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Full text of "University of Kansas publications, Museum of Natural History"

V 



HARVARD UNIVERSITY 




LIBRARY 



OF THE 



Museum of Comparative Zoology 



University of Kansas Publications 

MUSEUM OF natural HISTORY 



VOLUiME 17 • 1965-1968 



EDITORS 

E. Raymond Hall, Chairman 

Frank B. Cross 

Henry S. Fitch 

J. Kxox Jones, Jr. 



I IIMl/;ilV 



Museum of Natural History 

UNI\^RSITY OF KANSAS 

LAM'REXCE 

1972 



Museum of Natural History 

university of kansas 

lawrence 



/' (),inoy/,('(i;i,' 



PRINTED BY 

UNIVERSITY OF KANSAS 

PRINTING SERVICE 

LAWRENCE 



CONTENTS OF VOLUME 17 

1. Localities of fossil veitebiatos obtained from the Niobrara For- 
mation (Cretaceous) of Kansas. By David Bardack. Pp. 1-14. 
January 22, 1965. 

2. Chorda tympani branch of the facial nerve in the middle ear of 
tetrapods. B\- Richard C. Fox. Pp. 15-21. June 22, 1965. 

3. Fishes of the Kansas River System in relation to zoogeography 
of the Great Plains. By Artie L. Metcalf. Pp. 23-189, 4 figs. 
xMarch 24, 1966. 

4. Factors affecting growth and production of channel catfish, 
Ictalurus punctatus. By Bill A. Simco and Frank B. Cross. 
Pp. 191-256, 13 figs. June 6, 1966. 

5. A new species of fringe-limbed tree frog, genus Hyla, from 
Darien, Panama. By William E. Duellman. Pp. 257-262, 1 fig. 
June 17, 1966. 

6. Taxonomic notes on some Mexican and Central American hylid 
frogs. B\ \\'illiam E. Duellman. Pp. 263-279. June 17, 1966. 

7. Neotropical Inlid frogs, genus Smilisca. By ^^'illiam E. Duell- 
man and Linda Trueb. Pp. 281-375, piates 1-12, 17 figs. 
July 14, 1966. 

S. Birds from North Borneo. By Max C. Thompson. Pp. 377- 

433, 1 fig. October 27, 1966. 
9. Natural history of cottonmouth moccasin, Agkistrodon pisci- 

vorus (Reptilia). By Ray D. Burkett. Pp. 435-491, 7 figs. 

October 27, 1966. 

10. Systematic status of a South American frog, Allophryne ruth- 
venii Gaige. By John D. Lynch and Howard L. Freeman. 
Pp. 493-502, 3 figs." October 27, 1966. 

11. Genera of Leptodactylid frogs in Mexico. By John D. Lynch. 
Pp. 503-515, 5 figs. March 20^968. 

12. Middle American frogs of the Hyla microcephala group. By 
William E. Duellman and M. J. Fouquette, Jr. Pp. 517-557, 
pis. 13-16, 9 figs. March 20. 1968^ 

13. Description of new Hylid frogs from Mexico and Central 
America. By William E. Duellman. Pp. 559-578, pis. 17-19. 
April 5, 1968. 

14. Summer birds from the Yucatan Peninsula, Mexico. By Erwin 
Klaas. Pp. 579-611. 1 fig. April 24, 1968. 

,15. Morphological variation in a population of the snake, Tantilla 
gracilis Baird and Girard. By Laurence M. Hardy and Charles 
J. Cole. Pp. 613-629, 6 figs. May 14, 1968. 

Index. Pp. 631-659. 



;- !^f\- Aj_......£| 



University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 1, pp. 1-14 
— January 22, 1965 MUS. CO^^.. 



LIBRARY 

DEC 3 i 15c5 

HARVARD 

Localities of Fossil Vertebrates^NivERsiTY^ 
Obtained from the Niobrara Formation 
(Cretaceous) of Kansas 



BY 



DAVID BARDACK 



University of Kansas 

Lawrence 

1965 



University of Kansas Publications, Museum of Natural History 

Editors: E. Raymond Hall, Chairman, Henry S. Fitch, 
Frank B. Cross 



Volume 17, No. 1, pp. 1-14 
Published January 22, 1965 



University of Kansas 
Lawrence, Kansas 



PRINTED BY 

H/SRRY (BUD) TIMBERLAKE. STATE PRINTER 

TOPEKA. KANSAS 

1965 



30-3182 



Localities of Fossil Vertebrates 
Obtained from the Niobrara Formation l,br^r>| 

( Cretaceous ) of Kansas ^ ^ 

I DEC 3 1 ^- 

j DAVID BARDACK HARVARC 

~Rsn 

Fossil vertebrates have been obtained over the years from tBe 
Niobrara Formation in western Kansas since 1870. Most of the 
I specimens were obtained between 1875 and 1910. A list of the 
collectors includes such well-known persons as Bamum Brown, 

E. C. Case, E. D. Cope, O. C. Marsh, Handel T. Martin, Benjamin 

F. Mudge, Elmer S. Riggs, Charles H. Sternberg, George F. Stem- 
berg, and Samuel W. Williston. Vetebrates from the Cretaceous 
of Kansas are in museums throughout Europe and North America. 
Even the smaller museums display some specimens of Kansas fishes 
and reptiles. Extensive collections are to be found in the American 
Museum of Natural History, the Peabody Museum of Natural His- 
tory at Yale University, the Museum of Natural History at the 
University of Kansas, the United States National Museum, the 
Museum of Comparative Zoology at Harvard University, and the 
Fort Hays Kansas State College Museum. 

j j Abbreviations used in the text beyond for these museums are, 
^ ' respectively, as follows: AMNH, YPM, KU, USNM, MCZ, and FH. 

The Niobrara Formation is exposed in a band from west-central 
to north-central Kansas. The more extensive and paleontologically 
richer deposits are exposed in Logan, Gove and Trego counties. 
Graham and Rooks counties to the north and northeast, respectively, 
of Trego County also have produced fossil vertebrates. Few have 
been obtained from the scattered, smaller exposures of the Niobrara 
Formation in the nortli-central counties ( PhilHps, Smith and Jewell ) . 

The Niobrara Formation includes two members, the lower Fort 
Hays hmestone and the upper Smoky Hill Chalk. The term Kansas 
chalk is sometimes applied to the Niobrara Formation and some- 
times is hmited to the Smoky Hill Chalk member. Most vetebrate 
fossils are known from the Smoky Hill Chalk. 

Catalogue records for the majority of Kansas Cretaceous verte- 
brates, including nearly all of the type specimens, yield little or no 
data as to locality other than "western Kansas," "Kansas chalk," 
"Niobrara Formation," or a particular county. Nevertheless, cata- 

(3) 



4 University of Kansas Publs., Mus. Nat. Hist. 

logue records for many specimens, especially those in the collections 
of the University of Kansas Museum of Natural History, indicate 
by the name of a ranch or creek or number of miles from a town, 
the site at which a particular fossil was found. More precise lo- 
cality information would provide valuable data for studies of a 
stratigraphic or ecological nature. Differences in fossil assemblages 
from eastern and western exposures might be discovered. Evolu- 
tionary changes in certain groups during Niobrara time might be 
traced. 

In the course of studying anatomy and systematics of fossil fishes 
from the Niobrara Formation I have several times sought the loca- 
tion of such places as "Martin's Canyon," "Robber's Roost" or a site 
"3 miles North of Monument Rocks." In an effort to provide more 
adequate locality information I have consulted residents and county 
engineers of those counties in which chalk beds are exposed, talked 
with persons who have collected fossils from the chalk, examined 
U. S. Dept. of Agriculture aerial photographs and geologic maps of 
pertinent counties, studied old maps and correspondence of col- 
lectors and personally examined numerous chalk exposures. I have 
been able to determine range, township and in many cases section 
of the poorly defined localities in which early collections were made. 
It has been my primary interest to ascertain locaHties that have 
yielded specimens of fish. Because fossil reptiles and birds in the 
Niobrara Formation have generally come from the same sites as the 
fishes, localities of many reptilian and avian specimens may be 
determined from the information given below. 

Locahty records of fossils derived from pubhcations or museum 
catalogues are listed under the name of the museum in which the 
fossil material is housed. Each locality is numbered for reference 
purposes within this paper. The numerical listing by county for 
material at the University of Kansas corresponds to the numerical 
listing in the locality catalogue of the Division of Vertebrate Paleon- 
tology. The original description of a locahty is cited first in bold- 
face type followed by a statement of section(s), township, and 
range as determined in this study. Where exposures from which 
the fossils were obtained extend across more than one section, t\vo 
or more sections are cited. Comments on the geologic horizon 
and geographic position of a locality as originally cited are given 
if necessary. Estimates of mileage by the older collectors were in 
many, but not all, instances less tlian the true mileage. 

For assistance in determination of localities I am indebted to 
George F. Sternberg of Hays, Kansas, Albert O'Toole of Ransom, 



Localities of Fossil Vertebrates ( Cretaceous ) Kansas 5 

Kansas, Myrl V. Walker of Fort Hays Kansas State College Mu- 
seum, and Marion Bonner of Leoti. Examination of collections was 
made possible by National Science Foundation Grant No. GB 582. 

UiNivERSiTY OF Kansas Museum of Natural History 

Most of the Kansas Cretaceous vertebrates now in the Museum 
of Natiu-al History were obtained between the years 1895 and 1910 
through the efforts of S. W. Williston, H. T. Martin, C. E. McClung 
and several students including Barnum Brown, E. C. Case, Alban 
Stewart, Wyman Green, Roy Moodie and E. H. Taylor. The speci- 
mens at this museum represent results of the last large-scale effort 
at collecting Niobrara material for one museum. Locahty records 
for Niobrara specimens at the University of Kansas are more nearly 
complete than such records are for specimens at the American 
Museum of Natural History or the Peabody Museum of Natural 
History. Numbers assigned to localities under each county name 
correspond to numbers in the locality catalogue of the Division of 
Vertebrate Paleontology at the University of Kansas. 

A. Gove County 

1. Near Banner (Bana). See Trego Co., Locality No. 2. 

2. 6 mi. SE Gove City. Sections 23, 24, T. 13 S, R. 28 W. 

3. Sec. 5, T. 13 S, R. 35 W. Locality uncertain. R. 35 W would be in 
Logan County but chalk beds are not exposed in the section cited. R. 
25 W in Trego County also would be outside of the chalk beds. 

4. Plum Creek. This stream runs SE into the Smoky Hill River. Exposures 
near the junction of Plum Creek and the Smoky Hill River in T. 15 S, 
R. 28 W have been worked by several collectors according to Myrl V. 
Walker. 

5. Martin's Canyon. Sections 14, 15, 22, 23, 26, 27, 35, T. 14 S, R. 26 W. 
According to Albert O'Toole and George Sternberg this canyon (or pair 
of north-south canyons) carries the first tributary of the Smoky Hill River 
lying entirely within Gove County. The fossils probably came from the 
eastern ravine; it crosses sections 14, 23 and 26. 

6. Spencer's Canyon. Sections 17, 20, 29, T. 14 S, R. 25 W. This canyon 
is in Trego County according to Albert O'Toole. A broad, apparently 
unnamed canyon (eastern vertical tier of sections in T. 14 S, R. 26 W) 
is crossed by the Gove-Trego county line. This canyon might have been 
confused with Martin's Canyon or Spencer's Canyon by some collectors. 

7. 4 mi. SE Elkader. Sec. 13 and/or 24, T. 15 S, R. 32 W, in Logan County. 

8. 4 mi. SE Gove City. Sections 16, 17, T. 13 S, R. 28 W. 

9. 6 mi. SW Gove City. Sections 28, 29, T. 13 S, R. 29 W. 

10. 6 mi. W Castle Rock. Sec. 35, T. 13 S, R. 27 W. 

11. Spicer's Ranch, SW Castle Rock. Spicer's Ranch was in sec. 10, T. 13 S, 
R. 26 W. This is NW of Castle Rock. Several fossil vertebrates have 
been obtained from sec. 16, T. 13 S, R. 26 W on the Andrew Bird ranch. 

12. S side Smok-y Hill River. Locality cannot be determined. 

13. E Spencer's Canyon. This canyon is in Trego Co. See Gove Co., Locality 
No. 6. 



6 University of Kansas Publs., Mus. Nat. Hist. 

14. S side Smoky Hill River and Martin's Canyon. Numerous exposures in 
northeastern sections of T. 15 S, R. 26 W. 

15. Di mi. W Castle Rock. Sec. 3, T. 14 S, R. 26 W. 

16. 4 mi. N and E of Castle Rock, Perrington Ranch. Sec. 8 or 9, T. 14 S, 
R. 25 W. This locality is in Trego Co. on the Purinton Ranch. 

17. Monument Rocks. Sec. 34, T. 14 S, R. 31 W. 

18. 2 mi. E Monument Rocks. Sections 25, 26 or 34, 35, T. 14 S, R. 31 W. 

19. 6 mi. E Gove City on Hackberry Creek. See Gove Co., Locality No. 2. 
Hackberry Creek flows SE from Gove City. 

20. N of Monument Rocks. Locality cannot be determined. 

21. Hell Creek. Southeastern sections of T. 15 S, R. 31 W. Hell Creek is a 
tributary of the Smoky Hill River. Another tributary with broad chalk 
exposures lies just east of Hell Creek. These paleontologically rich 
exposures (sections 18, 19, 30, 31, T. 15 S, R. 30 W) are called Hell's 
Bar by George Sternberg. Specimens labelled Hell Creek may have 
come from HeU's Bar. 

22. 2 mi. SW Castle Rock. W)i sec. 13 and EM sec. 14, T. 14 S, R. 26 W. 

23. 35^ mi. SW Castle Rock. Sections 22, 23, T. 14 S, R. 26 W. 

24. 4 mi. W Castle Rock. Sec. 5, T. 14 S, R. 26 W and sec. 32, T. 13 S, 
R. 26 W. 

25. 3 mi. N Monument Rocks. Sections 15, 22, T. 14 S, R. 31 W. 

B. Graham County 

1. M mi. NW Hill City. Sec. 10, T. 8 S, R. 23 W. 

2. 1 mi. W Hill City. See Graliam Co., Locahty No. 1. 

3. IM mi. NW Hill City. NWll sec. 10, T. 8 S, R. 23 W. 

4. NW Hill Cit>'. Locality cannot be detennined. 

5. 6 mi. S Hill City. Sec. 14, T. 9 S, R. 23 W. Small exposure on bank of 
stream. 

6. 3 mL NW HiU City, near Solomon River. ?sec. 17, T. 8 S, R. 23 W. This 
exposvue is within a half mile of the river but W rather than NW of 
Hill City. 

7. Hill City. Locality cannot be determined. 

8. ( Pleistocene locality. ) 

9. « mi E Penokee. WJi sec. 13 and E'A sec. 14, T. 8 S, R. 24 W. This lo- 
cality is about 1 mi. E of Penokee. 

C. Logan County (formerly St. John County) 

1. Butte Creek. Locality of specimens cannot be determined. 

2. Near Twin Buttes. Twin Buttes is in NEJi sec. 1, T. 15 S, R. 37 W. 

3. Lisbon. Sec. 8, T. 12 S, R. 36 W. This town existed in the early days of 
tlie Kansas Pacific (now Union Pacific) Railroad. Sheridan, Kansas, was 
at approximately the same place. Rocks exposed in the vicinity of Lisbon 
are lower members of tlie Pierre Shale. 

4. NE Russell Springs. T. 13 S, R. 35 W. Small areas of chalk are ex- 
posed less than two miles northeast of the town. 

5. Near Russell Springs. T. 13 S, R. 35 W. Several exposures on the S 
side of the Smoky Hill River including one )i to ¥i mile in linear extent 
known locally as Hell's Half Acre {N)i sec. 27, T. 13 S, R. 35 W) occur 
near the town. 

6. 4/2 mi. NW Elkader, Logan Co. line. Elkader has moved several times, 
but the town has been at its present location (sec. 34, T. 14 S, R. 32 W) 
or within a mile of this site during the period in wliich most of the speci- 
mens at the University of Kansas were collected. The Gove-Logan 



Localities of Fossil Vertebrates ( Cretaceous ) Kansas 7 

county line lies NE of Elkader. The position of Locality No. 6 is there- 
fore uncertain. 

7. 7 mi. NW Elkader. Numerous small chalk exposures he NW of Elkader. 

8. S of Elkader. Locahty cannot be determined. 

9. SE of Elkader. Locahty cannot be determined. 

10. W of Russell Springs. Numerous exposures in T. 13 S, R. 35 W in 
sections lying W and SW of the town. See Logan Co., Locahty No. 5. 

11. 1 mi. N Elkader. ?sec. 22 or 27, T. 14 S, R. 32 W. 

12. Yi mi. E Beaver Creek, Elkader. T. 15 S, R. 32 W. Beaver Creek (now 
generally known as Ladder Creek) enters tlie Smoky Hill River in sec. 34, 
T. 14 S, R. 35 W. Several extensive exposures of Smoky Hill Chalk are 
in sec. 12, T. 15 S, R. 32 W. These are 1 to 2 mi. S of Elkader and 'A 
mi E of Beaver Creek. 

13. 2M mi. NE Elkader. Sec. 25, T. 14 S, R. 32 W. 

14. Elkader. Locality of fossils cannot be determined. 

15. 5 mi. SW Elkader, Beaver Creek. Sections 8, 17, T. 15 S, R. 32 W. 

16. (Phocene locahty.) 

17. (Pleistocene locahty.) 

18. 12 mi. W Russell Springs. Sec. 24, T. 13 S, R. 37 W. 

19. Near Elkader. Locality cannot be detennined. 

20. 10 mi. SW Russell Springs, Willow Canyon. Sections 11, 12, 13, T. 15 
S, R. 35 W. This locahty is S rather tlian SW of Russell Springs and 
appears to be identical with the locahty termed Cedar Canyon by several 
present-day collectors according to Marion Bonner. 

21. Beaver Creek. Locahty of fossils cannot be determined. 

22. 2 mi. NW Russell Springs. Sec. 14, T. 13 S, R. 35 W. Small exposures 
of chalk are along a stream flowing dirough Russell Springs and into the 
Smoky Hill River. 

23. Hell Creek. This stream is in Gove Co. See Gove Co., Locahty No. 21. 

24. 2J-S mi. N Elkader. ?sec. 15, T. 14 S, R. 32 W. 

25. 2]i mi. SE Elkader. ?sec. 11, T. 15 S, R. 32 W. 

26. Russell Springs. Locahty of fossils cannot be determined. 

27. 2 mi. W Russell Springs. Sec. 27, T. 13 S, R. 35 W. See Logan Co., 
Locahty No. 5. 

28. Smoky Hill River. Locality of fossils cannot be detennined. 

29. NWJ4 sec. 20, T. 11 S, R. 14 W, Benton Cretaceous. R. 14 W would be 
in Russell Coimty. 

30. McAllister. Sec. 24, T. 12 S, R. 37 W. Exposures in the vicinity of 
McAllister are in Pierre Shale. 

31. N point of bluff, E side of Beaver Creek. T. 15 S, R. 32 W. See Logan 
Co., Locahty No. 12. 

32. Near Lisbon. See Logan Co., Locality No. 3. 

D. Trego County 

1. SE Castle Rock. ?Near Wildcat Canyon. See Trego Co., Locahty No. 7. 

2. Banner. Sec. 29, T. 13 S, R. 25 W. Banner was a Trego Co. Post OflBce. 
The Banner Post Office at one time was in sec. 26 and at another time 
in sec. 32. Section 29 was the more likely reference point for other lo- 
cahties in Trego Co. during the period of field work by the University 
of Kansas. 

3. 2 mi. SE Banner. WJi sec. 3 and EM sec. 4, T. 14 S, R. 25 W. 

4. /2 mi. SE Banner. Chalk exposures are more than %. mi. from Banner. A 
small exposure (sec. 32, T. 13 S, R. 25 W and sec. 5, T. 14 S, R. 25 W) 
hes SW of Banner. 



8 University of Kansas Publs., Mus. Nat. Hist. 

5. SW Gove City. Gove City is in Gove Co. See Gove Co., Locality No. 9. 

6. Hackberry Creek. Locality of fossils cannot be determined. Several 
major exposures of chalk lie close to this creek. 

7. Cat Hills. SJ-S sec. 9 and WM sec. 15 and sec. 16, T. 14 S, R. 25 W. 
This locahty is better known as Wildcat Canyon. 

8. Martin's Canyon. See Gove Co., Locality No. 5. 

9. VA mi. S Banner, Cat Hills. See Trego Co., Locality No. 7. 

10. Bluffs W side Sec. 3. Sec. 3, T. 14 S, R. 25 W. See Trego Co., Locality 
No. 3. 

11. E of O'TooIe School. Sections 3 and 10 (along Gibson Creek), T. 15 S, 
R. 25 W. This school was in sec. 5, T. 15 S, R. 25 W. 

12. M mi. S O'TooIe's. Sec. 3, T. 15 S, R. 25 W. The O'Toole Ranch is 
in sec. 34, T. 14 S, R. 25 W. 

13. % mi. N Neuenschwander's Ranch, Banner. SWJi sec. 10, T. 14 S, R. 
25 W. The Neuenschwander Ranch is in sec. 15, T. 14 S, R 25 W. 

14. Near Banner, S of Neuenschwander's. W)2 sec. 15, T. 14 S, R. 25 W. 

15. Near Banner, M mi. SW Neuenschwander's. Sec. 16, T. 14 S, R. 25 W. 

16. VA mi. SE Neuenschwander's, Banner. NW}4 sec. 23, NEJ4 sec. 22, SEM 
sec. 15, SWA sec. 14, T. 14 S, R. 25 W. 

17. }i mi. S Neuenschwander's. WA sec. 15, T. 14 S, R. 25 W. 

18. 2/2 mi. E Banner. No chalk is exposed E of Banner. Locality is probably 
SE of Banner. See Trego Co., Locality No. 19. 

19. 2Ji mi. SE Banner. E'A sec. 4, WM sec. 3, T. 14 S, R. 25 W. Cf. Trego 
Co., Locahty No. 3. 

20. %. mi. N Neuenschwander's near Banner. EM sec. 10, T. 14 S, R. 25 W. 

21. 4 mi. S Banner, Wildcat HUls. Sec. 16, T. 14 S, R. 25 W. See Trego 
Co., Locahty No. 7. 

