ROBERT E. JENKIN,

and ERNEST A. LACHNFl

Criteria for Analysis

and Interpretation of

m the American Fish

Genera Nocomis Girard

and Hybopsis Agassiz i

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ERRATA: Smithsonian Contributions to Zoology, number 90

Lines on the indicated pages should read as follows:

Page 3, 1st column, 5th line from bottom —

"Hybopsis (sensu lato or sensu stricto)."

Page 3, 2nd column, 12th line from bottom --

"38 to 44, precaudal vertebrae number 20 to 24. De-"

Page 9, 1st column, 2nd line from top —

"ferences between the subspecies of Hybopsis (Eri-M

SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 90

Robert E. Jenkins and Criteria for Analysis

Ernest A. Lachner m

 J

and Interpretation of

the American Fish

Genera Nocomts Girard

and Hybopsis Agassiz

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971
ABSTRACT

Robert E. Jenkins and Ernest A. Lachner. Criteria for Analysis and Interpretation

of the American Fish Genera Nocomis Girard and Hybopsis Agassiz. Smithsonian

Contributions to Zoology, number 90, 15 pages, 1 figure, 1971.—Until recently

Nocomis Girard has been included as a subgenus in the polymorphic genus

Hybopsis Agassiz along with Couesius Jordan and six additional nominal American

cyprinid genera. Nocomis is diagnosed and recognized as a genus primarily on the

basis of sexual dimorphic features, its mound-nest building behavior, general

coloration, physiognomy, scale radii, and vertebral numbers. Nocomis and

Couesius, also given generic rank, are probably more closely related to the genus

Semotilus Rafinesque than to any group remaining in Hybopsis. Parexoglossum

Hubbs is merged with Exoglossum Rafinesque and these are implicated to be

related to Semotilus.

Characters previously employed to diagnose Hybopsis are widely shared with

other American cyprinids, such as the maxillary barbels which may be highly

variable. Certain subgenera of Hybopsis are probably more closely related to other

American cyprinid genera than to Hybopsis sensu stricto. Species of some subgenera of Hybopsis merit reallocation to other subgroups of Hybopsis or Notropis.

Additional nomenclatural changes in Hybopsis are not recommended until more

comprehensive studies are available.

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

in the Institution's annual report, Smithsonian Year.

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Robert E. Jenkins and

Ernest A. Lachner Criteria for Analysis

and Interpretation of

the American Fish

Genera Nocomis Girard

and Hybopsis Agassiz

Introduction

The group of nest-building barbeled cyprinid fishes

herein recognized as Nocomis Girard has had a history of instability regarding its level within the

genus-group category. A nomenclatural review of

Nocomis is given by Lachner and Jenkins (1971a)

and of Hybopsis by Reno (1969a).

Bailey (1951:192) submerged Nocomis and seven

additional nominal genera, including Couesins Jordan, in Hybopsis Agassiz. The nine nominal genera

are listed in Table 1. Bailey's arrangement has been

generally followed, with recognition of Nocomis as

a subgenus. However, Hubbs and Crowe (1956)

commented that "... the reexpanded genus Hybopsis

rests on an unsteady basis...." Lindsey (1956:771)

did not follow Bailey's actions, partly by reason of

the latter's failure to provide data for his contentions.

Bailey and Allum later (1962:36-37) attempted to

support lumping of Nocomis with Hybopsis. Recent

studies of brain and gustatory structures by Davis

and Miller (1967) and on the cephalic lateral-line

system by Reno (1967; 1969b), combined with our

work indicate that Hybopsis is an unnatural assemblage. Lachner and Jenkins (1967:558) regarded

Nocomis as a valid genus but reserved discussion of

this allocation for the present paper. Couesins is

herein recognized as a genus rather than a subgenus

of Hybopsis. Hybopsis, excluding Nocomis and

Robert E. Jenkins, Department of Biology, Roanoke College,

Salem, Virginia; Ernest A. Lachner, Department of Vertebrate

Zoology, National Museum of Natural History, Smithsonian

Institution, Washington, D.C.

Couesins, is still morphologically and biologically a

very diverse assemblage of about twenty species.

Our primary purpose is to define the genus

Nocomis as it relates to the remaining groups in the

currently recognized genus Hybopsis and to its apparent close relatives. Additionally, we discuss the

status of Hybopsis and make suggestions for future

realignment of some of the subgenera and species

included under it.

We have evaluated characters previously employed

in the classification of the genera and subgenera

treated herein with respect to their utility and importance. Most characters treated are external, but

some internal anatomical features have been considered. The morphology upon which some of the

statements given below are partly based will be presented for Hybopsis sensu stricto (the amblops

group) by Glenn H. Clemmer and for other groups of

Hybopsis and relatives by Jenkins and others. We

place more emphasis than some previous workers on

basic pigmentation patterns, aspects of physiognomy,

sexual dichromatism and dimorphism, behavior, and

ecology as useful group characters. Various aspects

of nuptial tuberculation, in particular, have been of

utmost importance as indications of relationship.

The systematic significance of tubercle characters

was discussed by Lachner and Jenkins (1967:567

and 1971a).

Acknowledgments

Technical assistance was received from former staff

members of the Division of Fishes, National Museum

1
2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 90

of Natural History: Kenneth Karb assisted in much

of the work on scale analysis; Dorothy Hubbs and

Martin L. Wiley aided considerably in taking and

reading some of the radiographs. Frank B. Cross,

University of Kansas, permitted use of his vertebral

counts for Hybopsis gracilis. We have profited in

our knowledge of Hybopsis through discussion with

Glenn H. Clemmer, Mississippi State University.

The Genus Nocomis

Nocomis is perhaps the most distinctive group that

has been relegated to Hybopsis. It is unique in two

and nearly unique in two others of the first four characters, which are concerned with reproduction, in

Table 1. It differs almost entirely in number of total

scale radii (Table 2) and considerably in precaudal

vertebral numbers (Table 4). Its cephalic lateralline system is unique only in that long canaliculi

develop in apparent compensation for increase in

thickness of facial integumental areas (Reno, 1967:

72, 107; 1969b:757). Nocomis shows homogeneity in

its lateral-line development, brain pattern, barbel

origin and morphology, and its taste-bud sensory

systems, but in these characters it is essentially similar to several other Hybopsis subgroups. It is a

well-marked group in having a robust body (not

shared with several other so-called chubs in Hybopsis sensu lato), specific contours of the head in

juveniles, and a characteristic body coloration.

