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Title: Fish Populations, Following a Drought, in the Neosho and Marais des Cygnes Rivers of Kansas
Author: Deacon, James Everett
Language: English
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                  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



     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



                Fish Populations, Following a Drought,
              In the Neosho and Marais des Cygnes Rivers
                              of Kansas

                                  BY

                         JAMES EVERETT DEACON



CONTENTS


                                                            PAGE
  INTRODUCTION                                               363

  DESCRIPTION OF NEOSHO RIVER                                366

  DESCRIPTION OF MARAIS DES CYGNES RIVER                     367

  METHODS                                                    368
    Electrical Fishing Gear                                  368
    Seines                                                   369
    Gill Nets                                                370
    Sodium Cyanide                                           370
    Rotenone                                                 370
    Dyes                                                     370
    Determination of Abundance                               371
    Names of Fishes                                          371

  ANNOTATED LIST OF SPECIES                                  371

  FISH-FAUNA OF THE UPPER NEOSHO RIVER                       405
    Description of Study-areas                               405
    Methods                                                  406
    Changes in the Fauna at the Upper Neosho Station,
      1957 through 1959                                      407
    Local Variability of the Fauna in Different Areas
      at the Upper Neosho Station, 1959                      409
    Temporal Variability of Fauna in the Same Areas          411
    Population-Estimation                                    412
    Movement of Marked Fish                                  416
    Similarity of the Fauna at the Upper Neosho Station
      to the Faunas of Nearby Streams                        418

  COMPARISON OF THE FISH-FAUNAS OF THE NEOSHO AND MARAIS
    DES CYGNES RIVERS                                        419

  FAUNAL CHANGES, 1957 THROUGH 1959                          420

  CONCLUSIONS                                                423

  ACKNOWLEDGMENTS                                            425

  LITERATURE CITED                                           425



TABLES


                                                                PAGE
   1. Stream-flow in Cubic Feet per Second (C. F. S.), Neosho
      River near Council Grove, Kansas                           364

   2. Stream-flow in Cubic Feet per Second, Neosho River near
      Parsons, Kansas                                            364

   3. Stream-flow in Cubic Feet per Second, Marais des Cygnes
      River near Ottawa, Kansas                                  364

   4. Stream-flow in Cubic Feet per Second, Marais des Cygnes
      River at Trading Post, Kansas                              365

   5. Numbers and sizes of long-nosed gar                        372

   6. Numbers and sizes of short-nosed gar                       374

   7. Length-frequency of channel catfish from the Neosho River  388

   8. Length-frequency of freshwater drum                        402

   9. Average number of individuals captured per hour            402

  10. Numbers of fish seen or captured per hour                  403

  11. Numbers of occurrences and numbers counted                 404

  12. Percentage composition of the fish fauna at the Upper
      Neosho station in 1957, 1958 and 1959, as computed
      from results of rotenone collections                       408

  13. Relative abundance of fish                                 410

  14. Changes in numbers of individuals                          411

  15. Data used in making direct proportion
      population-estimations                                     414

  16. Data on movement of marked fish                            416



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 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.

  =========================================================
  WATER-YEAR[A]  | Average flow |   Maximum  |   Minimum  |
  ---------------+--------------+------------+------------+
  1951           |     498.0    |   121,000  |     3.0    |
  1952           |      82.1    |     4,850  |      .7    |
  1953           |       5.37   |       202  |      .1    |
  1954           |       8.53   |     2,720  |      .1    |
  1955           |      31.2    |     6,480  |     0      |
  1956           |      10.1    |     5,250  |     0      |
  1957           |      68.5    |    12,300  |     0      |
  1958           |     131.0    |     5,360  |      .8    |
  1959           |     114.0    |     7,250  |     8.5    |
  ---------------+--------------+------------+------------+

  TABLE 2. STREAM-FLOW IN CUBIC FEET PER SECOND, NEOSHO RIVER
  NEAR PARSONS, KANSAS. DRAINAGE AREA: 4905 SQUARE MILES.

  =========================================================
  WATER-YEAR[B]  | Average flow |   Maximum  |   Minimum  |
  ---------------+--------------+------------+------------+
  1951           |    8,290     |  410,000   |  124.0     |
  1952           |    2,021     |   20,500   |   20.0     |
  1953           |      173     |    4,110   |     .3     |
  1954           |      430     |   27,900   |     .1     |
  1955           |      645     |   18,600   |    0       |
  1956           |      180     |    6,170   |    0       |
  1957           |    1,774     |   25,000   |    0       |
  1958           |    3,092     |   27,200   |   78.0     |
  1959           |    1,609     |   22,600   |  139.0     |
  ---------------+--------------+------------+------------+

  TABLE 3. STREAM-FLOW IN CUBIC FEET PER SECOND, MARAIS DES CYGNES
  RIVER NEAR OTTAWA, KANSAS. DRAINAGE AREA: 1,250 SQUARE MILES.

  =========================================================
  WATER-YEAR     | Average flow |   Maximum  |   Minimum  |
  ---------------+--------------+------------+------------+
  1951           |    2,113     |   142,000  |     25.0   |
  1952           |      542     |    12,000  |       .2   |
  1953           |       36.5   |     2,690  |       .2   |
  1954           |       73.6   |     5,660  |       .5   |
  1955           |       75.7   |     5,240  |       .7   |
  1956           |       26     |     1,590  |       .7   |
  1957           |      442     |    11,200  |       .7   |
  1958           |      775     |     9,130  |      5.6   |
  ---------------+--------------+------------+------------+

  TABLE 4. STREAM-FLOW IN CUBIC FEET PER SECOND, MARAIS DES CYGNES
  RIVER AT TRADING POST, KANSAS. DRAINAGE AREA: 2,880 SQUARE MILES.

  =========================================================
  WATER-YEAR     | Average flow |   Maximum  |   Minimum  |
  ---------------+--------------+------------+------------+
  1951           |     5,489    |   148,000  |   36.0     |
  1952           |     1,750    |    20,400  |    3.0     |
  1953           |       261    |     7,590  |    0       |
  1954           |       334    |    12,500  |    0       |
  1955           |       786    |    16,100  |     .2     |
  1956           |       202    |    10,000  |    0       |
  1957           |       871    |    14,700  |    0       |
  1958           |     2,453    |    20,400  |  120.0     |
  [C]1959        |       750    |    10,900  |    3.4     |
  ---------------+--------------+------------+------------+

  [A] (Oct. 1-Sept. 30, inclusive)

  [B] (Oct. 1-Sept. 30, inclusive)

  [C] The gaging station was moved a short distance downstream
      to the Kansas-Missouri state line.



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).

  [Illustration: 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, 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 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 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 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
3/4 inch to 2-1/2 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 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 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).

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.

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).

  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.

                                     Average total
  Location          Date   Number   length (inches)   Range

  Middle Neosho     1957    19            22.2        14-32
  Middle Neosho     1958    57            22.2        14-40
  Middle Neosho     1959    64            21.6        13-43
  Lower Neosho      1957    14            29.4         9-45
  Lower Neosho      1958     7            25.3        23-28
  Lower Neosho      1959   107            26.2        16-43


#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 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.