22. 2 mi. NW Wakeeney on Saline River. Chalk exposures are at least 4 mi. 
NW of Wakeeney. The Saline River is more than 5 mi. from Wakeeney. 
The locality cannot be determined. 

23. 6 mi. NW Wakeeney. Sections 16, 21, T. 11 S, R. 24 W. Other ex- 
posures are developed in sections 10, 15 and 22. This land is part of 
the E. J. Garrett Ranch. 

24. 3 mi. E Banner at Neuenschwander's. Neuenschwander's is not directly 
E of Banner. See Trego Co., Locality No. 13. The locality cannot be 
determined. 

25. 1 mi. N Smoky Hill River. The locality is probably in Spencer's Canyon. 
See Gove Co., Locality No. 6. 

26. 155^ mi. SW Banner. This site must be in Gove County. The locality 
cannot be determined. 

E. Wallace County 

1. Beaver Creek. See Logan Co., Locality No. 12. 

2. McAllister. See Logan Co., Locality No. 30. 

3. Eagle Tail Creek near Sharon Springs. T. 13 S, R. 40 W. Exposures 
of Pierre Shale. 

4. Near McAUister. See Logan Co., Locahty No. 30. 

Fort Hays Kansas State College Museum 

Locality records of fishes and reptiles (listed separately) in the 
State College Museum are cited. Most of these specimens were 
collected by George F. Sternberg, Myrl V. Walker and Marion 
Bonner. Locality numbers do not refer to the State College Mu- 



Localities of Fossil Vertebrates ( Cretaceous ) Kansas 9 

seum catalogue and such numbers are assigned here arbitrarily. 
Localities that are the same as those in the University of Kansas 
catalogue are so designated. 

Fishes 

1. Bluffs S of Castle Rock. Sec. 1, T. 14 S, R. 26 W, Gove Co. FH No. 25, 
type of Kansius sternbergi Hussakof. 

2. 11 mi. S Quinter. ?sec. 16, T. 13 S, R. 26 W, Gove Co. FH No. 1649, 
Cimolichthys sp. 

3. 2 to 3 mi. SE Glade. Sec. 12 or 13, T. 5 S, R. 18 W, Phillips Co. FH 
No. 1650, Saurodon sp. 

4. Ransom. Specimen probably from Trego Co. Pliocene exposiures occur 
in vicinity of Ransom. FH No. 1652, Ptychodus sp. 

5. Purinton Ranch. See Gove Co., Locality No. 16. FH No. 1659, Protos- 
phyraena sp. 

6. 7 mi. NW Ellis. T. 12 S, R. 21 W, Trego Co. FH No. 1660, Protos- 
phyraena sp. 

7. IM mi. SE Elkader. Sec. 11 or 12, T. 15 S, R. 32 W, Logan Co. FH 
No. 2024, Protosphyraena sp. 

8. N of OgaUah, Saline River. EVi sec. 15, WM sec. 14, T. 11 S, R. 22 W, 
Trego Co. FH No. 2025, Protosphyraena sp. 

9. 5 mi. SW Gove City. Central sections, T. 13 S, R. 29 W, Gove Co. 
FH No. 2081, Isurus sp. 

10. B}i mi. SE Elkader. Sec. 13 or 24, T. 15 S, R. 32 W, Logan Co. FH 
No. 2714, Isurus sp. 

11. 3 mi. N of Bogue, N side of S Fork Solomon River. Sec. 6, T. 8 S, 
R. 21 W, Graham Co. FH No. 5014, Cimolichthys sp. 

12. NW Bogue. See FH Locality No. 11. FH No. 8395, Xiphactinus sp. 

13. S Brakes, Butte Creek, Logan Co. Sec. 15 or 16 (according to Marion 
Bonner), T. 15 S, R. 34 W. FH No. 8567, Ichthyodectes sp. 

14. Marvin Albin Ranch about halfway between Castle Rock and Smoky Hill 
River. ?sec. 24, T. 14 S, R. 26 W, Gove Co. FH No. 10688-288, 
Ptychodus sp. 

Reptiles 

15. 6 to 7 mi. SE Russell Springs. T. 14 S, R. 34 W, Logan Co. FH No. 4, 
Tylosaurus sp. 

16. 5 mi. NW Elkader. T. 14 S, R. 32 W, Logan Co. FH No. 2085, 
Pteranodon sp. 

17. 2 mi. NE Penokee. Sec. 13, T. 8 S, R. 24 W, Graham Co. FH No. 2851, 
Pteranodon sp. 

18. Mendenhall Ranch, 4 mi. E Gove City. Sections 16, 17, T. 13 S, R. 
28 W, Gove Co. FH No. 2913, Platecarpus sp. 

19. Between Bogue and Hill City, 200 steps from the N side of the S Fork, 
Solomon River. Sec. 12, T. 8 S, R. 22 W, Graham Co. FH No. 5426, 
Pteranodon sp. 

20. 3 mi. NE Gove (City). Sec. 32, T. 12 S, R. 28 W, Gove Co. FH No. 
7109, Tylosaurus sp. 

21. Garrett Ranch. See KU Trego Co., Locality No. 23. FH No. 7937, 
Clidastes sp. 

22. N RusseU Springs. See KU Logan Co., Locality No. 5. FH No. 7960, 
plesiosaur. 

23. About 1 mi. SW Russell Springs. See KU Logan Co., Locality No. 5. 
FH No. 8400, Toxochelys sp. 



10 University of Kansas Publs., Mus. Nat. Hist. 

24. Horse Thief Canyon. W)^ sec. 9, T. 13 S, R. 26 W, Gove Co. FH No. 
10560, Pteranodon sp. 

25. 9 mi. NW Wakeenev, Garrett Ranch. See KU Trego Co., Locality No. 
23. FH No. 10598, Pteranodon sp. 

26. 7 mi. W Gove City, Emerson Ranch. ?sec. 36, T. 12 S, R. 30 W, Gove 
Co. FH No. 10856, mosasaur. 

27. Brakes of Hell Creek. See KU Logan Co., Locality No. 23. FH No. 
11311, Nyctosaurus sp. 

American Museum of Natural History 

Kansas Cretaceous vertebrates in the American Museum include 
the majority of type specimens from the chalk. Most of the ma- 
terial was gathered by E. D. Cope and his collectors in the 1870s. 
Specimens having locality records traceable to section, township 
and range are few. Localities from which specimens in the American 
Museum were obtained are cited in Cope ( 1875 ) . Several localities 
described by Cope are in areas lacking exposures of chalk. Other 
localities described as in yellow chalk are actually in gray or black 
Pierre Shale (Adams, 1898). Several sites are close to stations of 
the Butterfield Overland Stage. Cope probably followed parts of 
the stage route, which closely approaches chaUc exposures. He 
visited western Kansas in 1871. Starting from Fort Wallace (sec. 
29, T. 13 S, R. 38 W ) he went S and E exploring exposures in Logan 
County along the Smoky Hill River and Butte Creek. The locahties 
from which he described specimens of reptiles and fishes are listed 
with page references to Cope ( 1875 ) . 

1. S of Ft. Wallace on Butte Creek in the yellow^ chalk (p. 67). Although 
the upper drainage of Butte Creek lies S of Ft. Wallace, exposures of 
yellow chalk do not appear until one enters Logan Co. in R. 37 W. 
The locality is uncertain. 

2. Smoky Hill River 5 mi. W Ft. Wallace in yellow limestone (p. 70). 
Exposures W of Ft. Wallace are Pierre Shale. Small exposures of yellow 
chalk appear about 2 mi. SW of the Fort on the S side of the Smoky 
Hill River (sec. 32, T. 13 S, R. 38 W and sec. 5, T. 14 S, R. 38 W). 

3. Near mouth of N. branch of Smoky Hill River (p. 74). Material from 
small exposures of the Niobrara Formation and/or Pierre Shale in the 
vicinity of Forks of the Smoky Hill River Stage Station, NEJz T. 13 S, 
R. 36 W. 

4. Fossil Spring Canyon (p. 133, 144, 223). Cope obtained several specimens 
from this locality. Unfortunately I have been unable to identify the site 
definitely. Cope refers (Osbom, 1931:163) to a spring 4 mi. beyond 
Twin Buttes. At this point (sec. 4, T. 15 S, R. 36 W) a small tributary 
of Butte Greek is 2 mi. or less from the south bluffs of Butte Creek where 
extensive chaUc exposures are developed. 

5. Bluff of Butte Creek (p. 135). Probably in the vicinity of Locahty No. 4. 

6. 6 mi. S Sheridan (p. 138). Sheridan (W% sec. 8, T. 12 S, R. 36 W) was 
a terminus of the Kansas Pacific (now Union Pacific) Railroad in 1870-71. 
Six miles S of Sheridan would place the locahty along the N Fork of the 
Smoky Hill River where numerous exposures of Pierre Shale and one or 
two exposures of Smoky Hill ChaUc are developed. Cope cites (p. 157) 
a specimen from the yellow chalk 6 mi. S of Sheridan. 



Localities of Fossil Vertebrates ( Cretaceous ) Kansas 11 

7. Fossil Spring Canyon near mouth of Fox Canyon (p. 144). Cope (p. 18) 
described these canyons as follows: "Butte Creek valley, fifteen to 
eighteen miles to the south [presumably S of the Smoky Hill River], is 
margined by bluflFs of from twenty to one hundred and fifty feet in 
height. . . . The bluffs of the upper portion of Butte Creek, Fox 
and Fossil Spring ( five miles south [sic] ) Canons are of yellow chalk; and 
the reports of several persons stated that those of Beaver Creek, eight 
miles south of Fossil Spring, are exclusively of this material." This in- 
formation is inadequate to locate these canyons. Butte Creek lies about 
10 miles south of the Smoky Hill River. The locality of Fossil Spring 
is uncertain (see Locality No. 4). If Cope meant that Fossil Spring 
was five miles south of the Smoky Hill River, Fossil Spring would be in 
the flat valley of this river rather than in a canyon. Beaver Creek (now 
called Ladder Creek) is at least eight miles and most parts are 12 to 15 
miles south of Butte Creek. Rocks of the Niobrara Formation are ex- 
posed only near the mouth of Ladder Creek. 

8. N bank. Smoky Hill River, 30 mi. E Ft. Wallace (p. 149). This locality 
must lie in tlae south half of T. 13 S, R. 33 W along Twelve Mile Creek 
and in the vicinity of the Smoky Hill Stage Station ( SJz, EJz sec. 32, T. 13 
S, R. 33 W). 

9. Fox Canyon S of Ft. Wallace (p. 149). See AMNH locaHty Nos. 1 and 7. 

10. Gypsum Buttes (p. 150). Cope stated that tliese buttes lie 1 mi. SW of 
Sheridan (see AMNH Locality No. 6) in sec. 13, T. 12 S, R. 37 W. 
Cope also refers to Twin Buttes near Butte Creek. Twin Buttes lie in 
sec. 1, T. 15 S, R. 37 W. There are several chalk exposures near Twin 
Buttes. 

11. 1 mi. SE Sheridan on the North Fork of the Smoky Hill River (p. 157). 
Sec. 8 or 17, T. 12 S, R. 31 W. 

12. Low blufi^ on Butte Creek, 14 mi. S of Ft. Wallace (p. 160). Cope must 
have meant SE of the Fort. Exposures 14 mi. S of Ft. Wallace are 
beyond Butte Creek and in Tertiary rock. The locality cannot be de- 
termined. 

13. Fox Canyon near Ft. Wallace (p. 194). See AMNH Locality No. 7. 

14. Solomon River 160 miles from junction with Kansas River (p. 201). Near 
Hill City, Graham Co. 

15. Russel Spring on the Smoky Hill River (p. 209). Near the Russel Spring 
Stage Station. The fossils probably came from the SM T. 13 S, R. 35 W. 

16. 15 mi. S Ft. Wallace (p. 217). See AMNH Locality Nos. 1 and 12. 

17. Fossa Spring near Ft. Wallace (p. 223 and 225). See AMNH Locality 
No. 7. 

18. Near Stockton (p. 244). Stockton is in Rooks Coimty. Numerous small 
exposures of chalk are developed near Stockton. 

19. Spring Creek, Rooks County (p. 244d). Spring Creek flows north into 
the South Fork of the Solomon River in T. 8 and 9 S, R. 20 W. 

The following are localities of specimens of fish in the American 
Museum from tlie Kansas Cretaceous collected prior to 1940. 

20. Sand Creek, 2 to 3 mi. and 5 mi. above S Fork Solomon River. Sand 
Creek (T. 10 S, R. 19 and 20 W) Hes in Rooks County and flows into 
the Saline River. AMNH Nos. 1705, 1706, and 2062. 

21. 7 mi. SW Castle Rock, Trego Co. See KU Gove Co., LocaHty No. 5. 
AMNH No. 1747. 

22. Upper Fox Creek. PEquals Fox Canyon. See AMNH LocaHty No. 7. 
AMNH No. 1915. 

23. T. 13 S, R. 24 W, Trego Co. Exposures of the Niobrara Formation do 
not appear in tliis range and township. Possibly this locality should be 
T. 13 or 14 S, R. 25 W. AMNH No. 2003. 



12 University of Kansas Publs., Mus. Nat. Hist. 

24. 2 mi. E Sugarloaf, T. 6 S, R, 19 W, Rooks Co. This site is in sec. 12. 

AMNH No. 2307. 

25. Uncatalogued specimens of "berycoids" are recorded from: 

a) J^ mi. S Castle Rock. Sec. 1, T. 14 S, R. 26 W. 

b) 3 mi. SW Castle Rock. See KU Gove Co., Locality No. 5. 

c) 2.5 mi. E Castle Rock. See KU Trego Co., Locality No. 7. 

Peabody Museum of Natural History 

Extensive collections of reptilian and fish material in the Peabody 
Museum bear little or no record of locality. O. C. Marsh obtained 
the majority of Kansas Cretaceous fossils now at the Peabody 
Museum through the efforts of his collectors, primarily B. F. Mudge. 
Marsh's main contribution to our knowledge of Kansas Cretaceous 
fossils appears in his monograph (1883) on the so-called toothed 
birds. Localities cited, in that work, which can be identified to 
section, township and range are indicated below. 

1. S bank Smoky Hill River about 20 mi. E of Ft. Wallace (p. 195). Sec. 27, 

T. 13 S, R. 35 W. Near Russell Springs, Logan Co. 

2. 12 mi. E Ft. WaUace (p. 195, 198). Sec. 11 or 13, T. 13 S, R. 36 W. 
Near Forks of the Smoky Hill River Stage Station. 

3. Hackberry Creek, near Smoky Hill River, Gove Co. (p. 199). Sec. 31, 
T. 13 S, R. 25 W. Specimens were probably from near the Castle Rock 
Creek Stage Station. 

United States National Museum 

With the exception of the State College Museum collections at 
Fort Hays, the U. S. National Museum collections of Kansas Nio- 
brara vertebrates are less numerous than those of other museums 
discussed above. Specimens collected by Dr. David Dunkle during 
the past 15 years bear adequate locality data. Material collected 
earHer by Mr. George Sternberg and others is cited below and 
section, township and range are indicated or reference is made to 
identical localities of the University of Kansas Museum of Natural 
History. 

1. 10 mi. NW Wakeeney. This mileage was probably overestimated. George 
Sternberg collected fossils in this area from tlie Garrett Ranch. See KU 
Trego Co., Locality No. 23. USNM No. 21081. 

2. Hackberry Creek, 3 mi. below Gove City. See KU Gove Co., Locahty 
No. 8. USNM No. 12061. 

3. 12 mi. SE Russell Springs. Sec. 16, T. 15 S, R. 34 W. This locality is 
called "Robber's Roost" by George Sternberg. This site is on land that 
was part of the Berry Farm, a name which appears on labels of some 
specimens. USNM No. 12358. 

4. 25 mi. S of Oakley, S side of the Smoky Hill River. T. 15 S, R. 35 W. 

USNM No. 11554. 

5. 2J^ mi. S and 2 mi. E Oakley. Sec. 13 or 14, T. 15 S, R. 32 W. USNM 
No. 11650. 

6. 14 mi. S Quinter. Sec. 16, T. 13 S, R. 26 W. Andrew Bird Ranch. 
USNM No. 11557. 

7. 5 mi. SE RusseU Springs. Sec. 6, T. 14 S, R. 34 W. USNM No. 18224. 



Localities of Fossil Vertebrates ( Cretaceous ) Kansas 13 

Museum of Comp.\rative Zoology, Harvard University 

Many of tlie specimens at the Museum of Comparative Zoology 
are uncatalogued and locality records are scanty. St. John is a 
frequently-cited locality. Logan County originally bore the name 
St. John County. The specimen cited below is the only one having 
locality record adequate to determine section, township and range. 

1. 4 mi. N of the Pyramids (Monument Rocks). ?sec. 18, T. 14 S, R. 31 W, 
Gove Co. MCZ No. 5400. 

REMARKS 

It is hoped that clarification of locahties from which Kansas 
Niobrara Cretaceous vertebrates have been obtained will assist in 
future studies of these vertebrates. It would be premature to 
commence ecologic or stratigraphic investigations of Niobrara 
vertebrates from locality data alone. Intensive taxonomic study 
of each vertebrate group represented in the chalk is required be- 
cause earlier workers unnecessarily multiplied the number of 
specific names. Modern anatomical and taxonomic studies of 
vertebrate fossils from the Niobrara Formation of Kansas are just 
beginning. Dale Russell is reviewing the mosasaurs; T. H. Eaton 
and D. Bardack are revising several families of fishes. 

That geographic differences in the distribution of Kansas Nio- 
brara Cretaceous vertebrates do exist is demonstrated by two groups 
of fishes. Teeth of the elasmobranch Ptychodus are confined to 
eastern exposures of the Niobrara Formation in Gove and Trego 
counties and have not been found in Logan County where western 
exposures of the Niobrara Formation in Kansas are developed. 
Specimens of Protosphyraena are more abundant in eastern than 
in western chalk beds. Thus, Ptychodus and Protosphyraena are 
found only in the eastern or stratigraphically lower levels of the 
Smoky Hill Chalk or at best are more numerous in these levels. 
Distributional diflFerences among the reptiles may be more pro- 
nounced than among the fishes. This is suggested by Dale Russell 
( personal communication ) . 

Collections of Niobrara Cretaceous vertebrates from Kansas come 
primarily from areas of numerous, broad exposures in the west- 
central part of the State. Northeastern exposures of the chalk have 
scarcely been examined. Efforts to collect specimens in such areas 
might produce nev/ kinds or indicate ecologic or stratigraphic dif- 
ferences relative to the area of Kansas from which Cretaceous 
vertebrates are well known. 



14 University of Kansas Publs., Mus. Nat. Hist. 

BIBLIOGRAPHY 

Adams, G. I. 

1898. The Upper Cretaceous of Kansas: A historical review. Univ. 
Geol. Surv. Kansas. 4:15-27. 

Cope, E. D. 

1875. The vertebrata of tlie Cretaceous formations on the West. Rept. 
U. S. Geol. Surv. Territ. 2:1-303. 

HODSON, W. G. 

1960. Geology and ground-water resources of Gove County, Kansas. 

Univ. Kansas Publ. State Geol. Surv. BuU. 145:1-126. 
1963. Geology and ground-water resources of Wallace County, Kansas. 

Univ. Kansas Publ. State Geol. Surv. Bull. 161:1-108. 
Geology and ground-water resoiu-ces of Trego County, Kansas. 

Univ. Kansas Publ. State Geol. Surv. in press. 

Johnson, C. R. 

1958. Geology and ground-water resources of Logan County, Kansas. 
Univ. Kansas Publ. State Geol. Surv. Bull. 129:1-175. 

OSBORN, H. F. 

1931. Cope: Master NaturaUst. Princeton Univ. Press, xvi + 740 pp. 

PuEScoTT, G. C, Jr. 

1955. Geology and ground-water resources of Graham County, Kansas. 
Univ. Kansas Publ. State Geol. Surv. Bull. 110:1-98. 

Transmitted July 7, 1964. 



D 

30-3182 



M' ' '^l: :zj 



University of Kansas PuBLicATioiSjfy-'^- ^O'^" • - OC 

Kt M u LIBRARY 

Museum of Natural History 

OEC 3 1 1S65 

Volume 17, No. 2, pp. 15-21 

HARVARD 



June 22, 1965 



UNIVERSITY! 



Chorda Tympani Branch of the Facial Nerve 
In the Middle Ear of Tetrapods 



BY 



RICHARD C. FOX 



University of Kansas 

Lawrence 

1965 



University of Kansas Publications, Museum of Natur.\l History 

Editors: E. Raymond Hall, Chairman, Henry S. Fitch, 
Frank B. Cross 



Volume 17, No. 2, pp. 15-21 
Published June 22, 1965 



University of Kansas 
Lawrence, Kansas 



PRINTED BY 

HARRY (BUD) TIMBERLAKE. STATE PRINTER 

TOPEKA. KANSAS 

1 965 



30-7117 



Chorda Tympani Branch of the Facial Jg^^^Af^ 

In the Middle Ear of Tetrapods ^, ^ _^„ 

^ DEC 3 i bco 

BY 
RICHARD C. FOX HARVARD 

In recent years the middle ear of tetrapods has receivea the aFtemion of 
numerous morphologists in the attempt to clarify the liistories of amphibians, 
reptiles and mammals. Some workers have been more concerned with the 
comparatixe anatomy of the middle ear among Recent tetrapods than with its 
morphology in fossil genera. It should be noted, however, that paleontological 
and neontological approaches have seldom been mutually exclusive. Students 
of the middle ear of tetrapods who have approached the subject from a 
phylogenetic standpoint usually have invoked accessory evidence provided by 
Recent amphibians, reptiles and mammals to support conclusions; similarly, 
students relying primarily upon the morphology of the middle ear in Recent 
tetrapods have used some evidence provided by fossils. Neither "school" has 
been entirely successful in revealing the events that occurred in the evolution 
of the middle ear in the major groups of tetrapods. 

Both neontological and paleontological studies have failed to account satis- 
factorily for the position of the chorda tympani branch of the facial or seventh 
cranial nerve in relation to the stapes and its processes. The stapes of reptiles 
and their amphibian ancestors characteristically possesses three processes: 

1. A dorsal process, usually ossified, that extends upward and makes contact 
with the paroccipital process of the opisthotic. 

