N of Gainesville, 11

September 1947 (135). CU 19819: Tributary of Chestatee R.

1 mi SE of Murraysville, 26 March 1951 (19).

Lumpkin County.—USNM 166506: Chestatee R. tributary.

1.3 mi WSW of Dahlonega, 11 September 1947 (9). CU

21418: Cane Cr., 1.5 mi SW Dahlonega, 10 June 1952 (1).

CU 10990: Cane Cr., 1.6 mi WSW of Dahlonega, 10 April

1947 (1). TU 38362: Cane Cr., 2 mi W of Dahlonega, 21

June 1965 (4). UG 604: Cane Cr., at first bridge N of Dahlonega, 20 October 1958 (7). FSU 7523: Chestatee R., at

mouth of Yahoola Cr., 2.1 mi SE of Dahlonega, 17 July 1961

(3). UG 607: Yahoola Cr., at bridge behind Yahoola Church,

21 October 1958 (10). TU 30448: Chestatee R.. 4 mi SE of

Dahlonega, 14 April 1963 (6). TU 29773: Chestatee R., 10.7

mi NW of Cleveland, 17 August 1962 (2). TU 30423: Chestatee R., about 10 mi NW of Cleveland, 14 April 1963 (64).

UG 593: Dick's Cr., 0.8 mi above junction with Water's

Creek, 9 October 1958 (20). UMMZ 165559: Dick's Cr., tributary of Chestatee R., 15 mi W of Cleveland, 17 October

1940 (15).

White County.—TU 29764: Little Tesnatee Cr., 1.4 mi

NW of Cleveland, 17 August 1962 (14). USNM 161704:

Little Tesnatee Cr. tributary, 1 mi W of Cleveland, 3 April

1949 (1). TU 38325: Tesnatee Cr. tributary, 1.8 mi NW of

Cleveland. 21 June 1965 (67). TU 30437: Tesnatee Cr. tributary. 2 mi NW of Cleveland, 14 April 1963 (23). UMMZ

94590: Turner Cr., N of Cleveland, 13 November 1931 (2).

UG 592: Spoilcane Cr., 0.75 mi above its mouth, 8 October

1958 (1). UG 591: Spoilcane Cr., 1 mi above Chattahoochee

Management Area, 8 October 1958 (10). UMMZ 165618:

Chattahoochee R., 4 mi above Helen, 21 October 1940 (4).

Habersham County.—CU21404: Tributary, Chattahoochee

R. 2.9 mi SW of Cornelia, 8 June 1952 (2). CU 50304:

Tenner Branch near Cornelia, late spring 1951 (15). CU

18109: Grant Cr., tributary of Soque R. near its junction

with Chattahoochee R. near Cornelia, June 1949 (15). CU

17431: Tributary of Chattahoochee R., 2.7 mi E of Chattahoochee R. on Rte. 115, 1 April 1950 (11). CU 17439: Tributary of Soque R., 1 mi W Soque R. on Rte. 115, 1 April

1950 (1). FSU 5717: Soque R. tributary, 1.5 mi W of

Clarkesville, 16 July 1961 (5). USNM 168091: Tributary

below dam. 2.4 mi WNW of Clarkesville, 11 September 1954

(9). USNM 171663: Soque R. tributary, 5 mi NE of Clarkesville, 9 April 1947 (4). USNM 161687: Soque R., at Watt's

Mill, 9 mi N of Clarkesville, 3 April 1949 (2). UG 606:

Soque R., 0.75 mi upstream from Watt's Mill, 21 October

1958 (2). UG 597: Raper Cr., at Wickle's store, 13 October

1958 (2). UG 598: Raper Cr., (at the mines), 13 October

1958 (4). UG 588: Soque R. near Chimney Mountain Farm,

30 October 1958 (5). UG 587: Soque R., 0.25 mi above the

last bridge upstream, 30 September 1958 (13).

White County.—UG 622: Chickamauga Cr., 1 mi above

Hwy. 255, 22 October 1958 (6). UG 621: Sautee Cr., at

bridge off Hwy. 256, 22 October 1958 (9). UG 623: McClure

Cr., 055 mi above "lower" bridge, 22 October 1958 (20).

TU 29758: Dukes Cr., 1 mi SW of Nacoochee, 16 August

1962 (30).

White-Habersham County line.—UG 838: Chattahoochee

R. 24 October 1958 (9).

Nocomis leptocephalus leptocephalus (Girard)

N ew River: USN M 103979, 171707, 177352. 177356. 177359.