Bailey and Allum (1962:36-37) mentioned character trends associated with reproductive behavior

in Nocomis and other cyprinid genera and stated

that the resultant developments (such as large

tubercles) should not be over-emphasized in classification. We place considerable importance upon

the fundamental fact that all Nocomis build a

mound nest and all other Hybopsis do not. Elaborations of sexual dimorphic features apparently are

positively correlated with the nest building behavior,

and as such they may be regarded as forming a single

character complex. However, even if these characters are disregarded, there is still virtually no evidence for close relationship between Nocomis and

Hybopsis (sensu lato or sensu sricto).

Bailey and Allum (loc. cit.) apparently regarded

monotypy or oligotypy of phyletic groups to merit

reduced weight in classification. We do not subscribe to this opinion, and, in addition, note that a

greater amount of differentiation has occurred

within Nocomis than was indicated by Bailey and

Allum; three species groups have evolved.

No group remaining in Hybopsis s.l. approaches

Nocomis in its combination of morphology, coloration, and habits. In the event of further dismemberment of Hybopsis, Nocomis will obviously not be

associated with the name Hybopsis, because its relation to the H. amblops complex (the type species of

Hybopsis) is quite remote; there are no extant forms

transitional between the H. amblops complex and

Nocomis.

We regard Nocomis, which has recently been recognized as a genus by Douglas and Davis (1968),

Raney (1969), Scott and Crossman (1966), SmithVaniz (1968), and Suttkus and Ramsey (1967), as a

natural unit of generic rank, marked by the following characters: Nuptial males (1) construct a moundnest by transportation of stones in the mouth;

(2) have large to very large breeding tubercles on

the head and, in two species, on the body; (3) have

an expansive cephalic crest or swelling except in one

species group; (4) develop on the body elaborate

breeding coloration of pink-rosy, orange, and/or

bluish; (5) attain a moderate to large body size,

adult males 100 to 250 mm SL (Standard Length).

Other characters common to both sexes follow:

(6) coloration dark above, light below with broad

dark scale margins and post-opercular bar; (7) large

scales, the lateral line scales almost always fewer than

44 and body circumferential scales fewer than 37

(caudal peduncle scales 16-20); (8) mouth normal,

moderate to large in size, subterminal; (9) barbels

small, one terminal maxillary pair; (10) pharyngeal

tooth count 1,4-4,1; 4-4; or 3-3; (11) scales with

radii on posterior field only, the total number of

radii in adults 23-54; (12) total vertebrae number

38 to 43, precaudal vertebrae number 21 to 24. Descriptions of the species groups of Nocomis are given

by Lachner and Jenkins (1971a).

The closest relatives of Nocomis are probably the

nest-building species of Semotilus Rafinesque, S.

atromaculatus and S. corporalis. In addition to reproductive behavior (nest building), these species

are similar to at least the more primitive Nocomis in

their reddish nuptial colors and pattern, pectoral fin

tuberculation, head and body physiognomy, scale

characters, vertebral numbers, and general body

color pattern. The Couesius group shows affinity
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with Semotilus, particularly with Semotilus margarita, in the following characters: barbel position;

squamation pattern; opercular, branchiostegal, and

breast tuberculation; breeding color (see painting in

Raney, 1969); scale morphology; vertebral numbers;

and other features. The data indicate that if

S. margarita merits inclusion in the genus Semotilus

(Bailey and Allum, 1962:36-37), so does Couesius.

The mound-nesting behavior and other characters

of the Exoglossum group suggest that it, too, may

be related to Semotilus.

Scale Morphology

METHODS.—Scale samples were taken from adult or

subadult specimens. The scales were selected from

the dorsolateral portion of the body below the dorsal

fin. Only typical scales were studied, and those

showing any regeneration were discarded. Scales

were studied from about ten specimens of 19 of the

species of Hybopsis, the one species of Couesius, and

all species of Nocomis. In addition, the scales from

three specimens of each of 89 species of Notropis

were studied, as were scales of other American cyprinid genera.

Preliminary studies revealed that among the various characters of scale morphology, the best comparative character between the genera was the number

of radii. In Hybopsis and Nocomis the radii are

found only on the posterior field. The radii were

subdivided initially into two groups, primary and

secondary. The secondary radii were considered,

for convenience, as those one-half or less of the

length of the longest radii. All others were considered primary radii. The number of primary and/

or secondary radii were compared as a percentage of

the total number, and various differences among the

species were observed, but the most useful comparison was the total count (Table 2).

The use of scale morphology as a systematic character was initiated mainly by Agassiz in 1833 with

his work on fossil and recent fish scales. Later

studies in lepidology, such as those of Cockerell

(1911; 1913), Lagler (1947), Kobayasi (1951, 1952-

1955) and McCully (1961) reviewed scale characters

on a family or major group level. When our preliminary studies revealed possible meristic differences

in the number of scale radii, we undertook a comprehensive survey, although we were aware that conTABLE 2.—Total number of radii in the posterior

field of scales of Hybopsis, Couesius, and Nocomis

(Genus-group)

Hybopsis

(Hybopsis)

(Yuriria)

(Erimystax)

(Plalygobio)

(Macrhybopsis)

(Extrarius)

(Oregonichthys)

Couesius

Nocomis

Species

a. amblops

rubrifrons

hypsinotus

labrosa

sp.,cf. labrosa

storeriana

alta

d. dissimilis

x. x-punctata

i. insignis

cahni

monacha

harperi

g. gracilis

gelida

meeki

aestivalis

crameri

plumbeus

biguttatus

asper

effusus

platyrhynchus

raneyi

micropogon

leptocephalus

Mean

19.2

13.7

9.6

7.0

11.5

18.5

17.7

12.3

7.6

8.4

5.7

9.7

18.5

14.8

13.7

10.0

6.9

10.8

18.5

33.4

29.2

40.7

34.0

31.9

40.8

28.6

Range

15-25

9-19

8-14

5-9

8-15

14-24

16-21

9-17

6-16

7-10

5-7

7-11

13-26

13-18

11-20

7-13

4-12

7-13

15-22

26-40

23-36

25-19

24-48

25-36

24-54

24-35

Specimens

9

10

10

10

11

12

12

10

11

9

5

10

11

10

10

9

10

11

11

9

10

18

12

9

12

10

siderable variation in many aspects of scale structure

existed in any single species. Lagler (1947:153)

noted that the characters "least usable for diagnostic

purposes, even at the major taxonomic level of families, are absolute size, shape, numbers of various

countable structures, and position of the focus." Chu

(1935:21) in his studies of Chinese cyprinids, pointed

out that, "Granting sufficient allowance for variations, the number of radii in the various fields may

be stated as fairly characteristic for certain forms or

groups."