2. A tympanic process, usually cartilaginous, that extends outward and 
makes contact with the tympanum. 

3. A quadrate process, usually cartilaginous or ligamentous, that makes 
contact with the quadrate. 

In the embolomerous ancestors of reptiles the quadrate process is thought 
to have extended outward and downward from the stapes to the quadrate. 
Consequently, the long axes of the tympanic process and the quadrate process 
were in the same vertical plane. With the advent and evolution of reptiles, 
the stapes descended to a more ventral position in relation to the quadrate. 
The quadrate process, swinging upward approximately 90 degrees, extended 
outward and forward to the quadrate. The long axes of the tympanic process 
and quadrate process then lay in a horizontal plane. This stage in the evolu- 
tion of the middle ear had already been achieved in primitive pelycosaurs and 
captorhinomorphs. 

In Recent reptiles and birds the chorda tympani passes forward across the 
tympanic cavity in a path that is dorsal to the tympanic process of the stapes. 
Thereafter, the chorda tympani curves down below the quadrate process of 
the stapes and extends ventrally and anteriorly to the lower jaw. In Recent 
mammals, however, the position of the nerve relative to that of the stapes is 
indetenninate; the malleus and incus are interposed between the stapes and 
the tympanum. The nerve crosses the tympanum through the posterior mal- 
leolar folds, between the dorsal pars flaccida and the ventral pars tensa of the 
membrane. In frogs the nerve extends anteriorly across the tympanic cavit>' 

(17) 



18 University of Kansas Fuels., Mus. Nat. Hist. 

heneath tlic stapes before descending to the lower Jaw. The manner in which 
these differences are to be explained has resulted in controversy; the most 
recent and important attempts to resolve the seeming incompatibilities of the 
position of the nerve in the middle ear have been by Vaughn ( 1955 ) and 
Hotton (1960). 

It is generally thought that the primitive position of the chorda tympani was 
post-tympanic; in the first amphibians the nerve presumably passed forward 
across the tympanic cavity beneath the tympanic attachment of the stapes. 
The anuran condition has been supposed to be a retention of the primitive 
relationship between the nerve and the stapes ( see, for example, Hotton, 1960 ) , 
although deBeer ( 1937 ) would ascribe the position of the nerve in frogs and 
toads to peculiarities in the development of the chondrocranium in those 
animals. Additional support for the post-tympanic course of the nerve has 
Iieen provided by Hotton (1960), who has suggested that during the change 
of the rhipidistian hyomandibular to the tetrapod stapes the nerve was pushed 
foi-ward by the quadrate process as the hyomandibular swimg out from the 
anteroposterior rhipidistian orientation to the definitive transverse tetrapod 
orientation. To accomplish this, the nerve had to drape across the quadrate 
process of the stapes, and Hotton ( 1960 ) has suggested that it did so indeed. 

The descent of the tympanum from its dorsal labyrinthodont position to its 
ventral position among primitive captorhinomorphs and synapsids has been 
recognized in the attempts of both Vaughn (1955) and Hotton (1960) to 
explain the new position of the chorda tympani still found in Recent diapsids 
and birds. Vaughn suggested that during the amphibian-reptilian transition, 
with descent of the tympanum, the chorda tympani rose from its passage below 
the stapes to a new passage above the stapes. The vehicle of transfer in this 
scheme was a hypothetically thick tympanum tlirough wliich tlie chorda tym- 
pani passed distal to the tennination of the stapes. As Hotton (1960) has 
pointed out, such events are dependent upon the nerve passing across the 
tympanic cavity in a path close enough to the tympanum during its descent 
to permit the membrane to "seize" the nerve. Hotton rightly questions the 
supposition of Vaughn that the nerve was unusually close to the tympanum in 
embolomercs; nor is there any evidence to suggest that the membrane was 
unusually thick in these reptilian ancestors. 

Hotton's (1960) explanation of the change from a post-tympanic to pre- 
t>nipanic path followed by the chorda tympani during the embolomere- 
reptilian transition is more complex than that of Vaughn. Hotton believed, as 
noted above, that in the first amphibians the chorda tympani crossed the 
tympanic cavity by passing under the tympanic process and over the quadrate 
process of the stapes. With the loss of the otic notch of the embolomeres, the 
tympanic process underwent reduction and was finally lost as well. Concur- 
rently with these changes, the quadrate process of the stapes became increas- 
ingly ossified. With reduction of the tympanic process and loss of the otic 
notch the tympanum achieved a ventral attachment to the quadrate process. 
The continued emphasis of the quadrate process, now the functional stapes, 
led to the cradling of the tympanum by the dorsal edge of the process. The 
chorda tymiDani still retained its primitive relationship to the quadrate process 
by crossing above the bone. The appearance of reptiles more advanced than 
those represented by Dimetrodon, such as Captoihinus, involved most im- 
portantly the development of a new tympanic process of the stapes. If this 



Chorda Tympani in Middle Ear 19 

neoniorph arose distal to the path of tlic chorda tympani the rehition oi the 
nerve to the stapes found in living diapsids was achieved; if the ncoinorph 
arose proximal to the path of the nerve, the nerve and stapes were related 
in a fashion that explains the appearance of the mammalian condition. 

There is, however, considerable evidence that these events did not occur. 
In no living tetrapod known to me does the chorda tympani pass above or 
loop over the quadrate process of the stapes prior to the descent of the nerve 
to the lower jaw. For example, in Laccrta and Ctenosaura the chorda tym- 
pani passes below the stapedial-quadrate articulation (Goodrich, 1930; Oel- 
rich, 1956); this relationship of nerve to articulation is also present in Gecko 
(deBeer, 1937). Similarly, the developmental history of mammals reveals 
that the path of the chorda tympani is external to the stapedial-incus articula- 
tion; the passage of the chorda tympani across the medial face of the mam- 
malian tympanum corroborates this. Nor is there any phylogenetic necessity 
for supposing that the chorda tympani passed dorsal to the quadrate process 
in the history of tetrapods. During the change of the rhipidistian hyomandib- 
ular to the tetrapod stapes, the hyomandiliular branch of the seventh nerve, 
arising from a common root vdth the chorda tympani, remained in back of 
the quadrate process as the hyomandibular swung out to achieve its tetrapod 
orientation. I suggest that the chorda tympani also remained behind the 
hyomandibular while this change was taking place, alloviring the nerve to nm 
forward beneath both the tympanic and quadrate processes of the stapes. 

There is no evidence to suggest that the quadrate process of the stapes 
in primitive reptiles became heavily ossified and performed as a functional 
stapes. Indeed, Romer and Price ( 1940 ) refer to obvious scars on the ventral 
surface of the stapes and a pit on the medial face of the quadrate in 
Dimctrodon. They suggest that the scars and pit served as the proximal 
and distal attachments, respectively, of a typically ligamentous or cartilagenous 
quadrate process. 

Both Vaughn's (1955) and Hotton's (1960) approaches are handicapped 
by the limitations that are inherent in forcing the features of adult morphology 
into a phylogenetic scheme that is itself a recitation of the history of adult 
animals. Both workers presuppose spatial barriers (the stapes and its proc- 
esses) to the change in the path of the chorda tympani that may not have 
been present ontogenetically at the time that the nerve changed from a post- 
tympanic position to a pretympanic position. The descent of the tympanum 
from the dorsal position that it occupied in the adult embolomeres was paral- 
leled presumably by the descent of the tympanic diverticulum in the embryos 
of those animals involved in the amphibian-reptilian transition. The more 
ventral position of the tympanic diverticulum probably blocked the chorda 
tympani as it pursued its primitive course across the floor of the developing 
tympanic cavity. Freedom for the nerve's path was then found dorsal to 
the cavity. The subsequent appearance of the stapes was consequently ventral 
to the nerve. This ontogenetic sequence of the nerve and tympanic diverticu- 
lum is known among amniotes; it occurs, for example, in the developing 
chick (Patten, 1951). 

In the manner described above, the position of the chorda tympani in 
mammals is easily derived from the pretympanic position of the nerve in 
primitive reptiles. The events that have just been discussed presumably 
obtained in both primitive captorhinomorphs, ancestral to living diapsid rep- 



20 University of Kansas Publs., Mus. Nat. Hist. 



tiles, and primitive pelycosaurs, ultimately ancestral to mammals. In primi- 
tive pelycosaurs the quadrate was quite deep; the stapes extended postero- 
ventrally towards the quadrate. The chorda tympani passed over the tympanic 
process and under the quadrate process, both of which were developed to a 
degree that was generally characteristic of early reptiles. The dorsal process 
of the stapes arose close to the footplate, in a proximal position on the 
stapedial rod. 

The stapes of therapsids differed in several respects from the stapes of 
pelycosaurs. The bone was much smaller than in pelycosaurs; the reduction 
seems to have been most extreme in quadrate and tympanic processes. Thrin- 
axodon (Estes, 1961), Kmgoria (Cox, 1959), and Placerias (Camp and Welles, 
1956), for example, show the distal termination of the bony stapes to be close 
to the quadrate and thus close to the plane of the tympanum. Additionally, 
in these animals the dorsal process had lost the proximal position it had 
possessed among pelycosaurs and arose far out along the stapedial rod, close 
to its distal termination. The consequence of the new position of the dorsal 
process was a restriction of the space between the dorsal process and the 
tympanum through which the chorda tympani passed; what had been a wide 
channel dorsal to the stapes for passage of the chorda tympani was now a 
narrow slit. Continued decrease of this space through which the chorda 
tympani passed probably brought the nerve into contact with the tympanic 
membrane; in its passage above the stapes the nerve was trapped by the 
tympanum. 

At this point, however, the chorda tympani still maintained its position 
ventral to the attachment between the stapes and the quadrate. It should be 
noted also that the articulation between the quadrate and the articular was 
lateral to the passage of the nerve. The migration of the quadrate and 
articular into the middle ear consequently followed a path that moved up 
and over the chorda tympani, completely free spatially from the passage of 
the ners'e and in no way altering it. 

The speed with which these later changes occurred in the mammalian- 
reptilian transition must have been comparatively great. No known fossils 
exhibit the intermediate stages between the achievement of a double articula- 
tion of the jaw and the incorporation in typical mammalian fashion of the 
quadrate and articular bones into the mechanism of the middle ear. But 
the explanation of the above changes requires no reliance upon macroevolu- 
tionary phenomena. The rate of change undoubtedly quickened, but the 
magnitude of the single changes need not be thought of as extraordinary. 

As Hotton (1960) has noted, the captorhinomorphs and synapsids have 
given rise to all living mammals and reptiles, with the possible exception of 
turtles. The unified theory to explain the path of the chorda tympani in the 
higher classes of vertebrates supports Hotton's estimate of the ultimate origins 
of these groups, and argues against the classic sauropsid-theropsid dichotomy 
of living reptiles and mammals advocated by, among others, Goodrich (1930). 
The ancestry of turtles is obscure, in spite of the suggestion by Olson ( 1947 ) 
that they may be descended from diadectids. In turtles the chorda tympani 
is pretympanic. In diadectids the dorsal otic notch is retained, implying a 
retention of the embolomerous post-tympanic path of the chorda tympani. If 
turtles were derived from diadectids, the pretympanic path of the chorda 
tympani must have been achieved independently from all other living reptiles. 



Chorda Tympani in Middle Ear 21 

ACKNO WLE DGM E NTS 

I am appreciative for the financial assistance furnished me by National 
Science Foundation Grant (NSF-G8624), through Dr. Theodore H. Eaton, Jr., 
principal investigator, during the course of my study of primitive reptiles. This 
report is a part of tliat stud\'. 

LITERATURE CITED 

Camp, C. L. and Welles, S. P. 

1956. Triassic dicynodont reptiles. Mem. Univ. California, 13(4) :255- 
348, 62 figs., 4 pis. 

Cox, C. B. 

1959. On the anatomy of a new dicynodont genus with evidence of the 
position of the tympanum. Proc. Zool. Soc. London, 132(3) :321- 
367, 17 figs. 

deBeer, G. R. 

1937. The development of the vertebrate skull. Clarendon Press, Oxford, 
xxiii + 552 pp., 143 pis. 

ESTES, R. 

1961. Cranial anatomy of the cynodont reptile Thrinaxodon liorhiniis. 
Bull. Mus. Comp. Zool., 125(6) :165-18(), 3 figs., 2 pis. 

Goodrich, E. S. 

1930. Studies on the structure and development of vertebrates. Con- 
stable, London, xxx + 837 pp., 754 figs. 

IIOTTON, N., III. 

1960. The chorda tympani and middle ear as guides to origin and di- 
vergence of reptiles. Evolution, 14(2) :194-211, 4 figs. 

Oelrich, T. M. 

1956. The anatomy of the head of Ctenosaura pectinata (Iguanidae). 
Misc. Publ. Mus. Zool., Univ. Mich., No. 94, 122 pp., 59 figs. 

Olson, E. C. 

1947. The family Diadectidae and its bearing on the classification of 
reptiles. Fieldiana: Geology, 11( 1 ) :2-53. 

Patten, B. M. 

1951. Early embryology of the chick. McGraw-Hill, New York, xiv -f 
244 pp., 102 figs. 

RoMER, A. S. and Price, L. I. 

1940. Review of the Pelycosauria. Geol. Soc. Amer. Special Papers, No. 
28, X + 538 pp., 71 figs., 46 pis. 

^'aughn, p. p. 

1955. The Pennian reptile Araeoscelis restudied. Bull. Mus. Comp. Zool., 
113(5) :305-467, 15 figs., 2 pis. 

Transmitted March 18, 1965. 



D 

30-7117 



^ - Kjp - L~&>-^^*-ujodij^n • 



University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 3, pp. 23-189, 4 figs., 51 M^. COMP. zqoj 
March 24, 1966 t^fHRARY 



JUN I ^ i96g 

Fishes of the Kansas River Sy^tofesiTY 
in Relation to Zoogeography 
of the Great Plains 



BY 



ARTIE L. METCALF 



University of Kansas 

Lav^rence 

1966 



UNIVERSITY OF KANSAS PUBLICATIONS 
MUSEUM OF NATURAL HISTORY 

Institutional libraries interested in publications exchange may obtain this 
series by addressing the Exchange Librarian, University of Kansas Library, 
Lawrence, Kansas. Copies for individuals, persons working in a particular 
field of study, may be obtained by addressing instead the Museum of Natural 
History, University of Kansas, Lawrence, Kansas. When copies are requested 
from tile Museum, 25 cents should be included (for each 100 pages or part 
thereof) for the purpose of defraying the costs of wrapping and mailing. For 
certain longer papers an additional amount indicated below, toward the cost 
of production, is to ,b^ included. Materials published to date in this series 
are as foUows. \ '\ 

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

Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. 
"Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 

figures in text. April 9, 1948. 
•Vol. 3. Nos. 1-4 and index. Pp. 1-681. 1951. 

•Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 
figures in text. December 27, 1951. 
Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953. 
"Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. 
Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10. 1952. 
Vol. 7. Nos. 1-15 and index. Pp. 1-651, 1952-1955. 
Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956. 
Vol. 9. Nos. 1-23 and index. Pp. 1-690, 1955-1960. 

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 Natiu-al 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 developrnent in the prairie vole (Microtus ochro- 
gaster). By Henry S. Fitch. Pp. 129-161, 8 figures in text, 4 tables. De- 
cember 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, 84 plates, 8 figiu-es in text, 35 tables. November 7, 1958. 

7. Home ranges and movements of the eastern cottontafl 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 Michoac&n, Mexico. 
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. Duelknan. Pp. 599-610, 1 plate, 1 figure in text. May 2, 1960. 

Index. Pp. 611-626. 
Vol. 11. Nos. 1-10 and index. Pp. 1-703. 1958-1960. 

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

3. The baculvun 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 figwes in 
text. May 2, 1960. 

5. Natural history of the Bell Vireo. By Jon C. Barlow. Pp. 241-296, 6 figmes 
in text. March 7, 1962. 

6. Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C. 
Fox. Pp. 297-307, 6 figures in text. May 21, 1962. 

7. Vertebrates from the barrier island of Tamaulipas, Mexico. By Robert K. 
Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309- 
345, plates 5-8. June 18, 1962. 

8. Teeth of edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 fig- 
ures in text. October 1, 1962. 

9. Variation in the muscles and nerves of the leg in two genera of grouse 
(Tympanuchus and Pedioecetes ) . Bv E. Bruce Holmes. Pp. 363-474, 20 
figures. October 25, 1962. $1.00. 

(Continued on inside of back cover) 



University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 3, pp. 23-189, 4 figs., 51 maps 
March 24, 1966 



Fishes of the Kansas River System 

in Relation to Zoogeography 

of the Great Plains 



BY 



ARTIE L. METCALF 



University of Kansas 

Lav^^rence 

1966 



University of Kansas Publications, Museum of Natural History 

Editors: E. Raymond Hall, Chairman, Henry S. Fitch, 
Frank B. Cross 



Volume 17, No. 3, pp. 23-189, 4 6gs., 51 maps 
Published March 24, 1966 



MUS. COIVIP. ZOOL. 
LIBRARY 

JUN 1 ^ 1QIS6 

HaKVAialj 

.iJNIVERSITYxr 
University of Kansas 

Lawrence, Kansas 



PRINTED BY 

ROBERT R. (BOB) SANDERS. STATE PRINTER 

TOPEKA, KANSAS 

1 966 



30-8449 



Fishes of the Kansas River System 

in Relation to Zoogeography 

of the Great Plains 

BY 

ARTIE L. METCALF 

CONTENTS 

PAGE 

Introduction 27 

General Physical Features of the Kansas River System 27 

Physiography, Geology and the Fauna 29 

Glacial Till-plains 30 

Osage Cuestas 31 

Kansas City and Lansing Groups 31 

Douglas Lowlands 32 

Shawnee Escarpments 32 

Wabaunsee Group 33 

Fauna of Wakarusa River 33 

Flint Hills 34 

Great Plains 36 

Dissected High Plains 36 

Ogallala Formation 40 

Kansas River Basin in Nebraska 42 

Kansas Rr^er System Prior to Settlement 48 

Lower Mainstream 50 

Tributaries 53 

Summary 55 

Kansas River System and Its Fauna Subsequent to 

Settlement 56 

History of the High Plains and of the Ancient Plains 

Drainage 64 

Hudson Bay Component 71 

Deflection Southward of Ancestral Missouri System 72 

Mississippi Component 75 

Variation From Northeast to South\vest 81 

General Body Shape 82 

Mouth 84 

Eye 84 

(25) 



26 University of Kansas Publs., Mus. Nat. Hist. 



FACE 



Scales 84 

Adaptive Value 85 

Variation, Its Possible Causes and Its Taxonomic 

Interpretation 85 

Records and Collections of Fishes 89 

Accounts of Species 93 

Species Probably Occurring in the Kansas River Basin . 162 

Species Probably Not Occurring in the Kansas Rwer Basin, 164 

Summary 167 

Acknowledgments 170 

Maps 171 

Literature Cited 177 



INTRODUCTION 

Aims of this report are to (1) ascertain which species occur, (2) 
ascertain distributional patterns of the species and relate these to 
environmental conditions, and (3) relate the findings to the history 
and development of the Basin, and to the general zoogeography of 
fishes of the Great Plains and western part of the Interior Lowlands. 

General Physical Features of the Kansas River System 

Streams of the Kansas (or Kaw) River System arise in eastern 
Colorado and flow generally in an eastward direction, the main- 
stream debouching into the Missouri River at Kansas City (Figs. 
1 and 2). Linearly, from west to east, the system traverses ap- 
proximately 480 miles; the drainage area comprises 61,440 square 
miles. 

The Kansas River proper is formed at Junction City, Kansas, by 
union of its two largest tributaries, the Smoky Hill and Republican 
rivers. The Smoky Hill River heads in Cheyenne and Kit Carson 
counties, Colorado, but tributaries in Colorado are highly inter- 
mittent at present. In western Kansas, the mainstream receives a 
few permanently flowing tributaries. From Wallace County, Kan- 
sas, east to Lindsborg, Kansas, the Smoky Hill River maintains a 
fairly straight eastward course. Near Lindsborg the Smoky Hill 
River turns sharply northward (probably the result of a stream- 
capture to be discussed later). Near Salina it acquires its two 
longest tributaries — the Saline and Solomon rivers. The Saline 
River, one of the saltiest streams in the United States according to 
Parker (1911:220), flows a linear distance of approximately 220 
miles almost due east. The Solomon River, which is slightly longer 
than the Saline, consists of two parallel-flowing branches, North 
Fork and South Fork. 

The Republican River heads in eastern Colorado where it, unlike 
the Smoky Hill River, possesses a number of permanently flowing 
tributaries. The Republican River proper is formed by confluence 
of its North and South forks and of the Arikaree River in Dundy 
County, Nebraska. For approximately 225 miles (linearly) the 
mainstream of the Republican River flows parallel to and slightly 
north of the Kansas-Nebraska border. From the northwest the 
Republican River receives Frenchman Creek, Red Willow Creek, 

(27) 



28 



University of Kansas Publs., Mus. Nat. Hist. 



Medicine Creek and numerous shorter tributaries. Major south- 
western tributaries are Beaver, Sappa and Prairie Dog creeks, all 
parallel-trending streams that head in northwestern Kansas. In 
Nuckolls County, Nebraska, the Republican River turns sharply 
southward into Kansas, then southeastward to Junction City. Few 
tributaries enter the Republican River in Kansas other than White 
Rock Creek, which joins the Republican in Republic County. 

Near Manhattan, Kansas, the Kansas River receives another major 
tributary from the north — the Blue River, having an impoundment, 
Tuttle Creek Reservoir, north of Manhattan. 



102° 



9 go 



96° 



' (a)®® 6i)(n) 



40' 



(49) ® ® (38)® 



COLORADO^ 



0® 




® ® ® ®,® (5)@®(]])©0 



40° 



102° 



99° 



96° 



Fig. 1. Major streams of the Kansas River Basin. Names of streams to which 
numbers refer are indicated in Table 1. (Scale as in Figure 2). 



Table 1. Noteworthy Streams of the Kansas River Basin. Numbers Are 
ON Fig. 1 and the Streams to Which They Refer. 