177378. 177379, 177382, 177390, 177394, 177402. 177407,

194273, 194701-194710, 194806, 199736, 199738, 199740. C U

19415, 19456, 20452, 20940, 23848, 24749, 24968, 25318. 25352.

25787, 38202, 42132. T U 25496. 25531, 25597, 28001, 29663.

UMM Z 126229, 136636, 137969, 147651. D U F-110, F-113,

F-116.

Potomac River: USN M 177355, 194659-194661.

Rappahannock River: C U 46283, 52054.

York River: USN M 100224, 107725, 107739, 171617, 194277.

James River: USN M 100186, 100196, 100218, 103991,

105083, 107503, 163435, 166490, 171632, 171633, 171659.

171730, 194263, 194626, 194633, 194669, 194693-194697. 194699.

194700, 194731, 194732, 194735-194737, 199697. 199701, 199708.

199711, 199713, 199715 199717. C U 11358, 11595. 12222.

20949, 23828, 23875, 24162, 24167, 29156. UMM Z 171383.

174486, 174679. 174721. 174827. 175113.

Chowan River: USN M 101342, 171619, 177383. 194744,

194745, 194759-194761. C U 11883. 18603, 33164, 50437.

50981.

Roanoke River: USN M 100168, 100177, 101329, 101332,

102059, 102060, 103976, 104007. 104012, 104017. 104028.

105071. 107591, 107595, 107604, 107616. 107634, 107651.

116564, 132024, 162826-162830, 162882, 162883, 166388-166399.

166491, 168143. 171618, 171621-171623, 171626, 171636.

171637. 171639. 171649, 171652, 171655, 171656, 171658,

171661, 171676, 171678. 171683, 171684, 171694. 171696.

171702. 171704-171706, 171708, 171709, 171724-171726, 171731.

171736, 171738, 171742, 177374, 177377, 177390, 177399,

177405, 177406, 194256, 194264, 194711-194730, 194754-

194758, 194762-194771, 199870, 199963, 199980, 200017.

200089. C U 11497, 11914, 11959, 13875, 14001, 16833, 50402,

50824, 50960, 50971. K.U 3284. T U 25887. UMM Z 137586,

171360, 174707. 175141. ANSP 1857-70, 75732. D U F-129.

Tar River: USN M 171657, 171660. C U 19432, 19475,

19504, 19521, 25881. 25908, 26024, 31739. UMM Z 147581.

Neuse River: USN M 69799, 171605, 171616, 171634, 171635,

171650. 171653. 171667, 171729, 188554, 188737, 194269,

194270. C U 9535, 19842, 25868. 31686, 34483. D U F-107,

F-126, F-128, F-130.

Cape Fear River: USN M 40289, 93222, 166492, 171609,

171624, 171625, 188571. C U 31676, 31752. D U F-115, F-117,

F-125, F-127.

Pee Dee River: USN M 104002, 105048, 105069, 105073,

105078, 105081, 162925-162927, 162947, 164309, 166493,

171607. 171608, 171638, 171668-171670, 171677, 171682,
NUMBE R 9 2 35

171695, 171703, 171712. C U 3501. 10981, 11212, 11342, 11423.

11434, 11490, 11527, 14278. 17177, 19559, 19634, 19727. 21323.

21358, 25380. 29831, 29841, 30036. 31665. 31821. 37701, 37942,

38176. K U 8879. TV 29835. ANS P 1850-56. UMM Z

139432, 147635.

Santee River: USN M 40515, 87164, 92448, 92476, 93211,

100655, 100661, 119086, 119291, 162898, 162903-162905,

162939, 162953, 166403, 166427, 166494, 166496, 166498-166500,

168130, 171604, 171606, 171627, 171640, 171648, 171651.

171654. 171672, 171674, 171675, 171679, 171686, 171710,

171711, 171728, 171740, 171741, 171743, 192852, 192855,

192857. 199758-199768, 199771-199797. 199799, 199801,

199803-199811, 199813-199815, 199840, 199841, 199858-

199869, 199871-199898, 199900-199902, 199904-199928,

199930-199936, 199944, 199946-199952, 199954-199962,

199964-199968. 199970-199979, 199982-199984, 199992-200013,

200015, 200016, 200018-200024, 200026-200029, 200031-

200037, 200051-200088, 200090-200098, 200127-200132,

200134-200140. 200188-200200, 200755, 200758. T U 16498.