RESULTS.—Table 2 giving the average values for

the total number of radii in all the recognized species

of Hybopsis, Couesius, and Nocomis illustrates a

major difference among these genera. The highest

individual counts of only three species of Hybopsis

(amblops, storeriana, and harperi) approach or
NUMBER 90

RADII

• PRIMARY

O SECONDARY

• • •

o o

-o°o %.

o o o

0 2 0 40 60 80 100 120 140 160 180 200

STANDARD LENGTH , mm

FIGURE 1.—The relationship of the number of primary and secondary radii with increase in body

length in 91 specimens of S'ocomis micropogon, ranging in size from 16 to 207 mm in Standard

Length (SL).

slightly overlap the lowest individual counts in

Nocomis. Obviously, both higher and lower counts

in all species may be expected with increase in size

of sample, but modal values should remain about

the same.

In order to obtain some idea of ontogenetic differences and variations in number of radii, 91 specimens of N. micropogon ranging from about 16 to

207 mm SL were studied (Figure 1). This slightly

curvilinear relationship shows a fairly uniform increase in the number of primary and secondary radii

with increase in body length and also that, for any

size, the range of variation is moderate. The rate of

increase is greater in primary than in secondary radii.

The number of primary radii in relation to secondary radii was comparatively high in Nocomis

(Figure 1), and the relationship was similar among

all of the species. The average number of primary

radii, in percent of the total number in species of

Nocomis was: effusus 66, raneyi 71, platyrhynchxis 72,

micropogon 75, leptocephalus 15, asper 78, and

biguttatus 80. Mean values for species of Hybopsis

ranged from about 18 to 90 percent.

Cockerell and Allison (1909:161-162) distinguished Nocomis from Hybopsis on the basis of the

broader nuclear scale area in Nocomis, the "apical

radii almost parallel" (i.e., less convergent). We

cannot agree with this distinction. In species of

Hybopsis with high numbers of primary radii, the

convergence of the radii in the nuclear area is generally less than in species with fewer primary radii.

There are also differences in the convergence of the

radii in species with low numbers of radii, some

meeting on a much wider nuclear area than others.

The survey of the total number of radii in 89 species of Notropis (of the 101 species recognized as

valid) showed great differences among the species;

the average values ranged from about 5 to 20. These

values are very similar to those of Hybopsis (mean

values, 5.7 to 19.2) and unlike all species of Nocomis

(mean values, 29.2 to 40.8). Species of the subgenus

Luxilus as well as the Notropis species dorsalis,

anogenus, signipinnis, bairdi, heterolepis, atrocaudalis, petersoni, longirostris, and other Notropis species had mean values from about 13 to 20. Many

species, such as members of the subgenera Cyprinella

and Lythrnrus, had total radii counts ranging only

from about 5 to 13. Our data are in general agreement with many of the radii counts given by Cockerell

and Callaway (1909:190). Both sets of data indicate

the numbers of radii are lower characteristically in

Notropis than in Nocomis.

INTERPRETATION.—Cockerell and Allison (1909:

163) linked Nocomis with Semotilus, Couesius, and

Phenacobius on the basis of several scale characteristics. Phenacobins was certainly misplaced by

Cockerell, if one considers the scale characters he regarded significant. Phenacobius develops few radii

(about 8-15) and these converge broadly in the area

of the focus, whereas in S. atromaculatus and S. cor-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 3.—Total vertebral numbers in Hybopsis, Couesius, and Nocomis

Genus-group

Hybopsis

(Hybopsis)

(Yuriria)

(Erimystax)

(Platygobio)

(Macrhybopsis)

(Extrarius)

(Oregonichthys)

Couesius

Nocomis

species

34

a.amblops

rubrifrons

hypsinotus

labrosa

sp.,cf. labrosa

storeriana

alia

d. dissimilis

disxitnilis harry i

x. x-punctata

x-punctata trail tin an i

i. insignis

insignis eristigma

cahni

monacha

harperi

g. gracilis

gracilis .gracilis x gulonella

gracilis gulonella

gelida

meeki

aestivalis 3

era met i

plumb eus

biguttatus

asper

effusus

platyrhynchus

raneyi

micropogon

leptocephalus

35

6

13

8

36

44

2

2

9

29

18

37

20

20

18

5

1

2

33

26

Total vertebrae

38

2

23

4

32

6

1

12

12

6

4

9

1

1

24

39

1

11

2

12

24

5

14

4

33

4

13

15

1

46

1

2

5

188

40

19

6

20

11

6

6

19

12

1

124

12

10

59

27

168

134

41

1

1

1

8

3

3

18

18

6

16

19

3

47

94

204

278

10

42

3

15

37

14

14

18

1

7

111

70

43 44 45 46

2

1

5 15 6

37 8 2

1

26

2

8 1

poralis the radii are numerous (about 25-40) and

they converge comparatively sharply in the area of

the focus.

The scale morphology of Nocomis, in numbers of

radii and in scale configuration and development,

agrees closely with that of S. atrornaculatus and S.

corporalis. The small-scaled form Semotilns margarita has a moderate number of radii (15-21), but

otherwise its scale morphology generally agrees with

that of Nocomis and Semotilus. Couesius plumbeus

(15-22 total radii) is also close to this relationship in

scale morphology, particularly to S. margarita. The

scales of Ar. biguttatus, N. leptocephalus and N. micropogon were illustrated by Lachner (1952:442,

Figures 1-5); Ar. biguttatus by Lagler (1947, Plate 4,

Figure 21); N. effusus (as N. kentuckiensis) by

Cockerell (1913, Plate-figure 32:6); S. atromaculatus

and S. corporalis by Cockerell and Allison (1909,

Plate 3, Figures 4, 5); and S. atromaculatus by Ellis

(1914, Plate 12, Figure 61).