1. Kansas River 

2. Mill Creek 

3. Stranger Creek 

4. Captain Creek 

5. Washington Creek 

6. Delaware River 

7. Wakarusa River 

8. Soldier Creek 

9. Mission Creek 

10. Cross Creek 

11. Mill Creek 

12. Vermillion River 

13. Black Vermillion River 

14. Wildcat Creek 

15. Big Blue River 

16. Lyon Creek 



17. Little Blue River 

18. Turkey Creek 

19. Chapman Creek 

20. Lincoln Creek 

21. Gypsum Creek 

22. Beaver Creek 

23. Smoky Hill River 

24. Big Blue River 

25. Solomon River 

26. Big Sandy Creek 

27. Limestone Creek 

28. Republican River 

29. White Rock Creek 

30. North Fork Solomon River 

31. Thompson Creek 

32. South Fork Solomon River 



Fishes of Kansas River System 



29 



33. Saline River 

34. Big Creek 

35. Beaver Creek 

36. Deer Creek 

37. Sappa Creek 

38. Medicine Creek 

39. Prairie Dog Creek 

40. Red Willow Creek 
41. 



42. Blackwood Creek 

43. Chalk Creek 

44. Spring Creek (trib. of Stinking 
Water Creek) 

45. Turtle Creek 

46. Frenchman Creek 

47. North Fork Smoky Hill River 

48. Arikaree River 

49. North Fork Republican River 



Ladder Creek 

50. South Fork Republican River 

East of Manhattan the major northern tributaries of the Kansas 
River are the VermilHon River, the Delaware River and Stranger 
Creek; major southern tributaries are Mill Creek and the Wakarusa 
River. Along its lower course the Kansas River proper has not 
been impounded, although there is a low dam at Lawrence. 



PHYSIOGRAPHY, GEOLOGY AND THE FAUNA 

The landscape of the Kansas River Basin is mostly a product of 
Pleistocene and Recent geomorphic activity, although large ex- 
panses of only slightly modified Pliocene surface features exist in 




4 0' 



COLORADO. 



(ANSAS 



20 40 
I.I.I 

SCALE OF MILES 



40° 



102° 



99° 



96° 



Fig. 2. Important physiographic and geologic features of the Kansas River 
Basin bearing on the distribution of fishes. Above is a highly generalized 
cross-section (west-east) across the central part of the Basin in northern Kansas 
and eastern Colorado: T=: Tertiary; K == Cretaceous; P = Permian; ^ = 

Pennsylvanian. 
Below certain physiographic areas are indicated: HP = High Plains; NE = 
Escarpment formed on the Niobara Formation or Chalk Hills; GE := Escarp- 
ment formed on the Greenhorn Limestone Formation or Kearney Hills; DE = 
Escarpment formed on the Dakota Formation or Smoky Hills; FH = Flint 
Hills; OC = Osage Cuestas; LH = Loess Plains or Loess Hills; GT = Glacial 
Till-plains — NE, GE, and DE are all Cretaceous in age. 



30 University of Kansas Publs., Mus. Nat. Hist. 

the western part of the basin. All of the major glacial advances 
known to have developed in northeastern North America have left 
traces of their influence in the Basin, but ice of only two major 
glaciations, the Nebraskan and the Kansan, is thought actually to 
have reached the present confines of the Basin ( Frye and Leonard, 
1952:58). 

Glacial Till-Plains 

In Kansas, glacial till associated with the advance of Nebraskan 
ice has been identified with certainty only in the extreme north- 
eastern corner of the state where it is covered by till deposited 
during the subsequent Kansas Glaciation (Frye and Leonard, 
1952:57). Lugn (1935:40) indicated that, in eastern Nebraska, 
Nebraskan till is characteristically thicker than Kansan till. Never- 
theless, where not masked by Kansan till, the Nebraskan till has 
been eroded and covered by various sediments, chiefly glacial out- 
wash and loess, to such an extent that the till is of little consequence 
in a consideration of present physiography. 

Glacial till of Kansan age is, on the other hand, an important 
physiographic feature (Fig. 2). Kansan glacial till is found as 
far west as Marshall and Washington counties and as far south as 
central Wabaunsee, Shawnee and Douglas counties (Frye and 
Leonard, 1952:76, 77; O'Connor, 1960:50-52). In the southern part 
of the Big Blue Biver Drainage in Nebraska, Lugn (1935:P1. 1, 68, 
69) indicated the presence of Kansan till as far west as Fillmore 
and Jefferson counties, extending approximately to the valley of 
the Little Blue Biver. North of Crete, Nebraska, Lugn was not 
able to discern Kansan till west of Seward and Butler counties. 
Delimitation of the boundaries of the till area is difficult due to 
the large amount of dissection that has occurred around the pe- 
riphery of the till sheet. Erosion probably was facilitated by thin- 
ning of the original till sheet toward its margins. Factors other 
than erosion that have modified the Kansan till sheet since its 
deposition are soil-formation and further deposition, chiefly of 
loess, upon the till sheet. In a small area in the upper Big Blue 
Biver Basin in Nebraska thick loess overlies glacial till (Fig. 2). 
Several streams have headwaters in the Till Plains: the Big Blue 
River in Nebraska, the Black Vermillion River, the Vermillion Biver, 
the Delaware River and Big Stranger Creek. The lower parts of 
all these streams and lesser tributaries entering the Kansas River 
from the north have cut completely through the till mantie. After 
removal of the mantle some streams (especially in the area where 



Fishes of Kansas River System 31 

till covered the Shawnee Escarpment, discussed hereafter) have 
cut significant erosional valleys in the underlying bedrock (for 
example. Mud Creek, Nine-mile Creek and Wildhorse Creek). 
Above the point where these streams cut through the mantle to 
bedrock they have low gradient, bottoms of silt and sand and 
steeply-incised banks. Water is characteristically turbid. Below 
the point where these creeks cut through the till-mantle, water may 
remain turbid but gravel and rubble are common in stream-beds, 
gradient increases and rubbly riffles are formed. Concommitant 
with these physical changes are faunal changes. At seven collection- 
stations in the upper parts of the Delaware River and Big Stranger 
Creek above the point where glacial till is breached, only these 
species were found: Semotilus atromaculatus, Notropis I. lutrensis, 
Notropis stramineus mis^iriensis, Phenacobius mirabilis, Fime- 
phales promelas, Campostoma anomalum pullum and Ictalurtis 
melas. Below the breach in the till-mantle, the fauna is more 
diverse, including, in addition to the above species, Pimephales 
notaius, Notropis cornutus, Catostomus commersoni, Ictalunis 
natalis, Lepomis ctjanclliis and Etheostoma spectabile pulchellum. 

Osage Cuestas 

The Osage Cuestas ( Fig. 2 ) are formed on Upper Pennsylvanian 
strata that dip slightly to the west. The rocks are sedimentary, 
deposited in cyclothemic manner (Moore, 1949:25). Limestones 
and shales predominate but some sandstones are present, especially 
in the Douglas Group. Drainage is to the east, opposite to the 
direction of dip, resulting in east-facing cuesta-scarps formed on 
the more resistant strata. Such scarps are best developed where 
thick, resistant limestone beds overlie thick, nonresistant shales, as 
in the Oread Limestone over the Lawrence Shale; some scarps are 
at least 200 feet high (Schoewe, 1949:284). From east to west 
several escarpments are definable. 

Kansas City and Lansing Groups 

Of the Kansas City and Lansing groups of the Missourian Series, 
the Wyandotte, Plattsburg and Stanton limestones contribute to 
the formation of steep scarps along Kill Creek, Cedar Creek and 
Mill Creek in Johnson County, Kansas, and along Kaw Creek, Wolf 
Creek and several other short tributaries north of the Kansas River 
in Wyandotte and southeastern Leavenworth counties. These 
streams generally have permanent flow of clear water and have 



32 University of Kansas Publs., Mus. Nat, Hist, 

rubble bottoms and high gradients. The steep hillsides of their 
valleys have discouraged cultivation. Several species that are 
indicative of "good" (clear, unpolluted) water in the Kansas River 
Basin {Notropis cornutus, Etheostoma spectabile pulchellum and 
the crayfish Orconectes luteus) are found in these streams. Cedar 
Creek contains the most diverse fauna in this area. Mill Creek 
and small tributaries north of the Kansas River in Wyandotte 
County seem to have suffered some modification due to encroaching 
urban and industrial areas. 

Douglas Lowlands 

The Douglas Group of Pennsylvanian rocks is comprised largely 
of sandstone and shales. In northern Kansas these beds are weak, 
typically forming an area of low relief (herein termed "Douglas 
Lowlands"). Moore (1949:127) stated, "The outcrop belt of 
Douglas rocks is generally a rolling plain, bounded on the west by 
the east-facing Oread limestone escarpment. . . ." Tlie water- 
sheds of Little Wakarusa River, Spring Creek and Cole Creek are 
underlain, for the most part, by rocks of the Douglas Group, The 
Lawrence Shale Formation is the uppermost of the beds of the 
Douglas Group and forms the steep, continuous slope of the Oread 
Escarpment. Stteams possess low gradients, silty or sandy bottoms, 
and water is characteristically turbid. 

Shawnee Escarpments 

The Shawnee Group, of which the Oread Limestone is the lowest 
formation, includes four scarp-forming limestones, the Oread, 
Lecompton, Deer Creek and Topeka (herein termed "Shawnee 
Escarpments"). Moore (1949:139, 141) in discussing the Shawnee 
outcrop belt noted, "It is distinguished from the Douglas and 
Wabaunsee belts by the greater prominence of escarpments and 
generally more rugged topography ... In the vicinity of the 
Kansas River Valley, each of the Shawnee limestone formations 
makes a well-defined escarpment. . . ." 

The Oread Escarpment is one of the best-defined of the upper 
Pennsylvanian escarpments in northern Kansas, and is especially 
prominent in the vicinity of Lawrence (reliefs in excess of 100 feet 
are common). Limestones of tlie Shawnee Group form escarp- 
ments, not only along the Kansas River proper but also around tlie 
margins of the erosional valleys of the Wakarusa River and its 
tributaries in the western half of Douglas County and in eastern 



Fishes of Kansas River System 33 

Shawnee County. North of the Kansas River, Hmestones of the 
Oread Formation form similar rims around the valleys of Mud 
Creek, Wildhorse Creek, Nine-mile Creek, and the middle part of 
the Big Stranger Creek System. 

Wabaunsee Group 

Escarpments of the Wabaunsee Group of the Virgilian Series 
are less impressive than those of the Shawnee Group. Topograph- 
ically the area of outcrop in northern Kansas is predominantly a 
grassy, rolling upland. In its western part it is transitional to the 
Flint Hills in vegetation and topography. In tliis western part 
escarpments are formed by the Burlingame and Tarkio limestones. 
The watersheds of Snokomo, Mission and Soldier creeks are, for 
the most part, underlain by rocks of Wabaunsee age. The fish- 
fauna, as well as the flora, shows aflBnities to that of the Flint Hills 
directly to the west. Nofropis rubeUus, otherwise known only from 
the Flint Hills, was reported by Gilbert (1886:208) from Mission 
Creek (as N. rubrifrons) . 

Fauna of Wakantsa River 

The Wakarusa River traverses terrain formed mainly on rocks 
of the Wabaunsee, Shawnee and Douglas groups. The kinds of 
fishes found in Wakarusa tributaries of approximately the same 
size, originating in these tliree areas of outcrop are discussed below. 
Eight species, most of them widespread in the Kansas River Basin, 
were taken at stations in all three sections of the watershed: 
Semotilus atromaculattis, Notropis I. lutrensis, N. stramineus mis- 
suriensis, Pimephales promelas, P. notatus, Ictalurus melas, Lepomis 
cyanellus and L. humilis. Other species taken in the three areas 
are listed in Table 2. 

The lowest number of species (13) is from the Douglas Lowlands. 
The low relief developed on shales of the Douglas Group has made 
possible intensive cultivation of the area. Furthermore, the water- 
shed of Little Wakarusa Creek, east of its channel, drains a surface 
of exceedingly low reHef formed, according to O'Connor (1960:55), 
on deposits of Kansan glacial till and outwash. 

In the survey by Deacon and Metcalf (1961) made in 1959, no 
orange-throated darters {Etheostoma spectabile pulchellum) were 
taken in streams on the Douglas Lowland. In June, 1963, this 
species was common at some stations on Little Wakarusa River 
and Cole Creek in this area. Deacon and Metcalf (1961:320) 



34 



University of Kansas Publs., Mus, Nat. Hist. 



Table 2. — Fishes Taken in Three Sub-divisions 

Basin. 


OF THE WaKARUSA RiVER 




Douglas 

group 

lowland 


Shawnee 

group 

escarpments 


Wabaunsee 

group 

upland 


Carpiodes carpio 

Catostonms cotmnersoni 

Notemigonus crysolexicas 


X 
X 
X 
X 

X 


X 
X 

X 
X 
X 
X 
X 

X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 


X 
X 
X 


Hybopsis biguttata 




Notropis umbratilis 

Notropis cornutus 

Notropis topeka 

Phenacobius mirabilis 

Hybognathus placilus 

Campostoma anomalurn 

Idalurus pundatus 

Idalurus natalis 

Pylodidis olivaris 

Noturxis flavus 

Noturus exilis 

Percina caprodes 

Etheostoma nigrum 

Etheostoma spedabile 


X 
X 


Micropterus salmoides 

Lepomis macrochirus 


X 
X 


Lepomis megalotis 









theorized that E. s. pulchellum had been extirpated in much of the 
Wakamsa River System by drought in 1952-1956 and that in 1959 
it had not, due to a slow rate of dispersal, repopulated much of 
the Basin from its refugia in streams draining the Oread Escarp- 
ment, but by 1963 had dispersed to intermittent — less favorable but 
temporarily habitable — streams of the Douglas Lowlands. 

In the area of the Shawnee Escarpments 27 species were re- 
corded — more than twice the number found on the plain formed 
on rocks of the Douglas Group. Ten species were found exclu- 
sively in the zone of Shawnee Escarpments. On the upland plain 
formed on rocks of the Wabaunsee Group 15 species were collected, 
none exclusively in that part of the Wakarusa Basin. 



Flint Hills 

West of the Osage Cuestas the Kansas River traverses the Flint 
Hills, cuesta-scarps formed on limestones of Lower Permian age 
(Gearyan) (Fig. 2). Major southern tributaries are (east to west) 
Mill Creek, Antelope Creek, Deep Creek, McDowell Creek, Clarks 
Creek and Lyon Creek. From the north the Kansas River receives 
(east to west) Cross Creek, the Vermilhon River, the Blue River, 



Fishes of Kansas River System 35 

Wildcat Creek, Seven Mile Creek and Three Mile Creek, Much of 
the watershed of the Vermillion River is underlain by rocks of 
Pennsylvanian age (Wabaunsee Group) because of the presence 
of the buried Nemaha Mountains which have brought these deeper- 
lying rock strata to the surface in Pottawatomie and Nemaha 
counties in Kansas and in Pawnee and Richardson counties and 
northward in Nebraska. 

The cuestas of the Flint Hills generally possess greater relief 
than the Osage Cuestas because of the presence of thick beds of 
erosible shales separated at long intervals by especially resistant, 
often cherty limestones. The limestones are few in number. The 
Foraker, Grenola, Cottonwood, Wreford and Bameston formations 
are the principal scarp-makers. 

Streams originating in the Flint Hills have high gradients. In 
the process of degradation large quantities of limestone and shale 
rubble and gravel have been stripped away and transported by 
streams, this forming the dominant type of stream-bottom. Distance 
of transport is seldom great enough to reduce the sediments to 
the size of sand or less. The sharp relief of the escarpments and 
rockiness of the soil have hindered cultivation, while the luxuriant 
growth of tall prairie grasses has encouraged ranching. Lack of 
cultivation makes for less turbid streams, and the native grasses 
inhibit runoff, which also enhances clarity of the water and perma- 
nency of the streams. Streams of the Flint Hills support the most 
diverse fauna of the entire Kansas River Basin. Minckley (1959) 
in a survey of the Blue River in Kansas, a watershed situated almost 
entirely within the Flint Hills, recorded a total of 53 species of fishes. 
This compares favorably with the total of 83 species recorded here 
for the entire basin of the Kansas River. Fishes common in upland 
tributaries of the Blue River are Chrosomus erythrogaster, Semotilus 
atromaciilatus, Pimephales notatus, Pimephales promelas, Notropis 
cornutus, Catostomtis commersoni, Ictalurus melas, Ictalurus natalis, 
Etheostoma nigrum, Etheostoma spectabile pulchellum, Lepomis 
humilis, and Lepomis cyanellus. 

The clear streams of the Flint Hills, together with those draining 
the area of the Oread and other escarpments of the Shawnee Group, 
mentioned earlier, seem to have served as refugia for some species 
of fish and a crayfish that were formerly distributed more widely 
in the Kansas River Basin. Species that are still known to exist in 
both these refugia are Notropis topeka, Noturus exilis, Etheostoma 
n. nigrum, Percina caprodes and Lepomis megalotis. In recent 



36 University of Kansas Publs., Mus. Nat. Hist. 

years Hybopsis higuttata has been taken only from Mill Creek in 
the Flint Hills of Wabaunsee County but it was found as late as 
1924 in Rock Creek, Douglas County, in the area of the Oread 
Escarpment. 

Four species are, at present, known only from the Flint Hills 
segment of the Basin. These are Chrosomus enjthrogaster, Notropis 
ruhellus, Moxostoma erythrtirum and Percina maculata. Moxostoma 
erythrurum has been reported only from Mill Creek in Wabaunsee 
County, and recent records of N. rubeUus are only from Mill 
Creek, where it flourishes, and from one locality in the Blue River 
Drainage. 

In literature of the 1800's Notropis hudsoniiis, Etheostoma blen- 
nioides and Percopsis omiscomaycus were reported from the general 
area of the Flint Hills. 

Of the species mentioned in the preceding three paragraphs 
Chrosomus enjthrogaster, Hybopsis biguttata, Notropis topeka and 
Etheostoma n. nigrum were recorded in the 1800's from localities 
west of the Flint Hills. Recent changes in the distribution of N. 
topeka in the Kansas River System have been discussed by Minckley 
and Cross (1959). 

Great Plains 

All of the Kansas River Basin west of tlie FHnt Hills is witliin the 
Great Plains Physiographic Province of Fenneman (1931). Indis- 
pensable to aquatic life in this vast, semiarid region is the presence 
of water-bearing sediments to furnish water for permanent stream 
flow. There are few such strata, the most notable ones being (1) 
the Dakota Formation, (2) certain Pleistocene sands and gravels, 
widespread in southern Nebraska and scattered elsewhere, (3) the 
Pliocene Ogallala Formation where it is exposed in contact with 
the underlying aquicluding Niobrara Formation or Pierre Forma- 
tion both of Cretaceous age and (4) valley alluvium. The ground- 
water resources of the area are well known due to extensive work 
by the Kansas Geological Survey (reported mainly in Bulletins of 
the Kansas Geological Survey ) and by the United States Geological 
Survey in Nebraska (reported in various USGS Water Supply 
Papers ) . 

Dissected High Plains 

The eastern edge of the Great Plains in Kansas is an area of rolling 
hills that has been treated variously by physiographers. Fenneman 



Fishes of Kansas River System 37 

(1931:25) termed this region the Plains Border, recognizing it as 
an area formerly part of the High Plains, but in the process of being 
degraded to a lower base level. Emphasizing this process of deg- 
radation, Schoewe (1949:302) employed the name "Dissected 
High Plains" for this region. The boundary between the Flint 
Hills and the Dissected High Plains is ill-defined, mainly due to the 
presence of an intervening plain formed on rocks of the Wellington 
Formation of Middle Permian age. The bedrocks of the Dissected 
High Plains continue northward into Nebraska but north of the 
Republican River are covered by loess, glacial till or both. It seems 
best to consider the Republican River as the northern boundary 
of the Dissected High Plains Physiographic District, including the 
area to the north in the Loess Plains District (Fig. 2). 

The plain of the Wellington Formation in the Kansas River Water- 
shed ranges in width from approximately 10 miles in Washington 
and Marshall counties to 35 miles in southern Dickinson and south- 
eastern Saline counties. The principal tributaries of the Kansas 
River in this area are Holland and Turkey creeks from the south 
and Mud and Chapman creeks from the north. The Smoky Hill 
River here lacks the high bluffs characteristic of its erosional valley 
in the Flint Hills and Osage Cuestas to the east. According to 
Latta (1949:20) the Smoky Hill Valley is three to four miles wide 
where bordered by beds of the soft Wellington Shale but narrows 
to one to tsvo miles after entering the area of outcrop of the more 
resistant limestones of Gearyan ( Wolf campian ) age below Chap- 
man in Dickinson Count> . The gradient of streams in tliis area is 
low; sti-eams are turbid and have muddy bottoms. Bayne, Walters 
and Plummer (1959:57) characterized the Wellington Formation 
as a poor aquifer because of its low permeability. Presumably it 
yields little water to streams of the area. The fish-fauna is im- 
poverished, especially as compared to that of the Flint Hills im- 
mediately to the east. Hassler (Unpublished dissertation, Kansas 
State University, 1940) surveyed the fishes of Chapman Creek in 
Clay and Dickinson counties. He reported only 11 species; these 
I interpret to have been Lepisosteus osseus, Carpiodes c. carpio, 
Semotilus atromaculatus, Notropis stramineiis missuriensis (re- 
ported by Hassler as N. cornutus — possibly more than one Notropis 
was represented but N. stramineiis surely was the most abundant 
species present), N. lutrensis, Pimephales promelas, Campostoma 
anomalum pulliim, Ictalurus melus, I. punctatus, Lepomis cijanelhis 
and L. Jmmilis (listed by Hassler as Eupomotis gibbosus). 



38 University of Kansas Publs., Mus. Nat. Hist. 

My collections in Holland, Mud, and Turkey creeks have revealed 
a similar paucity of species and have not added any species to the 
list given from Chapman Creek. 

West of the plain formed on the Wellington Formation are three 
belts of hills in the Dissected High Plains District; the easternmost 
is commonly called the Smoky Hills, and the westernmost two 
comprise the Blue Hills (Schoewe, 1949:310). 

The highly irregular escarpment of the Smoky Hills has developed 
on resistant beds of the Dakota Formation (DE in Fig. 2). The 
line of outcrop of this formation trends southwest to northeast, out- 
cropping over large parts of Ellsworth, Lincoln, Ottawa, Cloud, and 
Washington counties (Schoewe, 1952: Fig. 1). Plummer and 
Romary (1942:327) described the Dakota Formation as consisting 
mostly of varicolored clays that include lenticular siltstones and 
channel sandstones that may appear at any stratigraphic level in 
the formation. These lenses are the most salient features of the 
formation topographically as they form the crests of the Smoky 
Hills. 

