[Cover]

 

University of Kansas Publications

Museum of Natural History

---

Volume 13, No. 9, pp. 359-427, pls. 26-30, 3 figs.

  August 11, 1961  

 

 

 

Fish Populations, Following a Drought,

In the Neosho and Marais des Cygnes Rivers

of Kansas

 

 

BY



JAMES EVERETT DEACON





(Joint Contribution from the State Biological Survey and

the Forestry, Fish, and Game Commission)

 

 

 

 

University of Kansas

Lawrence

1961

 

 

[Pg 359]

 

University of Kansas Publications

Museum of Natural History

---

Volume 13, No. 9, pp. 359-427, pls. 26-30, 3 figs.

  August 11, 1961  

 

 

 

Fish Populations, Following a Drought,

In the Neosho and Marais des Cygnes Rivers

of Kansas

 

 

BY

 

JAMES EVERETT DEACON



(Joint Contribution from the State Biological Survey and

the Forestry, Fish, and Game Commission)

 

 

 

 

 

 

University of Kansas

Lawrence

1961

[Pg 360]

University of Kansas Publications, Museum of Natural History



Editors: E. Raymond Hall, Chairman, Henry S. Fitch,

Robert W. Wilson

 

Volume 13, No. 9, pp. 359-427, pls. 26-30, 3 figs.



Published August 11, 1961

 

 

University of Kansas

Lawrence, Kansas

 

 

PRINTED IN

THE STATE PRINTING PLANT

TOPEKA, KANSAS

1961



28-7576

[Pg 361]

Fish Populations, Following a Drought,

In the Neosho and Marais des Cygnes Rivers

of Kansas

BY

JAMES EVERETT DEACON

CONTENTS

[Pg 362]

TABLES

[Pg 363]

INTRODUCTION

This report concerns the ability of fish-populations in the Neosho
and Marais des Cygnes rivers in Kansas to readjust to continuous
stream-flow following intermittent conditions resulting from the
severest drought in the history of the State.

The variable weather in Kansas (and in other areas of the Great
Plains) markedly affects its flora and fauna. Weaver and Albertson
(1936) reported as much as 91 per cent loss in the basal prairie
vegetative cover in Kansas near the close of the drought of the
1930's. The average annual cost (in 1951 prices) of floods in
Kansas from 1926 to 1953 was $35,000,000. In the same period the
average annual loss from the droughts of the 1930's and 1950's was
$75,000,000 (in 1951 prices), excluding losses from wind- and soil-erosion.
Thus, over a period of 28 years, the average annual flood-losses
were less than one-half the average annual drought-losses
(Foley, Smrha, and Metzler, 1955:9; Anonymous, 1958:15).

Weather conditions in Kansas from 1951 to 1957 were especially
noteworthy: 1951 produced a bumper crop of climatological events
significant to the economy of the State. Notable among these were:
Wettest year since beginning of the state-wide weather records in
1887; highest river stages since settlement of the State on the
Kansas River and on most of its tributaries, as well as on the Marais
des Cygnes and on the Neosho and Cottonwood. The upper
Arkansas and a number of smaller streams in western Kansas also
experienced unprecedented flooding (Garrett, 1951:147). This
period of damaging floods was immediately followed by the driest
five-year period on record, culminating in the driest year in 1956
(Garrett, 1958:56). Water shortage became serious for many
communities. The Neosho River usually furnishes adequate quantities
of water for present demands, but in some years of drought all
flow ceases for several consecutive months. In 1956-'57, the city
of Chanute, on an emergency basis, recirculated treated sewage for
potable supply (Metzler et al., 1958). The water shortage in many
communities along the Neosho River became so serious that a joint
project to pump water from the Smoky Hill River into the upper
Neosho was considered, and preliminary investigations were made.
If the drought had continued through 1957, this program might
have been vigorously promoted. Data on stream-flow in the Neosho
and Marais des Cygnes (1951-'59) are presented in Tables 1-4.

These severe conditions provided a unique opportunity to gain
insight into the ability of several species of fish to adjust to marked
[Pg 364]
changes in their environment. For this reason, and because of a
paucity of information concerning stream-fish populations in Kansas,
the study here reported on was undertaken.

Table 1. Stream-flow in Cubic Feet per Second, Neosho River near
Council Grove, Kansas. Drainage Area: 250 Square Miles

Table 2. Stream-flow in Cubic Feet per Second, Neosho River near
Parsons, Kansas. Drainage Area: 4905 Square Miles.

Table 3. Stream-flow in Cubic Feet per Second, Marais des Cygnes
River Near Ottawa, Kansas. Drainage Area: 1,250 Square Miles.

[Pg 365]

Table 4. Stream-flow in Cubic Feet per Second, Marais des Cygnes
River at Trading Post, Kansas. Drainage Area: 2,880 Square Miles.

[Pg 366]

DESCRIPTION OF NEOSHO RIVER

The Neosho River, a tributary of Arkansas River, rises in the
Flint Hills of Morris and southwestern Wabaunsee counties and
flows southeast for 281 miles in Kansas, leaving the state in the
extreme southeast corner (Fig.
1). With its tributaries (including
Cottonwood and Spring rivers)
the Neosho drains 6,285
square miles in Kansas and enters
the Arkansas River near
Muskogee, Oklahoma (Schoewe,
1951:299). Upstream from its
confluence with Cottonwood
River, the Neosho River has an
average gradient of 15 feet per
mile. The gradient lessens rapidly
below the mouth of the Cottonwood,
averaging 1.35 feet
per mile downstream to the State
line (Anonymous, 1947:12). The
banks of the meandering, well-defined
channel vary from 15 to
50 feet in height and support a
deciduous fringe-forest. The
spelling of the name originally
was "Neozho," an Osage Indian word signifying "clear water"
(Mead, 1903:216).

Fig. 1. Neosho and Marais des Cygnes
drainage systems. Dots and circles indicate collecting-stations.

Neosho River, Upper Station.—Two miles north and two miles
west of Council Grove, Morris County, Kansas (Sec. 32 and 33, T.
15 S., R. 8 E.) (Pl. 28, Fig. 2, and Pl. 29, Fig. 1). Width 20 to 40
feet, depth to six feet, length of study-area one-half mile (one
large pool plus many small pools connected by riffles), bottom of
mud, gravel, and rubble. Muddy banks 20 to 30 feet high.

According to H. E. Bosch (landowner) this section of the river
dried completely in 1956, except for the large pool mentioned
above. This section was intermittent in 1954 and 1955; it again
became intermittent in the late summer of 1957 but not in 1958 or
1959.

A second section two miles downstream (on land owned by Herbert
White) was studied in the summer of 1959 (Sec. 3 and 10,
[Pg 367]
T. 16 S., R. 8 E.) (Pl. 29, Fig. 2 and Pl. 30, Figs. 1 and 2). This
section is 20 to 60 feet in width, to five feet in depth, one-half mile
in length (six small pools with intervening riffles bounded upstream
by a low-head dam and downstream by a long pool), having
a bottom of gravel, rubble, bedrock, and mud, and banks of
mud and rock, five to 20 feet in height.

Neosho River, Middle Station.—One mile east and one and one-half
miles south of Neosho Falls, Woodson County, Kansas (Sec.
3 and 4, T. 24 S., R. 17 E.) (Pl. 26, Fig. 1). Width 60 to 70 feet,
depth to eleven feet, length of study-area two miles (four large
pools with connecting riffles), bottom of mud, gravel and rock.
Mud and rock banks 30 to 40 feet high.

According to Floyd Meats (landowner) this section of the river
was intermittent for part of the drought.

Neosho River, Lower Station.—Two and one-half miles west,
one-half mile north of Saint Paul, Neosho County, Kansas (Sec.
16, T. 29 S., R. 20 E.). Width 100 to 125 feet, depth to ten feet,
length of study-area one mile (two large pools connected by a
long rubble-gravel riffle), bottom of mud, gravel, and rock. Banks,
of mud and rock, 30 to 40 feet high (Pl. 26, Fig. 2).

This station was established after one collection of fishes was
made approximately ten miles upstream (Sec. 35, T. 28 S., R. 19 E.).
The second site, suggested by Ernest Craig, Game Protector, provided
greater accessibility and a more representative section of
stream than the original locality.

DESCRIPTION OF MARAIS DES CYGNES RIVER

The Marais des Cygnes River, a tributary of Missouri River,
rises in the Flint Hills of Wabaunsee County, Kansas, and flows
generally eastward through the southern part of Osage County
and the middle of Franklin County. The river then takes a southeasterly
course through Miami County and the northeastern part
of Linn County, leaving the state northeast of Pleasanton. With
its tributaries (Dragoon, Salt, Pottawatomie, Bull and Big Sugar
creeks) the river drains 4,360 square miles in Kansas (Anonymous,
1945:23), comprising the major part of the area between the watersheds
of the Kansas and Neosho rivers. The gradient from the
headwaters to Quenemo is more than five feet per mile, from
Quenemo to Osawatomie 1.53 feet per mile, and from Osawatomie
to the State line 1.10 feet per mile (Anonymous, 1945:24). The
total length is approximately 475 miles (150 miles in Kansas). The
[Pg 368]
river flows in a highly-meandering, well-defined channel that has
been entrenched from 50 to 250 feet (Schoewe, 1951:294). "Marais
des Cygnes" is of French origin, signifying "the marsh of the swans."

Marais des Cygnes River, Upper Station.—One mile south and
one mile west of Pomona, Franklin County, Kansas (Sec. 12, T.
17 S., R. 17 E.) (Pl. 27, Fig. 1). Width 30 to 40 feet, depth to
six feet, length of study-area one-half mile (three large pools with
short connecting riffles), bottom of mud and bedrock. Mud banks
30 to 40 feet high.

According to P. Lindsey (landowner) this section of the river
was intermittent for most of the drought. Flow was continuous
in 1957, 1958 and 1959.

There are four low-head dams between the upper and middle
Marais des Cygnes stations.

Marais des Cygnes River, Middle Station.—One mile east of
Ottawa, Franklin County, Kansas (Sec. 6, T. 17 S., R. 20 E.) (Pl.
27, Fig. 2). Width 50 to 60 feet, depth to eight feet, length of study-area
one-half mile (one large pool plus a long riffle interrupted by
several small pools), bottom of mud, gravel, and rock. Mud and
sand banks 30 to 40 feet high.

This section of the river was intermittent for much of the drought.
In the winter of 1957-'58 a bridge was constructed over this station
as a part of Interstate Highway 35. Because of this construction
many trees were removed from the stream-banks, the channel was
straightened, a gravel-bottomed riffle was rerouted, and silt was
deposited in a gravel-bottom pool.

Marais des Cygnes River, Lower Station.—At eastern edge of
Marais des Cygnes Wildlife Refuge, Linn County, Kansas (Sec.
9, T. 21 S., R. 25 E.). Width 80 to 100 feet, depth to eight feet,
length of study-area one-half mile (one large pool plus a long
riffle interrupted by several small pools), bottom of mud, gravel,
and rock. Mud banks 40 to 50 feet high.

This section of the river ceased to flow only briefly in 1956.

METHODS

Electrical Fishing Gear

The principal collecting-device used was a portable (600-watt,
110-volt, A. C.) electric shocker carried in a 12-foot aluminum
boat. Two 2 × 2-inch wooden booms, each ten feet long,
were attached to the front of the boat in a "V" position so they
[Pg 369]
normally were two feet above the surface of the water. A
nylon rope attached to the tips of the booms held them ten feet
apart. Electrodes, six feet long, were suspended from the tip and
center of each boom, and two electrodes were suspended from the
nylon rope. The electrodes extended approximately four feet into
the water. Of various materials used for electrodes, the most satisfactory
was a neoprene-core, shielded hydraulic hose in sections
two feet long. These lengths could be screwed together, permitting
adjustment of the length of the electrodes with minimum effort. At
night, a sealed-beam automobile headlight was plugged into a six-volt
D. C. outlet in the generating unit and a Coleman lantern was
mounted on each gunwale to illuminate the area around the bow
and along the sides of the boat (Pl. 3a). In late summer, 1959, a
230-volt, 1500-watt generating unit, composed of a 115-volt, 1500-watt
Homelite generator was used. It was attached to a step-up
transformer that converted the current to 230 volts. The same
booms described above were used with the 230-volt unit, with
single electrodes at the tip of each boom.

A 5.5-horsepower motor propelled the boat, and the stunned fish
were collected by means of scap nets. Fishes seen and identified
but not captured also were recorded. On several occasions fishes
were collected by placing a 25-foot seine in the current and shocking
toward the seine from upstream.

The shocker was used in daylight at all six stations in the three
years, 1957-'59. Collections were made at night in 1958 and 1959
at the middle Neosho station and in 1959 at the lower Neosho
station.

Seines

Seines of various lengths (4, 6, 12, 15, 25 and 60 feet), with
mesh-sizes varying from bobbinet to one-half inch, were used. The
4-, 12-, and 25-foot seines were used in the estimation of relative
abundance by taking ten hauls with each seine, recording all species
captured in each haul, and making a total count of all fish captured
in two of the ten hauls. The two hauls to be counted were chosen
prior to each collection from a table of random numbers. Additional
selective seining was done to ascertain the habitats occupied
by different species.

Trap, Hoop, and Fyke Nets.—Limited use was made of unbaited
trapping devices: wire traps 2.5 feet in diameter, six feet long,
covered with one-inch-mesh chicken wire; hoop nets 1.5 feet to three[Pg 370]
feet in diameter at the first hoop with a pot-mesh of one inch; and a
fyke net three feet in diameter at the first hoop, pot-mesh of one
inch with wings three feet in length. All of these were set parallel
to the current with the mouths downstream. The use of trapping
devices was abated because data obtained were not sufficient to
justify the effort expended.