                                            Average total
  Location                  Date   Number  length (inches)   Range

  Middle Neosho             1958    6          14.9        13.9-15.5
  Middle Neosho             1959    9          13.6        11.0-16.0
  Lower Neosho              1958    3          21.0        20.3-21.6
  Lower Neosho              1959    5          21.3        18.0-24.5


#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 communication)
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 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., 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.

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).

  [Illustration: FIG. 2. Length-frequency of river carpsucker
  in the Neosho River, 1958 and 1959.]


#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 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 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.


#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 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 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 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.

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.


#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 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 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.

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.)

                            June 2    June 7
                             1958      1958
  Length          Nov. 2   (shallow   (deep    Sept. 9   Sept. 11
  in inches        1957     riffle)  riffle)    1958       1959

   1.5                                             1
   2.0              3
   2.5             13         2                    1          2
   3.0              4        11                    3          4
   3.5              3        21          7         1         14
   4.0                       11         12                    9
   4.5                        4         10         1
   5.0                        2         11         2
   5.5                        1          7        26
   6.0                                            58          2
   6.5                                   1        32          5
   7.0                                            16          5
   7.5                                   1         4          5
   8.0                                                       22
   8.5                                                       45
   9.0                                                       81
   9.5                                                       41
  10.0                                                       21
  10.5                                                        8
  11.0                                                        4
  11.5                                                        1
  12.0                                                        3
  12.5                                                        1
  13.0                                                        1

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 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 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 station 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 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.

  [Illustration: 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 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 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. 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 Neosho 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.


#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.


#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 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 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.


#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.

  TABLE 8. LENGTH-FREQUENCY OF FRESHWATER DRUM FROM THE MIDDLE
  NEOSHO STATION IN 1957, 1958 AND 1959.

  Total length        Aug. 19    Aug. 19-26    July 27-Aug. 4
   in inches           1957         1958             1959

       2                             1
       3                  1
       4                  4
       5                             1
       6                            12
       7                            21                 1
       8                  3         14                 2
       9                  3          3                 2
      10                  4          6                 6
      11                  2          4                 1
      12                             2
      13                                               2
      14                                               1

  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.

  ======================================================================
                  | Morning    | Afternoon  | Early night | Late night |
                  | 5 hours    | 6 hours    | 18 hours    | 8 hours    |
     SPECIES      | of effort  | of effort  | of effort   | of effort  |
                  | expended   | expended   | expended    | expended   |
                  | 6:30 a.m.  | 12:30 p.m. | 6:30 p.m.   | 12:30 a.m. |
                  | 12:30 p.m. | 6:30 p.m.  | 12:30 a.m.  | 6:30 a.m.  |
  ----------------+------------+------------+-------------+------------+
  Long-nosed Gar  |  0         |  0.3 (2)   |  1.2 (21)   |  1.1 (9)   |
  Short-nosed Gar |  0.2 (1)   |  0         |  0.2 (3)    |  0.4 (3)   |
  Gizzard Shad    |  0.2 (1)   |  0.3 (2)   |  0.1 (1)    |  0.1 (1)   |
  Black Buffalo   |  0         |  0.2 (1)   |  0.1 (1)    |  0         |
  Small-mouthed   |            |            |             |            |
    Buffalo       |  0.4 (2)   |  0.3 (2)   |  0.8 (14)   |  0.8 (6)   |
  River           |            |            |             |            |
    Carpsucker    |  3.4 (17)  |  3.3 (20)  |  5.7 (102)  |  4.9 (39)  |
  Redhorse        |  0         |  0.2 (1)   |  0.6 (10)   |  0.6 (5)   |
  Carp            |  1.8 (9)   |  0.2 (1)   |  0.7 (12)   |  0.8 (6)   |
  Channel Catfish |  1.6 (8)   |  1.0 (6)   | 10.2 (183)  | 10.5 (84)  |
  Flathead        |  2.2 (11)  |  1.3 (8)   |  2.4 (43)   |  3.6 (29)  |
  Spotted Bass    |  0.4 (2)   |  0.5 (3)   |  0.3 (6)    |  0.1 (1)   |
  Green Sunfish   |  0.2 (1)   |  0.2 (1)   |  0.2 (3)    |  0.1 (1)   |
  Long-eared      |            |            |             |            |
    Sunfish       |  0         |  0         |  0.1 (2)    |  0.4 (3)   |
  Orange-spotted  |            |            |             |            |
    Sunfish       |  0.2 (1)   |  0         |  0          |  0         |
  White Crappie   |  0.2 (1)   |  0.2 (1)   |  0.2 (5)    |  0.4 (3)   |
  Freshwater Drum |  1.0 (5)   |  0.8 (5)   |  5.6 (101)  |  5.3 (42)  |
  Number captured |            |            |             |            |
    per hour      | 13.4       |  9.3       | 29.5        | 33.8       |
  ----------------+------------+------------+-------------+------------+

  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.