We present our scale data with a cautious attitude

in regard to its use as a systematic character. The difference, however, between the scale morphology of

Nocomis and that of Hybopsis s.l. and Notropis is of

such magnitude that it cannot be omitted as criteria

in the evaluation of these generic groups. The scale
NUMBER 90

TABLE 4.—Precaudal and caudal vertebral numbers in Hybopsis, Couesius, and Nocomis

Genus-group

Hybopsis

(Hybopsis)

(Yuriria)

(Erimystax)

(Macrhybopsis)

(Extrarius)

(Oregonichthys)

Couesius

Nocomis

species

16 17

a. amblops 2

rubrifrons

hypsinotus

labrosa

sp.,cf. labrosa

storeriana

alta

d. dissimilis

dissimilis harryi

x. x-punctata

x-punctata trautmani

i. insignis

insignis eristigma

cahni

monacha

harperi

gelida

meeki

aestivalis 8 46

crameri

plumbeus

biguttatus

asper

effusus

platyrhynchus

raneyi

micropogon

leptocephalus

18

37

14

1

2

5

4

20

12

Precaudal

19

33

29

15

22

34

1

3

7

24

7

1

38

20

4

12

20

3

8

7

3

8

2

I

8

9

26

3

13

2

3

8

5

1

21

8

36

15

3

2

1

2

10

2

1

2

16

33

5

22

1

13

7

5

15

131

3

2

93

34

324

117

23

18

55

13

37

19

160

61

35

24

1

19

11

16

1

6

10

13

17

22

3

7

4

1

1

10

5

19

87

11

28

4

7

151

108

Caudal

18

41

12

15

8

3

38

6

4

7

4

1

1

29

14

5

7

105

5

28

47

166

221

35

19

9

25

2

27

14

9

11

1

7

8

27

7

9

7

2

26

21

6

2

69

101

43

20

6

16

2

1

5

4

1

1

19

2

6

14

29

7

8

10

21

14

3

1

10

12

3

22 23

1

2

4

4 3

Difference1

PorC

P

C

P

P

P

P

P

P

P

P

P

C

C

C

P

P

P

P

P

P

P

P

P

=

=

1

1

—

1.5

3

2

2

1

1

1

=

1

—

1

1

2

3

1.5

4

4

6

5.5

3

5

4

5

1

 P = precaudal, C = caudal. Region with higher mode, if any, in left column; approximate difference between modal values

of precaudal and caudal vertebrae in right column.

data support our concept that Nocomis is unique

among the Hybopsis-Notropis complex and that its

relationships are close to the Semotilus group.

Vertebral Counts

METHODS.—Total and precaudal counts include

the Weberian vertebrae, counted as four, and urostylar vertebra counted as one. Of the 18 references

cited herein for cyprinid vertebral counts, only 4 excluded the Weberian vertebrae from the total: Olund

and Cross (1961), Al-Rawi and Cross (1964), Cross

(1967, [apparently excluded only for N. biguttatus

and the subspecies of H. (Platygobio) gracilis but

not in the total given for the latter species]), and

Lachner and Jenkins (1967). Their data, before

being used herein (Tables 1, 3, and 4), were adjusted

by adding four for the Weberian complex.

Counts were made from radiographs over a lightbox with an opaque plastic top. For smaller specimens a focusing magnifying lens mounted in a plastic

cylinder was used. A pointer mounted below the

lens in the cylinder facilitated counting.

An attempt was made to minimize possible environmental influence upon the data by using samples obtained from localities scattered throughout

the ranges of the species. This was not possible for

all species. Garside (1966) includes keys to, or sum-
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

maries of, the literature and a critical discussion of

the effect of environment upon vertebral number.

An attempt to distinguish between precaudal and

caudal vertebrae was also made. If a haemal spine

was distinctly longer (by about 25 percent or more),

than an adjacent anterior haemal element, the vertebra on which such a spine occurred was counted as

the first caudal vertebra. This method probably

yields a close approximation of the actual line of

division, which usually occurred in the region of the

tip of the anteriormost anal pterygiophore, the succeeding haemal spines being approximately equal in

length. Such a distinction was not obvious in some

specimens. Thus for some species the total of the

precaudal and caudal counts is less (Table 4) than

the total shown in Table 3. Precaudal and caudal

vertebral counts of American cyprinids appear to

have been published only by Suttkus and his collaborators (see Literature Cited); the present study indicates the importance of their consideration.

In Tables 3 and 4 the data are segregated by currently recognized subspecies except for those of three

species: H. (Extrarius) aestivalis, which shows a

complex pattern of variation (Hubbs and Ortenburger, 1929:23-28; Yerger and Suttkus, 1962:327;

Metcalf, 1966:108-110; Davis and Miller, 1967:16-

20); and H. (Erimystax) harperi and Couesius

plumbeus, in which the recognition of subspecies

was questioned by Howell (1961) and Lindsey (1956:

769-771), respectively. Hybopsis (Hybopsis) labrosa

and H. (Hybopsis) sp. cf. labrosa are closely related

to each other.

RESULTS.—Nocomis is relatively homogeneous with

respect to the number of vertebrae; among the seven

species, the modes for total vertebrae (Table 3)

range from 39 to 41 and the modes for total precaudal and caudal vertebrae (Table 4) range, respectively, from 22 to 23 and from 17 to 19. Total number of vertebrae average higher in Nocomis than in

most Hybopsis s.l., the differences being due to the

higher counts in the precaudal and lower counts in

the caudal regions of Nocomis. The precaudal

counts in Nocomis show complete separation from

11 of the 20 forms of Hybopsis and a high percentage

of separation of specimens from most of the remainder (excluding the single specimen of the 419

N. micropogon examined, with 20 precaudals). In

the 20 forms of Hybopsis the modal values for precaudal and caudal counts are about the same in 5

forms, the precaudal count is lower than the caudal

in 4, and the precaudal is higher in 11 forms (9 species). The magnitudes of the differences between

precaudal and caudal counts are pertinent. In the

11 Hybopsis forms whose precaudal count is higher

than the caudal, the differences between the modes

are about 1 vertebra in 7 forms, 2 in 3 forms, and

3 only in H. (Yuriria) aha. The differences between the precaudal and caudal modes in Nocomis

are greater than 3 in all but one species, 3 in N.