Gill Nets

Gill-netting was done mostly in 1959 at the lower Neosho station.
Use of gill nets was limited because frequent slight rises in the
river caused nets to collect excessive debris, with damage to the
nets.

Gill nets used were 125 feet long, six feet deep, with mesh sizes
of ¾ inch to 2½ inches. Nets, weighted to sink, were placed at right
angles to the current and attached at the banks with rope.

Sodium Cyanide

Pellets of sodium cyanide were used infrequently to collect fish
from a moderately fast riffle over gravel bottom that was overgrown
with willows, making seining impossible. The pellets were
dissolved in a small amount of water, a seine was held in place,
and the cyanide solution was introduced into the water a short
distance upstream from the seine, causing incapacitated fish to
drift into the seine. Most of these fish that were placed in uncontaminated
water revived.

Rotenone

Rotenone was used in a few small pools in efforts to capture
complete populations. This method was used to check the validity
of other methods, and to reduce the possibility that rare species
would go undetected. Rotenone was applied by hand, and applications
were occasionally supplemented by placing rotenone in a
container that was punctured with a small hole and suspended
over the water at the head of a riffle draining into the area being
poisoned. This maintained a toxic concentration in the pool for
sufficient time to obtain the desired kill. Rotenone acts more slowly
than cyanide, allowing more of the distressed fish to rise to the surface.

Dyes

Bismark Brown Y was used primarily at the upper Neosho station
to stain large numbers of small fish. The dye was used at a
dilution of 1:20,000. Fishes were placed in the dye-solution for[Pg 371]
three hours, then transferred to a live-box in midstream for variable
periods (ten minutes to twelve hours) before release.

Determination of Abundance

In the accounts of species that follow, the relative terms "abundant,"
"common," and "rare" are used. Assignment of one of these
terms to each species was based on analysis of data that are presented
in Tables 9-16, (pages 402, 403, 404, 405, 408, 410, 411, 414-415,
and 416). The number of fish caught per unit of effort with
the shocker (Table 10) and with seines (Table 11) constitute the
main basis for statements about the abundance of each species at all
stations except the upper Neosho station. Species listed in each Table
(10 and 11) are those that were taken consistently by the method
specified in the caption of the table; erratically, but in large numbers
at least once, by that method; and those taken by the method specified
but not the other method.

For the species listed in Table 10, the following usually applies:
abundant=more than three fish caught per hour; common=one
to three fish caught per hour; rare=less than one fish caught per
hour.

Tables 12-16 list all fish obtained at the upper Neosho station
by means of the shocker, seines, and rotenone.

Names of Fishes

Technical names of fishes are those that seem to qualify under
the International Rules of Zoological Nomenclature. Vernacular
names are those in Special Publication No. 2 (1960) of the American
Fisheries Society, with grammatical modifications required for
use in the University of Kansas Publications, Museum of Natural
History.

ANNOTATED LIST OF SPECIES

Lepisosteus osseus (Linnaeus)

Long-nosed Gar

The long-nosed gar was abundant at the lower and middle Neosho
stations and the lower Marais des Cygnes station. Numbers increased
slightly in the period of study, probably because of increased,
continuous flow. The long-nosed gar was not taken at
the upper Neosho station. At lower stations the fish occurred in
many habitats, but most commonly in pools where gar often were[Pg 372]
seen with their snouts protruding above the water in midstream.
Gar commonly lie quietly near the surface, both by day and by
night, and are therefore readily collected by means of the shocker.
Twice, at night, gar jumped into the boat after being shocked.

Young-of-the-year were taken at the middle and lower stations
on both the Neosho and Marais des Cygnes rivers, and all were near
shore in quiet water. Many young-of-the-year were seined at the
lower Neosho station on 18 June 1959, near the lower end of a
gravel-bar in a small backwater-area having a depth of one to three
inches, a muddy bottom, and a higher temperature than the mainstream.
Forty-three of these young gar averaged 2.1 inches in total
length (T.L.).

Comparison of sizes of long-nosed gar taken by means of the
shocker and gill nets at the lower and middle Neosho stations revealed
that: the average size at each station remained constant
from 1957 to 1959; the average size was greater at the lower than at
the middle station; and, with the exception of young-of-the-year,
no individual shorter than 13 inches was found at the middle station
and only one shorter than 16 inches was taken at the lower
station (Table 5).

Table 5. Numbers and Sizes of Long-nosed Gar Captured by Shocker
and Gill Nets at the Middle and Lower Neosho Stations in 1957, 1958
and 1959.

Because collecting was intensive and several methods were used,
I think that the population of gars was sampled adequately. Wallen
(Fishes of the Verdigris River in Oklahoma, 1958:29 [mimeographed
copy of dissertation, Oklahoma State University]) took
large individuals in the mainstream of the Verdigris River in Oklahoma
and small specimens from the headwaters of some tributaries.
Because I took young-of-the-year at the lower Neosho station,
it is possible that long-nosed gar move upstream when small
and then slowly downstream to the larger parts of rivers as the fish
increase in size. This pattern of size-segregation, according to size
of river, merits further investigation.[Pg 373]

Ripe, spent, and immature long-nosed gar (38 males and 10 females)
were taken in three gill nets, set across the channel, 150
to 500 yards below a riffle, at the lower Neosho station on June 16,
17, and 18, 1959. On 23 June, 1959, 12 males and two females were
taken in gill nets set 50, 150, and 400 yards above the same riffle.
Operations with the shocker between 24 June and 10 July, 1959,
yielded 29 males and three females. The fish were taken from
many kinds of habitat in a three-mile section of the river.

Direction of movement as recorded from gill nets shows that of
67 gar taken, 45 had moved downstream and 22 upstream into the
nets. Only ten of the above gar were taken from the nets set above
the riffle; six of the ten were captured as they moved downstream
into the nets.

On one occasion I watched minnows swimming frantically about,
jumping out of the water, and crowding against the shore, presumably
to avoid a long-nosed gar that swam slowly in and out of
view. I have observed similar activity when gar fed in aquaria.
Stomachs of a few gar from the Neosho River were examined and
found to contain minnows and some channel catfish.

Long-nosed gar have a relatively long life span (Breder, 1936).
This longevity and their ability to gulp air probably insure excellent
survival through periods of adverse conditions. The population of
long-nosed gar probably would not be drastically affected even in
the event of a nearly complete failure of one or two successive
hatches. Maturity is attained at approximately 20 inches, total
length.

Collections at the middle Neosho station in 1958 indicate that
the long-nosed gar is more susceptible to capture at night than in
daytime (Table 9, p. 402).

Lepisosteus platostomus Rafinesque

Short-nosed Gar

Only one short-nosed gar was taken in 1957, at the lower station
on the Neosho River. In 1958 this species was taken at the lower
station on the Marais des Cygnes and in 1958 and 1959 at the lower
and middle stations on the Neosho. More common in the Neosho
than the Marais des Cygnes, L. platostomus occurs mainly in large
streams and never was taken in the upper portions of either river.
Although short-nosed gar were about equally abundant at the
middle and lower stations on the Neosho, the average size was[Pg 374]
greater at the lower station (Table 6). This kind of segregation
by size is shared with long-nosed gar, and was considered in the
discussion of that species. Short-nosed gar were taken only in quiet
water. Both species were collected most efficiently by means of gill
nets and shocker. While shocking, I saw many gar only momentarily,
as they appeared at the surface, and specific identification
was impossible. The total of all gar seen while shocking indicated
that gar increased in abundance from 1957 to 1959 (see Tables 5
and 6). Judging from the gar that were identified, the increase
was more pronounced in short-nosed gar than in long-nosed gar.

At the lower Neosho station in 1959, two ripe females and one
spent female were taken in gill nets (16, 23 and 17 June, respectively)
and were moving downstream when caught. No males
were taken in the nets. Subsequently, by means of the shocker
(26 June-8 July), two spent and two ripe males were captured
in quiet water of the mainstream that closely resembled areas in
which the gill nets were set. No females were taken by means
of the shocker.

Table 6. Numbers and Sizes of Short-nosed Gar Captured by Shocker
and Gill Nets at the Middle and Lower Neosho Stations in 1958 and
1959.

Dorosoma cepedianum (LeSueur)

Gizzard Shad

Gizzard shad declined in abundance from 1957 to 1959. The
largest population occurred at the middle station on the Marais des
Cygnes in 1957. Shad were mainly in quiet water; often, when the
river-level was high, I found them predominately in backwaters
or in the mouths of tributary streams. Examination of nine individuals,
ranging in size from seven inches to 13.5 inches T. L., indicated
that maturity is reached at 10 to 11 inches T. L. Spawning
probably occurred in late June in 1959 ("ripe" female caught on
26 June); young-of-the-year were first recorded in mid-July.

Cycleptus elongatus (LeSueur)

Blue Sucker

The blue sucker was taken rarely in the Neosho River and not at
all in the Marais des Cygnes in my study. Cross (personal com[Pg 375]munication)
obtained several blue suckers in collections made in
the mainstream of the Neosho River in 1952; both young and adults
occupied swift, deep riffles. The species seemingly declined in
abundance during the drought, and at the conclusion of my study
(1959) had not regained the level of abundance found in 1952.

Ictiobus cyprinella (Valenciennes)

Big-mouthed Buffalo

Big-mouthed buffalo were found in quiet water at all stations,
but were rare. A ripe female, 21.5 inches long, was taken at the
lower station on the Neosho on 16 June, 1959.

Ictiobus niger (Rafinesque)

Black Buffalo

and

Ictiobus bubalus (Rafinesque)

Small-mouthed Buffalo

Black buffalo were not taken at the upper station on the Neosho
and were rare at other stations. Small-mouthed buffalo were taken
at all stations and were common in the lower portions of the two
streams. While the shocker was being used, buffalo were often
seen only momentarily, thereby making specific identification impossible;
both species were frequently taken together, and for this
reason are discussed as a unit. Both species maintained about
the same level of abundance throughout my study.

The two species were taken most often in the deeper, swifter
currents of the mainstream, but were sometimes found in pools,
creek-mouths and backwaters. On several occasions in the summer
of 1959, buffalo were seen in shallow parts of long, rubble riffles,
with the dorsal or caudal fins protruding above the surface. Ernest
Craig, game protector, said buffalo on such riffles formerly provided
much sport for gig-fishermen. He stated that the best catches
were made at night because the fish were less "spooky" then than
in daytime. In my collections made by use of the shocker, buffalo
were taken more frequently at night (Table 9, p. 402).

On 19 June, 1959, I saw many buffalo that seemed to be feeding
as they moved slowly upstream along the bottom of a riffle. The
two species, often side by side, were readily distinguishable underwater.
Small-mouthed buffalo appeared to be paler (slate gray)
and more compressed than the darker black buffalo. To test the[Pg 376]
reliability of underwater identifications, I identified all individuals
prior to collection with a gig. Correct identification was made
of all fish collected on 19 June. The smallest individual obtained
in this manner was 18.5 inches T. L. On 26 August, 1959, 16 small-mouthed
buffalo were captured and many more were seen while
the shocker was in use in the same riffle for one hour and ten minutes.
One small-mouthed buffalo was caught while the shocker
was being used in the pool below that riffle for one hour and
fifty minutes. No black buffalo were taken on 26 August.

Spawning by buffalo was not observed but probably occurred
in spring; all mature fish in my earliest collections (mid-June of
each year) were spent. Small-mouthed buffalo reach maturity at
approximately 14 inches T. L.

Carpiodes carpio carpio (Rafinesque)

River Carpsucker

River carpsucker were abundant throughout the study at all
stations. Adults were taken most frequently in quiet water, but
depth and bottom-type varied. The greatest concentrations occurred
in mouths of creeks during times of high water; occasionally,
large numbers were taken in a shallow backwater near the head of
a riffle at the middle Neosho station. River carpsucker feed on
the bottom but seem partly pelagic in habit. They were taken readily
by means of the shocker and gill nets at all depths. The population
of C. carpio in the Neosho River probably was depleted by
drought, although many individuals survived in the larger pools.

When stream-flow was restored, carpsucker probably moved rapidly
upstream but had a scattered distribution in 1957. Trautman
(1957:239) states that in the Scioto River, Ohio, river carpsucker
moved upstream in May and downstream in late August and early
September. Numbers found at the middle and lower Neosho stations
suggest similar movements in the Neosho River in 1957. In
midsummer they were common at the middle station but rare at
the lower station; however, they became abundant at the lower
station in November. The abundance in late fall at the lower
Neosho station might have resulted either from downstream migration
or from continued upstream movement into thinly populated
areas. No indication of seasonal movement was found in 1958 or
1959.

River carpsucker reach maturity at approximately 11 inches T. L.,[Pg 377]
and spawning occurs in May or June. A ripe male was taken from
a gravel-bottomed riffle, three
feet deep, at the middle station
on the Neosho station on 10 June
1959.

Fig. 2. Length-frequency of river
carpsucker in the Neosho River, 1958 and 1959.

The size-distribution of individuals
taken at the middle Neosho
station is presented in Fig.
2. The collection in early July
of 1958 indicates that one size-group
(probably the 1957 year-class)
had a median length of
approximately seven inches. The
modal length of this group was
nine inches in June, 1959. A
second, predominant size-group
(Fig. 2) seemed to maintain almost
the same median size
throughout all the collection periods,
although specimens taken
in the spring of 1959 were
slightly smaller than those obtained
in 1958. This apparent
stability in size may have been
due to an influx of the faster-growing
individuals from a
smaller size-group, coupled with mortality of most individuals more
than 14 inches in length.

Young-of-the-year were taken at every station. Extensive seining
along a gravel bar at the lower Neosho station indicated that the
young are highly selective for quiet, shallow water with mud bottom.
In these areas, young-of-the-year carpsucker were often
the most abundant fish.