In addition to its importance as a scarp-maker the Dakota Forma- 
tion is noteworthy for its role as an aquifer in north-central Kansas 
(Fishel et al, 1948:82 Repubhc County and northern Cloud County; 
Berry and Drumm, 1952:25 Lincoln County; Hodson, 1959:66 
Mitchell County). Water from this formation is variable in hard- 
ness. In Mitchell County, Hodson (1959:Table 5) found total 
hardness ranging from 27 ppm. to 816 ppm. and dissolved calcium 
from 7.1 to 296 ppm. 

Where the Dakota Formation crops out, stream-flow is more 
nearly permanent than in the Wellington Plain to the east or the 
Blue Hills to the west. 

The Blue Hills consist of two belts of hills, trending southwest 
to northeast. The easternmost escarpment is formed on the Green- 
horn Limestone ( GE in Fig. 2 ) ; the western escarpment, also called 
Chalk Bluffs, is formed on the Niobrara Chalk Formation (NE in 
Fig. 2). The Dakota Escarpment is separated from that formed 
on the Greenhorn Limestone by an area of littie relief formed on 
the Graneros Shale; another such area that has formed on the Carlile 
Shale separates the Greenhorn Escarpment from that formed on 
the Niobrara Chalk. All of these rocks of the Blue Hills are poor 
aquifers. Ground water studies in the counties in which the 
Graneros Shale and Carlile Shale crop out, consistently report that 
few or no wells obtain water from these formations. The aquatic 
life of streams originating in the area of the Greenhorn Limestone 



Fishes of Kansas River System 39 

Escarpment is meager. The only streams of consequence are those 
that originate to the west and flow through the escarpment such 
as the Smoky Hill, Saline, and Solomon. Tributaries are shallow, 
generally intermittent, and most have silty bottoms. 

The westernmost of the three escarpments of the Dissected High 
Plains, that formed on the Niobrara Chalk Formation (comprising 
the Fort Hays and Smoky Hill chalk members), does not form 
strong cuestas as do the other t\vo but tends, rather, to produce 
the well-known and often bizarre bluffs, pinnacles and buttes of 
the "Chalk Bluffs" of western Kansas. In the Ladder Creek Drain- 
age of Wallace, Greeley, Wichita, Logan, Scott, Gove, and Lane 
counties, Bradley et al (1957:16, 20) found that the Niobrara 
Formation was not an important aquifer but that it supplied water 
to a few wells in areas where no other water-bearing rocks were 
accessible. Hodson and Wahl (1960:64,65) noted that "The Fort 
Hays Chalk Member is not known to yield water to wells in Gove 
County . . ." and that "The Smoky Hill Chalk Member is a 
poor aquifer in Kansas and yields very little water to wells in Gove 
County." 

Streams draining the area of the Chalk Bluffs are highly inter- 
mittent. Twin Butte Creek and Chalk Creek in southern Logan 
County are streams 30 to 35 miles in length that, as their names 
suggest, traverse an area dominated by the Niobrara Chalk and 
the overlying Pierre Shale. Johnson (1958:17) reported tliat springs 
occur on Chalk Creek where its erosional valley is incised below 
the contact of the Ogallala and Niobrara formations. He reported 
springs as common in one central part of the stream but added 
that "running water in the channel farther downstream is infrequent 
during the hot, dry summers, despite tlie increments in flow from 
the springs." Johnson (1958:16,17) reported an average annual 
discharge probably of less than six cfs for Chalk Creek and tliree 
cfs for Twin Butte Creek. Bradley et al ( 1957:29) noted that during 
years of high precipitation there was some discharge in Twin Butte 
Creek but that in dry years it flowed only after heavy precipitation. 

In August, 1961, I was unable to find running water or any fishes 
in either Twin Butte Creek or Chalk Creek. On April 1, 1962, 
however, I found both running water and fishes in the lower part 
of Chalk Creek, although Twin Butte Creek was dry. Fishes taken 
in Chalk Creek at that time were Semotilus atromaculatus, Notropis 
stramineus missuriensis, Notropis I. lutrensis, Pimephales promelas, 
Campostoma anomalum pullum, Fundulus kansae, Ictalurus melas, 
Lepomis ctjanelliis, Lepomis humilis, and Etheostoma spectabile 
2—8449 



40 University of Kansas Publs., Mus. Nat. Hist. 

pulchellum. Specimens of S. atromacttlatus, C. a. ptillum and E. s. 
pulchellum were in breeding condition. The fish faunas of these 
creeks of the Chalk Bluffs Area of Gove and Logan counties is 
seemingly limited to species that ascend the streams from the some- 
what more permanent Smoky Hill River during periods when run- 
ning water is present. Few fish probably survive tlie summer in 
these streams. 

The Pierre Shale that overlies the Niobrara Chalk resembles the 
latter in being a poor aquifer but an excellent aquiclude. The 
aquicludal properties of the Niobrara Chalk and the Pierre Shale 
are important in relation to the overlying Ogallala Formation of 
Pliocene age that covers a large part of the High Plains. 

Ogallala Fonnation 

In much of the Basin of the Kansas River, west of a line from 
Ness to Smith counties, Kansas, to Adams County, Nebraska, the 
Ogallala Formation is present. Its thickness varies from a feather- 
edge in the east to at least 350 feet in western Kansas (Bayne, 
1956:25). According to Frye, Leonard, and Swineford (1956:8) 
the formation in northern Kansas is a heterogeneous complex of 
clastic deposits ranging from coarse gravels to clay, uncemented or 
variably and variously cemented by opaline or calcareous cements. 
Frye and Leonard (1959:12) interpreted this heterogeneity as the 
result of a "random intermixture of channel and flood-plain environ- 
ment of deposition." Further aspects of the environment of deposi- 
tion of this formation are discussed elsewhere. Along many stream 
valleys, particularly along that of the upper Smoky Hill River, the 
Ogallala Formation has been removed by erosion and Cretaceous 
bedrock is exposed. Much of the formation has also been mantled 
by loess, dune sand and Pleistocene sands and gravels. Frye and 
Leonard (1959:6) stated "The Ogallala formation is of great eco- 
nomic importance throughout the Great Plains, primarily because 
it constitutes the largest and most extensive groundwater reservoir 
of the region." By virtue of this quality, the formation is stressed 
here. 

The aquicludal nature of the underlying Niobrara Formation or 
Pierre Shale has been mentioned above. Springs occur at many 
localities where the contact between the water-bearing Ogallala 
formation and the underlying aquicludes is exposed. Bradley et at 
(1957:50-53) mentioned such springs in Rose Creek valley and 
along Eagle Tail Creek in Wallace County and along Ladder Creek 



Fishes of Kansas River System 41 

in Scott County State Park, where one spring, called Big Spring, 
is reported to yield 400 gpm. These authors further observed 
(p. 53) "Many springs rise in other tributaries of Smoky Hill River, 
particularly in Hell and Salt creeks in the eastern part of the area. 
The contribution from the Ogallala formation by such spring dis- 
charge to the flow of Smoky Hill River east of Ladder Creek is 
considerable." Walters (1956:30) described similar springs along 
Beaver Creek in Rawlins County: "Water moving laterally at the 
base of the Ogallala formation on the top of the impermeable 
Pierre shale flows or seeps out at the surface where the top of the 
Pierre shale is exposed, Beaver Creek, which is an effluent stream, 
has many inconspicuous seeps along its entire course and receives 
considerable water from the ground-water reservoir." The perma- 
nency of the water supply derived from the Ogallala Formation 
was indicated by Johnson (1958:48,49) in describing ground water 
conditions in Logan Count}', Kansas. In wells measured period- 
ically he found little or no influence on water-level by seasonal or 
year-to-year variations in precipitation. Long-term trends in pre- 
cipitation possibly were reflected in the water levels after a lag of 
about three and a half years. 

The springs and seeps deriving their discharge from the Ogallala 
formation are of paramount importance to aquatic Hfe in this semi- 
arid region. Areas in which springs are especially numerous are 
the upper Smoky Hill Valle>', where aquifer and aquiclude are ex- 
posed, and along the valley of the upper Republican River and the 
lower courses of its tributaries where dissection has been great 
enough to expose Cretaceous aquicludes (Bradley and Johnson, 
1958:599,603). Much of the lower course of Medicine Creek 
(Frontier Co., Nebraska) is cut in Cretaceous bedrock due to the 
fact that this stream parallels roughly the crest of the Cambridge 
Arch, which raises lower strata (Bradley and Johnson, 1958:599). 
Medicine Creek, then, receives water from springs at the contact 
of Pliocene and Cretaceous rocks and in this respect differs from 
Red Willow Creek to the west and from the streams that receive 
water from Pleistocene gravels to the east. Noteworthy among the 
springs in the Republican drainage are those along Rock Creek in 
Dundy County and along the North Fork Repubhcan River near 
Wray, Colorado; at both localities small fish hatcheries have been 
built. Table 3 indicates the species of fishes recorded for Wallace 
County, Kansas, and for Dundy County, Nebraska — areas in which 
springs issuing from the Ogallala Formation supply much water to 
streams. From the two areas a total of 30 species has been recorded. 



42 University of Kansas Publs., Mus. Nat. Hist. 

For this longitude in the High Plains this is a surprisingly large 
number of species. Collections from the area east of the Ogallala 
Escarpment in the Dissected High Plains District of Kansas rarely 
contain more than ten species of fish. Several collections made in 
Rose Creek and in the Smoky Hill River (combined species list) 
near Wallace, Kansas, have yielded the following numbers of 
species: 13 (collection by Taylor, 1950); 16 (collection by Cross 
and Nelson, 1958); and 15 (collection by me, 1961). 

Notropis blennius and Hybognathus hankinsoni have been taken 
almost exclusively in the High Plains part of the Kansas River 
Basin where spring-fed, permanently-flowing streams occur. 

Kansas River Basin in Nebraska 

Most of that part of the Kansas River Basin in Nebraska cannot 
be placed in the physiographic subdivisions that have been em- 
ployed for Kansas, although the basin is almost entirely wdthin the 
Great Plains Physiographic Province (Fenneman, 1931:4). This 
difficulty is mainly due to the greater degree of mantling by glacial 
till, loess, dunesand, and fluviatile sands and gravels that occurred 
in Nebraska during Pleistocene time. To my knowledge no detailed 
study of the physiography of Nebraska has been undertaken. 

Loess Plains (Loess Hills) — In the Kansas River Basin the Illi- 
noian and Wisconsinan glacial ages were times marked by the 
deposition of fine, in great part eolian, sediments. There are three 
stratigraphic imits of the post-Kansan Pleistocene of which 'loesses" 
or silts are prominent components. These are, oldest to youngest, 
the Loveland Silt of lUinoian age (Frye and Leonard, 1951:293-295) 
and the Peoria and Bignell silts of Wisconsinan age (Frye and 
Leonard, 1951:298-303). All have been ranked as members of the 
Sanborn Formation by Frye and Fent (1947:42). 

Although the origin of "loess" has been a subject of controversy, 
there seems to be agreement that the late Pleistocene loesses of 
Kansas and Nebraska are almost entirely eolian in origin (Lugn, 
1935:130fiF., 161ff.; Swineford and Frye, 1951:317). Thicknesses of 
loess are great enough in most of south-central Nebraska between 
the Platte and Republican rivers to have physiographic importance 
(LH in Fig. 2). 

Most of the Kansas River Basin west of the Big Blue River and 
north of the Republican River seems best assigned to the Loess 
Plains District. (A small part of the Big Blue River Basin in 
Seward, Lancaster, Saline, Gage, and Jefferson counties, Nebraska, 



Fishes of Kansas River System 



4S 



Table 3. — Species of Fish Taken in ( 1 ) Tributaries and Mainstream of 
Smoky Hill River in Wallace County, Kansas, (2) Tributaries ani> 
Mainstream of Republican Rh'er in Dundy County, Nebraska, and (3) 
Southward-flowing Tributakies on the North Side of the Republican 
RrvER in Franklin and Webster Counties, Nebraska. ( 1 ) and ( 2 ) Are 
Fed by Springs Issuing From the Base of the Ogallala Formation; 
(3) Drains an Area Underlain by Pleistocene Sands and Gravels. 





(1) 
W allace 
Co., Ks. 


(2) 
Dundy Co. 

Nebraska 


(3) 
Republican 
tributaries 


diinrimts carvio 


X 
X 

X 

X 
X 
X 
X 
X 
X 
X 
X 
X 
X 

X 
X 
X 
X 
X 
X 
X 

X 
X 

X 


X 
X 
X 

X 
X 

X 
X 
X 
X 
X 

X 
X 
X 
X 
X 

X 
X 
X 
X 
X 
X 
X 
X 
X 


X 


fifmotilus atroitiaculatus 


X 


H iihonsis aracilis 


X 


H uhnnftis biauttata 




Huhnnsis sloreriancL 




Huhovsis aestivaiis 




Pikenacohius mirabilis 

Notropis cornutus 

Notropis lulrensis 


X 
X 


Notropis blennjus 




Notropis stramineus 


X 


Hybognathus hankinsoni 

Hybognathus placitus 

Pimephales nolatus 


X 
X 


Pimephales promelas 

Campostoma anomalum 

Carviodes cwrinus 


X 
X 


Carpiodes carpio 


X 


Catostomus commersoni 


X 


Ictalurus punctatus 


X 


Ictalurus natalis 


X 


Ictalurus melas 


X 


Noturxis flavus 


X 


Fundulus kansae 

Micropterus salmoides 

Lepomis cyanellus 


X 
X 


Lepomis humilis 

Lepomis macrochirus 

Ponioxis nigromaculatus 

Eiheostoma spedabile 


X 
X 


Total 


23 


24 


20 



can be assigned to the Glacial Till Plains District discussed hereto- 
fore.) Fenneman (1931:21,22) discussed the Loess Plains District 
only briefly, applying the term to an area east of the Sandhills 
District and adding, "To the south the loess overlaps the flat uplands 
of the typical High Plains, thinning out in northern Kansas where 
it merges with the general soil sheet . . ." That a loess mantle 
is also widespread over much of the upland surface of northwestern 
Kansas has been pointed out by Hibbard et al (1944:6). In a 
generalized areal mantierock map of Nebraska, Condra and Reed 
(1959: Fig. 3) indicated that nearly all of the Kansas River Basin 



44 University of Kansas Publs., Mus. Nat. Hist. 

west of the Big Blue River in Nebraska is covered by loess. Excep- 
tions are areas in which stream erosion has removed the loess cover 
or where dunesand has been deposited rather than loess. These 
areas of dunesand in the Kansas River Basin are mainly in Dundy, 
Chase, Perkins, Lincoln and Hayes counties in southwestern Ne- 
braska. They are surrounded by plains of loess, are scattered in 
occurrence, and seem better refen-ed to the Loess Plains District 
than to the Sandhills District. To the south and west the mantle 
of loess gradually becomes thinner over northwestern Kansas and 
eastern Colorado and is, consequently, of less importance physio- 
graphically than in Nebraska, even though overlying most of the 
surface of the Ogallala Foiination. 

The surficial mantle of the Loess Plains overlies various other, 
often intervening, sediments in addition to die Ogallala Formation 
of the High Plains; in some cases these underlying sediments are 
of more importance to the regimen of streams in the area than is 
the loess cover itself. 

In parts of the drainage of the Big Blue River in Saline, Jefferson, 
Polk, Butler and Seward counties, Nebraska, and notably in the 
drainages of Lincoln Creek, Beaver Creek, and North Branch Blue 
River, loess overlies glacial till. Neither loess nor glacial till is 
effective as an aquifer; thus, streams receive little discharge from 
this doubly thick mantle. Streams have done little valley-cutting 
in the area, gradients are only a few feet per mile, and bottoms are 
of fine, tenacious silt derived from the loess mantle. Water in these 
streams is characteristically highly turbid; banks are low but steep 
and the streams resemble miniature canyons winding across the 
loess mantle. These streams are some of the poorest within the 
basin in terms of condition of water and diversity of faima. The 
only species found in September, 1961, were Pimephales promelas 
and Ictalurus melas, which were common, and Lepomis cyanelliis, 
which was rare. The crayfish Orconectes nais was abundant at 
some stations. Even in the 1800's the fauna in this area seems to 
have contained few species. Evermann and Cox (1896:349), at 
four stations in this area, obtained only the following species: 
Semotilus atromaculahis, Notropis I. lutrensis, Notropis stramineiis 
missuriensis, Pimephales promelas, Ictalurus melas, Noturus flavus 
(USNM 76123), and Lepomis humilis. At Seward, Nebraska, on 
Lincoln Creek, Evermann and Cox secured only one species — 
Notropis I. lutrensis. They noted that the water at all stations was 
muddy. Downcutting by streams west of an area where loess 



Fishes of Kansas River System 45 

overlies water-bearing sands and gravels of Pleistocene age (dis- 
cussed below) generally has not been deep enough to remove 
the thick deposits of loess and of the Ogallala Formation so as to 
expose the Ogallala-Cretaceous contact, which is often spring- 
bearing. Bradley and Johnson (1958:600) reported that both the 
Loveland Loess and Peoria Loess in uplands of this area may 
reach tliicknesses of 60 feet each. The overlying Bignell Loess is 
as thick as 40 feet in places, and the underlying Ogallala Formation 
as thick as 400 feet. Bradley and Johnson (1958:611) noted that 
recent alluvium is the common aquifer for stock and domestic wells 
in the smaller tributary-valleys. These wells are shallow and often 
go dry. Presumably such Recent alluvium also furnishes scant 
amounts of water to sti-eams. The area is poorly watered. Red 
Willow Creek in July, 1961, after flowing for a distance of more 
than 50 miles, seemed to carry less water than did creeks such as 
Ehn Creek in Webster County (draining an area of Pleistocene 
sands and gravels) that were less than 20 miles long. 

In addition to separating the level of drainage-channels from that 
of the water table, the great thicknesses of loess further modify 
streams by contributing fine silt, which renders the water turbid and 
forms stream-beds of fine, tenacious silt. Streams such as Red 
Willow Creek and Stinking Water Creek resemble those farther 
east where loess overlies glacial till in the valley of the upper Big 
Blue River. A Secchi disc reading in Medicine Creek above the 
area of Cretaceous and Pliocene outcrops on August 22, 1961, was 
eight inches. The fauna is impoverished; fishes taken in Red 
Willow Creek were Cyprinus carpio, Hyhopsis gracilis, Notropis I. 
lutrensis, Notropis stramineiis tnisstiriensis, Lepomis cyanellus, 
Lepomis humilis and Pomoxis annularis. 

In extreme western Nebraska and in much of the Basin in eastern 
Colorado, as in northwestern Kansas, the mantle of loess thins and 
the Loess Plains gradually merge into the typical High Plains in 
which the surface of the Ogallala Formation is relatively lightly 
mantled. 

Pleistocene Sands and Gravels — To the west of the glaciated area 
described above the loess mantle lies upon another mantle com- 
posed of fluviatile sands and gravels of Pleistocene age. Lugn 
(1935:88) placed these fluviatile sediments west of the glacial till 
area in four formations: Holdrege, Fullerton, Grand Island, and 
Upland, all comprising the "Platte series," which is distributed over 
15,000 to 20,000 or more square miles in south-central Nebraska. 



46 University of Kansas Publs., Mus. Nat. Hist. 

Frye and Leonard (1952:Fig. 2) ranked (1) the Holdrege and 
Fullerton as members making up the Blanco Formation and (2) the 
Grand Island and Sappa ( = Upland Formation of Lugn, 1935 ) as 
members making up the Meade Formation. Of chief interest here 
are the Holdrege and Grand Island members, which in Nebraska 
are, according to Lugn (1935:92,103), predominantly sands and 
gravels of quartz and other granitic and metamorphic crystalline 
minerals and rock fragments. Both members are coarse toward the 
base and become finer toward the top. The Fullerton and Sappa 
members, which succeed in Nebraska the Holdrege and Grand 
Island members, respectively, consist of clay, silts, and fine sands. 
Frye and Leonard (1952: Fig. 1) indicated a time of deposition for 
the Holdrege Formation from the time of maximum Nebraskan 
glaciation until late in the Nebraskan Age. They indicated a similar 
correspondence of the Grand Island Member and the Kansan glacia- 
tion (p. 104, Fig. 1), with deposition of the Sappa Member extend- 
ing into early Yarmouthian time. 

The great geographic extent of these gravels and their unique 
history seems first to have been properly appreciated and explained 
by Lugn (1935:81-127). As visualized by Lugn these gravels and 
sands accumulated as "inwash-outwash" sediments west of the 
glaciated areas in eastern Nebraska and Kansas. That is, around 
the glacial margins, barriers must have been formed, either con- 
temporaneously or successively, by glacial ice, moraines, and other 
features of glacial till. Drainage was to the west and south away 
from these barriers (especially during the time of glacial melting). 
At the same time streams swollen by pluvial conditions and, in some 
cases, receiving in their upper parts meltwaters from glaciers of 
the Rocky Mountains brought great amounts of sediments into 
south-central Nebraska from the west and north. Thus both inwash 
and outwash contributed to accumulation of deposits of sands and 
gravels. During Holdrege time sands and gravels were laid down 
in pre-existing valleys to be overlain in turn by finer sediments 
during Fullerton time. As sediments (Grand Island and Sappa) 
continued to be added, valleys were filled, transgressed, and a 
coalescent sheet of elastics covered a great part of south-central 
Nebraska (Lugn, Fig. 5). 

In some of the areas discussed above a stratum of heterogeneous 
clastic materials was laid down above the silts and clays of the 
Sappa Formation. This unit has been termed the Crete Member 
by Frye and Leonard (1951:293), who considered it to be lUinoian 
in age. The Crete sands and gravels are far less widespread than 



Fishes of Kansas River System 47 

are those of the Holdrege and Grand Island members and are rela- 
tively thin. According to Liign (1935:129) the Crete Member 
attains thicknesses of at least 35 feet. It is locally significant as a 
source of ground water. 

The thickest fluviatile deposits occur in the counties west of the 
glacial till border in Nebraska. Sands and gravels extend as far 
west as Medicine Creek, but thin considerably towards the west 
in the Repubhcan River Valley. According to Lugn (1935:104) 
the Grand Island member "ranges in thickness from 30 to perhaps 
150 feet, but averages about 75 feet." In the area directiy north 
of the Republican River the fluviatile sheets of sands and gravels 
are not coalescent but break up and almost fill narrow, steep-sided 
valleys ("outlet valleys") that existed along the north side of the 
ancestral Republican River in pre-Holdrege time (Lugn, 1935:113). 
These outlet valleys are now occupied by short, permanently- 
flowing tributaries of the Republican River that obtain their water 
from the gravels considered here. Contemporaneous fluviatile 
gravels extend into adjacent northern Kansas in northern Republic 
County (Lugn, 1935:196) and along the valleys of Big and Little 
Blue rivers and Mill Creek in Marshall and Washington counties. 