  ====================================================================
                     |         Marais des Cygnes River               |
                     |-----------------+-----------------+-----------+
     SPECIES         |      Upper      |     Middle      |   Lower   |
  -------------------+-----+-----+-----+-----+-----+-----+-----+-----+
                     | 1957| 1958| 1959| 1957| 1958| 1959| 1957| 1958|
  -------------------+-----+-----+-----+-----+-----+-----+-----+-----+
  Gar                |  .7 | 1.3 | 1.2 |  .6 | 2.7 | ... | 2.2 | 9.4 |
  Gizzard Shad       |  .9 |  .2 | ... | 9.9 | 2.5 | ... | ... |  .5 |
  Buffalo            | 2.0 | 3.7 |  .6 |  .8 | 2.0 | ... | 5.7 | 6.4 |
  River Carpsucker   | 4.0 | 4.9 |  .6 | 6.5 | 2.2 | 2.0 | 1.8 | 3.9 |
  Shortheaded        |     |     |     |     |     |     |     |     |
    Redhorse         | 3.3 |  .9 |  .6 |  .8 |  .2 | ... | ... | ... |
  Carp               |10.6 | 6.4 | 2.4 | 8.6 | 5.0 | 3.5 | 6.0 |10.4 |
  Black Bullhead     | ... | ... | ... | 3.9 |17.2 | ... | ... | ... |
  Channel Catfish    |  .5 |  .9 | ... | 4.7 | 2.5 | ... | 1.8 |  .7 |
  Flathead           |  .2 | ... | 2.4 |  .5 | ... | ... | 1.8 |  .5 |
  Largemouth         | 1.0 | ... | ... |  .3 |  .2 | ... | ... | ... |
  White Crappie      | 1.7 | 5.1 |  .6 | 1.3 |  .7 | ... | ... |  .2 |
  Freshwater Drum    |  .9 | 1.6 |  .6 |24.5 | 2.2 | ... |  .7 |  .2 |
                     |     |     |     |     |     |     |     |     |
  Hours shocked      |4-1/2|4-1/2|1-2/3| 4   | 4   | 2   |2-5/6|4-1/2|
  -------------------+-----+-----+-----+-----+-----+-----+-----+-----+
                     |                 Neosho River                  |
                     |-----------------------------------------------|
                     |         Middle        |         Lower         |
                     |-------+-------+-------+-------+-------+-------+
                     | 1957  | 1958  | 1959  | 1957  | 1958  | 1959  |
  -------------------|-------+-------+-------+-------+-------+-------+
  Gar                |   3.2 |   4.2 |   3.8 |   5.3 |   4.9 |   8.4 |
  Gizzard Shad       |    .5 |    .2 |    .4 |   1.9 |   1.0 |    .4 |
  Buffalo            |   2.9 |   1.8 |   1.2 |   6.2 |    .9 |   1.5 |
  River Carpsucker   |   5.5 |   7.4 |   2.9 |   7.5 |  13.3 |   6.3 |
  Shortheaded        |       |       |       |       |       |       |
    Redhorse         |   1.9 |    .6 |   1.6 |    .7 |   ... |   1.6 |
  Carp               |   2.1 |   2.1 |   1.4 |   3.4 |   1.2 |   1.1 |
  Channel Catfish    |   2.6 |   8.8 |    .9 | 107.0 |    .5 |    .7 |
  Flathead           |   7.6 |   3.7 |   2.7 |  10.8 |    .2 |   1.2 |
  Bass               |   1.6 |    .4 |    .1 |    .2 |    .2 |    .1 |
  White Crappie      |   ... |    .9 |    .2 |   1.8 |    .7 |    .1 |
  Freshwater Drum    |   3.9 |   3.3 |    .8 |  15.9 |   2.8 |    .7 |
                     |       |       |       |       |       |       |
  Hours shocked      | 5-2/3 | 55-5/6| 48-1/2| 4-1/6 |   4   | 16-5/6|
                     |       |       |       |       |       |       |
  -------------------+-------+-------+-------+-------+-------+-------+

  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.

  ======================================================================
                     |    Marais des Cygnes stations    |    Neosho
                     +-----------+----------+-----------+---------------
    SPECIES          |   Upper   |  Middle  |   Lower   | Lower station
                     +-----+-----+----+-----+-----------+------+--------
                     |1957 |1959 |1957| 1959| 1957 |1959| 1957 |  1959
  -------------------+-----+-----+----+-----+------+----+------+--------
  Golden Shiner      | ... | ... |--- | ... |  ... | ...| ...  | ...
  -------------------+-----+-----+----+-----+------+----+------+--------
  Creek Chub         | ... | --- |... | ... |  ... | ...| ...  | ...
  -------------------+-----+-----+----+-----+------+----+------+--------
  Silver Chub        | ... | ... |... | ... |  --- | ...| ...  | ...
  -------------------+-----+-----+----+-----+------+----+------+--------
  Gravel Chub        | ... | ... |... | ... |  ... | ...| ...  |   3.0
                     |     |     |    |     |      |    |      |  _2.3_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Sucker-mouthed     | --- |  6  |... |  3  |  ... |  1 |    2 |  10.0
    Minnow           |     |     |    | _1_ |      |    |      | _43.0_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Red-finned Shiner  | ... | ... |... |  1  |  2.5 |  2 |  ... |   4.7
                     |     |     |    |     | _5.0_|    |      |  _2.3_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Blunt-faced Shiner | ... | ... |--- | ... |  ... | ...|  ... | ...
  -------------------+-----+-----+----+-----+------+----+------+--------
  Red Shiner         |21   | 15  | 8  | 19  | 16.0 | 15 |   27 |  20.0
                     |_6_  |     |_4_ |_22_ |_69.0_|_22_|_1119_|_102.0_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Mimic Shiner       | ... | ... |... | ... |  ... | ...|  --- | ...
  -------------------+-----+-----+----+-----+------+----+------+--------
  Ghost Shiner       | 7.5 |  1  |... |  1  |  9.5 |  2 |   17 |  11.7
                     |     |     |    |     |_96.5_|    |  _54_| _76_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Sand Shiner        | --- |  7  |... |  8  |  1.5 |  3 |  ... |   1
                     |     |     |    | _2_ |      |    |      |   _.3_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Mountain Minnow    | ... | ... |... | ... |  ... | ...|   12 |   9.3
                     |     |     |    |     |      |    |  _25_| _13.6_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Blunt-nosed Minnow | --- |  2  |... |  8  |  1.0 |  1 |    6 |  14.0
                     |     |     |    |     |  _.5_|    |   _4_|  _7.6_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Parrot Minnow      | ... | ... |... | ... |  ... | ...|   12 |  19.0
                     |     |     |    |     |      |    |   _6_| _28.6_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Fat-headed Minnow  |10.5 |  4  | 5  |  7  |  ... | ...|   ...|   8.3
                     |_1.5_|     |_2_ | _1_ |      |    |      |  _3.0_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Stoneroller        | --- |  6  |--- | ... |  ... | ...|  --- |   2.3
                     |     |     |    |     |      |    |      |  _1.0_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Black Bullhead     | ... | ... |... | ... |   .5 | ...|  ... | ...
  -------------------+-----+-----+----+-----+------+----+------+--------
  Channel Catfish    | 4.5 |  2  | 1  | 13  |  5.0 | 10 |   12 |   6.3
                     |_1.5_|     |_1_ | _7_ | _1.0_| _6_|   _5_| _41.6_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Flathead           | --- |  1  |--- | --- |  1.0 | ...|  --- |    .3
  -------------------+-----+-----+----+-----+------+----+------+--------
  Stonecat           | ... | ... |--- | ... |  6.0 | ...|  --- |   1.0
                     |     |     |    |     |  _.5_|    |      |
  -------------------+-----+-----+----+-----+------+----+------+--------
  Neosho Madtom      | ... | ... |... | ... | ...  | ...|  ... |   3.3
                     |     |     |    |     |      |    |      |  _2.0_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Brook Silversides  | ... | ... |... | ... |   .5 | ...|  ... |   1.7
                     |     |     |    |     | _1.0_|    |      |
  -------------------+-----+-----+----+-----+------+----+------+--------
  Black-striped      | ... | ... |... | ... |  1.0 |  2 |  ... |   1.0
    Topminnow        |     |     |    |     | _1.0_|    |      |   _.7_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Spotted Bass       | ... | ... |... | ... | ...  | ...|    2 |   3.7
                     |     |     |    |     |      |    |      |   _.3_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Largemouth         | ... | ... | 1  |  3  | ...  | ...|    1 | ...
                     |     |     |_1_ | _1_ |      |    |   _2_|
  -------------------+-----+-----+----+-----+------+----+------+--------
  Green Sunfish      | 9   |  8  | 9  | 17  | 11.0 |  3 |    7 |  10.0
                     |_7.5_|     |_3_ | _3_ |_12.0_| _1_|   _2_|  _3.6_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Long-eared Sunfish | ... | ... |... | ... |   .5 | ...|    6 |   4.3
                     |     |     |    |     |      |    |      |   _.7_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Orange-spotted     | 4.5 | --- | 2  |  3  |  2.5 | ...|   12 |  12.0
    Sunfish          |_6_  |     |_4_ |     |      |    |   _5_|  _5.0_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Bluegill           | 1.5 |  1  |... |  6  |  3.5 |  1 |    1 |    .3
                     |     |     |    | _1_ |      |    |      |   _.3_
  -------------------+-----+-----+----+-----+------+----+------+--------
  White Crappie      | ... | ... | 4  |  4  |  ... | ...| ...  | ...
                     |     |     |_7_ |     |      |    |      |
  -------------------+-----+-----+----+-----+------+----+------+--------
  Logperch           | ... | ... |... | ... |  ... | ...|    1 |    .3
                     |     |     |    |     |      |    |      |   _.7_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Slender-headed     | --- | 13  |... |  2  |  6.5 |  3 |    1 |   8.3
    Darter           |     |     |    |     |_15.0_| _1_|      |  _3.0_
  -------------------+-----+-----+----+-----+------+----+------+--------
  Orange-throated    | --- |  7  |... | ... |  ... | ...|    1 | ---
    Darter           |     |     |    |     |      |    |      |
  -------------------+-----+-----+----+-----+------+----+------+--------
  Seining units      | 2/3 |  1  | 1  |  1  |  2   |  1 |    1 |   3
  -------------------+-----+-----+----+-----+------+----+------+--------