River carpsucker were collected more readily by use of the
shocker after dark than in daylight (Table 9, p. 402).

Carpiodes velifer (Rafinesque)

High-finned Carpsucker

A specimen of Carpiodes velifer taken at the lower station on the
Neosho in 1958 provided the only record of the species in Kansas
since 1924. Many specimens, now in the University of Kansas[Pg 378]
Museum of Natural History, were taken from the Neosho River
system by personnel of the State Biological Survey prior to 1912.
The species has declined greatly in abundance in the past 50 years.

Moxostoma aureolum pisolabrum Trautman

Short-headed Redhorse

The short-headed redhorse occurred at all stations. It was
common at the middle and lower stations on the Neosho, rare at
the upper station on the Neosho, abundant at the upper station on
the Marais des Cygnes in 1957, and rare thereafter at all stations
on the Marais des Cygnes. Short-headed redhorse typically occur
in riffles, most commonly at the uppermost end where the water
flows swiftly and is about two feet deep. An unusually large concentration
was seen on 13 June, 1959, in shallow (six inches), fast
water over gravel bottom at the middle station on the Neosho River.

Thirty-nine individuals were marked by clipping fins at the
middle Neosho station in 1959. Four were recovered from one to
48 days later: two at the site of original capture (one 48 days
after marking), one less than one-half mile downstream, and one
about one mile downstream from the original site of capture.

At the middle Neosho station in 1958, this species was taken more
readily by use of the shocker at night than by day (Table 9, p. 402).

Moxostoma erythrurum (Rafinesque)


Golden Redhorse

The golden redhorse was abundant at the upper Neosho station,
rare at the middle Neosho station, and did not occur in collections
at other stations. This species was taken most frequently over
gravel- or rubble-bottoms in small pools below riffles, and was
especially susceptible to collection by means of the shocker.

Twenty-nine golden redhorse of the 1957 year-class, taken at
the upper Neosho station on 9 September 1958, were 6.2 to 8.6
inches in total length (average 7.4 inches); 26 individuals of the
same year-class caught on 21 August 1959 were 9.3 to 13.5 inches in
total length (average 10.9 inches).

Cyprinus carpio Linnaeus


Carp

The carp decreased in abundance from 1957 to 1959 at the upper
and middle Marais des Cygnes station and at the middle and lower
Neosho stations. Carp were more abundant in the Marais des[Pg 379]
Cygnes than in the Neosho, although the largest number in any
single collection was found in one pool at the upper Neosho station
in 1958.

Carp were taken most commonly in quiet water near brush or
other cover. At the middle Neosho station, collecting was most
effective between the hours of 6:30 a.m. and 12:30 p.m. and
least effective between 12:30 p.m. and 6:30 p.m. (Table 9, p. 402).
Ripe males were taken as early as 19 April (16.1 inches, 19.4 inches
T. L.) and as late as 30 July (16 inches T. L.) at the middle Neosho
station. Ripe females were taken as early as 19 April at the middle
Neosho station (19.2 inches T. L.) and as late as 7 July at the
lower Neosho station (16 inches T. L.). Young-of-the-year were
taken first at the middle Marais des Cygnes on 8 July 1957. They
were recorded on later dates at the upper Marais des Cygnes and
at the lower and middle Neosho stations.

Notemigonus crysoleucas (Mitchill)


Golden Shiner

The golden shiner was taken rarely at the upper Marais des
Cygnes station in 1958 and 1959 and at the middle Marais des
Cygnes station in 1957 and 1958. At the middle Neosho station
Notemigonus was seined from a pond that is flooded frequently
by the river, but never was taken in the mainstream.

Semotilus atromaculatus (Mitchill)


Creek Chub

The creek chub was taken only at the upper stations on both
rivers. It increased in abundance at the upper Neosho station from
1957 to 1959, and was not taken in the upper Marais des Cygnes
until 1959.

Hybopsis storeriana (Kirtland)


Silver Chub

A single specimen from the lower Marais des Cygnes station
provides the only record of the species from the Marais des Cygnes
system in Kansas, and is the only silver chub that I found in either
river in 1957-1959. The species is taken often in the Kansas and
Arkansas rivers.[Pg 380]

Hybopsis x-punctata Hubbs and Crowe


Gravel Chub

The gravel chub, present only at the lower and middle Neosho
stations, occupied moderate currents over clean (free of silt) gravel
bottom. The gravel chub was not taken in 1957, was rare at both
Neosho stations in 1958, became common at the lower Neosho
station in part of 1959, but was never numerous at the middle
Neosho station. Dr. F. B. Cross recorded the species as "rare"
in 1952 at a collection site near my middle Neosho station, but larger
numbers were taken then than in any of my collections at that
station. The population was probably reduced by drought, and
recovery was comparatively slow following restoration of flow.

Young-of-the-year and adults were common in collections from
riffles at the lower Neosho station from 1 July through 8 July, 1959.
I obtained only one specimen in intensive collections in the same
area on 25, 26, and 27 August. Seemingly the species had moved
off shallow riffles into areas not sampled effectively by seining.

Phenacobius mirabilis (Girard)


Sucker-mouthed Minnow

The sucker-mouthed minnow was common at the middle Marais
des Cygnes station but was not taken at the upper and lower stations
until 1959, when it was rare. At the middle and lower Neosho
stations this fish increased in abundance from 1957 to 1959; at the
upper station, sucker-mouthed minnows were not taken until 1959
when collections were made on the White farm. There, the species
was common immediately below a low-head dam, but was not
taken in extensive collections on the Bosch Farm in 1959.

The species was most common immediately below riffles, or in
other areas having clean gravel bottom in the current. On 5 June,
1959, many individuals were taken at night (11:30 p.m.) on a
shallow gravel riffle (four inches in depth) where none had been
found in a collection at 5:00 p.m. on the same date.

Young-of-the-year were taken at the lower Neosho station on 24
June, 1959, and commonly thereafter in the summer.

Notropis rubellus (Agassiz)


Rosy-faced Shiner

In 1958, the rosy-faced shiner was taken rarely at the lower stations
on both streams. This species is common in smaller streams
tributary to the lower portions of the two rivers, and probably[Pg 381]
occurs in the mainstream only as "overflow" from tributaries. Possibly,
during drought, rosy-faced shiners found suitable habitat in
the mainstream of Neosho and Marais des Cygnes rivers, but re-occupied
tributary streams as their flow increased with favorable
precipitation, leaving diminishing populations in the mainstream.

Notropis umbratilis (Girard)


Red-finned Shiner

The red-finned shiner, most abundant at the upper Neosho station,
occurred at all stations except the upper Marais des Cygnes.
This fish seems to prefer small streams, not highly turbid, having
clean, hard bottoms. It is a pool-dwelling, pelagic species.

Notropis camurus (Jordan and Meek)


Blunt-faced Shiner

The blunt-faced shiner was taken only in 1957, at the middle
Neosho station, where it was rare. This species, abundant in clear
streams tributary to the Neosho River (field data, State Biological
Survey) may have used the mainstream as a refugium during
drought. The few specimens obtained in 1957 possibly represent a
relict population that remained in the mainstream after flow in
tributaries was restored by increased rainfall.

Notropis lutrensis (Baird and Girard)


Red Shiner

The red shiner, abundant in 1952 (early stage of drought), was
consistently the most abundant fish in my collections in the Marais
des Cygnes and at the lower and middle Neosho stations. However,
the abundance declined from 1957 to 1959 at the two Neosho
stations. At the upper Neosho station the species was fourth in
abundance in 1957, and third in 1958 and 1959 (Table 12).

The red shiner is pelagic in habit and occurs primarily in pools,
though it frequently inhabits adjacent riffles. Collections by seining
along a gravel bar at the lower station showed this fish to be most
abundant in shallow, quiet water over mud bottom, or at the head
of a gravel bar in relatively quiet water. At the lower end of the
gravel bar in water one to four feet deep, with a shallow layer of silt
over gravel bottom and a slight eddy-current, red shiners were
replaced by ghost shiners or river carpsucker young-of-the-year as
the dominant fish.

Fifty-nine dyed individuals were released in an eddy at the lower[Pg 382]
end of a gravel bar at the middle Neosho station on 5 June, 1959.
Some of these fish still were present in this area when a collection
was made 30 hours later. No colored fish were taken in collections
from quiet water at the upper end of the gravel bar. A swift riffle
intervening between the latter area and the area of release may
have impeded their movement. Forty-six individuals, released at
the head of the same gravel bar on 10 June, 1959, immediately
swam slowly upstream through quiet water and were soon joined
by other minnows. These fish did not form a well-organized school,
but moved about independently, with individuals or groups variously
dropping out or rejoining the aggregation until all colored
fish disappeared about 50 feet upstream from the point of release.

Evidence of inshore movement at night was obtained on 8 June,
1959, in a shallow backwater, having gravel bottom, at the head
of a gravel bar at the middle Neosho station. A collection made in
the afternoon contained no red shiners, but they were abundant
in the same area after dark.

In Kansas, red shiners breed in May, June, and July. Minckley
(1959:421-422) described behavior that apparently was associated
with spawning. Because of its abundance, the red shiner is one
of the most important forage fishes in Kansas streams, and frequently
is used as a bait minnow.

Notropis volucellus (Cope)


Mimic Shiner

The mimic shiner was taken only rarely at the two lower Neosho
stations. This species, like N. camurus, is normally more common
in clear tributaries than in the Neosho River, and probably frequents
the mainstream only during drought.

Notropis buchanani Meek


Ghost Shiner

Field records of the State Biological Survey indicate that the
ghost shiner was common in the mainstream of the lower Neosho
River during drought. In 1957, the species was abundant at the
lower and middle stations on the Neosho River and at the lower
Marais des Cygnes station.

Collections at all stations show that the species has a definite
preference for eddies—relatively quiet water, but adjacent to the
strong current of the mainstream rather than in backwater remote[Pg 383]
from the channel. The bottom-type over which the ghost shiner
was found varied from mud to gravel or rubble.

Notropis stramineus (Cope)


Sand Shiner

The sand shiner was taken rarely in the Neosho and commonly
in the Marais des Cygnes in 1952. In my study the species occurred
at all stations, but not until 1959 at the upper and lower Neosho
stations. Sand shiners were found with equal frequency in pools
and riffles. Spawning takes place in June and July.

Pimephales tenellus tenellus (Girard)


Mountain Minnow

The mountain minnow was common at the lower and middle
Neosho stations throughout the period of study, and increased
in abundance from 1957 to 1959. It was taken only in 1959 at the
upper Neosho station, where it was rare. This species does not
occur in the Marais des Cygnes River. The largest numbers were
found in 1959 at the lower Neosho station, where this fish occurred
most commonly in moderate current over clean gravel bottom.
The mountain minnow, like Hybopsis x-punctata, was common in
late June and early July but few were found in late August, 1959.
The near-absence of this species in collections made in late August is
responsible for the apparent slight decline in abundance from 1957
to 1959, as shown in Table 11. Metcalf (1959) found mountain
minnows most commonly in streams of intermediate size in Chautauqua,
Cowley and Elk counties, Kansas. The predilection of this
species for permanent waters resulted in an increase in abundance
during my study. With continued flow, this species possibly will
decrease in abundance in the lower mainstream of the Neosho
River. I suspect that the species is, or will be (with continued
stream-flow), abundant in tributaries of intermediate size in the
Neosho River Basin.

Pimephales vigilax perspicuus (Girard)


Parrot Minnow

The parrot minnow was not taken in the Marais des Cygnes
River and was absent at the upper Neosho station until 1959.
This species was common at the lower and middle Neosho stations
throughout the period of study and increased in abundance from
1957 to 1959.[Pg 384]

At the lower Neosho station, this fish preferred slow eddy-current
over silt bottom, along the downstream portion of a gravel bar.
The parrot minnow was taken less abundantly in the latter part
of the summer, 1959, than in early summer, but the decline was
less than occurred in the mountain minnow.

Pimephales notatus (Rafinesque)


Blunt-nosed Minnow

The blunt-nosed minnow was common, and increased in abundance
in both rivers from 1957 to 1959. The largest numbers were
found at the upper Neosho station in 1959, and a large population
also was present at the lower Neosho station in 1959.

Pools having rubble bottom, bedrock, and small areas of mud
were preferred at the upper Neosho station. At the lower Neosho
station the fish was most common in quiet water at the lower end
of a gravel bar. The parrot minnow also was common in this
general area; nevertheless, these two species were seldom numerous
in the same seine-haul, indicating segregation of the two. The
blunt-nosed minnow was taken frequently in moderate current over
clean gravel bottom, especially in late summer, 1959, when P. notatus
increased in abundance as the mountain minnow decreased.

Pimephales promelas Rafinesque


Fat-headed Minnow

The fat-headed minnow was taken at all stations except at the
lower one on the Marais des Cygnes, and was most abundant at
the upper Neosho station. Intensive seining at the lower Neosho
station indicated that this species preferred quiet water and firm
mud bottom.

In the Neosho River in 1957 to 1959, habitats of the species of
Pimephales seemed to be as follows: Pimephales tenellus (mountain
minnow) occurred primarily in moderately flowing gravel
riffles in the downstream portions of the river. Pimephales vigilax
(parrot minnow) was mostly in the quiet areas having mud bottom
at the downstream end of gravel bars, and less commonly on adjacent
riffles, at the lower station. Pimephales notatus (blunt-nosed
minnow) had a wider range of habitats, occurring in quiet areas
and moderate currents both upstream and downstream. Pimephales
promelas (fat-headed minnow) occurred throughout both rivers
but was most abundant in the quiet water at the uppermost stations.[Pg 385]

Campostoma anomalum (Rafinesque)


Stoneroller

The stoneroller was most abundant at the upper Neosho station
and was not taken at the lower Marais des Cygnes station. This
fish increased in abundance from 1957 to 1959, but was never
common at the middle Marais des Cygnes or the middle and lower
Neosho stations.