Due to the efficiency of the Holdrege and Grand Island gravels 
and sands as aquifers, streams in this area contrast greatly with 
those just described in the area where loess mandes glacial till. 
Lugn (1935:203-207) pointed out that the configuration of the 
level of the water table of the area discussed here is inclined down- 
ward toward the south, thus allowdng subsurficial flow of water 
from the Platte River Valley southward. The water emerges ir 
tributaries of the Little Blue River and the Republican River. Not all 
streams in the area are perennial, however. The deptli to the water 
table is variable and dependent on the thickness of the overlying 
loess strata, dissection of the land, and nature and extent of the 
water-bearing sediments. Johnson and Keech (1959:20) noted 
that the water table is approximately 75 feet below the valley of 
the West Fork of the Big Blue River north of Hastings, Nebraska. 
Westward and southward from these tributaries of the Big Blue 
River a greater amount of perennial stream flow exists in the basin 
of the Litle Blue River and in south-flowing tributaries of the Re- 
publican River in Nuckolls, Webster, Franklin, and Harlan counties, 
Nebraska. However, the upper parts, even of these streams, are 
above the water table and are highly intermittent. Where the level 
of the stream channel intersects the level of the water table streams 
become effluent and, in some cases, pick up considerable volume 



48 University of Kansas Publs., Mus. Nat. Hist. 

over a relatively short distance. Thus, on July 19, 1961, I noted 
that Elm Creek in Webster County changed in a distance of less 
than two miles from a dry stream bed to a flowing stream (with 
fish) carrying several cubic feet per second. On September 9, 1961, 
the upper portion of the Little Blue River was highly intermittent 
to completely dry above Holstein (Adams County, Nebraska) but 
accrued a considerable volume of flow in a distance of a few miles 
south of Roseland, Adams County. 

Effluent parts of streams of this area resemble greatly, due to 
their permanency of flow, those streams that are fed from permanent 
springs issuing from the base of the Ogallala Formation. Typically 
the streams are shallow and braided, with clear, cool water flowing 
briskly over sandy bottoms. The streams have not been affected 
greatly by settlement or cultivation, because they are fed from 
springs generally in or along the stream-bed itself; runoff from 
upland areas occurs only in times of heavy rainfall. Cultivation is 
not extensive because streams have dissected the non-resistant loess 
mantle and formed deep, steeply-walled valleys. 

The fauna of the streams resembles that found in the spring-fed 
streams of Wallace County, Kansas, and Dundy County, Nebraska. 
A comparison of species that have been taken in the three areas is 
presented in Table 3. The crayfish Orconectes n. neglectus has 
been taken in Cottonwood Creek, Franklin County, Nebraska, and 
in Elm Creek, Webster County, Nebraska. Elsewhere in the Kansas 
River Basin it has been found in recent years only in streams drain- 
ing areas of contact between Pliocene and Cretaceous rocks in 
Dundy County, Nebraska, and Yuma County, Colorado, and in the 
Flint Hills area of Kansas. Amphipods (species undetermined) 
occur in springs of this area, in Dundy County, Nebraska, in Wallace 
County, Kansas, along the Shawnee Escarpments in the Osage 
Cuestas, and in the Flint Hills. 

To the west the Pleistocene gravels become thinner. It has been 
noted previously that much of the lower course of Medicine Creek 
was cut down to Cretaceous and Pliocene bedrock due to the 
presence of the Cambridge Arch, West of Medicine Creek, 
Pleistocene gravels are limited mainly to the valley of the Repub- 
lican River, and in the area north of the River loess deposits or sand 
dunes lie directly on bedrock of the Ogallala Formation. 

KANSAS RIVER SYSTEM PRIOR TO SETTLEMENT 

Extensive archeological work in the Basin has failed to reveal 
much information concerning fishes. Concerning the Pawnee In- 



Fishes of Kansas River System 49 

dians, Wedel (1936:62) noted that "Fish appear to have been used 
but little, as their bones are seldom present in the refuse-filled 
caches." The Pawnee Indians were inhabitants of the middle 
reaches of the Kansas River Basin, north and west of Big Blue 
Ri\er. Whether the more sedentary Kansa Indians that inhabited 
the better-watered lower part of the Basin made greater use of 
fish is not known, but Wedel ( 1946; 1961 ) did not list remains of 
fishes or artifacts connected with fishing in his discussions of their 
archeology. Explorers who visited the Kansa Indians in the 1800's 
such as Thomas Say (Thwaites, 1905c), Frederick Chouteau 
(Adams, 1904), and Father Pierre Jean DeSmet (Thwaites, 1906a), 
although discussing foods of the tribe, did not mention the use of 
fishes. 

Reports of their use by white pioneers are equally rare, although 
Carey (1954:89) quoted a letter of George O. Willard from Juniata, 
on the Big Blue River a few miles north of present Manhattan, Riley 
County, Kansas: "The river is filled with fish weighing from one 
to one hundred pounds. I ate a portion of one caught in the 
Kansas, which weighed 76 pounds." Similar accounts of large 
fishes in the rivers of the lower part of the basin in the 1800's have 
been given by Snow (1875) and Dyche (1914). 

The condition of streams in the Kansas River Basin prior to the 
time of the Louisiana Purchase is virtually unknown. Wedel 
(1941:24-27) inferred from archeological and sedimentational ob- 
servations that droughts, some sufficient to depopulate the western 
plains, had occurred repeatedly in prehistoric times. Among the 
earliest recorded observations are those of Lewis and Clark 
(Thwaites, 1905b: 35) as they passed the mouth of the Kansas River 
during their explorations of the newly-acquired Louisiana Purchase. 
The first explorer to leave a fairly detailed journal concerning parts 
of the Kansas River Basin was Zebulon Pike, who twice crossed 
the Smoky Hill, Saline and Solomon rivers while traveling from 
southern Kansas to and from a Pawnee Indian village on the Re- 
publican River. In 1819 and 1820 a party of exploration under the 
command of Major S. H. Long traveled across the Great Plains to 
the Rocky Mountains and back. Accompanying Long on this ex- 
pedition was Thomas Say, noted American naturalist. Say was 
a member of a detachment that left the main party to explore the 
lower Kansas River Valley as far as its confluence with the Big 
Blue River Valley. Frederick Chouteau was a trader with the 
Kansa Indians from about 1829 to 1853. During part of this 
period, he plied up and down the Kansas River between a trading 



50 University of Kansas Publs., Mus. Nat. Hist. 

outpost near Mission Creek in present Shawnee County and a set- 
tlement near the mouth of the Kansas River. His reminiscences 
have been recorded by Adams (1904). Maximihan, Count of Wied- 
Neuwied, a rhenish German interested in natural history, par- 
ticipated in a voyage up the Missouri River in 1833. In 1842 and 
1843 Col. John C. Fremont traversed parts of the Kansas River 
Basin while carrying out his famed explorations of the West. 

Lower Mainstream 

These explorers and observers recorded a number of impressions 
about the lower mainstream of the Kansas River. Concerning width, 
Fremont (Smucker, 1856:74) found the river near present Topeka, 
Kansas, on June 14, 1842, to be swollen from recent rains and ap- 
proximately 230 yards wide. Joel Palmer (Thwaites, 1906b: 38) also 
found the river at Topeka swollen and "about two hundred and fifty 
yards in width." 

Observations regarding current and depth of water indicate con- 
siderable temporal variation, tlien as now. Lewis (Thwaites, 
1905b: 35) noted tliat the current was "gentle," whereas Fremont 
found the river at the date and place noted above to be "sweeping 
with an angry current." Clark (Thwaites, 1905a:385) noted that his 
party on September 15, 1806 ". . . passed the enterance of the 
Kanzas river which was very low." Maximilian (Thwaites, 1905d: 
251 ) also found the river to be "very shallow" at its mouth in April, 
1833. Chouteau (Adams, 1904:428) indicated extreme fluctuation 
in depth of the Kansas River in the 1830's: "Going down it some- 
times took a good many days, as it did going up, on account of low 
water. I have taken a month to go down from my trading-house at 
American Chief ( or Mission ) creek, many times lightening the boat 
with skiffs; other times going down in a day." The crafts in which 
Chouteau made these journeys were pirogues — boats sixty or 
seventy feet long and about four feet wide, two of which were 
lashed together; they were capable of carrying 10 to 15 tons (Adams, 
1904:428). Thomas Says impression (Thwaites, 1905c: 185) of the 
Kansas River near present-day Topeka in August, 1819, was of a 
river "so shoal as at almost any point to admit of being forded 
without difficulty." Several explorers, however, found the river too 
swollen with floodwaters to be forded when observed by them. 
Mention has been made above that Fremont found the river high at 
Topeka in June, 1842. As far upstream as the confluence of the 



Fishes of Kansas River System 51 

Republican and Kansas rivers, Fremont on June 8, 1843, found it 
impossible to ford the river (Smucker, 1856:193). In July, 1844, 
wliile camped along the Smoky Hill River (Smucker, 1856:490) a 
heavy rainstorm caused rapid flooding of the Fremont campsite, 
ruining many of the perishable collections of the party. 

Turbidity seems also to have been variable in the lower main- 
stream before settlement of the watershed. Lewis wrote (Thwaites, 
1905b: 35), probably from second-hand information since the party 
did not ascend the Kansas River, tliat in summer and autumn the 
waters of the Kansas River were transparent. Long (Thwaites, 
1905c: 173) noted that the Kansas was less turbid than the Missouri 
except at times of high floods. Maximilian (Thwaites, 1905d:251) 
described efl^uent from Kansas River: "Its clear green water was 
distinguished by a well-defined, undulating line, from the muddy 
stream of the Missouri." However, Fremont observed (Smucker, 
1856:74) on June 14, 1842, that the river was "yellow and turbid as 
the Missouri." 

Lewis (Thwaites, 1905b:35) wrote that the bed of the river was 
"composed of soft loam, gravel and sand." More detailed descrip- 
tions of the bottom were recorded in the survey of Lt. Joseph L. 
Tidball, summarized below. 

Concerning banks and streamside flora of the lower mainstream, 
Thomas Say (Thwaites, 1905c: 185) noted that "Willow islands, 
moving sand-bars, and falling-in banks, are as frequent as in the 
Missouri. The Hne of forest which skirts the banks, including the 
bed of the river, is about half a mile wide, but not entirely unin- 
terrupted. The course of the river is remarkably serpentine, forming 
woodland points alternately on both sides." Major Long wrote 
(Thwaites, 1905c: 173,174), "Its valley, like that of the Missouri, has 
a deep and fertile soil, bearing similar forests of cottonwood, syca- 
more, &c., interspersed with meadows; but, in ascending, trees 
become more and more scattered, and at length disappear almost 
entirely, the country, at its sources, being one immense prairie." 
Fitch and McGregor (1956) in an examination of early records 
concerning forested habitat in the area of Lawrence, wrote ( p. 126 ) 
"In the Kansas River flood plain and small tributary valleys, rich 
mesophytic forest of predominantly oak-hickory type was present." 

Most informative of the early accounts concerning the lower 
mainstream is that of Lt. Joseph L. Tidball, who, in August, 1853, 
made a survey of the Kansas River from Ft. Riley, Geary County, 
to its mouth, in order to determine the navigabihty of the river. 



52 University of Kansas Publs., Mus. Nat. Hist. 

He found the water too turbid to ascertain depth visually and 

made recourse continually to a sounding rod. He stated (Langs- 

dorf, 1950:149): 

"This process, though vexatious and wearisome, was attended with the 
advantage of giving a more accurate knowledge of the general depth of the 
water tlian could have resulted from less frequent soundings. This system of 
soundings showed the general depth of water in the main channel, for a 
distance of fifty miles, or thereabout, [downstream from Ft. Riley] to be 
from two to seven feet; that is, it varied between these limits, more frequently 
exceeding the greater than falling below tlie less . . ." 

Tidball noted all areas in which the depth averaged less than 
two feet and found nine such areas between Ft. Riley and the 
mouth of the Blue River. Along this part of the river he found 
the average width to be approximately 80 yards, occasionally 
widening to 120 or more yards. Between the mouth of the Blue 
River, which was observed to be 60-80 yards wide and 2.5-4 feet 
deep, and the mouth of Soldier Creek the river widened. Between 
Soldier Creek and the mouth of the Kansas River the stream again 
became narrower. Tidball frequently mentions the presence of 
large sandbars; for instance, approximately 12 miles above the mouth 
of the Vermillion River he observed (Langsdorf, 1950:151): 

". . . series of little bars, disposed like ribs across the channel, wdth 
not more than eight inches of water on some of them, while below and between 
them it was not unfrequently six or seven feet deep." 

Tidball mentions another area below the present location of St. 
Marys in which shoals as shallow as 12 inches prevailed for ap- 
proximately one mile. Regarding the bottom, Tidball wrote, 
(Langsdorf, 1950:154): 

"Except in a few places to which allusion has been made, at the rapids 
and in their vicinity, the bed of the river is an easily yielding quicksand, and 
its surface broken." 

In regard to the banks, he noted (Langsdorf, 1950:155): 

"The banks of the upper portion of the river are fonned almost entirely 
of sand, occasionally mingled with clay. Lower down, this is seen in some- 
what greater abundance, sometimes in thin strata alternating with sand; oc- 
casional beds of gravel and in a few places, for short distances, rocky develop- 
ments occur." 

Tidball surmised that he descended the river at a time of 'low 
water." He found evidence and received testimony from stream- 
side residents that the river-level had been 5 to 8 feet higher earlier 
in the same year. He formed the opinion that high water in spring 
and early summer was sufficiently characteristic of the stream to 
allow navigation during that part of the year. 



Fishes of Kansas River System 53 

Tributaries 

References to tributaries of the Kansas River before settlement 
are few. 

Thomas Say (Thwaites, 1905c: 186) crossed the Vermillion River 
in present Pottawatomie County, Kansas, remarking that it was 
four feet deep and approximately twenty yards wide. Concerning 
Blue River a mile or two above its mouth, Say (Thwaites, 1905c: 
200) reported a width of 25 yards and a maximum depth of three 
feet. Fremont (June 20, 1842) found the lower Blue River to be 
(Smucker, 1856:79), "... a clear and handsome stream, about 
one hundred and twenty feet wide, running with a rapid current 
. . ." Proceeding northwestward in the Blue River Basin, Fre- 
mont described some creeks as "handsome" with clear water and 
sandy beds, and mentioned others that were dry. After encounter- 
ing several of the latter, Fremont wrote (Smucker, 1856:80), 
". . . after a hard day's march of twenty-eight miles, encamped, 
at 5 o'clock, on the Little Blue, where our arrival made a scene of 
the Arabian Desert. As fast as tiiey arrived, men and horses rushed 
into the stream, where they bathed and drank together in common 
enjoyment." Fremont found Little Blue River at this place to be 
approximately 50 feet wide and three to four feet deep. 

Zebulon Pike traversed central Kansas in September and October, 
1806, and noted (Hart and Hulbert, 1932:73-76) various tributaries 
that were dry. Pike judged (p. 73 ) Smoky Hill River near present- 
day Bridgeport, Saline County, Kansas, to be 25 or 30 yards wide, 
and navigable in flood seasons. Returning east from his Expedition 
of 1843-44 in July, 1844, Fremont descended Smoky Hill River, 
describing the journey as follows (Smucker, 1856:489): 

"On the evening of the 8th we encamped in a cottonwood grove on the 
banks of a sandy stream-bed, where there was water in holes suflBcient for the 
camp. Here several hollows, or dry creeks with sandy beds, met together, 
forming the head of a stream which afterwards proved to be the Smoky Hill 
fork of the Kansas River. 

"As we traveled down the valley, water gathered rapidly in the sandy bed 
from many little tributaries; and at evening it had become a handsome stream, 
fifty to eighty feet in width, with a lively current in small channels, the water 
being principally dispersed among quicksands. 

"Gradually enlarging, in a few days' march it became a river eighty yards 
in breadth, wooded with occasional groves of cottonwood." 

Previously enroute westward, Fremont found (Smucker, 1856: 
196) Prairie Dog Creek to be ". . . forty feet wide and four 
feet deep" on June 23, 1843. On June 26, 1843, approximately 80 



54 University of Kansas Publs., Mus. Nat. Hist. 

miles farther west, Fremont came to the South Fork of Repubhcan 
River (Smucker, 1856:196) : ". . . whose shallow waters, with a 
depth of only a few inches, were spread out over a bed of yellowish 
white sand 600 yards wide." 

Regarding the appearance of valleys of tributaries, Say remarked 
(Thawites, 1905c: 200) that along the Big Blue River near present- 
day Manhattan, Kansas, "The soil supports but a thin growth of 
grass, and the timber is far from abundant, consisting principally of 
different sorts of oak, confined to the margin of the creek, its ravines 
and tributaries." Of the Little Blue River Fremont wrote (Smucker, 
1856:81), "Our route the next morning lay up the valley, which, 
bordered by hills with graceful slopes, looked uncommonly green 
and beautiful. The stream . . . fringed by cotton-wood and 
willow, with frequent groves of oak, tenanted by flocks of turkeys." 
Fremont found Prairie Dog Creek (Smucker, 1956:196) to have "a 
dense border of wood, consisting principally of varieties of ash" but 
approximately 100 miles farther west on the Republican River (pp. 
196, 197 ) : "With the exception of one or two distant and detached 
groves, no timber of any kind was to be seen; and the features of 
the country assumed a desert character, with which the broad river, 
struggling for existence among the quicksands along the treeless 
banks, was strikingly in keeping." None of these reports of condi- 
tions in the tributaries prior to settlement mention silty bottoms 
although several mention sand or "quicksand." 

In October and November, 1869, exploration pursuant to planning 

a railroad route was undertaken in the Solomon River Valley by a 

party under the leadership of Mr. Robert McBratney. Entries from 

McBratney's diary have been published by Caldwell (1937). In 

reading these comments it should be borne in mind that McBratney 

was interested in promoting settlement of the Solomon Valley. On 

October 21, 1869, McBratney noted (Caldwell, 1937:64, 65): 

"Crossed the S. B. of S. [South Fork of the Solomon] and struck out for the 
Waconda or great spirit spring of the Indians, which is about four miles below 
the forks. Three miles below camp crossed the main Sol. and have thus crossed 
aU branches. Find about the same amt. of water in all, that is a stream about 
50 ft. wide and from two to three feet deep, at this its dryest. Water clear 
and pure, & excellent for drinking." 

On this same date McBratney explored several tributaries of the 
Solomon near Waconda Springs (Mitchell County), making the 
following notations (Caldwell, 1937:65, 66): 



Fishes of Kansas Ri\'er System 55 

Carr Creek — "Water in creek stopped by Beaver dams. Cr. well timbered 
with hard timber, and growing better up. Width of bottom ii to Yi mile, water 
good." 

Unnamed Creek — "Examined Cr. for six to seven mile. Running water." 
Oak Creek — ". . . examined Oak Cr. for seven to 9 miles. Find it 
nearly as large as a Br. of Solomon, with bottom VA miles. Well wooded with 
hard wood. . . . Water of Cr. running, but not strong." 

Farther upstream on the North Fork of the Solomon in the vicinity 
of the present town of Kirwin, McBratney noted ( Caldwell, 1937:69 ) 
"The water of die Solomon and its tributary is clear, pure and 
hard. . . ." Returning from the expedition, the South Fork of 
the Solomon in present Osborne County was found to be ( Caldwell, 
1937:73) a ". . . veiy much larger stream than the N. B. North 
Fork of the Solomon & running three times as much water." 

Settlement of the western part of the Kansas River Basin lagged 
greatly behind that of the eastern part. This was due partly to 
hostihty of Indian tribes of the area, chiefly the Cheyennes. Holding 
an important place in the history of the area was Fort Wallace, 
which was an active estabhshment until 1882 (Montgomery, 1928: 
278). In 1870 the acting assistant surgeon at the post, M. M. 
Shearer, wrote, as reported by Montgomery (1928:242), "At this 
point the Smoky Hill is a comparatively beautiful stream of running 
\\'ater, due to one of its tiibutaries, which joins it about three miles 
west of the fort; beyond that point the river consists of occasional 
ponds with intervening sand beds, through which the water takes a 
subterranean course." Prof. Edward Drinker Cope sojourned at 
Fort Wallace while collecting in the Cretaceous beds of western 
Kansas. In September, 1871, he wrote (Osborn, 1931:163), "We 
found a high bluflF near the Twin Buttes which we determined to 
search. We found Butte Creek totally dry and had to go four 
miles further and camp by a spring in a narrow ravine." The stream 
mentioned by Shearer may have been Rose Creek, which is still a 
permanent stream unhke the Smoky Hill River. The "Butte Creek" 
or Twin Butte Creek mentioned by Cope was obser\'ed by me to be 
dry at several places in September, 1961, and April, 1962. 

Summary 

The accounts cited above and others suggest that the Kansas 
River and its larger tributaries have long been subject to fluctuation 
in amount of discharge. There seems also to have been considerable 
fluctuation in the past, as now, in the degree of turbidity, especially 

3—8449 



56 University of Kansas Publs., Mus. Nat. Hist. 

in the larger streams. That the Kansas River was not, in the middle 
1800's, consistently a deep, clear stream is well-documented by Tid- 
ball's navigability survey (Langsdorf, 1950) and by numerous in- 
stances reported by Greene (1906) of steamboats going aground on 
sandbars during the period 1854-1866. Even for small boats such as 
the pirogues used by Chouteau (Adams, 1904: 428) or the grain boat 
employed by James R. Mead (Mead, 1906:9) water was often too 
shallow for navigation on the lower Kansas River. 

That all upland tributaries were not permanently flowing streams 
is attested by early observers such as Pike (Hart and Hulbert, 
1932:73,76), Fremont (Smucker, 1856:489), Shearer (Montgom- 
ery, 1928:242), Cope (Osborn, 1931:163) and Mead (1906:15). 
There is some evidence, however, that streams in areas now ex- 
tensively tilled were more nearly permanent and were clearer than 
at present. McBratney's (Caldwell, 1937) enthusiastic accounts of 
the clear, permanent tributaries of the Solomon suggest this, as 
do other descriptions quoted. 