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 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.

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.

  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.

  ============================================================
               SPECIES              |  1957  | 1958  | 1959
  ----------------------------------+--------+-------+--------
  Big-mouthed Buffalo...............| ...... |  T[D] |   T
  Small-mouthed Buffalo.............| ...... | ..... |   T
  River Carpsucker..................|    T   |  0.8  |   1.8
  Golden Redhorse...................|    T   |  3.0  |   5.7
  Creek Chub........................| ...... |  T    |   0.8
  Red-finned Shiner.................|  1.3   |  3.0  |   0.8
  Red Shiner........................|  6.5   | 13.1  |  12.1
  Ghost Shiner......................|    T   |  T    | ......
  Blunt-nosed Minnow................|    T   |  T    |   T
  Fat-headed Minnow.................|    T   |  T    |   1.4
  Stoneroller.......................|  0.8   |  1.5  |   3.5
  Black Bullhead....................| 40.8   | 30.5  |  32.0
  Yellow Bullhead...................|  5.3   |  8.8  |   2.5
  Channel Catfish...................| 28.4   | 15.5  |  18.5
  Flathead..........................|    T   |  T    |   T
  Stonecat..........................|    T   |  T    |   1.4
  Spotted Bass......................|    T   |  T    |   0.8
  Largemouth........................|    T   |  T    |   T
  Green Sunfish.....................|  3.1   |  6.8  |   6.4
  Long-eared Sunfish................|  8.8   |  3.7  |   1.9
  Orange-spotted Sunfish............|  3.1   |  8.9  |   2.5
  Bluegill..........................|    T   |  T    |   T
  White Crappie.....................|    T   | ..... |   T
  Logperch......................... |    T   |  2.1  |   0.8
  Slender-headed Darter.............|  0.6   |  0.6  |   3.1
  Orange-throated Darter............| ...... |  T    |   2.5
  Total number of fish..............|  786   |  965  | 513
  Size of sample-area in acre-feet..| .002   |  .33  |    .33
  ----------------------------------+--------+-------+--------

  [D] T denotes less than one-half of one per cent of the population.

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.


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 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.

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.

  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).

  ======================================================================
                          | Area | Area | Area | Area |          |  All
                          |   1  |   3  |   5  |   6  | Rotenone | areas
  ------------------------+------+------+------+------+----------+------
  Big-mouthed Buffalo     | .... | .... | T[E] | .... |    T     |  T
  Small-mouthed Buffalo   | .... | .... |   .6 | .... |    T     |  T
  River Carpsucker        | .... |  T   | 10.6 |  T   |    1.8   |   .8
  River Carpsucker (yy)[F]| .... |   .8 |  T   |  3.7 |    ....  |  1.0
  Short-headed Redhorse   | .... | .... |   .6 | .... |    ....  |  T
  Golden Redhorse         |   .8 |  1.0 |  3.2 | .... |    5.7   |  T
  Carp                    | .... | .... | .... | .... |    ....  |  T
  Golden Shiner           | .... | .... | .... | .... |    ....  |  T
  Creek Chub              |  1.6 |  T   |  T   |  T   |     .8   |  T
  Sucker-mouthed Minnow   | .... | 11.2 |  T   |  3.4 |    ....  |  1.4
  Red-finned Shiner       | .... | .... | .... |  4.0 |     .8   |  8.1
  Red Shiner              | 18.2 | 24.0 |  7.8 | 20.1 |   12.1   | 35.9
  Sand Shiner             | .... |  5.2 | .... |  1.1 |    ....  |  T
  Pimephales (yy)         | .... | .... | .... | .... |    ....  |  6.7
  Mountain Minnow         | .... | .... | .... |  T   |    ....  |  T
  Blunt-nosed Minnow      | .... |   .8 |  4.1 | 11.7 |    T     |  3.4
  Parrot Minnow           | .... | .... | .... | .... |    ....  |  T
  Fat-headed Minnow       |  T   |  T   |  3.4 | 12.1 |    1.4   |  2.6
  Stoneroller             | 27.7 | 17.4 |   .6 |  5.8 |    3.5   |  5.1
  Black Bullhead          |  2.1 |  T   |  7.3 |  T   |   32.0   |   .6
  Yellow Bullhead         |  T   |  T   | .... |  T   |    2.5   |  T
  Channel Catfish (j)[G]  |  5.8 |  7.6 | 41.3 |  T   |   14.6   |  4.2
  Channel Catfish (yy)    |  9.5 |  7.0 |  T   |  4.3 |    3.9   |  2.5
  Flathead (j)            | .... |   .8 |  2.1 |  T   |    T     |  T
  Flathead (yy)           |  1.6 |  T   | .... | .... |    ....  |  T
  Stonecat                | 10.3 |  1.4 | .... | .... |    1.4   |   .7
  Spotted Bass            | .... |  T   |   .6 |  T   |     .8   |  T
  Largemouth              | .... | .... |  T   | .... |    T     |  T
  Green Sunfish           | 11.2 |  3.5 |  5.9 | 12.2 |    6.4   | 10.1
  Long-eared Sunfish      |  5.4 |  6.0 |  5.1 | 14.6 |    1.9   | 12.8
  Orange-spotted Sunfish  |  T   |  T   |  1.4 |  1.8 |    2.5   |   .5
  Bluegill                | .... | .... |  1.0 | .... |    T     |  T
  White Crappie           | .... | .... | .... | .... |    T     |  T
  Logperch                |  T   |  T   |  T   |  T   |     .8   |  T
  Slender-headed Darter   |  T   | 11.4 |  1.1 |  1.6 |    3.1   |  1.3
  Orange-throated Darter  |   .8 |  1.8 |  T   |   .5 |    2.5   |  T
  Freshwater Drum         | .... | .... |  T   | .... |    ....  |  T
  Total number of fish    |  242 |  484 |  727 |  924 |    513   |17,796
  Area in square feet     |  840 | 6324 |12500 |10000 |    ....  | ....
  Volume                  | .... | .... | .... | .... |   1/3    |
                          |      |      |      |      |acre-foot |
  ------------------------+------+------+------+------+----------+------

  [E] "T" designates species that comprised less than 0.5 per cent
      of the population.