The stoneroller prefers fast, relatively clear water over rubble
or gravel-bottom.

Ictalurus punctatus (Rafinesque)


Channel Catfish

The abundance of channel catfish was greatly reduced as a result
of the drought of 1952-1956. With the resumption of normal stream-flow
in 1957, the small numbers of adult channel catfish present in
the stream produced unusually large numbers of young. These
young of the 1957 year-class, which reached an average size of
about nine inches by September 1959, will provide an abundant
adult population for several years.

The reduction in number of channel catfish in streams can be
related to the changed environment in the drought. When stream
levels were low in 1953 (Tables 1-4), fish-populations were crowded
into a greatly reduced area. An example of these crowded conditions
was observed by Roy Schoonover, Biologist of the Kansas
Forestry, Fish and Game Commission, in October, 1953, when he
was called to rescue fish near Iola, Kansas. The Neosho River
had ceased to flow and a pool (less than one acre) below the city
overflow dam was pumped dry. Schoonover (personal communication)
estimated that 40,000 fish of all kinds were present in the
pool. About 30,000 of these were channel catfish, two inches to
14 inches long, with a few larger ones. Fish were removed in the
belief that sustained intermittency in the winter of 1953-1954
would result in severe winterkill. These conditions almost certainly
were prevalent throughout the basin.

In addition to winterkill, crowding probably resulted in a reduced
rate of reproduction by channel catfish, and by other species
as well. This kind of density-dependent reduction of fecundity is
known for many species of animals (Lack, 1954, ch. 7). In fish, it
is probably expressed by complete failure of many individuals to
spawn, coupled with scant survival of young produced by the adults
that do spawn. Reproductive failure of channel catfish in farm[Pg 386]
ponds, especially in clear ponds, is well known, and is often attributed
to a paucity of suitable nest-sites (Marzolf, 1957:22; Davis,
1959:10).

In the Neosho and Marais des Cygnes rivers, the intermittent
conditions prevalent in the drought resulted in reduced turbidity
in the remaining pools. Many spawning sites normally used by
channel catfish were exposed, and others were rendered unsuitable
because of the increased clarity of the water. In addition, predation
on young channel catfish is increased in clear water (Marzolf;
Davis, loc. cit.), and would of course be especially pronounced in
crowded conditions. The population was thereby reduced to correspond
to the carrying capacity of each pool in the stream bed.

The return of normal flow in 1957 left large areas unoccupied
by fish and the processes described above were reversed. The
expanded habitat favored spawning by nearly the entire adult
population, and conditions for survival of young were excellent.
As a result, a large hatch occurred in the summer of 1957. (Several
hundred small channel catfish were sometimes taken by use of the
shocker a short distance upstream from a 25-foot seine, set in a
riffle). Subsequent survival of the 1957 year-class has been good.
By 1959, few of the catfish spawned in 1957 had grown large enough
to contribute to the sport fishery, but they are expected to do so in
1960 and 1961.

The 1957 year-class was probably the first strong year-class of
channel catfish since 1952. Davis (1959:15) found that channel
catfish in Kansas seldom live longer than seven years. The 1952
year-class reached age seven in 1959. The extreme environmental
conditions to which these fish were subjected in drought caused a
higher mortality than would occur in normal times. The adult
population in the two rivers probably was progressively reduced
throughout the drought, and the reduction will continue until the
strong 1957 year-class replenishes it. For these reasons, fishing
success was poor in 1957-1959.

Juvenile channel catfish were more abundant in the Neosho than
in the Marais des Cygnes in 1958 and 1959, although both streams
supported sizable populations. In the Marais des Cygnes the upper
station had fewer channel catfish than the middle and lower stations.
In the Neosho, populations were equally abundant both upstream
and downstream. The habitat of channel catfish in streams
has been discussed by Bailey and Harrison (1948).

I found adults in various habitats throughout the stream, but[Pg 387]
most abundantly in moderately fast water at the lower and middle
Neosho stations. At the upper Neosho station where riffles are
shallow, yearlings and two-year-olds were numerous in many of
the small pools over rubble-gravel bottom. Cover was utilized
where present, but large numbers were taken in pools devoid of
cover. Young-of-the-year were nearly always taken from rubble- or
gravel-riffles having moderate to fast current at both upstream
and downstream stations.

Collections showed that young of 1957 were abundant on riffles
throughout the summer and until 17 November, 1957. Subsequent
collections were not made until 11 May, 1958, at which time 1957-class
fish still were abundant on riffles at the lower Neosho station;
on that date, the larger individuals were in deeper parts of the
riffles than were smaller representatives of the same year-class.

In a later collection (2 June, 1958), numbers present on the
riffles were greatly reduced and the larger individuals were almost
entirely missing. Some of the smaller individuals were still present
in the shallower riffle areas. Table 7 compares sizes of the individuals
obtained on 2 June with sizes collected from deep riffles
at the middle Neosho station on 7 June, 1958. The larger size of
the group present in deep riffles is readily apparent. The yearlings
almost completely disappeared from subsequent collections on
riffles.

A bimodal size-distribution of young-of-the-year was noted also
in 1958 and 1959; but, no segregation of the two sizes occurred
on riffles in summer. Marzolf (1957:25) recorded two peaks in
spawning activity in Missouri ponds. Two spawning periods may
account for the bimodal size distribution of young-of-the-year
observed in my study.

In 1959, young-of-the-year began to appear in the latter part of
June and became abundant by the first part of July. Individuals
as small as one inch T. L. were taken in gravel-bottomed riffles
on 1 July, 1959.

Yearling individuals at the lower and middle Neosho stations
showed a pronounced tendency to move into shallow, moderately
fast water over rubble or gravel bottom at night, where they were
nearly ten times more abundant than in daytime (Table 9). Adults
probably have the same pattern of daily movement as yearlings,
except that at night the adults move to deeper riffles. Bailey and
Harrison (1948:135-136) demonstrated that channel catfish feed
most actively from sundown to midnight.[Pg 388]

Channel catfish (especially two-year-olds and adults) were abundant
on a rubble-riffle during the day in some collections at the
lower Neosho station in 1959.

Table 7. Length-frequency of Channel Catfish from the Neosho River,
1957, 1958 and 1959. (Numbers in Vertical Columns Indicate the
Number of Individuals of a Certain Size Collected on That Date.)

Near the end of the spawning season in 1959, I found spawning
catfish at the lower Neosho station. Ripe females were taken between
9 June and 30 June, 1959; and, on 19 June I found a channel
catfish nest with eggs (water temp. 79° F.). The nest-site was a
hole in the base of a clay bank; the floor was clean gravel with a
small mound of gravel at the entrance. The nest-opening, five to
six inches in diameter, widened almost immediately into a chamber
about two and one-half feet long and one foot wide. Normally
the water was about six inches deep in the mainstream as it ran
over a riffle adjacent to the catfish nest. When I put my hand into
the opening the fish bit vigorously, but became quiescent when
I stroked its belly. I then felt the rounded gelatinous mass of eggs
on the bottom of the nest. On June 22 (water temp. 86° F.) the
fish was removed, struggling, from the nest, and returned to the
stream. The next day (23 June 1959, water temp. 84° F.) the eggs
had hatched and the young were in a swarm in the nest. The[Pg 389]
adult did not attempt to bite but left as soon as I put my hand into
the hole.

Marzolf (1957:25) reports that young remain in the nest from
seven to eight days after hatching. My seining records show a
marked increase in abundance of small young-of-the-year on the
first of July. Probably the time of hatching of the nest described
above correlated well with hatches of other nests.

One and sometimes two channel catfish were found in other
holes in the stream-bank or bottom. The fish occasionally attacked
my hand vigorously, but at other times remained quiet or left without
attacking. No other channel catfish eggs were found, although
one hole under a rock in the middle of the river had one or two
individuals in it each time it was checked until 11 July, 1959. A
local fisherman informed me of his belief that these holes are occupied
only in the spawning season.

Observations that I made in a pond owned by Dr. E. C. Bryan
of Erie indicated that channel catfish, when disturbed in the early
stages of guarding the eggs, either eat the eggs and abandon the
nest or leave the nest exposed to predation by other animals. In
the later stages of nesting, the fish, if removed, will return to guard
the nest. After the eggs hatch the guarding response probably
diminishes and the fish leaves the nest readily.

At the lower Neosho station, several "artificial" holes were dug
into the clay bank and two pieces of six-inch pipe were forced into
the bank. Nearly all these holes were occupied by catfish for a
short period in June; many of the holes were enlarged, either by
the current or by fish. I suspect that fish enlarged some holes, because
in the spawning season several males were observed that had
large abrasions atop their heads, around their lips, and to a lesser
extent on their sides. These could have been caused by butting
and scraping the sides, roof and floor of a hole. I found it possible
to enlarge the holes by rapidly moving my hand while it was inside
a hole.

The growth-rate of channel catfish in the Neosho was approximately
the same at all stations, and the large 1957 year-class grew
to an average size of about nine inches by mid-September, 1959
(Table 7). Channel catfish mature at a total length of 12 to 15
inches. Thus, most individuals of the 1957 year-class in the Neosho
River probably will mature in their fourth or fifth summer (1960
or 1961 spawning season).

The sizes attained by young-of-the-year in 1957 differed in the[Pg 390]
two rivers. Six hundred and thirty-three young taken in the Marais
des Cygnes River attained an average size of 4.7 inches (range
two to six inches) by mid-September. (Age was determined by
length-frequency and verified by examining cross-sections of fin-spines
from the larger individuals). One hundred and fifty young
from the Neosho River averaged 3.0 inches (range 2 to 3.7 inches)
on 2 November. Gross examination of the riffle-insect faunas indicated
a larger standing crop in the Neosho than in the Marais des
Cygnes River. Thus, the slower growth of young channel catfish
in the Neosho seemed not to be correlated with food supply. Bailey
and Harrison (1948:125-130) found that young channel catfish in
the Des Moines River, Iowa, fed almost exclusively on aquatic
insect larvae. My observations indicate that this is true in the
Neosho and Marais des Cygnes rivers also.

Young produced in 1958 in the Neosho River attained an average
total length of three inches by 26 August, and young produced in
1959 attained an average size of 3.5 inches by 11 September. Both
groups probably continued growth until October, and may have
averaged four inches total length at that time.

The 1958 and 1959 year-classes were much less abundant than
were the 1957 young. Therefore, it seems likely that the growth
of the 1957 young in the Neosho River was depressed because of
crowding. The 1959 year-class was larger than the small 1958
year-class, thus conforming to a general expectation that strong
year-classes will be followed by weak year-classes.

Reproduction by channel catfish in 1957 seemed greater in the
Neosho River than in the Marais des Cygnes River (Table 10); this
coincided with a greater change in volume of flow in the Neosho
River than in the Marais des Cygnes River (Tables 1-4). The
1957 year-class seemed more crowded, and grew more slowly, in the
Neosho than in the Marais des Cygnes River.

Ictalurus natalis (LeSueur)


Yellow Bullhead

Yellow bullhead were taken only at the middle station on the
Marais des Cygnes and upper station on the Neosho. The yellow
bullhead is more restricted to streams than is the black bullhead.
Both species decreased in abundance during a period of continuous
flow (1957 to 1959) following drought at the upper Neosho station.
Collections in 1958-'59 indicated an increase in average size.
Of four individuals marked and released at the upper Neosho sta[Pg 391]tion
in 1959, one was recaptured about three hours after being released.
It had not moved from the area of release.

Ictalurus melas (Rafinesque)


Black Bullhead

The black bullhead was abundant at the upper stations on each
river, especially in backwaters having mud-bottom. The species
was not taken in the mainstream of the lower and middle Neosho
stations, but was taken at the middle Neosho station in a pond
that is often flooded by the river. Although the fish was common
or abundant in nearly all pools at the upper Neosho station, it was
most abundant in one pool that had a bottom predominately of mud.

At the middle Marais des Cygnes station, 109 individuals were
collected and fin-clipped on 8, 9 and 24 July 1957. Three of the
19 marked on 8 July were recaptured in the same area on 9 July.
The area was poisoned on 13 September, 1957, and 130 black
bullhead were taken, none of which had been marked.

In 1959, 96 black bullhead were taken at the upper Neosho station
(five in Area 1 and 91 at the White Farm). In these collections,
25 were marked (fin-clipped or dyed) and six were recaptured.
Four of the six had not left the area of capture one and two
days after being released. The fifth fish recaptured was one of
five individuals that had been displaced one pool downstream.
When recaptured seven days later, this fish had moved upstream
over two steep riffles (two to three inches deep, 75 feet and 166
feet long) past the site of original capture to the next pool. The
sixth fish, marked at the same time but returned to the original
pool, was recaptured nine days after original capture and had
moved upstream over a long riffle (two to three inches deep, 166
feet long) and a short riffle into the second pool above the original
site of its capture.

Rotenone was applied to a small (.04 acre-feet) backwater ditch
having a soft mud bottom at the upper Marais des Cygnes station
on 25 July, 1957; 1526 black bullhead, one green sunfish and one
white crappie were collected. A sample of 60 bullhead averaged
4.6 inches T.L. (range 3.5 to 6.6 inches) and 540 individuals averaged
.7 ounce each. These fish probably represented the 1956 year-class.