KANSAS RIVER SYSTEM AND ITS FAUNA 
SUBSEQUENT TO SETTLEMENT 

A simplified categorization of man's effects on the rivers of the 
Kansas River Basin might include tliat which is (1) taken from 
and (2) added to the streams. 

The first category is probably the less deleterious in regard to 
fishes of the Basin. Water is taken directly from the rivers and 
indirectly from groundwater supplies along the river for domestic 
and industrial use and for irrigation. 

Substances added directly or indirectly to streams as a result of 
man's activities include (1) domestic and industrial wastes, and 
(2) silts and other sediments that enter in increased amounts be- 
cause of modification of the land. The Kansas State Board of 
Health, in cooperation with municipal and industrial organizations, 
has carried on an effective program that has ameliorated or re- 
moved the worst aspects of the domestic and industrial pollution 
problem. Colby et al (1956:64) noted that "Throughout most of 
the Basin . . . municipal and industrial pollution has been to 
a large extent controlled. Only in the Kansas City area do serious 
problems still exist" where ". . . the last four miles of the 
Kaw's course . . . approximates an open sewer." In the cen- 
tral part of the Basin, oil-field brine was formerly led into streams. 
As some rivers in the same area ( for example the Saline and Smoky 



Fishes of Kansas River System 57 

Hill rivers) were naturally highly saline, a condition inimical both 
to man and to some aquatic organisms was created. At present, 
however, most brine is reinjected into deep-lying rock strata. Al- 
though parts of the Kansas River System may long have been at 
least periodically turbid, intensive cultivation of the land in many 
parts of the Basin seems to have increased turbidity and siltation 
of streams. 

The highly variable nature of precipitation patterns in the Kansas 
River Basin with recurrent droughts is interrelated with the condi- 
tions discussed above. In hot, dry years more water is needed by 
tlie public at the same time that streamflow is minimal; increasing 
population in the Basin intensifies this problem. In the drought 
year, 1954, according to Colby et al (1956:61), sixty-one municipal 
water supply systems in that part of the Basin in Kansas were in- 
adequate in providing necessary amounts of water. Furthermore, 
reduced flow may decrease the ability of streams to transport wastes 
and sediments and to recover from pollutants. 

In July, 1885, O. P. Hay traveled in northwestern Kansas, collect- 
ing fishes at several localities. His published account (1887) con- 
tains several references to stream conditions at that time. Of a 
small tributary of the Solomon River immediately west of Beloit, 
Mitchell County, Kansas, Hay wrote (1887:242, 243) "This stream, 
like all others in this part of the State, flows in a deep ravine, is very 
sluggish, and so filled with ooze as to make the dragging of a seine 
a very diflBcult and disagreeable task." Along the North Fork 
Solomon River at Kirwin, Phillips County, Kansas, Hay (1887:247) 
noted, "The collection obtained at this point was made in a muddy, 
ooTX, stream. . . . Only nine species were secured as the fruits 
of a day's hard work." This is the same Solomon Valley described 
by McBratney (Caldwell, 1937 — discussed heretofore), who had ex- 
plored it only 16 years before the collections were made by Hay. 
At the time of McBratney's exploration there were virtually no white 
settlements in the valley and McBratney's party was accompanied 
by troops to fend oflF possible attack by Indians. It would seem 
that in this short period of time some siltation of stream bottoms had 
begun, if one interprets Hay's "muddy ooze" as evidence of erosion 
due to agriculture. 

Farther west, however, Hay did not mention siltation, describing 
the North Fork Solomon River at Lenora, Norton County, Kansas 
(1887:248) as ". . . small, and in most places a shallow, stream, 
flowing with sufficient rapidity to carry away the finer materials and 



58 University of Kansas Publs., Mus. Nat. Hist. 

leave for itself a clean bed of sand." Likewise the Saline River, 5 
miles north of WaKeeney, Trego County, Kansas, was found (1887: 
250) to be a ". . . shallow, rather rapid prairie brook, with a 
clean, sandy bed. Its depth, when visited varied from a few inches 
to 3 or 4 feet." Hay found the Smoky Hill River near Wallace, 
Wallace County, Kansas (1887:251), to be ". . . much like the 
Saline at WaKeeney, but smaller and more rapid." 

Hay's observations suggest that siltation was notable only in 
eastern parts of the streams of northwestern Kansas. Today siltation 
seems to have progressed farther upstream. Hay's description of the 
North Fork Solomon River at Kirwin and the fauna he obtained 
there approximate the situation found today at Lenora, 50 miles to 
tlie west. 

My observations along North Fork Solomon River in June, 1963, 
are recorded below (west to east, mileage approximate and Hnear, 
not adjusted for irregularities in the course of the river) : 

Mile one. Ten miles north of Hoxie, Sheridan Co., Kansas. Stream com- 
pletely dry. Probably the stream west of this place was dry to its source, 
a distance, linearly, of approximately 50 miles. 

Mile ten. Tluee miles SW AUison, Sheridan Co. Upland brook, up to 15 
feet wide with clear water (Secchi disc reading, approximately 3 feet). 
Bottom of clean sand. Fringe forest of ehn, ash, cottonwood and willow. 
Water temperature 71° F. at 11:30 a.m. Beaver dams ponding stream. 
Water here is obtained from springs along the contact zone of Pliocene 
and Cretaceous rocks. 

Mile 30. Lenora, Norton Co. Creek up to 25 feet wide. Water turbid 
( Secchi disc reading 5 inches ) . Bottom of sand and silt. Fringe forest 
present. Water temperature 81° F. at 2:30 p. m. 

Mile 55. Three miles east of Logan, PhilHps Co. River up to 100 feet vnde, 
shallow, braided. Bottom of sand and tenacious silt. Water very turbid 
(Secchi disc reading 3 inches). Water temperature 87° F. at 4:00 p. m. 

In addition to and probably in conjunction with the changes ob- 
served in the character of the North Fork Solomon River, there seem 
to have been changes in the nature of the fish fauna. As indicated 
in Table 4, Hay took 16 species at Lenora in 1885, whereas the two 
collections of recent years taUied only seven and eight species. 
Etheostoma spectabile, taken by Hay at Lenora, seemingly no longer 
occurs there, but a population still exists 20 miles upstream in the 
clearer, colder headwaters at "Mile 10" mentioned above. 

In Table 5 the faunal list of Hay's collection from Wallace, 
Wallace County, is compared with lists made at that locality in the 
past 25 years. 

Species taken by Hay, but not subsequently at any of his stations, 
are Lepisostem osseus, Htjbopsis higuttata, Chrosomus erythro- 
gaster, Notropis topeka, Moxostoma macrolepidotum and Etheo- 



. 



Fishes of Kansas River System 



59 



Table 4. — Species of Fishes Taken at Lenora, Norton County, Kansas, 
ON Three Occasions (USNM Nijmbers Given Where Available). 





Hay 

1885 


F. B. Cross 
1958 


Metcalf 
1963 


Semotilus atromaculains 

Chrosomns erythrogaster 

Phenacobius mirabilis 


(USNM 38233) 
(USNM 38241) 
(USNM 37962) 
(USNM 38232) 

X 

X 
(USNM 37950) 

(USNM 38240) 

(USNM 37939) 

(USNM 38236) 

X 

(USNM 37963) 

X 

X 

X 
(USNM 37951) 


X 
X 

X 
X 
X 
X 

X 

X 


X 


Noiropis umhratilis 




Notropis cornutus 




Notropis lutre7isis 


X 


Notropis stramineus 

Hyhognathns placihis 


X 
X 


Pimephales promelas 


X 


Pimephales notatus 




Campostoma anomahim 

Catostotnus cominersoni 


X 


Ictalurus melas 


X 


Noturus flavus 




Fundulus kansae 




Lepomis cyanellus 

Etheostoma nigrum 

Etheostoma spectabile 








Total 


16 


8 


7 







Stoma n. nigrum (and, indirectly, Notropis heterolepis if, as inter- 
preted by Hubbs, 1951a, it was one of the parental species of the 
hybrid, "Notropis germanus" Hay). Species taken by others in 
subsequent years but not by Hay include only Ictalurus natalis and 
Cijprinus carpio. Hay's "Ictiohus velifer" was probably Carpiodes 
c. carpio. The only carpsucker taken by Hay that I have been able 
to locate (USNM 37936 from Beloit) is Carpiodes c. carpio. 

Considering the records of Hay and other early records in light 
of present distribution of fishes, at least the following species seem 
to have suffered depletion in numbers, restriction of range, or ex- 
tirpation in the Kansas River Basin in historic time: ( * — doubt 
exists as to native occurrence in the Basin). 



Ichthyomyzon castaneus 
Hiodon alosoides 
Hyhopsis biguttata 
Chrosomus erythrogaster 
Notropis cornutus 
Notropis h. hudsonius ' 
Notropis heterolepis " 



Cycleptus elongatus 
Carpiodes velifer 
Percopsis omiscomaycus " 
Stizostedion v. vitreum * 
Percina maculata 
Etheostoma blennioides * 



Etheostoma n. nigrum 
Notropis topeka 

M Further evidence of changes that have occurred in the aquatic 
fauna of the Kansas River Basin since settlement is provided by 
certain gastropods. Aughey (1877) hsted three species of pleu- 



60 



University of Kansas Publs., Mus. Nat. Hist. 



Table 5. — Fishes Taken at Two Localities in Wallace County, Kansas. 
All Collections Contain Fish From the Smoky Hill River Near Wal- 
lace. In Addition, the Collections of Cross, Taylor and Metcalf Con- 
tain Fishes From Nearby Rose Creek. USNM Numbers Are Given Where 

Available. 



Species 


Hay 
1885 


Taylor 
July, 
1950 


Cross 

June, 

1958 


Metcalf 

August, 

1961 


Cyprinus carpio 


X 
X 
X 
X 

(USNM 37947) 
X 

(USNM 37934) 
(USNM 37949) 
(USNM 38237) 

X 

X 

X 
(USNM 37935) 

X 
X 

(USNM 37957) 

(USNM 37953) 

(USNM 37937) 
(USNM 37944) 


X 
X 

X 
X 
X 

X 

X 

X 
X 
X 

X 
X 

X 


X 
X 

X 
X 
X 
X 

X 
X 
X 

X 

X 

X 
X 
X 

X 
X 


X 


Semotilus atromaculatus .... 

Hyhopsis biguttata 

Phenacobnis mirabilis 

Notropis cornutus 


X 
X 


Notropis lidrensis 


X 


Notropis stramineus 

Notropis topeka 

"Notropis germanus Hay" . . 
Hybognathxis hankinsoni .... 

Hybognathus placitus 

Pimephales promelas 

Campostoma anomalum 

Catostomus commersoni 

Ictalurus natalis 


X 

X 
X 
X 
X 


Ictalurus nielas 


X 


Noturus flavus 


X 


Fundulus kansae 


X 


Micropterus salmoides 

Lepomis cyanellus 


X 


Lepomis: cyanellus x 

ynacrochirus 

Lepornis humilis 

Etheostoma spectabile 


X 
X 


Total 


19 


13 


16 


15 







rocerid snails from the Blue River in Nebraska. According to 
Goodrich (1939:4) one of these snails was probably Pleurocera 
acuta Rafinesque; the other two seemingly were misidentifications. 
Probably, then, at least one species of pleurocerid gastropod 
formerly inhabited the Blue River System, altliough, to my knowl- 
edge, no pleurocerids have been reported anywhere in the Kansas 
River Basin in this centurv. Pleurocera acuta, seems to be intolerant 
of the sUtation of streams that has accompanied cultivation of the 
land (Leonard, 1959:41). I have noted during the past seven years 
the extirpation of what may be one of the last colonies of this 
snail in the Kansas part of the Marais des Cygnes River directly 
south of the Kansas River. This colony seemed to be flourishing 
in 1956 in an area of gravelly rijBBes three miles east of Ottawa, 
Kansas. In 1958 and subsequent years, however, I was unable to 
find any living specimens, although dead shells remained. During 



Fishes of Kansas Rn'ER System 61 

this period channel-straightening, dike construction, clearing of trees 
from banks, and other modifications took place due to a program 
of flood control upstream and to construction of a divided highway 
across the area. 

In addition Aughey (1877:701) reported viviparid snails from 
the Blue River. It seems probable that these were of the genus 
Campeloma. Call (1885:52; 1886:184; 1887:17) and Hanna (1909: 
96) also ascribed Campeloma to the Blue, Delaware, Wakarusa and 
Kansas rivers and to Soldier Creek. However, Branson (1963:73) 
suggested that a few streams in extreme soutlieastem Kansas 
". . . may be the last ecological situations available for such 
clear-water species in the state." Aughey (1877:702) and Call 
(1885:52) reported the genus Amnicola from a number of localities 
in the Kansas River Basin. At the present time this genus seems 
rare in Kansas (Leonard, 1959: Fig. 20). 

The present disjunct distribution and the existence of older 
records from intervening areas suggest that the crayfish Orconectes 
neglectus Faxon was once more widespread in the cential and 
western part of the Kansas River Basin than at present (Williams, 
1954:573; Metcalf and Distler, 1961:355). 

It seems probable that the presence or absence of beavers, espe- 
cially in tributaries of the western part of tlie Basin, may bear 
some relation to the kinds and numbers of fishes present. Following 
is a statement by James R. Mead (1906:15), one of the first settlers 
in the Saline River Valley and namer of many of the tributaries of 
the Saline River, concerning conditions in 1859: 

"We crossed the Saline below some salt springs from which the river derives 
its saline properties, and traveled north. We soon found a large, dry, sandy 
creek coming from the hills in the distance; following this up we came to 
beaver dams and water. The beaver held back all the water in the dr>- 
season. Further along were plenty of buflFalo, and where the stream came 
out of the bufiFs were groves of beautiful oak timber. The canyons were full 
of large cedars and no sign of an ax or of white man's presence in any of it. 
I had found a stream unknown. 

As we drove into this beautiful spot I exclaimed, 'Boys, we have got into 
paradise at last!' — and that name it bears to this day, and the town of Paradise 
is near the spot of our first camp." 

The town of Paradise is in northwestern Russell County, and 
Paradise Creek, which flows nearby, again supports colonies of 
beaver according to Henderson (1960:79). Beaver were exceed- 
ingly rare in Kansas (Henderson, 1960) at least from 1900 to 1920 
and probably for an even longer time. By 1930 populations were 
slowly increasing, especially in the northwestern part of the state. 



62 University of Kansas Publs., Mus. Nat. Hist. 

Henderson (1960:14) suggested that a higher population per suit- 
able stream mile may exist in the western than in the eastern part 
of the Kansas River Basin due to the greater availability of cotton- 
wood and willow, preferred food of the beaver, along these western 
streams. In 1961, 1962, and 1963, I observed many beaver dams 
along the tributaries of the Kansas River in northwestern Kansas 
( as at "Mile ten" on North Fork Solomon River, mentioned above ) . 
These dams seem to contribute to ponding and conservation of 
water in these small, shallow streams of low gradient, which other- 
wise possess few deep pools. Henderson (1960:27,28) pointed out 
that beaver impoundments provide excellent habitat for fishes on 
these small western streams. Probably repopulation by beavers 
on these streams has enhanced piscine life in general and been 
especially beneficial for those species that are less tolerant of 
adverse conditions. Hall (1946:484) pointed out the beneficial 
effects of ponding by beavers on aquatic and streamside life in 
Nevada, and Hanson and Campbell (1963:148) found in north- 
central Missouri that "Beaver activity appeared to improve the 
carrying capacity of the stream for warm-water fishes and to pro- 
vide conditions suitable for a large variety of fishes. Thus their 
activity helped oflFset the deleterious effects of floods and of man's 
activities, especially in the headwaters." Possibly the more wide- 
spread occurrence of fishes such as Etheostoma spectabile pulchel- 
lum and Noturus fiavus in streams of northwestern Kansas in collec- 
tions made between 1961 and 1963 than in tliose made by John 
Breukelman in 1938 (Breukelman, 1940a) reflects improved stream 
conditions partly brought about by increased ponding by beavers. 
A recent and more striking kind of ponding has resulted from 
man's construction of large reservoirs, especially in the western part 
of the Basin. The principal impoundments completed or under con- 
struction are as follows: 

1. Bonny Yuma County, South Fork Republican River 

Colorado 

2. Enders Chase County, Frenchman Creek 

Nebraska 

3. Trenton Hitchcock County, Republican River 

Nebraska 

4. Harry Strunk . . . Frontier County, Medicine Creek 

Nebraska 

5. Norton Norton County, Prairie Dog Creek 

Kansas 

6. Harlan County . Harlan County, Republican River 

Nebraska 



Fishes of Kansas River System 63 

7. Cedar Bluff .... Trego County, Smoky Hill River 

Kansas 

8. Webster ....... Rooks County, South Fork Solomon River 

Kansas 

9. Kirwin Phillips County, North Fork Solomon River 

Kansas 

10. Wilson Russell County, Saline River 

Kansas 

11. Glen Elder Mitchell County, Solomon River 

Kansas 

12. Lovewell Jewell County, White Rock Creek 

Kansas 

13. Kanopolis Ellsvv^orth County, Smoky Hill River 

Kansas 

14. Milford Geary Coimty, Republican River 

Kansas 

15. Tuttle Creek . . . Riley County, Big Blue River 

Kansas 

16. Perry Jefferson County, - Delavi'are River 

Kansas 

As concerns fishes in the Basin, the stabilization of flow and filter- 
ing effect with consequent clarification of water afforded by ponding 
of streams are surely of importance, although the effect on distribu- 
tion of fishes below and above reservoirs of the Basin has not been 
elucidated. A more easily observed effect has been the introduction 
of both native and non-native fishes into impoundments and streams 
of the Basin. 

Graham (1885a: 78) hsted 13 species that had been "planted in 
the waters of Kansas." Of these, six (Salmo gairdneri, Esox lucitis, 
Cyprinus carpio, Pomoxis annularis, Stizostedion v. vitreum and 
Perca flavescens) occur in streams and impoundments in the Basin 
at present. Additional species, deemed non-native, that have been 
placed in waters of the Basin by man are Salmo truita, Carassius 
auratus, Ictalurus nehtdosus, Micropterus dolomieui and Micropterus 
punctulatiis. Evidence is equivocal as regards native occurrence of 
Fundulus sciadicus, Chaenohnjttus gulostis, Ambloplites nipestris 
and Pomoxis nigromaculatiis. Following is a Hst, kindly supplied by 
Mr. Roy Schoonover, Chief, Fisheries Division, Kansas Forestry, 
Fish and Game Commission, of species that have been stocked in 
reservoirs of the Kansas River Basin in Kansas by the Forestry, Fish 
and Game Commission: Esox lucius, Ictalurus punctatus, Ictalurus 
melas, Roccus chrysops, Micropterus dolomieui, Micropterus 
salmoides, Micropterus punctulatus, Lepomis macrochirus, Pomoxis 
nigromacuhtus, Pomoxis annularis, Stizostedion v. vitreum, Perca 
flavescens and Aplodinotus grunniens. 



64 University of Kansas Publs., Mus. Nat. Hist. 

HISTORY OF THE HIGH PLAINS 
AND OF THE ANCIENT PLAINS DRAINAGE 

Knowledge of the Miocene liistory of the Great Plains is not 
great. Chaney and Elias ( 1936:26) noted that "The floras as known, 
and the associated mammalian remains, point toward climatic 
conditions which favored the development of broad grasslands, 
with forests restricted to the stream borders." 

In the basin of the Kansas River there are widespread deposits of 
PHocene and Pleistocene age that allow insight into some aspects of 
the history of the Basin. Knowledge of the PHocene and Pleistocene 
history of the region is, in great part, due to the efforts of Maxim K. 
Elias, John C. Frye, A. Byron Leonard, Ada Swineford, Claude 
Hibbard and tlieir associates. Lugn (1935) presented an important 
summary of the Pleistocene History of Nebraska and Condra and 
Reed (1950) wrote concerning the correlation of tlie Pleistocene 
deposits of Nebraska. 

As underlying Miocene or earlier Tertiary sediments are absent 
throughout most of the central and southern Great Plains, and 
Pliocene deposits are widespread, it seems probable that a regimen 
marked by erosion may have been gradually replaced by one of 
deposition at the beginning of Pliocene time. 

The lowermost or Valentine Member of the Ogallala Formation of 
Phocene age occupies the lower portions of west-east trending 
valleys cut in Cretaceous bedrock by previous erosional activity 
(Merriam and Frye, 1954:57), Frye, Leonard and Swineford ( 1956: 
56, 61) noted that during the succeeding Ash Hollow and Kimball 
time, alluvium continued to be laid down in the pre-existing valleys 
until it began to overlap the sides of the valleys and to coalesce. 
These authors believed that the upper part of the Ash Hollow FoiTna- 
tion (p. 56) "transgressed most, if not all, former divides and formed 
a virtually continuous coalescent sheet of alluvial material through- 
out northwestern Kansas and soudiwestern Nebraska." In regard to 
the Kimball FoiTnation these authors stated (p. 61) "The Kimball 
was the culmination of Ogallala alluviation, and overlapped most if 
not all of the former bedrock divides. Its surface extended as a vast 
coalescent plain of regional alluviation. During Kimball deposition 
the former bedrock valleys were no longer effective in controlling 
the position or direction of flow of major streams, which became free 
to shift in response to load, volume, and gradient." From the above 
descriptions one visualizes late Pliocene streams as shallow, sedi- 
ment-laden, at least periodically, and meandering over a terrain 
with little gradient. 



Fishes of Kansas River System 65 

After filling and transgressing tlie shallow valleys and coalescing, 
the sediments of the Ogallala Formation are thought to have come to 
an "erosional-depositional equilibrium" (Swineford, Leonard and 
Fr\'e, 1958 : 114 ) . These authors suggested that the period of surface 
stability was "long," Nvith a climatic trend towards desiccation that 
resulted in formation of a tliick cahche zone and ultimately in the 
pisolitic or "algal" limestone that occurs at the top of the Ogallala 
Formation. 

Frye, Leonard and Swineford (1956:57) and Bayne and Fent 
(1963:363) found no evidence tliat the east-flowing streams of 
Ogallala time crossed the Flint Hills in Kansas and suggested rather 
that drainage was to the south, through a major valley west of the 
Flint Hills into the area occupied by the present Arkansas River, 
Supporting this view, Fiye (1955:80) pointed out that high terrace 
deposits, predominantly of chert derived from the Flint Hills and 
probably of Pliocene age, are found east of the Flint Hills in Kan- 
sas, although no sediments of the kinds found in the Ogallala 
Formation are found east of the Flint Hills. 