platyrhynchus, 4 in N. biguttatus and N. micropogon, 5 in N. raneyi and N. leptocephalus, 5.5 in

N. effusus, and 6 in N. asper.

With respect to vertebral count, Couesius and

Semotilus have modal values similar to or distinctly

higher than those of Nocomis: The modal value for

Couesius is 42; for the three species of Semotilus the

ranges are 40-42 for S. margarita, 42-44 for S. atromaculatus (42-45 for Kansas material, Cross, 1967),

and 44-46 for S. corporalis. Couesius and Semotilus

have the large precaudal-caudal differences characteristic of Nocomis; the precaudal has modally from

4 to 6 vertebrae more than the caudal in these two

groups. These data again support a possible relationship among Nocomis, Couesius and Semotilus.

A survey was made of vertebral counts published

for American cyprinids. The references and groups

follow: Miller and Hubbs, 1960 (Plagopterini); Miller, 1963 (Gila crassicauda); Snyder, 1908 (Ptychocheilus); Barbour and Contreras, 1968 (Algansea);

Jordan and Evermann, 1896 (Gila, Ptychocheilus,

Orthodon, Pogonichthys, Semotilus); Fingerman

and Suttkus, 1961, and Al-Rawi and Cross, 1964

(Hybognathus); Minckley and Craddock, 1962

(Phenacobius). The following presented counts for

various species or groups of Notropis: Bailey and

Suttkus (1952), Suttkus (1955), Suttkus and Raney

(1955), Suttkus (1959), Bailey and Allum (1962),

Gilbert and Bailey (1962), Gilbert (1964), Snelson

(1968), Suttkus and Clemmer (1968). Cross (1967)

gave the range for all cyprinids of Kansas, presumably based upon Kansas specimens, thus making

available counts for six groups in addition to those

listed above: Notemigonus, Chrosomus, Opsopoeodus, Dionda, Pimephales, and Campostoma. Counts

for Hemitremia, Semotilus, certain species of Phenacobius, and the Notropis boucardi group were obtained by the authors.

Distinctive specific differences exist between some
NUMBER 90

species of Hybopsis, and there also appear to be differences between the subspecies of Hybopsis (Eimystax) dissimilis. Of particular interest is H.

(Platygobio) gracilis, in which two subspecies were

recognized by Olund and Cross (1961) on the basis

of several characters in addition to the vertebrae

numbers. Bailey and Allum (1962:44-45) did not

recognize these subspecies, addressing their remarks

apparently only to the difference in the number of

vertebrae. They gave counts for 476 specimens from

South Dakota, and their interpretation of the data

was that these differences in number of vertebrae

strongly indicated environmental modification. This

may be true, as is evidenced by the fact that the

greatest and second greatest ranges in total vertebrae,

7 and 6, respectively, appear in H. gracilis and the

rather polytypic H. aestivalis, species that apparently

occur in the widest range of ecological conditions.

Metcalf (1966:103-104) summarized additional information indicating that differences in vertebral

counts among subspecies of H. gracilis are not solely

due to phenotypic responses to temperature; he also

gave a logical zoogeographic hypothesis concerning

the origin and intergradation of the subspecies.

The vertebrae count of a broad sample of American cyprinids ranges from 33 to 47. Notropis tends

to have lower counts (more than 75 percent of the

34 species surveyed have modal values less than 40),

whereas the elongate Phenacobius, the larger sized

Semotilus, and a number of western groups have

higher counts. Apparent trends in body form and

size, and vertebrae count in percids (Bailey and

Gosline, 1955) seem to be roughly paralleled in

cyprinids.

Scope of the Genus Hybopsis

The characters most frequently used to describe

Hybopsis sl.l. are: (1) pharyngeal teeth in major

row 4-4 (in minor row 0-2); (2) upper jaw protrusile, no frenum; (3) mouth subterminal to inferior;

(4) barbels present, their origin terminal or near

terminal on maxillae; (5) intestine not whorled

(except one species); (6) scale radii restricted to

posterior field (Jordan and Evermann, 1896:314;

Bailey, 1951:216; Moore, 1968:67). To evaluate

these characters we compared Hybopsis with a large

majority of other American cyprinid genera, particularly Notropis. On the basis of our studies and

through reference to the general treatments of Jordan and Evermann (1896) and Moore (1968), we

found that characters 1, 2, 3, 5, and 6, are widely

shared with other cyprinid groups, and (except tor

character 3) by about 50 to 75 percent of the extant

American cyprinid species, so that emphasis upon

any one of these characters, or all in combination, as

a principal support for merging genus-groups under

Hybopsis appears questionable.

As for item 4, the possession of a barbel pendant

from or very close to the lip angle (terminal maxillary position) seems to have been a chief requisite

for inclusion of fishes in Hybopsis. Histologically,

this basically gustatory structure in Hybopsis and

Nocomis is fleshy and lacks osseous, cartilaginous,

and muscular support, according to Davis and Miller

(1967:13), who give (p. 15) for most species, the

number and size of taste buds on barbels. The barbels have, in effect, been the key character for distinguishing between Hybopsis and Notropis, species

lacking these structures being assigned to the latter

genus. However, the barbel in some American

cyprinid species and genus-groups is now thought to

be a variable and labile character. Their occurrence

in certain groups is probably an advanced rather

than primitive condition that has been developed

through several evolutionary pathways. The observed variation in barbels and lips indicates o p

portunity for their selective change as environmental

conditions may warrant. We cite some of the known

cases that support these statements:

(1) In specimens of some species of Hybopsis (in

part, Moore, 1968) and other typically barbeled

groups (e.g., Rhinichthys, Hubbs and Miller, 1948:

16-17; Semotilus, Gould and Brown, 1968, and our

studies; Nocomis and Exoglossum, our studies), the

barbel or barbels are occasionally absent.

(2) In some species of Hybopsis (particularly the

subgenus Erimystax) and at least one other group

(including Phenacobius) a small flap or fold of skin

occurs, extending from beneath the anterior end of

the lachrymal bone to the posterior end of the

maxilla. Sometimes part of this flap is excised or

nearly so, somewhat resembling a Semotilus-type

barbel.