The upper Neosho station had a large population of black bullhead,
strongly dominated by fish less than four inches T. L. (range
1.5 to 3.8 inches), in the spring of 1957. Most were approximately[Pg 392]
two inches T. L. and probably represented the 1956 year-class.
Growth, according to length-frequency, following restoration of
stream-flow, shows a regular increase in length of this dominant
1956 year-class (Fig. 3). A scarcity of young, especially in 1958
and 1959, is apparent in Fig. 3. This may be due to the fact that
a strong year-class usually is followed by one or several weak
year-classes. However, it more probably reflects the fact that
black bullhead are characteristically pond fish, and as such are
not so well adapted to reproduction in flowing streams as are many
other species. Metcalf (1959) found this species most abundantly
in the intermittent headwaters of Walnut River and Grouse Creek
in Cowley County, Kansas.

Fig. 3. Length-frequency of black bullhead at the upper Neosho station, 1957,
1958 and 1959.

Pylodictis olivaris (Rafinesque)


Flat-headed Catfish

The flathead is the largest sport-fish occurring in Kansas. Several
weighing more than 40 pounds are caught from streams each year,
and the species reportedly attains sizes in excess of one hundred
pounds. Several aspects of the biology of the flathead in Kansas
have been discussed by Minckley and Deacon (1959).

The abundance of flathead declined slightly from 1957 through
1959, counting fish of all sizes. This trend is attributable to a
large hatch in 1957; the 1957 year-class strongly dominated the[Pg 393]
population throughout my study. Natural mortality in that year-class
was compensated by increased average size of the individuals
(to six inches in autumn, 1958, and 11 inches in autumn, 1959).

The numbers of flathead caught at the upper stations on the
Neosho and Marais des Cygnes rivers differed from the general
trend in that the species was rare in 1957 and increased slightly
by 1959. Flathead are most numerous in large streams, and in the
drought they probably were almost extirpated from the headwaters.
After 1957, continuous flow and increased volume of flow
were accompanied by a gradual increase in numbers of flathead
in the upstream parts of the two rivers. The species was most
abundant at the middle and lower Neosho stations, where 10.5
per cent of all fish shocked in 1957 and 1958 were P. olivaris.

The habitat of the flathead varied with size of the individuals.
Young-of-the-year inhabited swift riffles having rubble bottom;
individuals four to 12 inches in total length were distributed
throughout the stream; those more than 12 inches in total length
were most commonly in pools in association with cover (rocks, or
drifts of fallen timber).

Male flathead mature at 15 to 18 inches total length, females at
18 to 20 inches. The spawning season in 1959 probably began
in early June and extended to mid-July. I attempted to find spawning
fish on 19 June and for one month thereafter. On 19 June
nine holes were dug into a 75-yard section of a clay bank adjacent
to a long, shallow, rubble riffle. A flathead was first found in one
of these holes on 22 June, and others were frequently found in
this and one other hole until mid-July. Although channel catfish
were often found in nearby holes, that species was never present
in the two holes used by flatheads. The holes occupied by flathead
(as well as those used by channel catfish) characteristically had
silt-free gravel bottoms and a ridge of clean gravel across the entrance.

A nest containing a flathead and eggs was located on 11 July.
In checking the hole I first put my foot into the entrance, then slowly
advanced my hand into the hole, feeling along the bottom with
my fingers until they entered the open mouth of a large catfish.
I backed off slowly and then felt beneath the fish. The fish was
directly above the egg-mass, seemingly touching the eggs with its
belly. As I touched the front of the egg-mass the fish struck
viciously, taking my entire fist into its mouth. It continued striking
until I removed my hand from the hole after obtaining a small[Pg 394]
sample of eggs, which proved to be in an early stage of development
(no vascularization evident).

When the nest was checked again on 13 July the eggs and fish
were gone. As in the case of channel catfish, I suspect that disturbance
of a flathead in the early stages of guarding the nest
results in destruction of the nest either by the guardian fish or by
predation resulting from its absence.

The hole occupied by the above fish was one that I had dug
seven to nine inches in diameter and extending two and one-half to
three feet into the bank. At the time this fish occupied the hole
its depth was approximately the same as originally, but the entrance
had been enlarged to 14 inches in diameter, and the chamber
widened to 32 inches. The holes were checked later in the summer
and all were heavily silted or had been undercut by action of the
current.

The number of flathead of catchable size was not reduced as
severely during my study as was the number of large channel catfish.
Flathead have a longer life-span than channel catfish; therefore,
it is not surprising that, of flathead and channel catfish that
survived the drought, a higher proportion of flathead persisted
throughout the next three years, in which my study was made. In
drought, when fish were concentrated in residual pools, the piscivorous
(fish eating) habit of flatheads may have favored their
survival.

The growth rate of flathead taken from the Neosho River in
1957 and 1958 was reported by Minckley and Deacon (1959:351-352).
Individuals hatched in 1955 and 1956 and collected in 1957
had attained average sizes of 9.5 inches and 4.8 inches, respectively,
by the end of the 1956 growing-season.

Flatheads of the 1956 and 1957 year-classes attained average
sizes of 8.7 and 3.2 inches, respectively, by the end of the 1957
growing season. These data indicate that growth was retarded
in the summer of 1957. Many species, including P. olivaris, had an
exceptionally large hatch in 1957, associated with increased water
levels in that year. Despite the great increase in amount of water,
I suppose that young-of-the-year and yearlings were subjected to
crowding resulting from exceptional hatches. This caused reduction
in growth of young flathead, and probably in several other
species.

Food of flatheads 4.0 inches and shorter was nearly all insect
larvae; that of fish 4.1 to 10 inches was insect larvae, fishes and
crayfish; and that of larger flatheads was mostly fish and crayfish.[Pg 395]
The specific kind of food eaten was correlated with abundance
of the food item in the stream (Minckley and Deacon, 1959:350-351).

Noturus flavus Rafinesque


Stonecat

The stonecat was not taken at the upper Marais des Cygnes station,
and was less abundant at the middle Marais des Cygnes station
than at other stations. The abundance of the stonecat was greatest
at the lower Marais des Cygnes station in 1957 and at the upper
Neosho station in 1959. The species increased in abundance from
1957 to 1959 in the Neosho River, where the principal habitat was
riffles over rubble bottom.

Thirty-three stonecats were marked at the upper Neosho station
in 1959. Five of these were recaptured three hours after release,
all near the point of release. One individual was taken from
a riffle, fin-clipped, and released at the foot of the next riffle downstream.
When recaptured four days later, this fish was still in the
area of release. Young-of-the-year were taken on July 1, 1959,
at the lower Neosho station.

Noturus gyrinus (Mitchill)


Tadpole Madtom

Trautman (1957:444-445) describes the habitat of the tadpole
madtom as "low-gradient lowland streams, springs, marshes, oxbows,
pothole lakes, and protected harbors and bays of Lake Erie, where
conditions were relatively stable, the water was usually clear, the
bottom was of soft muck which generally contained varying amounts
of twigs, logs, and leaves, and where there usually was an abundance
of such rooted aquatics as pondweeds and hornwort. The
species seemed to be highly intolerant to much turbidity and rapid
silting,..." The tadpole madtom was obtained only at the
middle Marais des Cygnes station in a small, deep, mud-bottomed
pool in 1957 after water levels, and probably turbidity, had been
low for five years. The occurrence provides the westernmost record
station in Kansas. Cross and Minckley (1958:106) reported the
species from the lower part of the Marais des Cygnes in Kansas.

Noturus nocturnus Jordan and Gilbert


Freckled Madtom

The freckled madtom was taken only at the middle Neosho station
on 19 April, 1958. This species occurs most frequently in
small streams, and individuals living in the mainstream of the Neo[Pg 396]sho
probably are "strays" from nearby tributaries. This species may
have utilized the mainstream as a refugium in the drought of
1952-'56.

Noturus exilis Nelson


Slender Madtom

The slender madtom was taken only at the middle Marais des
Cygnes station in the fall of 1957. This species prefers permanent
riffles of clear streams (Deacon and Metcalf, 1961:317). My specimen
possibly strayed from a nearby tributary; or, it was a relict
from a population living in the mainstream during drought.

Noturus sp.


Neosho Madtom

A description of this species, which is endemic to Neosho River,
has been prepared but not yet published by Dr. W. Ralph Taylor.
I found the Neosho madtom only at the middle station in 1958 and
1959, and at the lower station in 1959, where the species was common
in shallow water having moderate current over clean gravel
bottom. Specimens were most effectively collected by digging into
the gravel above the seine and allowing the gravel to wash into the
seine. In 1952, Cross (1954:311) found this species in abundance
in riffles at the confluence of the South Fork and Cottonwood River,
and at several other localities in the Neosho mainstream (personal
communication). The Neosho madtom is nearly restricted to gravel
riffles having moderate flow; therefore, it may be drastically reduced
by intermittency of flow. I found none in 1957 and few in 1958. By
1959, the third summer of continuous flow, the Neosho madtom
was again common.

Fundulus notatus (Rafinesque)


Black-striped Topminnow

The black-striped topminnow was rare in the mainstream at the
lower Marais des Cygnes and the middle and lower Neosho stations,
where it was found in quiet water near shore.

Near the middle Neosho station, a large population was present
in an oxbow lake that is frequently flooded by the river.

Labidesthes sicculus (Cope)


Brook Silversides

The brook silversides occurred rarely at the lower Marais des
Cygnes and at the middle and lower Neosho stations.[Pg 397]

Micropterus dolomieui Lacépède


Small-mouthed Bass

One individual was taken at the lower Neosho station in 1957.

Micropterus punctulatus punctulatus (Rafinesque)


Spotted Bass

The spotted bass occurs in Kansas only in the southeastern part
of the state—in southern tributaries of the Osage system, in Spring
River drainage, and in relatively clear streams of the Flint Hills.
At my stations on the Neosho River, this fish was more abundant
in 1957 than in 1958 or 1959.

Spotted bass were taken most frequently over rubble bottom or
near boulders in moderate current. Collections made in the evening
or early morning more often contained spotted bass than collections
made at other times of day (Table 9). Data from a few
specimens that were marked, released, and recaptured indicated that
the species is relatively sedentary; therefore, the greater abundance
in the morning and evening collections probably indicates increased
activity during these periods, possibly in connection with feeding.
The spawning season in 1957 may have continued as late as 10
July when a ripe female 11.3 inches T. L. was taken. Young-of-the-year
were taken on 24 June in moderate current over gravel
bottom and in quiet water over mud bottom.

Spotted bass normally form a small part of the game-fish fauna
in the lower Neosho River. The species attains greater abundance
in smaller, clear streams of the Arkansas River Basin in Kansas
(Cross, 1954, and unpublished data of State Biological Survey
of Kansas). During the drought, the lower Neosho probably assumed
many characteristics of a smaller stream in normal times.
Flow was reduced or entirely interrupted and turbidity was lessened.
These conditions resulted in faunal changes in which
spotted bass were more prominent than in years of normal flow.
During this period of reduced flow, some fishermen turned from
catfishing to bass-fishing; I think this constitutes evidence for an increase
in numbers of bass, accompanied by a decrease in numbers
of channel catfish. With the return of continuous flow and a consequent
rise in turbidity, bass declined in abundance in the mainstream.[Pg 398]

Micropteras salmoides salmoides (Lacépède)


Large-mouthed Bass

The large-mouth was rare at all stations. It prefers quiet water
near cover; to become abundant, the large-mouth probably requires
clearer water than is afforded by most Kansas streams. This
species, like spotted bass, declined in abundance during the period
of study. Nevertheless, young-of-the-year were taken in 1957 and
1958 (earliest date of capture, 7 June in 1958).

Lepomis cyanellus Rafinesque


Green Sunfish

Green sunfish were taken at all stations, but most abundantly
at the upper Neosho station where the number captured increased
slightly from 1957 to 1959. Young-of-the-year and adults were most
common in shallow backwater. At the upper Neosho station green
sunfish inhabit quiet pools, where recaptures of marked fish indicated
that the species is notably sedentary in habit. Hasler and
Wisby (1958) have shown that green sunfish exhibit a homing
reaction.

This fish provides some sport for fishermen, especially in the
smaller streams, but I found few green sunfish that were larger
than six inches T. L. at any station.

Lepomis megalotis (Rafinesque)


Long-eared Sunfish

Long-eared sunfish were taken at all stations but were notably
more abundant in the Neosho River, where the largest population
occurred at the upper station. In all three years of the study, large
samples were obtained by means of rotenone in the same pool at the
upper Neosho station. There were fewer long-eared sunfish present
each year, and average size increased slightly. Collections in
other pools at this station indicated that long-eared sunfish maintained
a high level of abundance throughout my study.

Long-eared sunfish occurred in pools having bottoms of gravel
or bedrock at the upper Neosho station, or near shore over rubble
or gravel in slow to moderate current at the middle Neosho station.

Lepomis humilis (Girard)


Orange-spotted Sunfish

The orange-spotted sunfish occurred at all stations; it was most
abundant in the Neosho River, especially at the uppermost station.
This fish was taken in a variety of habitats, but was most[Pg 399]
common in areas where the current was slack, often over mud or
silt bottom.

Lepomis macrochirus Rafinesque


Bluegill

Bluegill were taken at all stations but were rare. This species
occurred exclusively in pools, usually near cover (brush or trees in
the water). Bluegill are predominately pond-fish in Kansas, and
populations in rivers may consist partly of individuals that escaped
from ponds in time of overflow. I know of no stream in Kansas
that has a population large enough to contribute significantly to
the sport fishery.

Pomoxis nigromaculatus (LeSueur)


Black Crappie

This species was represented by only one specimen, taken at the
lower Neosho station in 1957.