  [F] (yy) signifies young-of-the-year.

  [G] (j) signifies yearlings or two-year-olds.


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.

  ======================================================================
                  |     Area 1      |     Area 3      |     Area 6
       SPECIES    +-----+-----+-----+-----+-----+-----+-----+-----+-----
                  |  3  |  4  |  8  |  9  |  10 |  15 |  16 |  18 |  20
  ----------------+-----+-----+-----+-----+-----+-----+-----+-----+-----
  Golden Redhorse |   2 |   2 | ... |   5 |   5 |   2 | ... | ... |   3
  Creek Chub      |   4 |   3 |   7 |   1 | ... | ... |   1 |   2 | ...
  Sucker-mouthed  |     |     |     |     |     |     |     |     |
    Minnow        | ... | ... | ... |  54 |  42 |  25 |  31 |   7 |   6
  Red-finned      |     |     |     |     |     |     |     |     |
    Shiner        | ... | ... |   1 | ... | ... |   4 |  31 |  13 |   4
  Red Shiner      |  44 |   7 | 211 | 117 | 170 | 438 | 186 | 209 |  62
  Blunt-nosed     |     |     |     |     |     |     |     |     |
    Minnow        | ... | ... | ... |   4 |  10 |  19 | 108 |  91 |  13
  Fat-headed      |     |     |     |     |     |     |     |     |
    Minnow        |   1 | ... | ... |   1 |   2 |   3 | 112 | 156 |  48
  Stoneroller     |  67 |  39 |  49 |  84 | 107 |  55 |  54 |  67 |  22
  Black Bullhead  |   5 | ... |   1 |   2 |   1 | ... | ... |   3 |   7
  Yellow Bullhead |   1 |   1 | ... |   2 |   1 | ... |   1 | ... |   3
  Channel Catfish |  14 |   7 | ... |  36 |  16 | ... |   3 |   1 |  23
  Channel         |     |     |     |     |     |     |     |     |
    Catfish(yy)[H]|  23 |  16 |  17 |  34 |  34 |  22 |  40 |  23 |  28
  Flathead        | ... | ... | ... |   4 |   8 |   1 |   2 | ... |   1
  Flathead(yy)    |   4 |   1 |   1 |   2 |   1 |   1 | ... | ... | ...
  Stonecat        |  25 |   8 |  12 |   7 |   7 |   5 | ... | ... | ...
  Green Sunfish   |  27 |  17 |  12 |  13 |  16 |  17 |  62 |  62 |  74
  Long-eared      |     |     |     |     |     |     |     |     |
    Sunfish       |  13 |  12 |   1 |   6 |   3 |   3 |  10 |  22 |  31
  Logperch        |   1 | ... | ... |   2 | ... | ... | ... | ... | ...
  Slender-headed  |     |     |     |     |     |     |     |     |
    Darter        | ... |   1 |   2 |  55 |  45 |  23 |  15 |   1 |   1
  Orange-throated |     |     |     |     |     |     |     |     |
    Darter        |   2 |   1 |   2 |   9 |  11 |   8 |   5 | ... |   1
  ----------------+-----+-----+-----+-----+-----+-----+-----+-----+-----
  Total           | 233 | 115 | 316 | 438 | 480 | 626 | 661 | 657 | 347
  ----------------+-----+-----+-----+-----+-----+-----+-----+-----+-----

  [H] (yy) means young-of-the-year only.

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.

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.


_Area 1_

The order of abundance at Area 1, in terms of the estimated population
per 500 square feet, was as follows: stoneroller (47.6), stonecat
(29.4), channel catfish (young-of-the-year) (20.6), green sunfish
(19.4), red shiner (18.2), long-eared sunfish (9.4), channel catfish
(yearlings and older) (6.5), golden redhorse (1.2). Insufficient data
make inclusion of other species unreliable.

A comparison of the order of abundance between the estimated total
population and the percentage composition in the first collection from
each area shows significant correlations. The percentage-composition of
the fish fauna at Area 1 was calculated as follows: stoneroller (27.7%),
red shiner (18.2%), green sunfish (11.2%), stonecat (10.3%), channel
catfish (young-of-the-year) (9.5%), channel catfish (yearlings and
older) (5.8%), long-eared sunfish (5.4%), golden redhorse (0.8%). It can
be seen that the stoneroller, green sunfish, long-eared sunfish and
golden redhorse follow each other in the same order in both
calculations. The stonecat is shown to be more common than channel
catfish (young-of-the-year) in both calculations, but both species
appear to be more abundant than green sunfish and red shiner in
calculations of the total population and less abundant in the
percentage-composition in the first collection. I think that the order
of abundance as shown by percentage-composition is the more accurate
figure for Area 1. The abundance of the red shiner is known to have been
affected by mortality in collecting. Furthermore, as will be shown
later, the species is so mobile that its abundance often changes
markedly in a short time. Therefore, it is not surprising to find the
red shiner in widely varying positions of relative and absolute
abundance. However, the green sunfish maintains stable populations and
should remain in about the same position of abundance in relation to
other species (such as the stonecat and channel catfish
young-of-the-year) that also maintain stable populations. The
differences in order of abundance obtained by the two methods for green
sunfish and channel catfish young-of-the-year are not great. However, in
the estimation of total population the abundance of the stonecat seems
significantly greater, in relation to other species, than in the
calculation of percentage-composition. I believe that this difference
can be attributed to the relatively low number of marked fish
recaptured, which is probably due to a slow rate of dispersal from the
point of release. Stonecats were released in relatively quiet water, and
if they remained there they might be missed in subsequent collections,
because they lack air-bladders and tend to remain on the bottom when
shocked. Therefore, the calculated total population of the stonecat in
Area 1 may be too high.


  TABLE 15. DATA USED IN ESTIMATING TOTAL POPULATIONS, BY DIRECT
  PROPORTIONS, IN AREAS 1, 3, AND 6 AT THE UPPER NEOSHO STATIONS.