(3) Sometimes there occurs along the ventral edge

of this flap an elongate, apparently compound, taste

bud that resembles a tiny barbel and may function

as such. A series of elongate pendent "barbels" oc-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

curs along the flap in some specimens of Hybupsis

(Erimystax) x-punctata; these were roughly sketched

and termed sensory bristles by Branson (1962).

(4) Occasional specimens of Nocomis, Semotilus,

the subgenus Erimystax, and of species of other

groups were found with a deeply cleft barbel or two

barbels, completely separated, usually small and on

one side of the mouth.

(5) The individuals of certain populations of

Hybopsis (Extrarius) aestivalis typically possess two

well-developed pairs of barbels rather than one

(Hubbs and Ortenburger, 1929:23-28; Yerger and

Suttkus, 1962:327). The evidence presented by the

former authors indicates that the tetranemus condition, unique to aestivalis among American cyprinids,

evolved within this species independently from Old

World groups (and perhaps more than once).

(6) Discrete differences exist in details of barbel

origin and form among species-groups of Hybopsis

s.l., some of which were mentioned by Branson

(1962). These data indicate that barbels may have

arisen independently several times.

(7) Notropis bairdi, a turbid-river dweller, shows

a tendency toward prolongation of the lip angle into

a barbel-like structure whereas its two close relatives

apparently do not show this condition (Cross,

1953:254).

(8) Nuptial males of Pimephales notatus develop a

transitory barbel-like structure derived from tissue

at the lip angle. It apparently is at least partly associated with increase in cephalic epidermal gibbosity

in nuptial males that forces the lip angle downward.

This is the only species of Pimephales known to

possess the structure and was considered on the basis

of trophic, nuptial, and other characters to be the

most specialized in its group (Hubbs and Black,

1947:9-13).

(9) The presence of a barbel in Plagopterus is regarded as an advanced state in the Plagopterini.

This genus seems to be the most highly specialized

plagopterin (Miller and Hubbs, 1960:6-11), most of

its modifications apparently being related to life in

turbid waters. The ancestral plagopterin was suggested by Miller and Hubbs to be a species of Gila,

all known members of which lack a barbel, or "a

similar, relatively unspecialized cyprinid genus."

(10) The possibility of close relationship between

the non-barbeled Notropis amnis and usually barbeled Hybopsis (Hybopsis) amblops winchelli was

suspected by Hubbs (1951:5; and more so in Moore,

1968:72). Study of this possibility by Glenn H.

Clemmer shows that these forms are intimately related: they are regarded as congeneric by Clemmer

and by us.

We conclude that, for groups under consideration

herein, the barbel alone (its occurrence, position,

and morphology) is not necessarily of decisive significance in classification. It is conceivable that a

genus may include both barbeled and non-barbeled

forms. The only current New World examples of

such groups are Notropis, to which the barbeled

boucardi group was recently referred by Cortes

(1968), and Exoglossum, to which we assign the barbeled Parexoglossum.

The presence of barbels in some but not all members of a group or species reduces the systematic

value of this character, although contrary to Cortes

(1968) and the opinions of others, its usage at the

generic level has not been invalidated. Barbels remain important in the determination and delimitation of phyletic lines and generic groups, particularly

when considered with other details of morphology,

sensory functions, and group life history. We concur with a statement by Hubbs and Miller (1948:16-

17) that, "The capacity to develop a barbel... is of

more significance than its actual presence or

absence."

This review of six important diagnostic characters

given for Hybopsis s.l., the first five of which are

trophic ones, reveals that the group is probably insecure; the lack of other cohesive characters, such as

melanophore pigmentation patterns, head and body

physiognomy, nuptial features, and life histories

further substantiates this point. No additional specialized feature is known that is common to most

species. The subgroups of Hybopsis comprise an

extremely heteromorphic assemblage, one that Jordan and Evermann (1896:314) regarded to show

"... a range of variation as wide as that in Notropis,

though with a [considerably] smaller number of species." Hybopsis, at that time, did not include

Platygobio and Couesius. The geographic range of

Hybopsis extends from Arctic Canada southward

into Mexico and from the Atlantic to the Pacific

slope; if this were a natural group it would be the

most wide-ranging American cyprinid genus.

On the basis of several studies in progress, some

tentative summaries concerning relationships among
NUMBER 9 0 11

and within groups of Hybopsis and other cyprinid

genera now can be made. Davis and Miller (1967)

and Reno (1967) restricted their studies to Hybopsis

s.l. and therefore could not determine the wider

systematic significance of their results; their findings

concerning divergence in sensory structures among

and within previously established subgroups, however, usually correlate with studies of other character complexes.

We have already indicated that Couesius probably

is closely related to Semotilus, particularly through

the species Semotilus margarita. The subgenera

Platygobio, Macrhybopsis, and Erimystax are near

allies of each other and are probably closer to

Phenacobius than to Hybopsis s.s. Hybopsis (Hybopsis) storeriana is misplaced in the subgenus Hybopsis and appears to be a close relative of Platygobio.

Hybopsis (Erimystax) harperi is probably referrable

to Notropis. Hybopsis (Hybopsis) labrosa and Hybopsis (Hybopsis) sp., both from southeastern

United States, are related to the Notropis (Cyprinella) group. The relations of the latter four species

with Hybopsis s.s. are distant. Also, Hybopsis s.s. is

probably more closely related to a Notropis lineage

than to the remaining subgenera of Hybopsis s.l.

There is too great a diversity engendered by the

groups just mentioned (including Nocomis and its

closest relatives) to place them into a single genus.

At least three rather distantly related phyletic lines

are represented in Hybopsis s.l., only one of which

seems to have an (apparently advanced) end in

Hybopsis s.s. The eventual dismemberment of Hybopsis s.l. and some of its subgroups, and indication

of their relationship with other genera, will lead to

considerable nomenclatural changes, a necessary reaction to earlier uncritical study and understanding.

Although we are certain that changes will be necessary, we do not recommend nomenclatural alterations until further studies are completed.