Pomoxis annularis Rafinesque


White Crappie

White crappie were taken at all stations, but were common only
at the upper and middle stations on the Marais des Cygnes and
the upper Neosho station. At the last station, this fish was abundant
in a single large pool that contained much more water during
drought than any other area at this station. There was little dispersal
into several smaller pools, below the large pool, which were
sampled in 1957, 1958 and 1959. White crappie were not taken
in the lower pools until 1959, and then were rare. Most crappie
were taken in quiet water near cover or near shore.

Young-of-the-year were found in 1957, 1958 and 1959, but never
abundantly. At the lower Neosho station in 1959, ripe individuals
were collected on 19 June, a spent female on 24 June, and young-of-the-year
on 1 July. The young were present in quiet, shallow
water over mud bottom at the lower end of a gravel bar. Large
white crappie (10-14 inches T. L.) were common at the middle
and lower Neosho stations in 1957 and in April, 1958. Large fish
were almost entirely absent from later collections. Average size,
maximum size and abundance declined during the period of study.

Percina phoxocephala (Nelson)


Slender-headed Darter

The slender-headed darter was taken at all stations but was
more abundant in the Neosho than in the Marais des Cygnes. The
lower Marais des Cygnes, however, was the only station with a[Pg 400]
relatively large population in 1957. Slender-headed darters were
rare in the Neosho River in 1957 and did not become common
until 1959.

The largest population was found at the upper Neosho station
in 1959. This darter occurs most frequently in swift water over
gravel bottom, but was taken in various habitats, including an intermittent
pool at the upper Neosho station on 7 September, 1957.

At the middle and lower Neosho stations, considerably greater
numbers were taken in June, July, and early August than in May or
late August. The abundance in my collections diminished from a
peak in early July, to scarcity in late August.

Young-of-the-year were taken at the lower Neosho station on
1 July, 1959 (and subsequently), in moderately fast water over
gravel. On 21 August, 1958, a ripe female (eggs stripped easily)
was the only slender-headed darter present in a collection from
riffles at the middle Neosho station.

Percina caprodes (Rafinesque)


Logperch

Logperch were not taken in the Marais des Cygnes. They were
rare in the Neosho, where they were taken most frequently at the
upper station in water two to three feet deep, over gravel bottom,
in moderate to slight current. This species was present in intermittent
pools at the upper Neosho station in 1957.

Percina copelandi (Jordan)


Channel Darter

One specimen was taken at the lower Neosho station in 1959.
Because no others ever have been found in the mainstream of the
Neosho River, I suspect that my specimen is a "stray" from one of
the smaller tributaries, where channel darters are locally common.

Etheostoma flabellare Rafinesque


Fan-tailed Darter

The fan-tailed darter is represented in my collections by one
specimen, obtained in the mainstream of the Neosho River at the
lower station in 1957. Records of this species in Kansas are almost
confined to the smallest, clear, permanent streams of the southeastern
part of the state. My specimen may represent a small
population that retreated to the mainstream of the Neosho during
drought.[Pg 401]

Etheostoma spectabile (Agassiz)


Orange-throated Darter

Orange-throated darters were common at the upper Marais des
Cygnes and upper Neosho stations in 1959, rare at the middle and
lower Neosho stations, and absent from the middle and lower
Marais des Cygnes stations. The species was found almost exclusively
on upstream riffles over gravel-rubble bottom. The population
in the upper Neosho was decimated by drought, and the fish
did not become common until the summer of 1959, the third year
after resumption of normal stream-flow.

Deacon and Metcalf (1961:320) indicated that long periods of
intermittency result in depletion or elimination of populations of
the orange-throated darter in the Wakarusa River, Kansas. A limited
number of orange-throated darters probably survived in the
few permanent pools in the upper Neosho and provided the brood-stock
necessary to repopulate this section of the stream.

Aplodinotus grunniens Rafinesque


Freshwater Drum

Drum were taken at all stations, but were most abundant at
the middle and lower Neosho stations. A high level of abundance
also was found in 1957 at the middle Marais des Cygnes station.
The abundance of drum declined from 1957 to 1959, but the average
size increased because of a dominant 1957 year-class that was moderately
reduced by natural mortality in 1958-'59. Although the
population was composed largely of young-of-the-year and adults
in 1957, it was dominated by yearling individuals in 1958. By 1959
the number had declined considerably and the population consisted
mostly of juveniles and adults. Fish of the 1957 year-class reached a
length of approximately ten inches by mid-summer of 1959 (Table
8).

Adults were taken in a variety of habitats, but most often in
quiet water. On the other hand, yearlings were extremely abundant
in 1958 near shore in shallow, moderately fast water over
rubble bottom at night. Drum were rare in the same areas in daylight
(Table 9). Young-of-the-year occur in shallow, quiet water,
usually over mud-bottom.

The freshwater drum matures at about 12 inches T. L. Ripe
males were taken as late as 23 June 1959; however, the height of
the spawning season probably is in May.[Pg 402]

Table 8. Length-frequency of Freshwater Drum from the Middle
Neosho Station in 1957, 1958 and 1959.

Table 9. Average Number of Individuals Captured per Hour, Using the
Shocker, at Different Times of the Day and Night at the Middle Neosho
Station in 1958. Numbers in Parentheses Indicate Total Number
Captured.

[Pg 403]

Table 10. Numbers of Fish Seen or Captured per Hour by Use of the
Shocker. Excludes Fish Taken by Shocking into a Seine on Riffles;
Young-of-the-year Channel Catfish and Flathead Catfish Predominated
in Samples Taken by that Method.

[Pg 404]

Table 11. Number of Occurrences (Roman type) and Number Counted
(Italic type) per Seining Unit. One Seining Unit Equals 30 Seine-Hauls
(ten each with the 4-foot, 12-foot and 25-foot seine) of Which Six Randomly-chosen
Hauls Were Counted. Dashes Signify That the Species Occurred
in Uncounted Collections Only.

[Pg 405]

FISH-FAUNA OF THE UPPER NEOSHO RIVER

Collections at the upper Neosho station were more intensive than
at any other station, especially in 1959. Rotenone was used in
the summers of 1957, 1958 and 1959, to obtain large samples of
the population in one section of the stream. In September, 1959,
the shocker was used in other sections in order to estimate populations
in particular pools and riffles, to measure variability in the
fauna between areas having slightly different habitat, and to record
movement of marked individuals in a short section of the stream.

Description of Study-areas

Two sections of the stream, each about one-half mile long (See p. 366),
were studied. Additional description of particular areas is presented below.
Area 1 and the pools in which rotenone was used are on the Bosch Farm approximately
two miles upstream from the White Farm where Areas 2, 3, 4,
5, 6 and 7 are situated.

Area 1 has a length of 210 feet, an average width of four feet, and a
maximum depth of two feet. The upper half is a swift, rubble riffle four inches[Pg 406]
in average depth; the lower half is one and one-half feet in average depth and
has a slow current (Pl. 29, Fig. 1).

Area 3 has a length of 186 feet, an average width of 34 feet, and a maximum
depth of two and one-half feet. This area includes a shallow riffle at both
upstream and downstream ends of a pool 73 feet long and approximately
one foot in average depth (Pl. 29, Fig. 2).

Area 5 has a length of 250 feet, an average width of 50 feet, and a maximum
depth of two and one-half feet. This is a shallow, quiet pool over rubble
and bedrock bottom except for a small area of mud bottom (backwater) above
the point where a short riffle drains into this pool from Area 6 (Pl. 30, Fig. 1).

Area 6 has a length of 200 feet, an average width of 50 feet, and a maximum
depth of one and one-half feet. This is a shallow, quiet pool over bedrock
bottom, except for a small area of mud bottom at one side of the upper end
of the pool. A short, steep, rubble-riffle is included in this area at the upstream
end (Pl. 30, Fig. 2).

Areas 2, 4, and 7 resemble at least one of the areas described above but
were sampled less intensively. Data from areas 2, 4, and 7 are included in
discussion of the total fauna of the upper Neosho river but are excluded from
the discussion of representative parts of that fauna.

Methods

Rotenone

Rotenone was applied to an intermittent pool in 1957. In 1958 and 1959
rotenone was applied to the upper end of a pool and mixed by agitating the
water. The concentration in the pool was maintained by slowly introducing
part of the rotenone into the riffle at the head of the pool. This was the most
effective means of obtaining a large sample of fish from the deeper, slowly flowing
water of the upper Neosho. Pools in which rotenone was used had areas of
as much as one-half acre and depths in excess of six feet.

Shocker

In 1959 the shocker was used extensively in several areas of the upper
Neosho. Because of the small size of the stream, "tennis-racket" electrodes
were used effectively by two men—one carrying the electrodes and one picking
up fish and placing them in a live-box. In fast water, many fish floated into a
seine placed across the lower end of the area. A large segment of the population
was collected in this manner. Areas in which fish were collected by means
of the shocker included riffles, and pools having flowing water no more than
three feet in maximum depth. The bottom-type was usually gravel, rubble
or bedrock, but a small amount of mud bottom was present in many pools.

Because of the necessity of wading, we could not use the shocker effectively
in water more than three feet deep. In addition, turbidity of the water prevented
effective collection of stunned fish in the deeper pools. Therefore,
rotenone was more effective in deep water than was the shocker. In shallow,
swift riffles and pools, the shocker yielded more reliable samples than did
rotenone, because of difficulty in maintaining adequate concentrations of
rotenone where flow was swift.

[Pg 407]

The relative abundance of each species in the upper Neosho was calculated
from cumulative results obtained by use of the shocker in seven areas in 1959.
Population estimates were made by collecting fish with the shocker, marking
them by clipping fins or staining them in Bismark Brown Y at a concentration
of 1:20,000 (Deacon, 1961), returning them to the stream, and making a second
collection three hours (Areas 1 and 3) or 24 hours (Area 6) later. The
same area was shocked again within two to eight days. Collections throughout
the one-half-mile section yielded information on movement.

Changes in the Fauna at the Upper Neosho Station,
1957 Through 1959.

The following discussion is based principally on collections made
with rotenone in 1957, 1958 and 1959 (Table 12). Other supplementary
data aid in understanding the changes that occurred after
the resumption of normal flow at the upper Neosho station.

The population in 1957 was strongly dominated by black bullhead
and young-of-the-year channel catfish. Other common species
were long-eared sunfish, red shiner, yellow bullhead, orange-spotted
sunfish and green sunfish. This fauna, with the exception of young-of-the-year
individuals, was a fauna produced during the years of
drought. Deacon and Metcalf (1961:318-321) found a similar fauna
in streams of the Wakarusa River Basin that had been seriously
affected by drought.

The black bullheads taken in 1957 were predominately yearlings.
It is likely that by 1956 the total fish population in the upper Neosho
had been decimated by drought. The ponded conditions prevalent
in that year were conducive to production and survival of young
black bullheads. Fig. 3 shows that this dominant 1956 year-class
reached an average length of approximately 6.5 inches by August,
1959.

Reproduction by black bullheads was limited in 1957, 1958, and
1959, and slight reduction in relative abundance occurred from
1957 to 1958. The relative abundance in 1959 remained nearly
stable. If stream-flow remains essentially continuous for the next
few years, the number of black bullheads probably will decline
as individuals of the 1956 year-class reach the end of their life-span.

Reference has been made to the large hatch of channel catfish
in 1957, in a discussion of that species. Conditions for survival of
young channel catfish at the upper Neosho station in 1957 were
good because there was continuous flow over many gravel-rubble
riffles, which were largely unoccupied by other fish, in the spring
and summer of 1957.[Pg 408]

Table 12. Percentage-composition of the Fish-fauna at the Upper
Neosho Station in 1957, 1958 and 1959, as Computed from Collections
Obtained by Using Rotenone.

Channel catfish also showed a slight decline in relative abundance
after 1957, resulting from mortality in the 1957 year-class. With
continuous flow, channel catfish will probably remain abundant,
although annual reproductive success probably will be less than in
1957.

The big-mouthed buffalo, small-mouthed buffalo, creek chub
and orange-throated darter were not taken in 1957, but appeared
in collections in 1958. The river carpsucker, golden redhorse, red
shiner, fat-headed minnow, stoneroller, stonecat, and slender-headed
darter also increased in abundance between 1957 and 1959. The
increased abundance of all these species in 1958 and 1959 resulted
in a more diversified fauna, with lesser predominance by any single
species, than in 1957 (Table 12); this change is related to the increased,
permanent flow in 1958 and 1959.[Pg 409]

Local Variability of the Fauna in Different Areas at the
Upper Neosho Station, 1959

The shallow areas in which the shocker was used in 1959 are
the prevalent habitat in the upper Neosho River. The relative
abundance of fishes found in these areas is presented in Table 13.
The red shiner was most abundant and was followed (in decreasing
order) by long-eared sunfish, minnows of the genus Pimephales,
green sunfish, red-finned shiner, channel catfish, and stoneroller.
Other species combined comprise less than ten per cent of the
population.

Table 13 also shows the variability in relative abundance of different
species among areas that have the same general kind of
habitat. The species composition is similar in all areas. The sample
obtained with rotenone in 1959 is included in Table 13 to show
differences in the fauna of deep, slowly flowing areas and shallower
areas with stronger current. The differences in relative abundance
indicate the kind of habitat that each species is able to utilize most
fully.

Golden redhorse and black bullhead were most abundant in
large, deep, quiet pools (5.7 per cent and 32 per cent of the total
population) and were more abundant in Area 5 (3.2 per cent and
7.3 per cent respectively) than in any of the other shallow areas.
Area 5 has greater average depth, more mud bottom, and less riffle
area than areas 1, 3 and 6.

The golden redhorse and black bullhead have specific habitat
preferences that are not evident in the above discussion. My collections
indicate that the golden redhorse prefers deep water having
some current, whereas the black bullhead prefers little or no current.