  ======================================================================
                        |     Number     |    Number    |     Number
                        | captured first |  marked and  |captured second
          SPECIES       |   collection   |   released   |   collection
                        +----+-----+-----+----+----+----+----+-----+----
                        |  1 |  3  |  6  | 1  | 3  | 6  | 1  |  3  |  6
  ----------------------+----+-----+-----+----+----+----+----+-----+----
  Golden Redhorse       |  2 |   5 |   0 |  2 |  5 |  0 |  2 |   5 |   0
  Sucker-mouthed Minnow |  0 |  54 |  31 |  0 | 51 | 15 |  0 |  42 |  12
  Red Shiner            | 44 | 116 | 186 | 22 |106 | 86 |  7 | 165 | 202
  Sand Shiner           |  0 |  25 |  10 |  0 | 25 |  7 |  0 |  35 |  10
  Blunt-nosed Minnow    |  0 |   4 | 108 |  0 |  3 | 28 |  0 |  10 |  91
  Fat-headed Minnow     |  1 |   1 | 112 |  1 |  1 |101 |  0 |   2 | 156
  Stoneroller           | 67 |  84 |  54 | 58 | 79 | 33 | 39 | 107 |  67
  Channel Catfish(j)[I] | 14 |  37 |   3 |  9 | 32 |  3 |  7 |  16 |   1
  Channel Catfish(yy)[J]|  3 |  34 |  40 | 22 | 33 | 39 | 16 |  34 |  23
  Stonecat              | 25 |   7 |   0 | 25 |  7 |  0 |  8 |   7 |   0
  Green Sunfish         | 27 |[K]--|  62 | 27 | -- | 62 | 17 |  -- |  62
  Long-eared Sunfish    | 13 |   6 |  10 | 13 |  6 | 10 | 12 |   3 |  22
  ----------------------+----+-----+-----+----+----+----+----+-----+----
  ======================================================================
     Number of  |    Estimated    |   Percent of   |      Number
    marked fish |      total      |   marked fish  |      per 500
    recaptured  |    population   |    recovered   |    square feet
  ----+----+----+----+-----+------+-----+-----+----+------+------+------
    1 | 3  | 6  | 1  |  3  |   6  |  1  |  3  | 6  |   1  |   3  |  6
  ----+----+----+----+-----+------+-----+-----+----+------+------+------
    2 |  5 |  0 |  2 |   5 |    0 | 100 | 100 | -- |  1.2 |   .4 |  0
    0 | 17 |  0 |  0 | 126 |   -- |  -- |  33 |  0 |    0 | 10.0 | --
    5 | 18 | 14 | 31 | 972 | 1284 |  23 |  17 | 11 | 18.2 | 77.1 | 64
   -- | 12 |  1 |  0 |  73 |   -- |  -- |  48 | -- |    0 |  5.8 | --
    0 |  1 |  8 |  0 |  -- |  319 |  -- |  33 | 28 |    0 |   -- | 16
    0 |  0 | 19 | -- |  -- |  830 |   0 |   0 | 19 |   -- |   -- | 41.5
   28 | 35 |  8 | 81 | 242 |  276 |  48 |  44 | 24 | 47.6 | 19.2 | 13.8
    6 | 13 |  0 | 11 |  39 |   -- |  67 |  41 |  0 |  6.5 |  3.1 | --
   10 | 11 |  1 | 35 | 102 |   -- |  45 |  33 |  3 | 20.6 |  8.1 | --
    4 |  1 | -- | 50 |  -- |    0 |  16 |  14 | -- | 29.4 |   -- |  0
   14 | -- | 22 | 33 |  -- |  175 |  52 |  -- | 35 | 19.4 |   -- |  8.8
   10 |  3 |  6 | 16 |   6 |   37 |  76 |  50 | 60 |  9.4 |   .5 |  1.9
  ----+----+----+----+-----+------+-----+-----+----+------+------+------

  [I] (j) Denotes juveniles only.

  [J] (yy) Denotes young-of-year only.

  [K] A dash denotes incomplete or insufficient data.


_Area 3_

The order of abundance of the species at Area 3, in terms of the
estimated population per 500 square feet, was as follows: red shiner
(77.1), stoneroller (19.2), sucker-mouthed minnow (10.0), channel
catfish (young-of-the-year) (8.1), sand shiner (5.8), channel catfish
(yearlings and older) (3.1), long-eared sunfish (0.5), golden redhorse
(0.4). Insufficient data make inclusion of other species unreliable.

For comparison with the estimates of total population, the
percentage-composition in the first collection gives the following
results: red shiner (24.0%), stoneroller (17.4%), sucker-mouthed minnow
(11.2%), channel catfish (yearlings and older) (7.6%), channel catfish
(young-of-the-year) (7.0%), long-eared sunfish (6.0%), sand shiner
(5.2%), and golden redhorse (1.0%).

For the most part, the species have the same order of abundance in both
methods of analysis. Those that are apparently out of order are channel
catfish (yearlings and older) and long-eared sunfish. The first species
is mobile (excepting young-of-the-year) and commonly fluctuates widely
in numbers in the same area; the second species was treated differently
in that only adults were considered in the population-estimation
whereas both young and adults were considered in calculating
percentage-composition. (I found that I could not confidently
distinguish between young-of-the-year of green sunfish, long-eared
sunfish and orange-spotted sunfish after staining.)


_Area 6_

The order of abundance of the species at Area 6, in terms of the
estimated population per 500 square feet, was as follows: red shiner
(64.0), fat-headed minnow (41.5), blunt-nosed minnow (16.0), stoneroller
(13.8), green sunfish (8.8), long-eared sunfish (1.9). Insufficient data
make inclusion of other species unreliable.

Calculations of percentage-composition give the following results: red
shiner (20.1%), long-eared sunfish (14.6%), green sunfish (12.2%),
fat-headed minnow (12.1%), blunt-nosed minnow (11.7%), stoneroller
(5.8%). The two species of sunfish form a more significant part of the
population in the latter analysis because young are included. Only
adults were considered in the estimation of total population.

The fact that estimates of the total population and the
percentage-composition agree in most respects lends support to the
validity of both methods of analysis. It should be re-emphasized that
differences in the order of abundance in the various areas reflect the
ability of each species to utilize each particular kind of habitat.


Movement of Marked Fish

  TABLE 16. DATA ON MOVEMENT OF MARKED FISH AT THE UPPER NEOSHO
  STATION, SEPTEMBER, 1959.

  ======================================================================
                          | Number |   Number   |  Number  |   Number
           SPECIES        | marked | recaptured |  moved   |   moved
                          |        |            | upstream | downstream
  ------------------------+--------+------------+----------+--------------
  Golden Redhorse         |    24  |    16      |     0    |     2
  Sucker-mouthed Minnow   |    68  |    27      |     7    |     0
  Red-finned Shiner       |    74  |     0      |     0    |     0
  Red Shiner              |  1326  |   152      |    48    |    25
  Blunt-nosed Minnow      |   136  |    32      |     1    |    10
  Fat-headed Minnow       |   151  |    40      |     0    |     0
  Stoneroller             |   177  |    90      |     1    |     0
  Black Bullhead          |    25  |     6      |     2    |     0
  Channel Catfish (j)[L]  |   294  |    36      |     4    |     7
  Channel Catfish (yy)[M] |   145  |    34      |     2    |     0
  Stonecat                |    33  |     6      |     0    |     0
  Green Sunfish           |   124  |    68      |     1    |     0
  Long-eared Sunfish      |    33  |    21      |     0    |     0
  Slender-headed Darter   |    70  |     1      |     0    |     0
  Orange-throated Darter  |    13  |     0      |     0    |     0
  ------------------------+--------+------------+----------+------------

  [L] (j) denotes juveniles only.