Materials

A list of the material of Hybopsis and Couesius used

for vertebral data is given below. The samples of

Nocomis are from localities scattered throughout

the ranges of the species and are reported in papers

by Lachner and Jenkins (1967; 1971a) and Lachner

and Wiley (1971). Numbers in parentheses refer to

the specimens on which counts were made. Abbreviations for institutions are: Cornell University (CU);

University of Kansas (KU); Tulane University (TU);

University of Michigan, Museum of Zoology

(UMMZ); National Museum of Natural History

(USNM). The locality data and identifications for

all series of the Hybopsis (Platygobio) group were

listed by Olund and Cross (1961) or are from their

original data sheets.

Hybopsis (Hybopsis) a. amblops (Rafinesque)

White R. drainage: USNM 42681, Missouri, Texas Co., N. Fk.

White R., 1889 (10); USNM 59310 and 59311, Arkansas,

Washington Co., Mid. Fk. White R. (3 and 5); USNM 59313,

Arkansas, Washington Co., White R. (1); USNM 59312,

Arkansas, Newton Co., Little Buffalo R. (3).

Tennessee R. drainage: USNM 63084, Alabama, Colbert Co.,

Tuscumbia, 1909 (11); USNM 188710, North Carolina, Yancy

Co., Cane Cr., 1964 (5); USNM 40535, North Carolina, Madison Co., Spring Cr., 1888 (6); USNM 194857, Virginia, Smyth

Co., Mid. Fk. Holston R., 1958 (6); USNM 190960, Virginia,

Smyth Co., N. Fk. Holston R., 1959 (5).

Green R. drainage, Kentucky: USNM 107208, Mammoth Cave,

1929 (5); USNM 163099, Metcalfe Co., trib. Russell Cr., 1952

(7); USNM 163101, Adair Co., Russell Cr., 1952 (3); USNM

163100, Casey Co., trib. Green R., 1952 (2).

H. (Hybopsis) rubrifrons (Jordan)

Savannah R. drainage: USNM 165721, Georgia, Madison Co.,

trib. Broad R., 1952 (5); USNM 168122, South Carolina,

Oconee-Pickens Co. line, Keowee R., 1954 (6); USNM 168103,

South Carolina, Oconee Co., Chauga Cr., 1954 (2).

Altamaha R. drainage, Georgia: USNM 165729, Walton Co.,

trib. 1 mi NE Monroe, 1952 (4); TU 26141, Gwinnett Co.,

Yellow R., 1962 (29).

H. (Hybopsis) hypsinotus (Cope)

Pee Dee R. drainage: USNM 104100, Virginia, Carroll Co.,

Lovels Cr., 1937 (10); USNM 162921, North Carolina, Wilkes

Co., trib. Yadkin R., 1952 (1); USNM 166285, North Carolina,

Wilkes Co., trib. Yadkin R., 1947 (1).

Santee R. drainage, South Carolina: USNM 192851, Greenville

Co., Oil Camp Cr., 1964 (5); USNM 162954, Spartanburg Co.,

Fair Forest Cr., 1952 (7).

H. (Hybopsis) labrosa (Cope)

Santee R. drainage: USNM 168129, North Carolina, Iredell

Co., Buffalo Cr., 1956 (2); USNM 192853, South Carolina,

Spartanburg Co., N. Pacolet R., 1954 (2); TU 26207, South

Carolina, Anderson Co., Brushy Cr., 1962 (27).

H. (Hybopsis) sp., cf. labrosa

Pee Dee R. drainage: USNM 162928, North Carolina, Surrey

Co., Mitchell R., 1952 (2); USNM 162929, North Carolina,

Wilkes Co., Bugaboo Cr., 1952 (1); USNM 162251, North

Carolina, Wilkes Co., trib. Yadkin R., 1947 (10); USNM

105075, Virginia, Carroll Co., Ararat R., 1933 (2).

Santee R. drainage: USNM 162356, South Carolina, Greenville
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Co., N. Saluda R., 1947 (10); USNM 67996, North Carolina,

Burke Co., Johns R., and McDowell Co., Bucks Cr., 1888 (7);

USNM 40551, North Carolina, Burke Co., Johns R., 1888 (7).

H. (Hybopsis) storeriana (Kirtland)

Mobile R. drainage: USNM 167997, Alabama, Choctaw Co.,

Tombigbee R., 1955 (7).

Tennessee R. drainage, Alabama: USNM 108552, Lauderdale

Co., Shoal Cr., 1938 (2); USNM 108558, Wheeler Res., 1938

(10).

H. (Yuriria) alta (Jordan)

Mexico: USNM 48905 and 125131, Rio Lerma, Salmanca (5

and 3); USNM 55757, Aguas Calientes (7); USNM 37819 and

41819, Guanajuato (3 and 1); TU 30663, Zacatecas state, Rio

Juchipila, 1963 (32).

H. (Erimystax) d. dissimilis (Kirtland)

Tennessee R. drainage, Virginia: USNM 195832, Smyth Co.,

N. Fk. Holston R., 1958 (8).

Wabash R. drainage, Indiana: USNM 66892, Marshall Co.,

Tippecanoe R., 1890 (6).

Upper Ohio R. drainage, Pennsylvania: USNM 116420, Westmoreland Co., Loyalhaune Cr., 1893 (6); USNM 161903,

Mercer Co., French Cr., 1947 (4).

H. (Erimystax) dissimilis harryi Hubbs and Crowe

White R. drainage, Missouri: CU 24242, Barry Co., White R.,

1956 (5); CU 38345, Stone Co., Indian Cr., 1958 (1); CU

32874, Ripley Co., Current R., 1953 (2).

H. (Erimystax) x. x-punctata Hubbs and Crowe

Ouachita R. drainage, Arkansas: USNM 36428, Clark Co.,

Ouachita R., 1884 (7); USNM 36462, Saline Co., Saline R. (7).

H. (Erimystax) cahni Hubbs and Crowe

Tennessee R. drainage, Tennessee: UMMZ 157709, Claiborne

Co., Powell R., 1939 (4); UMMZ 103462-63 Anderson Co.,

Clinch R., 1936 (4); USNM 70580 and 201454, Clinch R.,

Walkers Ford, 1893 (5 and 6).