25939, 26199, 27485. 27492, 27494, 27513, 28231, 28234, 29445.

ANSP 1826-40. K U 5270. 5272, 8890. C U 9935. 10064.

10123, 10535, 11756, 11938, 11962, 17070. 17074, 17091, 17114,

17201, 17305, 17540, 17722, 19225, 19288. 19331. 19340. 19399,

19552. 19659, 19674, 19688, 19690, 19704, 19711, 19714, 19716.

19741, 19788, 21243, 21371, 21382, 21386, 21394, 25167, 25859.

25972, 25978, 26060, 28359, 31857, 37841, 38063. 38076. 38108,

46430, 50883. UMM Z 163859. 165641. D U F-118. Univ.

Minn. 17673.

Nocomis leptocephalus bellicus Girard

MOBILE BAY DRAINAGE (ALABAMA-TOMBIGBEE

RIVERS)

UAIC 58. 59, 427. 432, 435, 492. 523. 532, 533, 678, 680.

685, 814, 819, 822, 829, 888. 928. 934, 959. 960, 961, 963, 965.

966, 971, 972, 1036, 1038, 1039, 1064. 1066-1069, 1098, 1102,

1232, 1241, 1249, 1250. 1282, 1284. 1307-1310. 1312, 1315,

1317-1319, 1354. 1359-1363, 1375, 1376, 1378, 1379, 1381,

1382, 1387, 1479, 1496, 1497, 1500, 1504, 1514, 1593, 1668,

1771, 1829. 1863, 1864. 1876, 1891, 1892, 2005.

T U 2628, 3421, 9932, 12083, 15282, 18702, 24555, 25992,

27340, 27551, 27555, 29903, 30187, 30227, 30950, 32434, 32490,

32510, 32527, 32539, 32581, 32599, 32656, 32673, 32690, 32694,

32740, 34405, 35071, 35161. 35174, 35319, 35324, 35373, 38749,

38771, 40595, 40650, 40664.

UMM Z 113917, 124065, 124100, 146514, 146530, 166359,

166379, 168662, 168667, 168761. 175830, 177740, 177752.

C U 13781, 13997, 16021, 19256, 40212, 42314, 42357.

USN M 161705, 166009, 199543, 200756.

API 404-2, 501-1, 714.

PASCAGOULA BAY DRAINAGE

T U 8032, 8041, 8053. 8065, 9899. 15439. 39390.

UMM Z 157784.

C U 11618, 12581, 16252, 16257.

PEARL RIVER DRAINAGE

T U 980, 1545, 3605, 3845, 7505, 7600, 13998. 14100. 15151.

15203. 15524, 16736, 17525. 17943. 19806. 23385. 23423. 23500.

23748, 26837, 26869. 26683, 26886, 27182, 27280. 27336, 27481.

28703, 28780. 28830, 28869, 30130, 31672, 31686, 36090, 40101.

C U 11859, 12487, 15684, 16266. 16613. 37393, 46419.

UMM Z 113775, 155350, 161151, 175388.

USN M 131905.

EASTERN \fississippi RIVER TRIBUTARIES

UAIC 1268. 1269, 1587.

FSU 9186, 10727.

T U 2925, 7214, 7897, 11998, 15130, 16621, 19776. 19843,

23889, 23953, 28896, 30154. 30888, 30897, 31790, 32343. 32923,

40281, 40370.

UMM Z 161199, 161208.

USN M 172027. 172034.

C U 37581.

MGFC 4468. 4520. 5754. 5755.

TENNESSEE RIVER DRAINAGE

UAIC 1760. 1776. 1886.

TU 40533.

Nocomis leptocephalus intergrade

interocularis x leptocephalus

EDISTO RIVER DRAINAGE (SOUTH CAROLINA)

Aiken County.—USNM 200201: Chinguapin Cr., 6.4 mi SSW

of Leesvillc at South Carolina Hwy. 391 crossing, 5 July 1956

(50). CU 29851: Chinguapin Cr. tributary, 6.7 mi S of

Batesburg, Rte. 391, 28 March 1956 (28). USNM 203918:

Chinguapin Cr., about 4 mi S of Batesburg, 0.25 mi W of

Rte. 391, 20 April 1967 (40). USNM 203920: Chinguapin

Cr., about 6 mi S of Batesburg, 20 April 1967 (10). USNM

203919: Chinguapin Cr., about 3.5 mi SSW of Batesburg, 21

April 1967 (27).

•if U.S. GOVERNMENT PRINTING OFFICE: 1971

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