  [M] (yy) denotes young-of-year only.

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.

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 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 restricted 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 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 minnow, 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 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 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.


PLATE 26

  [Illustration: FIG. 1. Neosho River, Middle Station, Sec. 3 and 4,
  T. 24 S., R. 17 E., looking upstream, July, 1958.]

  [Illustration: FIG. 2. Neosho River, Lower Station, Sec. 16,
  T. 29 S., R. 20 E., along gravel bar, July, 1959.]


PLATE 27

  [Illustration: FIG. 1. Marais des Cygnes River, Upper Station,
  Sec. 12, T. 17 S., R. 17 E., looking downstream, June, 1960.]

  [Illustration: FIG. 2. Marais des Cygnes River, Middle Station,
  Sec. 6, T. 17 S., R. 20 E., looking downstream, June, 1960.]


PLATE 28

  [Illustration: FIG. 1. Electrical fishing gear used at night.]

  [Illustration: 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

  [Illustration: FIG. 1. Area 1, upper Neosho station, Sec. 33,
  T. 15 S., R. 8 E., looking upstream, June, 1960.]

  [Illustration: FIG. 2. Area 3, upper Neosho station, Sec. 10,
  T. 16 S., R. 8 E., looking downstream, June, 1960.]


PLATE 30

  [Illustration: FIG. 1. Area 5, upper Neosho station, Sec. 3,
  T. 16 S., R. 8 E., looking upstream, June, 1960.]

  [Illustration: FIG. 2. Area 6, upper Neosho station, Sec. 3,
  T. 16 S., R. 8 E., looking upstream, June, 1960.]



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 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.



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.



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BREDER, C. M., JR.

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DAVIS, J.

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DEACON, J. E.

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WALTON, G., CHANG, S. L., CLARKE, N. A., PALMER, C. M., and
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MINCKLEY, W. L.

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----, and DEACON, J. E.

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SCHELSKE, C. L.

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  _Transmitted March 30, 1961._


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          11. A new subspecies of pocket mouse from Kansas. By E.
              Raymond Hall. Pp. 587-590. November 15, 1954.

          12. Geographic variation in the pocket gopher, Cratogeomys
              castanops, in Coahuila, Mexico. By Robert J. Russell and
              Rollin H. Baker. Pp. 591-608. March 15, 1955.

          13. A new cottontail (Sylvilagus floridanus) from northeastern
              Mexico. By Rollin H. Baker. Pp. 609-612. April 8, 1955.

          14. Taxonomy and distribution of some American shrews. By
              James S. Findley. Pp. 613-618. June 10, 1955.

          15. The pigmy woodrat, Neotoma goldmani, its distribution and
              systematic position. By Dennis G. Rainey and Rollin H.
              Baker. Pp. 619-624, 2 figures in text. June 10, 1955.

          Index. Pp. 625-651.

  Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956.

  Vol. 9.  1. Speciation of the wandering shrew. By James S. Findley.
              Pp. 1-68, 18 figures in text. December 10, 1955.

           2. Additional records and extension of ranges of mammals
              from Utah. By Stephen D. Durrant, M. Raymond Lee, and
              Richard M. Hansen. Pp. 69-80. December 10, 1955.

           3. A new long-eared myotis (Myotis evotis) from northeastern
              Mexico. By Rollin H. Baker and Howard J. Stains.
              Pp. 81-84. December 10, 1955.

           4. Subspeciation in the meadow mouse, Microtus
              pennsylvanicus, in Wyoming. By Sydney Anderson.
              Pp. 85-104, 2 figures in text. May 10, 1956.

           5. The condylarth genus Ellipsodon. By Robert W. Wilson.
              Pp. 105-116, 6 figures in text. May 19, 1956.

           6. Additional remains of the multituberculate genus
              Eucosmodon. By Robert W. Wilson. Pp. 117-123, 10 figures
              in text. May 19, 1956.

           7. Mammals of Coahuila, Mexico. By Rollin H. Baker.
              Pp. 125-335, 75 figures in text. June 15, 1956.

           8. Comments on the taxonomic status of Apodemus peninsulae,
              with description of a new subspecies from North China.
              By J. Knox Jones, Jr. Pp. 337-346, 1 figure in text,
              1 table. August 15, 1956.

           9. Extensions of known ranges of Mexican bats. By Sydney
              Anderson. Pp. 347-351. August 15, 1956.

          10. A new bat (Genus Leptonycteris) from Coahuila. By Howard
              J. Stains. Pp. 353-356. January 21, 1957.

          11. A new species of pocket gopher (Genus Pappogeomys) from
              Jalisco, Mexico. By Robert J. Russell. Pp. 357-361.
              January 21, 1957.

          12. Geographic variation in the pocket gopher, Thomomys
              bottae, in Colorado. By Phillip M. Youngman. Pp. 363-387,
              7 figures in text. February 21, 1958.

          13. New bog lemming (genus Synaptomys) from Nebraska. By J.
              Knox Jones, Jr. Pp. 385-388. May 12, 1958.

          14. Pleistocene bats from San Josecito Cave, Nuevo León,
              México. By J. Knox Jones, Jr. Pp. 389-396. December 19,
              1958.

          15. New subspecies of the rodent Baiomys from Central America.
              By Robert L. Packard. Pp. 397-404. December 19, 1958.

          16. Mammals of the Grand Mesa, Colorado. By Sydney Anderson.
              Pp. 405-414, 1 figure in text, May 20, 1959.

          17. Distribution, variation, and relationships of the montane
              vole, Microtus montanus. By Sydney Anderson. Pp. 415-511,
              12 figures in text, 2 tables. August 1, 1959.

          18. Conspecificity of two pocket mice, Perognathus goldmani
              and P. artus. By E. Raymond Hall and Marilyn Bailey
              Ogilvie. Pp. 513-518, 1 map. January 14, 1960.

          19. Records of harvest mice, Reithrodontomys, from Central
              America, with description of a new subspecies from
              Nicaragua. By Sydney Anderson and J. Knox Jones, Jr.
              Pp. 519-529. January 14, 1960.

          20. Small carnivores from San Josecito Cave (Pleistocene),
              Nuevo León, México. By E. Raymond Hall. Pp. 531-538,
              1 figure in text. January 14, 1960.

          21. Pleistocene pocket gophers from San Josecito Cave, Nuevo
              León, México. By Robert J. Russell. Pp. 539-548, 1 figure
              in text. January 14, 1960.

          22. Review of the insectivores of Korea. By J. Knox Jones,
              Jr., and David H. Johnson. Pp. 549-578. February 23, 1960.

          23. Speciation and evolution of the pygmy mice, genus Baiomys.
              By Robert L. Packard. Pp. 579-670, 4 plates, 12 figures in
              text. June 16, 1960.