H. (Erimystax) monacha (Cope)

Tennessee R. drainage: USNM 36664, Alabama, Lauderdale

Co., Shoal Cr., 1884 (2); USNM 190644, Tennessee, RheaRoane Co. line, White Cr., 1959 (7); USNM 70580, Tennessee,

Clinch R., 1893 (1); USNM 70581, Tennessee, Indian Cr.,

Cumberland Gap (2); USNM 40499, North Carolina, Madison Co., Spring Cr., 1888 (1); USNM 177367, Tennessee,

Hawkins-Sullivan Co. line, N. Fk. Holston R., 1954 (1);

CU 46397, Tennessee, Sullivan Co., N. Fk. Holston R., 1963

(2); USNM 40475, Virginia, Smyth Co., N. Fk. Holston R.,

1888 (3).

H. (Erimystax) harperi (Fowler)

Florida: TU 20358, Wakulla Co., Wakulla R., 1959 (36); USNM

133245, Lake Co., Juniper Springs, 1946 (10).

H. (Platygobio) g. gracilis (Richardson)

UMMZ 166845 (9); KU5127 (3); KU5128 (2); UA uncataloged, Simonette R. (2); UA uncataloged, Peace R. (5);

ROMZ 14500 (5).

H. (Platygobio) gracilis: gracilis x gulonella

KU 1841 (10); MSC 1878 (7); MSC 2021 (6) ; ROMZ 3885 (2) ;

UMMZ 166803 (9); MSC 1960 (5) ; KU 2140 (2); KU 4863

(12); UMMZ 134813 (9); UMMZ 135280 (10); UMMZ

135786 (7); UMMZ 114644 (8); UMMZ 127681 (15).

H. (Platygobio) gracilis gulonella (Cope)

KU 4742 (5); KU 4769 (5); MSC 2010 (5); KU 4266 (5);

KU 4269 (6); KU 3409 (10); KU 4235 (5); WU 2084 (4);

WU 2095 (3).

H. (Macrhybopsis) gelida (Girard)

UMMZ 164849, Illinois, Jackson Co., Mississippi R. (4).

Missouri R. drainage: UMMZ 161909, South Dakota, Pennington Co., Cheyenne R., 1950 (7); USNM 125133, Nebraska,

Hall Co., Platte R., (5); USNM 76049, Wyoming, Sheridan

Co., 1893 (4); USNM 59137 and 174890, Wyoming, Sheridan

Co., Powder R., (10 and 2); USNM 174889 Wyoming, Converse Co., Platte R., (1).

H. (Macrhybopsis) meeki Jordan and Evermann

Missouri R.: USNM 35889, Missouri, Buchanan Co., (1);

KU 9675, Missouri, St. Louis Co., 1963 (10); KU 9783,

Missouri, Lafayette Co., 1963 (8); KU 9688, Missouri, ,St.

Charles Co., 1963 (9); CU 30323, South Dakota, Yankton Co.,

1952 (5); UMMZ 166966, South Dakota, Charles Mix and

Gregory Co., 1952 (7).

H. (Extrarius) aestivalis (Girard)

Mexico: USNM 55727, Nuevo Le6n state, trib. San Fernando

R., (4); CU 24390, San Fernando R., Tamaulipas, 1952 (7).

Colorado R. drainage: USNM 36582, Texas, Travis Co., Rio

Colorado, 1884 (9).

San Antonio R. drainage: USNM 46229, Texas, Comal Co.,

Rio San Marcos (7).

Red R. drainage: USNM 173317, Louisiana, Bossier Parish,

1956, (4); USNM 36342, Arkansas, Hempstead Co., Red R.,

1884 (10).

Ouachita R. drainage, Louisiana: USNM 172285, Union

Parish, Ouachita R., 1956 (2); USNM 172361, 172381, and

172394, Ouachita Parish, Ouachita R., 1956 (2, 2, and 1).

Arkansas R. drainage, Arkansas: USNM 36374, Sebastian Co.,

Arkansas R., 1884 (5); USNM 16136, Oklahoma, Payne Co.,

Cimarron R., 1935 (10).

Missouri R. drainage, Nebraska: USNM 76047 and 125132,

Buffalo Co., 1893 (3 and 5).

Wabash R. drainage, Indiana: CU 32229, Posey Co., Wabash

R., 1958 (7).

Green R. drainage, Kentucky: CU 46651, Green Co., Green R.,

1963 (3).

H. (Oregonichthys) crameri Snyder

Umpqua R. drainage, Oregon: USNM 58116, Douglas Co.,

S. Umpqua R., (1); USNM 58132, S. Umpqua R., (4).

Williamette system of Columbia R. drainage, Oregon: UMMZ

166728, Lane Co., Cottage Grove Reservoir, 1950 (7);
NUMBER 90 13

CU 20988 and 31026, Benton Co., McFaddens Pond, 1946 and

1951 (4 and 3); CU 43623, Benton Co., Willamette R.,

1952 (7).

Couesius plumbeus (Agassiz)

Alberta: USNM 125198 (4).

Saskatchewan R. drainage, Montana: USNM 104725, Upper

Waterton L., 1934 (7).

L. Superior drainage, Michigan: USNM 193063, Houghton

Co., Otter R., 1953 (7).

Missouri R. drainage, Idaho: USNM 125200, Bonner Co., Pend

Oreille R., 1892 (1).

Labrador: USNM 165435, Northwest R., 1951 (3).

Maine: USNM 57004, Telos L. (9).

Literature Cited

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1833. Recherches sur les poissons fossiles. Neuchatel, 1833-

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1964. Variation in the Plains Minnow, Hybognathus placitus Girard. Transactions of the Kansas Academy of

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Bailey, R. M.

1951. A Checklist of the Fishes of Iowa with Keys for Identification. Pages 187-238 in Iowa Fish and Fishing, by

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1962. Fishes of South Dakota. Miscellaneous Publications,

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1955. Variation and Systematic Significance of Vertebral

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1952. Notropis signipinnis, a New Cyprinid Fish From

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1968. Algansea monticola, a New Cyprinid Fish From the

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1962. Observations on the Breeding Tubercles of Some

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1966. Developmental Rate and Vertebral Number in Salmonids. Journal of the Fisheries Research Board of

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14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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1971b. Systematics, Distribution and Evolution of the

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with a description of a new species from the Ozark

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NUMBER 9 0 15

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1968. Freshwater Fishes of Alabama. Auburn University

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