Species that prevailed in or near riffles were: creek chub, sucker-mouthed
minnow, stoneroller, channel catfish (young-of-the-year
only), flathead (young-of-the-year only), stonecat, slender-headed
darter, and orange-throated darter. Of these species, the sucker-mouthed
minnow, slender-headed darter and orange-throated darter
reached their greatest abundance at Area 3, where the riffle is
shallow, slow, and has a bottom composed of flat limestone rubble.

The riffle at Area 1 is, for the most part, deeper and faster than
at Area 3 and has a bottom composed of gravel and small rocks.
The creek chub, stoneroller, channel catfish (young-of-the-year),
flathead (young-of-the-year), and stonecat reached their greatest
[Pg 410]
abundance in Area 1. All species that showed a preference for
riffles were rare or absent in Area 5 where no riffle-habitat was
sampled. The riffle-dwelling species that were present in collections
made with rotenone in the deeper pools were taken from the riffle
into which rotenone was introduced.

Table 13. Relative Abundance of Fish (Per Cent of Total Population
Made Up by Each Species), in the First Collection Made in Each of
Four Different Shallow Areas by Means of the Shocker, is Shown in
Vertical Columns 1-4. Results of the Use of Rotenone in a Fifth,
Deeper Area are Shown in Column 5. Column 6 Combines Data from
All Collections Made by Using the Shocker in Seven Shallow Areas
(Including Columns 1-4).

[Pg 411]

The river carpsucker, blunt-nosed minnow, fat-headed minnow,
channel catfish (yearlings and two-year-olds), flathead (yearlings
and two-year-olds), green sunfish and long-eared sunfish showed a
preference for shallow, quiet water. All of these species were
more common in collections from Areas 5 and 6 than in collections
from other areas.

Temporal Variability of Fauna in the Same Areas

The variability of the population in successive collections from
the same area is presented in Table 14. Supplementary data obtained
in Areas 2, 4 and 7 support conclusions discussed below
for Areas 1, 3 and 6. The abundance of some species maintained
a constant level, whereas that of others varied.

Table 14. Numbers of Individuals Collected by Means of the Shocker
at Varying Intervals in September, 1959. The Number at the Top of
Each Column is the Date When the Collection was Made.

[Pg 412]

Stoneroller, channel catfish (young-of-the-year), green sunfish,
and long-eared sunfish formed the most stable element of the population,
in that the numbers of these species varied less in successive
collections than did numbers of other species.

The number of orange-throated darters remained constant at
Areas 1 and 3, and the number of stonecats changed little in successive
collections from Area 3. I suspect that an apparent decline in
stonecats at Area 1 on September 4 was due to a slow rate of dispersal
from the point of release (see pages 413, 414).

Some species (sucker-mouthed minnow, red-finned shiner, slender-headed
darter, and fat-headed minnow) decreased significantly
in successive samples from the same area because of mortality in
handling or movement out of the area of initial capture.

The decrease in abundance of the sucker-mouthed minnow may
have been due to some mobility of the species. Evidence for
mortality caused by handling was obtained for the red-finned shiner
and probably accounts for the reduction of this species in Area 6.
The red-finned shiner is also probably a mobile species. The reduction
in abundance of the slender-headed darter seems unexplainable
because no evidence was obtained for either movement or mortality.

Fat-headed minnows also declined markedly in successive collections
from Area 6, the only area in which the species was common.
No marked fat-headed minnows were taken outside the
area of release, indicating low mobility of the species. I cannot
certainly account for their decline; possibly there was latent mortality
due to shocking.

The numbers of red shiners, blunt-nosed minnows, and juvenile
channel catfish varied erratically in successive collections, probably
as a result of movement. This problem is discussed for all species
in a later section.

Population-Estimation

The direct-proportion method was used to estimate fish populations
in Areas 1, 3 and 6. Reliable results could not be obtained
for all species because of scarcity, mortality in handling, mobility,
or other factors.

A high rate of mortality due to handling was observed in Area
1 for the red shiner and in Area 6 for river carpsucker (young-of-the-year),
sucker-mouthed minnows, red-finned shiner, red shiner, blunt-nosed
minnow, and stoneroller. In Area 3, in contrast, there was
little mortality in the same species during the twelve-hour interval
that fish were held in traps prior to release as marked individuals.[Pg 413]

The following species were common in at least one area, but
probably are sufficiently mobile (see page 416) to invalidate estimates
of static populations in small areas: red shiner, red-finned
shiner, and channel catfish (yearlings and older). Other species
were rare and are indicated as "T" in Table 13.

Those species for which population-estimates seem warranted
include: golden redhorse, sucker-mouthed minnow, red shiner,
sand shiner, fat-headed minnow, stoneroller, stonecat, channel catfish
(young-of-the-year), green sunfish, long-eared sunfish, slender-headed
darter, and orange-throated darter. I consider the estimate
valid if a high percentage of the marked fish is recaptured. Results
are presented in Table 15, and ordinarily will not be referred to in
the following discussion of the population in each of the three areas.

Table 15. Data Used in Estimating Total Populations, by Direct
Proportions, in Areas 1, 3, and 6 at the Upper Neosho Stations.

Movement of Marked Fish

Table 16. Data on Movement of Marked Fish at the Upper Neosho
Station, September, 1959.

Some measure was gained of the amount of movement exhibited
by several species of fish. Results are biased in favor of a conclusion
that a species is sedentary because a large percentage of the
recaptures were made in collections taken in the same immediate
area three hours after release of marked fish, the total area checked
was not large (one mile), and collecting was limited to an eleven-day
period. Nevertheless, some species were shown to be definitely
mobile and others exhibited pronounced sedentary tendencies.
The results of experiments on movement are presented in Table
16. Marked fish (dyed and fin-clipped) were taken as long as seven
days after being marked. Only those species in which more than
ten individuals were marked are included.[Pg 417]

Blunt-nosed minnow, red shiner, and channel catfish (yearlings
and older) are more mobile than other species.

The mobility of channel catfish has been discussed by Muncy
(1958) and Funk (1957). My records show that of 36 marked
channel catfish that were recaptured, 11 were taken in areas other
than the one into which they had been returned. A pronounced
mobile tendency on the part of the red shiner and blunt-nosed minnow
is shown by the fact that of 152 marked red shiners recaptured,
73 had moved from the area of release; and of 32 marked blunt-nosed
minnows recaptured, 11 had moved from the area of release.
The fact that the habitat occupied by these species is not precise
(ranging from swift riffles to quiet pools) supports a conclusion
that the species are mobile.

The fat-headed minnow, stoneroller, channel catfish (young-of-the-year),
green sunfish and long-eared sunfish form a sedentary
element of the population. With the exception of the fat-headed
minnow, the sedentary group also maintained relatively stable numbers
in Areas 1, 3 and 6 throughout the study (Table 14). It is
interesting to note that, in contrast to the mobile group, the species
forming the sedentary group have rather well-defined habitat preferences.

A third group of species, represented by the red-finned shiner,
stonecat, slender-headed darter and orange-throated darter, was
characterized by having a low rate of recapture. I suspect that
mortality is a factor contributing to the failure to recapture red-finned
shiners, because in one collection only four of 31 red-finned
shiners captured were successfully marked and released, in another
case 70 of 818. The red-finned shiner occurs most often in pools but
is also taken in other areas, is pelagic, and probably is a mobile
species.

The stonecat, slender-headed darter and orange-throated darter
are generally restricted to riffle-habitats, and are probably sedentary.
The low number of recaptures for these three species probably
is due either to a slow rate of dispersal from the point of release
or to latent mortality resulting from shock. Table 14 shows that
these three species maintain comparatively stable populations, but
there seems to be a tendency for a reduction in numbers with continued
collecting, even though all fish captured were returned to
the stream.

Golden redhorse showed a high rate of recapture. All individuals
marked were recaptured three hours after release in Areas 1 (two[Pg 418]
fish) and 3 (five fish). Nine individuals were taken from Area 4
on 11 September; seven of these were marked and released in the
next pool downstream (Area 3). On 15 September, two fish were
retaken in Area 3 and two were retaken in Area 2, the next pool
downstream. The species was common in Area 5 also where five
of eight marked individuals were recaptured two days after release.
It seems that the golden redhorse is somewhat restricted in movement,
at least for short periods.

The sucker-mouthed minnow and black bullhead showed some
movement—less than such mobile species as red shiners and channel
catfish, but more than the sedentary group. Seven of 27 marked
sucker-mouthed minnows were taken in areas adjacent to the
one to which they had been returned. Two of six black bullheads
that were recaptured had moved. The black bullhead moved the
greater distance. The extent of short-term movement by several
of the species in the Upper Neosho correlates well with redistribution
subsequent to drought in the Wakarusa River, discussed by
Deacon and Metcalf (1961).

Similarity of the Fauna at the Upper Neosho Station to the
Faunas of Nearby Streams

The fauna that I found to be characteristic at the upper Neosho
station has affinity with the upland tributary-fauna described by
Metcalf (1959) for Chautauqua, Cowley and Elk Counties, Kansas.
The primary difference is a nearly complete absence at my
station of the Ozarkian element of the population. Some species
(red-finned shiner, long-eared sunfish, and spotted bass) listed by
Metcalf as characteristic of the mainstream of smaller rivers occur
at the upper Neosho station in greater abundance then elsewhere
in the Neosho. This difference is probably due to the fact that the
upper Neosho station is somewhat larger and slightly more turbid
than Metcalf's "upland tributaries."

Hall (1952) reported on the distribution of fishes in the vicinity
of Fort Gibson Reservoir, an impoundment on the Grand (Neosho)
River in Oklahoma. He separated the fishes into three groups
according to habitat-preference: species restricted to upland tributaries
on the east side of Grand (Neosho) River, species restricted
to lowland tributaries on the west side of Grand (Neosho) River,
and species occurring in the Grand River proper and/or tributaries
on one or both sides.

Several species found in the upper Neosho River also occur in
the area studied by Hall. Of these, only the creek chub was re[Pg 419]stricted
to upland tributaries on the east side of Grand (Neosho)
River. The sucker-mouthed minnow and red-finned shiner were
restricted to the lowland tributaries on the west side of Grand
(Neosho) River in the Fort Gibson Reservoir Area. Golden redhorse,
stoneroller, yellow bullhead, spotted bass, green sunfish,
long-eared sunfish, and orange-throated darter were present in collections
from the Grand River proper and/or tributaries on both
sides of the river, most commonly in tributaries.

Hall's data show that black bullhead, large-mouthed bass, white
crappie, and logperch occurred most frequently in or near the quiet
water of the reservoir. In my study these fish were most common
in the larger, quiet pools at the upper Neosho station.

COMPARISON OF THE FISH FAUNAS OF THE

NEOSHO AND MARAIS DES CYGNES RIVERS

The Marais des Cygnes River has less gradient (especially in
the upstream portions), fewer and shorter riffles, and more mud
bottom than does the Neosho River. Stream-flow during drought
was reduced to a proportionately greater degree in the Neosho
River than it was in the Marais des Cygnes River. Average flow of
the Neosho River near Parsons (drainage area: 4905 square miles),
Kansas, was less than average flow of the Marais des Cygnes
River at Trading Post (drainage area: 2880 square miles), Kansas,
in 1953, 1955 and 1956. In normal times the Neosho River carries
a larger volume of water than the Marais des Cygnes. The Neosho
River has a greater variety of habitat-conditions and a more diversified
fish-fauna than the Marais des Cygnes.

The following species were taken in the Neosho River but not
in the Marais des Cygnes River: blue sucker, high-finned carpsucker,
golden redhorse, gravel chub, mimic shiner, mountain minnow,
parrot minnow, Neosho madtom (the only endemic in either
river), mosquitofish, spotted bass, smallmouth, black crappie, logperch
and fan-tailed darter. Most of the above species are usually
found in association with gravel-bottom, which is prevalent in Neosho
River. The blue sucker, high-finned carpsucker, gravel chub,
mountain minnow, and parrot minnow normally occur in the larger
streams in Kansas. The last three species became more abundant
in the Neosho River following resumption of flow. The golden
redhorse also increased in abundance from 1957 to 1959, but was
most numerous at the upper Neosho station, whereas the other
species occurred mainly at the lower stations.

The mimic shiner, spotted bass, smallmouth, and fan-tailed darter[Pg 420]
are characteristic of upstream habitats with clear water (tributaries,
rather than the mainstream), and were taken in the Neosho River
only in 1957 or became less abundant from 1957 to 1959.

The silver chub, slender madtom and tadpole madtom were taken
in the Marais des Cygnes River only in 1957 and were not taken in
the Neosho River.

The following species, common to both rivers, were more abundant
in the Neosho: long-nosed gar, short-nosed gar, river carpsucker,
creek chub, sucker-mouthed minnow, red-finned shiner,
red shiner, ghost shiner, blunt-nosed minnow, fat-headed minnow,
stoneroller, yellow bullhead, channel catfish, flathead, stonecat,
largemouth, long-eared sunfish, slender-headed darter, and freshwater
drum. These species, collectively, reflect the more diversified
habitats (more gravel-bottom, more riffle-areas, more gradient,
greater range of stream-size sampled) in the Neosho River.

The following species, common to both rivers, were more abundant
in the Marais des Cygnes: gizzard shad, carp, sand shiner,
black bullhead and white crappie. These species (with the exception
of sand shiner) emphasize the fact that the Marais des Cygnes
is a sluggish stream with large areas of mud bottom. Differences
in the abundance of the sand shiner in the two rivers are part of
taxonomic and distributional studies being conducted by Mr. Bernard
C. Nelson.

The following species were not consistently more abundant in
one river than the other: big-mouthed buffalo, black buffalo, small-mouthed
buffalo, short-headed redhorse, green sunfish, orange-spotted
sunfish and orange-throated darter. These species, excepting
the orange-throated darter and short-headed redhorse, occurred
in a wide variety of habitats.