          Index. Pp. 671-690.

  Vol. 10. 1. Studies of birds killed in nocturnal migration. By
              Harrison B. Tordoff and Robert M. Mengel. Pp. 1-44,
              6 figures in text, 2 tables. September 12, 1956.

           2. Comparative breeding behavior of Ammospiza caudacuta and
              A. maritima. By Glen E. Woolfenden. Pp. 45-75, 6 plates,
              1 figure. December 20, 1956.

           3. The forest habitat of the University of Kansas Natural
              History Reservation. By Henry S. Fitch and Ronald R.
              McGregor. Pp. 77-127, 2 plates, 7 figures in text,
              4 tables. December 31, 1956.

           4. Aspects of reproduction and development in the prairie
              vole (Microtus ochrogaster). By Henry S. Fitch. Pp.
              129-161, 8 figures in text, 4 tables. December 19, 1957.

           5. Birds found on the Arctic slope of northern Alaska. By
              James W. Bee. Pp. 163-211, plates 9-10, 1 figure in text.
              March 12, 1958.

           6. The wood rats of Colorado: distribution and ecology. By
              Robert B. Finley, Jr. Pp. 213-552, 34 plates, 8 figures
              in text, 35 tables. November 7, 1958.

           7. Home ranges and movements of the eastern cottontail in
              Kansas. By Donald W. Janes. Pp. 553-572, 4 plates,
              3 figures in text. May 4, 1959.

           8. Natural history of the salamander, Aneides hardyi. By
              Richard F. Johnston and Gerhard A. Schad. Pp. 573-585.
              October 8, 1959.

           9. A new subspecies of lizard, Cnemidophorus sacki, from
              Michoacán, México. By William E. Duellman, Pp. 587-598,
              2 figures in text. May 2, 1960.

          10. A taxonomic study of the Middle American Snake, Pituophis
              deppei. By William E. Duellman. Pp. 599-610, 1 plate,
              1 figure in text. May 2, 1960.

          Index. Pp. 611-626.

  Vol. 11. 1. The systematic status of the colubrid snake, Leptodeira
              discolor Günther. By William E. Duellman. Pp. 1-9,
              4 figures. July 14, 1958.

           2. Natural history of the six-lined racerunner, Cnemidophorus
              sexlineatus. By Henry S. Fitch. Pp. 11-62, 9 figures,
              9 tables. September 19, 1958.

           3. Home ranges, territories, and seasonal movements of
              vertebrates of the Natural History Reservation. By Henry
              S. Fitch. Pp. 63-326, 6 plates, 24 figures in text,
              3 tables. December 12, 1958.

           4. A new snake of the genus Geophis from Chihuahua, Mexico.
              By John M. Legler. Pp. 327-334, 2 figures in text.
              January 28, 1959.

           5. A new tortoise, genus Gopherus, from north-central Mexico.
              By John M. Legler. Pp. 335-343. April 24, 1959.

           6. Fishes of Chautauqua, Cowley and Elk counties, Kansas.
              By Artie L. Metcalf. Pp. 345-400, 2 plates, 2 figures in
              text, 10 tables. May 6, 1959.

           7. Fishes of the Big Blue River Basin, Kansas. By W. L.
              Minckley. Pp. 401-442, 2 plates, 4 figures in text,
              5 tables. May 8, 1959.

           8. Birds from Coahuila, México. By Emil K. Urban.
              Pp. 443-516. August 1, 1959.

           9. Description of a new softshell turtle from the
              southeastern United States. By Robert G. Webb. Pp.
              517-525, 2 plates, 1 figure in text. August 14, 1959.

          10. Natural history of the ornate box turtle, Terrapene ornata
              ornata Agassiz. By John M. Legler. Pp. 527-669, 16 pls.,
              29 figures in text. March 7, 1960.

          Index Pp. 671-703.

  Vol. 12. 1. Functional morphology of three bats: Eumops, Myotis,
              Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates,
              24 figures in text. July 8, 1959.

           2. The ancestry of modern Amphibia: a review of the evidence.
              By Theodore H. Eaton, Jr. Pp. 155-180, 10 figures in text.
              July 10, 1959.

           3. The baculum in microtine rodents. By Sydney Anderson.
              Pp. 181-216, 49 figures in text. February 19, 1960.

           4. A new order of fishlike Amphibia from the Pennsylvanian
              of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou
              Stewart. Pp. 217-240, 12 figures in text. May 2, 1960.

          More numbers will appear in volume 12.

  Vol. 13. 1. Five natural hybrid combinations in minnows (Cyprinidae).
              By Frank B. Cross and W. L. Minckley. Pp. 1-18.
              June 1, 1960.

           2. A distributional study of the amphibians of the Isthmus of
              Tehuantepec, México. By William E. Duellman. Pp. 19-72,
              pls. 1-8, 3 figures in text. August 16, 1960.

           3. A new subspecies of the slider turtle (Pseudemys scripta)
              from Coahuila, México. By John M. Legler. Pp. 73-84,
              pls. 9-12, 3 figures in text. August 16, 1960.

           4. Autecology of the Copperhead. By Henry S. Fitch. Pp.
              85-288, pls. 13-20, 26 figures in text. November 30, 1960.

           5. Occurrence of the Garter Snake, Thamnophis sirtalis, in
              the Great Plains and Rocky Mountains. By Henry S. Fitch
              and T. Paul Maslin. Pp. 289-308, 4 figures in text.
              February 10, 1961.

           6. Fishes of the Wakarusa River in Kansas. By James E. Deacon
              and Artie L. Metcalf. Pp. 309-322, 1 figure in text.
              February 10, 1961.

           7. Geographic Variation in the North American Cyprinid Fish,
              Hybopsis gracilis. By Leonard J. Olund and Frank B. Cross.
              Pp. 323-348, pls. 21-24, 2 figures in text. February 10,
              1961.

           8. Descriptions of Two Species of Frogs, Genus Ptychohyla;
              Studies of American Hylid Frogs, V. By William E.
              Duellman. Pp. 349-357, pl. 25, 2 figures in text.
              April 27, 1961.

           9. Fish populations, following a drought in the Neosho and
              Marais des Cygnes rivers of Kansas. By James Everett
              Deacon. Pp. 359-427, pls. 26-30, 3 figs. August 11, 1961.

          More numbers will appear in volume 13.

  Vol. 14. 1. Neotropical Bats from Western México. By Sydney Anderson.
              Pp. 1-8. October 24, 1960.

           2. Geographic Variation in the Harvest Mouse. Reithrodontomys
              megalotis, on the Central Great Plains and in Adjacent
              Regions. By J. Knox Jones, Jr., and B. Mursaloglu.
              Pp. 9-27, 1 figure in text. July 24, 1961.

           3. Mammals of Mesa Verde National Park, Colorado. By Sydney
              Anderson. Pp. 29-67, pls. 1 and 2, 3 figures in text.
              July 24, 1961.

          More numbers will appear in volume 14.





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