FAUNAL CHANGES, 1957 THROUGH 1959

The following species increased in abundance from 1957 to 1959
(Tables 10 and 11): long-nosed gar, short-nosed gar, river carpsucker,
creek chub, gravel chub, sucker-mouthed minnow, mountain
minnow, blunt-nosed minnow, parrot minnow, stoneroller,
stonecat, Neosho madtom, green sunfish, slender-headed darter, and
orange-throated darter.

These species can be separated into three groups, characteristic
of different habitats but having in common a preference for permanent
flow. One group, composed of long-nosed gar, short-nosed
gar, river carpsucker, gravel chub, mountain minnow, parrot min[Pg 421]now,
and Neosho madtom, prefers streams of moderate to large
size.

A second group composed of creek chub, sucker-mouthed minnow,
stoneroller, and orange-throated darter occurs most abundantly
in small, permanent streams. The green sunfish may be included
here on the basis of its abundance at the upper Neosho station;
however, this is a pioneer species and does not require permanent
flow.

The third group is characteristic of continuously flowing water,
but in both upstream and downstream situations. The species in
this group (blunt-nosed minnow, stonecat, and slender-headed darter),
increased in response to a resumption of permanent flow,
but did not respond as quickly as did channel catfish, flatheads and
freshwater drum, which are discussed subsequently.

The fact that riffle-insects were abundant throughout my study
convinces me that food was not a limiting factor in the re-establishment
of the fish-fauna on riffles of the Neosho River.

The following species decreased in abundance during my study
(Tables 10 and 11): gizzard shad, carp, rosy-faced shiner, blunt-faced
shiner, red shiner, mimic shiner, black bullhead, yellow bullhead,
channel catfish, flathead, slender madtom, tadpole madtom,
freckled madtom, spotted bass, largemouth, black crappie, fan-tailed
darter, and freshwater drum.

Among the species that decreased, three groups, characteristic
of different habitats, can be distinguished. The first group occurs
most commonly in ponded conditions or in slowly flowing streams.
Species in this group are: shad, carp, black bullhead, tadpole
madtom, largemouth, black crappie, and white crappie. Bullhead,
bass and crappie commonly occur in farm ponds and lakes
in Kansas and seem less well adapted to streams. It is therefore
not surprising to find that these species decreased in abundance
when flow was resumed.

A second group, composed of rosy-faced shiner, blunt-faced
shiner, mimic shiner, slender madtom, freckled madtom, spotted
bass, and fan-tailed darter, normally is characteristic of clear
tributaries rather than the mainstream of rivers. These species
probably used the mainstream as a refugium during drought; with
the resumption of flow, conditions became unsuitable for these
populations in the mainstream. At the same time, conditions probably
became favorable to the re-establishment of these species in
tributaries. Metcalf (1959:396) listed the rosy-faced shiner, blunt-faced
[Pg 422]
shiner and mimic shiner as species that were characteristic
of upland tributaries in the Flint Hills and Chautauqua Hills of
Chautauqua, Cowley and Elk counties in Kansas. The slender
madtom and fan-tailed darter are more common in clear streams of
southeast Kansas than in other areas of the state (Cross, personal
communication and data of the State Biological Survey of Kansas).
Both species are recorded by Hall (1952:57-58) only in upland
tributaries on the east side of Grand (Neosho) River in the Fort
Gibson Reservoir area of Oklahoma. Neither species was taken
in faunal studies of the Verdigris River in Oklahoma (Wallen,
1958), in the Verdigris and Fall rivers in Kansas (Schelske, 1957),
or by Metcalf (1959).

The spotted bass is not so restricted in its distribution and its
habitat-requirements as are other species in this group; but, in
Kansas, spotted bass are most abundant in clear creeks in the southeast
part of the state.

The freckled madtom was taken in most of the studies cited above
and is most common in the smaller streams of the southeast one-fourth
of Kansas and the northeast one-fourth of Oklahoma.
Schelske (1957:47) reports that the freckled madtom was taken only
in March, April, October and November in the Verdigris River,
Kansas. My only record of this species was obtained in the Neosho
River in April, 1958.

The third group is composed of channel catfish, flathead, and
freshwater drum. This group represents that element of the population
that responded most quickly to the resumption of continuous
flow. The fact that adult channel catfish and flatheads live in pools
and do not require flowing water to spawn gives these species a
survival advantage as well as a reproductive advantage over obligatory
riffle fishes (such as most darters) in the highly variable
conditions found in Kansas streams. These factors resulted in
unusually high reproductive success in 1957. Subsequent survival
of fry was excellent; however, some mortality in the highly-dominant
1957 year-class became apparent in the 1958 and 1959
collections, accounting for a numerical decline in these species.
The ability to respond immediately to increased flow is an adaptive
feature that allows these species to maintain high levels of abundance
in the highly fluctuating streams of Kansas.

The continuous flow that occurred in 1957 in the Neosho and
Marais des Cygnes rivers, for the first time in four years, provided
the necessary habitat for survival of young catfish hatched in that
[Pg 423]
year. The nearly complete absence of other species on the riffles,
and the abundant populations of riffle-insects that I observed in
the summer of 1957, were undoubtedly factors contributing to the
survival of young.

The decrease in abundance of the red shiner may be partially
due to an increase in the numbers of other species that are well
adapted to conditions of permanent flow. At the completion of my
study, the red shiner was still the most abundant minnow in both
rivers. In 1957 this species was common in many habitats, including
swift riffles, that were later occupied by madtoms, darters,
the gravel chub, mountain minnow and sucker-mouthed minnow.

The basic pattern of change was clearly an increase in the species
that are characteristic of permanently flowing waters, and a decrease
in the species that are characteristic of ponds or small, clear
streams.

CONCLUSIONS

The fauna of the Neosho and Marais des Cygnes rivers is capable
of a wide range of adjustment in response to marked environmental
changes. As these rivers become low and clear they take on many
of the faunal characteristics of smaller tributaries and ponds. Species
such as black bullhead, spotted bass, largemouth, white crappie,
red shiner, rosy-faced shiner, blunt-faced minnow, mimic shiner,
and slender madtom assume a more prominent position in the
total population. Other species such as channel catfish, flathead,
freshwater drum, blue sucker, and such riffle-dwelling species as
the gravel chub, Neosho madtom, and slender-headed darter hold
a less prominent position in the total population.

When permanent flow is re-established the more mobile and the
more generalized species (with respect to habitat) are able to
utilize the available space immediately. As a result, these species
increase rapidly in numbers. This increase occurs both by movement
from more permanent waters and by reproduction. Channel
catfish, flathead, freshwater drum, and river carpsucker are mobile
species (Funk, 1957; Trautman, 1957) and long-nosed gar probably
are mobile. Individuals that move supplement those that survive
in residual pools, and provide brood stock adequate to produce a
large year-class in the first year of permanent flow.

The five species last mentioned are found in diverse kinds of
streams, indicating that they are adaptable to varying habitats.
A sixth species, the red shiner, although probably less mobile, is
able to utilize opportunistically nearly any kind of habitat in
[Pg 424]
Plains streams. Although this species seldom is abundant in riffles,
it was, in 1957, abundant in both pool and riffle situations at all my
stations. These riffles were almost unoccupied by other species in
1957 until mid-summer, when hatches of channel catfish and flatheads
occurred. Although adult channel catfish and flatheads live
well in pools, the young occupy mainly riffles. This age- and size-segregation,
in different habitats, was an advantage to the rapid re-establishment
of these species in the Neosho and Marais des Cygnes
rivers in 1957.

Species that occupy restricted habitats, especially riffle-dwellers
such as the Neosho madtom, gravel chub, and slender-headed darter,
were slowest to increase following drought. These species seem
less capable of adapting to the variable conditions prevalent in the
Neosho and Marais des Cygnes rivers than species that have more
generalized habitat-requirements.

In the Neosho and Marais des Cygnes rivers nearly all species
that were found in years just prior to the drought of 1952-1956
were again found in the last year of my survey; however, some
species that live in a restricted habitat may eventually be extirpated
in these two rivers. The high-finned carpsucker Carpiodes velifer,
common shiner Notropis cornutus, horny-headed chub Hybopsis
biguttata, and johnny darter Etheostoma nigrum all have specific
habitat requirements and have disappeared or become restricted
to one tributary in the Wakarusa River System (Deacon and Metcalf,
1961). The disappearance or reduction of these species implies
long-term changes in the environment.

Suckers, minnows and catfishes constitute the main fauna of the
Neosho and Marais des Cygnes rivers, because these families contain
many species that have generalized habitat-requirements.
Many of these fish are able to live successfully in either ponds or
flowing waters and others are capable of long migrations. Because
these fish predominate in the streams of Kansas, attempts should
be made to utilize them more effectively.

In years such as 1957, large numbers of young channel catfish
could be collected and used to stock new ponds and lakes. So
doing would not affect the numbers of adults produced in the
stream, and, if enough young could be removed, those remaining
in the streams might grow faster.

Suckers and carp are abundant in the two rivers and mostly are
unused at present, because current regulations preclude the use
of methods effective for the capture of these species.

[Pg 425]

ACKNOWLEDGMENTS

The investigation here reported on was supported jointly by the
Kansas Forestry, Fish and Game Commission and the State Biological
Survey of Kansas.

I thank Messrs. W. L. Minckley, D. A. Distler, J. McMullen,
A. L. Metcalf, L. J. Olund, M. Topping, B. Nelson and Claude Hastings
for assistance in the field, and Mr. Ernest Craig, Game Protector,
Erie, Kansas, for valuable suggestions and co-operation. I
am especially grateful to Associate Professor Frank B. Cross for
his pre-drought data, guidance, and criticism throughout the course
of the work. I thank the many landowners who allowed me access
to streams, and am especially indebted to Mr. and Mrs. Floyd
Meats and Mr. and Mrs. Oliver Craig for their hospitality and
assistance.

Assistant Professor Kenneth B. Armitage and Associate Professor
Ronald L. McGregor read the manuscript and gave helpful advice.
Mrs. Maxine Deacon typed the manuscript and assisted in other
ways.

FOOTNOTES

LITERATURE CITED

Anonymous.

Bailey, R. M., and Harrison, H. M., Jr.

Breder, C. M., Jr.

Cross, F. B.

——, and Minckley, W. L.

Davis, J.

[Pg 426]

Deacon, J. E.

——, and Metcalf, A. L.

Foley, F. C., Smrha, R. V., and Metzler, D. F.

Funk, J. L.

Garrett, R. A.

Hall, G. E.

Hasler, A. D. and Wisby, W. J.

Lack, D.

Marzolf, R. C.

Mead, J. R.

Metcalf, A. L.

Metzler, D. F., Culp, R. L., Stoltenberg, H. A., Woodward, R. L., Walton,
G., Chang, S. L., Clarke, N. A., Palmer, C. M., and Middleton, F. M.

Minckley, W. L.

——, and Deacon, J. E.

Muncy, R. J.

[Pg 427]

Schelske, C. L.

Schoewe, W. H.

Trautman, M. B.

Weaver, J. E., and Albertson, F. W.

    Transmitted March 30, 1961.

PLATE 26

Fig. 1. Neosho River, Middle Station, Sec. 3 and 4, T. 24 S., R. 17 E.,
looking upstream, July, 1958.



Fig. 2. Neosho River, Lower Station, Sec. 16, T. 29 S., R. 20 E., along gravel
bar, July, 1959.

PLATE 27

Fig. 1. Marais des Cygnes River, Upper Station, Sec. 12, T. 17 S., R. 17 E.,
looking downstream, June, 1960.



Fig. 2. Marais des Cygnes River, Middle Station, Sec. 6, T. 17 S., R. 20 E.,
looking downstream, June, 1960.

PLATE 28

Fig. 1. Electrical fishing gear used at night.



Fig. 2. Pool at the upper Neosho station in which rotenone was used, Sec. 33,
T. 15 S., R. 8 E., looking downstream, June, 1960.

PLATE 29

Fig. 1. Area 1, upper Neosho station, Sec. 33, T. 15 S., R. 8 E., looking
upstream, June, 1960.



Fig. 2. Area 3, upper Neosho station, Sec. 10, T. 16 S., R. 8 E., looking
downstream, June, 1960.

PLATE 30

Fig. 1. Area 5, upper Neosho station, Sec. 3, T. 16 S., R. 8 E., looking
upstream, June, 1960.



Fig. 2. Area 6, upper Neosho station, Sec. 3, T. 16 S., R. 8 E., looking
upstream, June, 1960.

28-7576

UNIVERSITY OF KANSAS PUBLICATIONS



MUSEUM OF NATURAL HISTORY

Institutional libraries interested in publications exchange may obtain this
series by addressing the Exchange Librarian, University of Kansas Library,
Lawrence, Kansas. Copies for individuals, persons working in a particular
field of study, may be obtained by addressing instead the Museum of Natural
History, University of Kansas, Lawrence, Kansas. There is no provision for
sale of this series by the University Library, which meets institutional requests,
or by the Museum of Natural History, which meets the requests of individuals.
However, when individuals request copies from the Museum, 25 cents should
be included, for each separate number that is 100 pages or more in length, for
the purpose of defraying the costs of wrapping and mailing.

* An asterisk designates those numbers of which the Museum's supply (not the Library's
supply) is exhausted. Numbers published to date, in this series, are as follows:

Vol. 1.

*Vol. 2.

Vol. 3.

*Vol. 4.

Vol. 5.

*Vol. 6.

Vol. 7.

Vol. 8.

Vol. 9.

Vol. 10.

Vol. 11.

Vol. 12.

Vol. 13.

Vol. 14.