The Flint Hills may have acted as a watershed bai-rier to streams 
from the west due to the dip of their resistant beds to the west. In 
the Flint Hills of southern Kansas today, streams \\ath high gradient 
are eroding from the east against the regional dip of the rocks. One, 
the Cottonwood River, has breached the band of hills. Streams ap- 
proaching the hills from the west, however, have low gradients and 
do not erode up-chp eastward into the Flint Hills, but typically turn 
southward, parallehng the Hne of hills until they circumvent them 
or are captured by a stream, such as the Cottonwood, that has 
breached the hills fi-om the east. 

x\t the end of Pliocene time, then, a master stream (Preglacial 
Plains Stream) is thought to have flowed north to south across the 
present state of Kansas west of the Flint Hills ( Fig. 3 ) . This stream 
seems to have been fed almost entirely by tributaries from the west. 
The extent of this stream system, and directions taken north and 
south of Kansas, are not clearly ascertainable. To the north, Lugn 
(1935:35-37) noted the presence in southern Nebraska, west of the 
FHnt Hills, of three wdde, buried, preglacial valleys or "Basins" that 
trended northwest to southeast. In central Nebraska the preglacial 
topography was higher, suggesting that these streams had their head- 
waters in the general area presently occupied by the Loup rivers. 
Lugn (p. 153) wrote ". . . the waters of the early Pleistocene, 
and before that even, the pre-Pleistocene North Platte River found 
escape southeastward across the inwash-outwash plain of the south 



66 University of Kansas Publs., Mus. Nat. Hist. 

central part of the state, that is through the Holdrege or even tlie 
Hastings basin." Lueninghoener (1947:60) suggested that the Loup 
rivers might be former upper tributaries of a system of which the 
Blue River is now a lower remnant. Sharp turns to the northeast 
where each tributary of Loup River joins the mainstream suggest 
capture from a former southeasterly course such as might have 
existed preglacially. In summary, probably the upper Platte and 
the Loup rivers, but possibly not the upper Niobrara, were once part 
of tlie Preglacial Plains Stream (Fig. 3). Horberg and Anderson 
(1956: Figs. 1 and 2) held a different view and tentatively indicated 
outflow of both Preglacial Platte and Niobrara eastward into the 
Ancestral Mississippi. 

To the south, widespread fluvial deposits along the southern 
boundary of Kansas ( Harper to Sumner counties ) indicate that the 
main outwash-carrying channels of the Ancestral Plains Stream of 
the early Pleistocene (and probably the Preglacial Plains Stream) 
entered Oklahoma in this area; its direction thereafter is unknown. 
Quinn (1958:42) theorized that the Arkansas River first breached an 
Ozark-Ouachita upland and emptied eastward into the Mississippi 
at some time in the Pleistocene prior to Sangamonian time. Prob- 
ably, then, the Preglacial Plains Stream continued at least as far 
soutli as the Ancestral Red River. Faunal evidence (discussed 
below) suggests that drainage at some time had connections with 
the Ancient Red, Brazos and Colorado rivers of Texas. The presence 
of quartzite cobbles in high-level terraces near Dallas, Texas, older 
than Kansan in age (reported by Slaughter et al, 1963:9), suggested 
that drainage in that area once came from the north, crossing the 
area presently occupied by the valley of the Red River. The nearest 
outcrops of igneous rocks are in the Wichita Mountains in Oklahoma 
to the north. However, Frye and Leonard (1963:10-11) in describ- 
ing the Hardeman Alluvial Terrace of Kansan age along the Red 
River east of the High Plains inferred that, by Kansan time, the Red 
River occupied essentially its present valley. Subsequently, Dal- 
quest (1964:351; 1965:74) showed, on the basis of carbon 14 dates, 
that sediments at the type locality and at several other exposures 
of the "Hardeman Alluvial Terrace" were all less than 30,000 years 
in age, hence no older than Wisconsinan. 

Southwestward there is some zoological evidence (Koster, 1957:1) 
that a part of the plains drainage was captured by growth headward 
and northward of the Pecos River and subsequent capture of east- 
ward-flowing streams of the Ancestral Canadian System. 



Fishes of Kansas River System 



67 




Fig. 3. Idealized preglacial drainage patterns in the 
central United States. Dotted lines represent hypo- 
thetical streamcourses; solid Hnes are based upon exist- 
ing geologic information, but are, in part, also hypotheti- 
cal. For sources see text. ( 1 = Drainage of Plains 
Stream System; 2 = Hudson Bay Drainage; 3 = Drain- 
age of Teays/Mississippi System). 

Evidence about the fauna of the Preglacial Plains Drainage ( Fig. 
j 3 ) can be derived from fossils found in beds of Pliocene age in the 
High Plains, and from climatic conditions that are thought to have 
existed then. 

Hibbard (1960:11) pointed out that temperature and precipita- 
tion must be considered separately and not in terms of their present 
relationships in the Great Plains. He inferred, chiefly from paleon- 
tologic evidence, that in lower Pliocene time (p. 11) "The climate 
of the Interior Plains was mild-subtropical as far north and west as 
Shannon County, South Dakota" and that (p. 13-14) the early 
Middle Pliocene climate di£Fered from that of the Lower Pliocene by 
a slight decrease in moisture. Hibbard characterized the Great 
Plains of the Lower Pliocene as ( p. 13 ) ". . . a moist subhumid, 
subtropical, savanna with forests and tall grasses along the river 
valleys, with chiefly shrubs and tall grasses on the valley walls and 
on the low divides. Some short grasses may have occurred on the 
higher and well drained divides." In the Middle Pliocene he noted 
that there is evidence of large rivers having broad flood plains or 



68 Univ'ersity of Kansas Publs., Mus. Nat. Hist. 

valleys with oxbow lakes in which diatomaceous marl was deposited. 
On the basis of fossil plant seeds and vertebrates found in Phocene 
deposits in Beaver County, Oklahoma, Chaney and Elias (1936:34) 
visualized extensive grasslands or savannas in that area, during the 
Pliocene, traversed by streams with wooded valleys. 

Leonard and Frye (1956:31) made a strong case for a gradual 
increase in aridity during the Pliocene of the Great Plains. This 
was substantiated by a decrease in numbers and ultimate disappear- 
ance in the fossil record of branchiate snails, the ascendancy of 
prairie grasses and the decrease of arboreal elements. In addition 
there were periods of caliche formation and a fining of sediments in 
the Late Pliocene, all of which suggested, according to these authors, 
a lowering of the water table. 

A list of Pliocene fishes known from the High Plains has recently 
been compiled, augmented and discused by C. L. Smith (1962); 
only two of the species listed (Ictahirus pimctattis and Lepomis 
ctjanelliis) are common in the High Plains Region today. The 
range of two other species (for which the fossil identification to 
species is indicated by Smith as doubtful) extends approximately 
to the eastern edge of tlie High Plains; these are Ictiobus cf. bubaltis 
and Aplodinotus cf. grunniens. C. L. Smith (1962:512) suggested, 
with reservations, that Funduhis detillai Hibbard and Dunkle from 
the Middle Pliocene (Ogallala Formation) may be related to 
Fundulus kansae, a species now especially characteristic of streams 
of the High Plains. Recent counterparts of other species reported 
by Smith (1962) no longer occur naturally in the High Plains but 
are found farther to the east in the Mississippi Valley. 

Despite the fragmentary nature of the fossil record of Phocene 
fishes from the Great Plains, the known fauna (Hubbs, 1942:399; 
C. L. Smith, 1962: Table 1) indicates that catfishes of the genus 
Ictahirus and various sunfishes were probably common and that 
bowfin, gars, deep-bodied suckers, some cyprinids, cyprinodontids 
and an atherinid were present at one or more times. In North 
America today these groups, with the possible exception of the 
cyprinids, are best represented in the waters of the Atlantic and 
Gulf coastal plains and in the Mississippi Embayment. Hibbard's 
(1960) interpretation of Pliocene climate noted above suggests 
ecological conditions in which centrarchids, cyprinodontids, lepi- 
sosteids and large ictalurids might flourish if their ecological re- 
quirements approximated those observed for many members of 
these groups today. 



Fishes of Kansas River System 69 

Another line of evidence, indirect in nature, concerning the prc- 
glacial fauna of the present High Plains Region is oflFered by con- 
sideration of those species that now inhabit the western parts of 
the three rivers — Platte, Kansas and Arkansas — for which evidence 
best indicates preglacial interconnections (Fig. 3). Some of these 
connections continued into the Pleistocene until late Illinoian time 
or possibly somewhat later (Frye and Leonard, 1952:196-197). It 
is, however, difficult to ascertain which of the fishes now common 
to the tliree systems had their origin in the Preglacial Plains System, 
and which ones might more likely have reached the area after south- 
ward deflection of the Preglacial Hudson Bay Drainage (to be 
considered in a later section). 

The zoogeographic importance of a Preglacial or Pleistocene 
interconnection of the western parts of the Arkansas, Kansas and 
Platte rivers was first suggested by WilHams (1954:573) who theo- 
rized that the crayfish Orconectes n. neglectus dispersed into these 
systems by way of former stream connections west of the Flint 
Hills. D. A. Distler (personal communication) makes a similar 
case for dispersal of a subspecies of the orange-throated darter — 
Etheosfoma spectahile piilchelhim — via such connections. The 
native range of E. s. pulcheUum extends from the Platte River south- 
ward to the Brazos and Colorado drainages of Texas, lending sup- 
port to the view that the Preglacial Plains Drainage may have had 
connections to the south at one or more times in the past. 

The distribution of the plains killifish, Fundiilus kansae, coupled 
with that of the closely related (Miller, 1955:10, 11) or conspecific 
fish Fundulus zehrinus, resembles, in regard to stream systems 
involved, the distribution of E. s. pulcheUum, except for the presence 
of the killifish in the Pecos River Drainage. Both F. kansae and 
E. s. pulcheUum range from the Brazos and Colorado systems north- 
ward definitely as far as the Platte. There are scattered records 
of F. kansae north of the Platte, probably best attributed to intro- 
duction according to Miller (1955:11, 12). As noted above fossil 
evidence suggests that a progenitor of F. kansae inhabited the Pre- 
glacial Plains Stream. 

The sucker-mouthed minnow Phenacobius mirabilis is found 
(disjunctly) in the Colorado River of Texas (Clark Hubbs, 1957: 
93), and from the Red River northward to the Platte and some 
nearby drainages in northwestern Iowa and doubtfully in South 
Dakota (Bailey and Allum, 1962:54). The possible invasion east- 
ward of this species from an area of origin in the Great Plains is 
discussed in the appropriate species account. 



70 University of Kansas Publs., Mus. Nat. Hist. 

Several species depart from the pattern just discussed only in a 
greater degree of penetration northward. Two such species, 
Hybognathus placitus and Notropis I. lutrensis, are, in the main, 
trans-mississippian in distribution. Three others, Carpiodes c. car- 
pio, Lepomis cyanellus and L. humilis, now range farther east but 
reasons for suspecting a western origin are presented in the accounts 
of these species. 

The western subspecies of the sand shiner (Notropis stramineits 
7mssuriensis) occurs from the upper Missouri River Basin south- 
ward to the Arkansas River System and there is evidence of inter- 
gradation between this subspecies and another, eastern subspecies 
in the western part of the Red River System in Oklahoma. 

The range of the southwestern subspecies of the flat-headed chub 
(Hybopsis gracilis gulonella) does not extend so far south as the 
Red River but does include the Arkansas River System and the 
upper Pecos Drainage and the western part of the Missouri River 
Drainage. (Possibly stocks of this fish were extirpated in the Red 
River and other rivers to the south that lack mountain-headwaters 
during dry periods in the past such as the one suggested by Frye 
and Leonard [1963:19] for the upper part of the Red River Basin 
in Illinoian time.) 

Notropis topeka ranges from the Arkansas River northward to 
the Missouri River (Bailey and Allum, 1962:Fig. 6). This species 
probably inhabited the Preglacial or Pleistocene Plains Stream and 
in post-glacial time extended its range into Minnesota, Iowa and 
Missouri. The southwestern limits of the range of the Topeka 
shiner seem to be retreating northeastward; N. topeka is now rare 
or absent west of the Flint Hills in the Arkansas and Kansas river 
systems, although it seemingly was common there in historic time 
(Minckley and Cross, 1959:210-211). 

Possibly there has been widespread extirpation of southern popu- 
lations of Fundulus sciadicus, which seems also to have originated 
in the Preglacial Plains Stream or the Preglacial Missouri System, 
judging from its western distribution at present. 

A number of other species are wide-ranging in the Mississippi 
Valley and their areas of origin are obscure; however, they prob- 
ably inhabited the Preglacial Plains Stream (as well as other 
ancient drainages) because: 

( 1 ) they are widespread in southwestern stream systems, including the 
Pecos System; 

( 2 ) southwestern or western populations differ morphologically from eastern 
populations, suggesting a long sojourn there rather than recent invasion. 



Fishes of Kansas River System 71 

Species meeting one or both of these specifications are: Hybopsis 
aestivalis, Semotilus afromaculatus, Pimephales promelas, Campo- 
stoma anomaltim, Carpiodes ctjprinus, Catostomus comniersoni and 
Ictalurus melas. Also deserving mention here are Notropis atheri- 
noides, western representatives of which have, until recently ( Bailey 
and Allum, 1962:56-60), been deemed specifically distinct, and 
Notropis blennius, for which a southwestern subspecies has been 
recognized by Hubbs and Bonham (1951:103). 

HUDSON BAY COMPONENT 

Former outflow of parts of the upper Missouri River System to 
Hudson Bay was suggested by Todd (1914, 1923), Flint (1955:148, 
1957:Fig. 10-2), Horberg and Anderson (1956:106) and Meneley, 
Christiansen and Kupsch (1957). Todd inferred a pre-Nebraskan, 
northward drainage of Missouri River tributaries as far south as 
the White River in South Dakota (1914:267). Fhnt (1955:148) 
suggested that a drainageway, at least as old as pre-Illinoian, com- 
prising Cheyenne, Grand and Moreau rivers, led north from north- 
eastern South Dakota into the Hudson Bay Drainage. The evidence 
consists of high-level abandoned valleys and areas of collapsed 
glacial drift east of the present Missouri River in South Dakota. 
Flint interpreted the northern point of the Coteau des Prairies as 
a headland between the "Ancient Cheyenne" mentioned above and 
the "Ancient Minnesota" — these two streams having their conflu- 
ence north of this headland and proceeding thence to an Arctic 
egress (Fig. 3). In a study of bedrock topography, Meneley, 
Christiansen and Kupsch (1957) found evidence of a "Pre-glacial 
Missouri" that trended northeastward from the vicinity of Poplar, 
Montana, across the northwestern comer of North Dakota into 
Saskatchewan where it was joined by a "Preglacial Yellowstone," 
also trending northeastward from the area of present WiUiston, 
North Dakota. Together these streams coursed eastward across 
southern Saskatchewan into Manitoba where they joined a "Pre- 
glacial Red River" and then proceeded northward to the present 
region of Hudson Bay (Fig. 3). 

Evidence indicates, then, that the Preglacial Missouri System was 
a major waterway draining, in the United States, a large area in- 
cluding parts of Montana, Wyoming, North and South Dakota, 
Minnesota, and possibly other areas to the east. 

There is little published evidence concerning the fishes that in- 

4—8449 



72 University of Kansas Publs , Mus. Nat. Hist. 

habited the Preglacial Hudson Bay Drainage and the extent to 
which its fauna differed from faunas of the Preglacial Plains Stream 
System and the Preglacial Teays/Mississippi System. 

Deflection Southward of Ancestral Missouri System 

One can reasonably assume (as did Alden, 1924:413) that in 
the northern Great Plains during Pleistocene glaciation, water 
must have escaped to the south, west of the glaciers, (Fig- 4) 
granted, of course, that small amounts of water may have escaped 
supra-glacially, intraglacially or percolated through gravels under 
the glaciers (Flint, 1957:158-159). During each of the glacial 
maxima water from the Upper Missouri Region was occluded from 
flowing northward to Hudson Bay, a region considered (Flint, 
1957:316-318) to have become glaciated early in all glacial ages. 
Connections to the Hudson Bay Drainage would seem to have 
been in preglacial times or during interglacial stages. 

Widespread and thick accumulations of Holdrege gravels in 
Nebraska ( Lugn, 1935 ) suggest a great inflow of glacial water into 
that state from north and west during Nebraskan glaciation. At 
the time of the Kansan glacial maximum, drainage from a vast 
area of the northern Great Plains, between the glacial front and 
the Rocky Mountains, must have been funneled south into and 
across Nebraska (Fig. 4). It is, then, not suq^rising that massive 
gravel and sand deposits, of the magnitude of the Grand Island 
Member, were deposited in central Nebraska in Kansan time. Lugn 
(1935:117) stated in reference to the Holdrege and Grand Island 
gravels, "The thickest fluviatile deposits occur in the counties west 
of the till border, where the Missouri-Niobrara drainage flowed 
southward along the ice margins . . ." Lugn (1935:168) also 
considered the ancestral North Platte River an important bearer, 
in Kansan time, of water and sediment from part of the High 
Plains and Rocky Mountains. He thought that much of the 
Grand Island inwash sediment in south-central Nebraska was 
borne by the ancestral North Platte. 

South of Nebraska during the Kansan maximum, water must 
have flowed south, west of the glacial ice and the Flint Hills, ap- 
proximately along the course of the long-used master-stream of 
Pliocene antecedents tlirough the Pleistocene McPherson Valley 
and southward at least as far as the Ancestral Arkansas River 
Drainage (Fig. 4). This view has been taken by Lohman and 
Frye (1940:852) and Frye and Leonard (1952:190, 191), chiefly 



Fishes of Kansas Rr'er System 



73 




Fig. 4. Drainage pattern of the Ancestral Plains 

Stream System as it may have been at the time 

of maximum extent of the Kansan Glacier. 

from the presence of sediments interpreted as glacial outwash from 
the northeast in McPherson Channel, However, Bayne and Fent 
(1963:372) recognized no outlet of Kansan glacial meltwaters 
to the southwest and considered drainage of the Solomon River area 
to have been to the east through the Flint Hills throughout Pleis- 
tocene time (p. 367). This interpretation of an early Pleistocene 
through-flowing Kansas River seems not to be borne out by the 
studies of gravel lithology reported by Davis (1951:183-189). 

As north-flowing streams of the Ancestral Missouri System 
were deflected southward by encroaching glaciers, fishes indigenous 
to the area probably were also deflected southward. It could, 
of course, be argued that climatic conditions in advance of the 
oncoming glaciers were so severe, or glacial advance so rapid, 
that aquatic life in the upper Missouri was extirpated. This view 
seems extreme, however, as several authors have presented evidence 
that cHmatic effects on the biota, even a few miles in advance of 
the glaciers, did not result in a general extinction. Hibbard 
(1960:20, 21) suggested that during late Kansan time southwestern 
Kansas enjoyed a more equable and/or warmer climate than today. 
Frye and Leonard (1952:161) inferred that western Kansas had 
more rainfall, a slightly lower mean temperature, and more equable 



74 University of Kansas Publs., Mus. Nat. Hist. 

temperatures during late Kansan time that at present. They noted 
that occurrences of fossil gastropods suggested presence of more 
permanent waters and woodlands that at present, but indicated 
no widespread forests. Deevey (1949:1394), in discussing the re- 
populating of deglaciating areas by aquatic animals, concluded 
that streams carrying glacial meltwater were not excessively cold 
and tliat the glacial lakes, though colder than their descendants 
today, were not arctic. — 

Thienemann (1950:335) noted that the central European aquatic 
fauna, even when compressed in a region of tundra between north- 
ern and Alpine glaciers, contained species still found in the same 
region. 

It seems likely, then, that faunal components of the Preglacial 
Missouri River System did not suffer extinction but persisted locally 
and were brought into contact with stream systems to the south. 
These may have included the western headwaters of some streams 
of the Teays/Mississippi System (Fig. 3). Horberg and Anderson 
(1956: Fig. 2), for instance, have tentatively mapped the head- 
waters of some streams of an Ancient Iowa River System and some 
other tributaries of the Teays/Mississippi as extending west into 
the Great Plains. (I have pointed out above that western tribu- 
taries of the Ancient Platte more likely flowed southward into the 
Preglacial Plains Stream.) It seems that the upper Missouri and 
other streams heading west of the glacial fronts of the Nebraskan 
and Kansan advances were deflected into the south-flowing Plains 
Stream west of the Flint Hills in Kansas (Fig. 4). Thus the first 
and second and possibly later glaciers probably caused a mingling 
of stocks of the Preglacial Missouri and Preglacial Plains Stream 
and possibly a lesser mingling with stocks of some western head- 
waters of the Teays/Mississippi. Bailey and Allum (1962:120) 
listed six kinds of fishes (Semotilus margarita nachtriebi, Chro- 
somus neogaeus, C. eos, Hybopsis plumbea, Catostomus cato- 
stomus and Pantosteus platyrhynchus) that they beheved capa- 
ble of enduring the last glaciation ( s ) in the upper Missouri in 
South Dakota. I suspect that several other species, originally 
derived from the Preglacial Missouri System or from western tribu- 
taries of the Teays/Mississippi System, persisted in the Middle 
or Upper Missouri during the Pleistocene for the following reasons: 

(1) Certain species exhibit racial variation from north to south 
in the Missouri System. These differences may be residual evi- 
dence of originally different races in the Preglacial Missouri 
and Preglacial Plains drainages. The existence of such differences 
argues for a northwestern origin rather than colonization in post- 



Fishes of Kansas Rrter System 75 

glacial times from a southeastern source in the lower Missouri 
River Basin. Penetration after the last glacial advance into the 
middle Missouri River Basin by way of connections to the upper 
Mississippi River System has been suggested for several species 
of fishes in eastern South Dakota by Bailey and Allum (1962:122). 
However, most of the fishes so categorized by Bailey and Allum 
seem localized east of the Missouri River. Tlie species treated 
here are not so locahzed. Probably they endured at least one 
glacial advance in the upper and middle Missouri River Basin. 

(2) The species to which my argument pertains ai'e widespread 
in Canada, at least as far north as Hudson Bay, and also penetrate 
west into the cooler headwaters of the Missouri System in Wyoming 
and central Montana. Species possessing a tolerance that has 
enabled their dispersal far northward might have survived Pleisto- 
cene conditions in the area of the present Upper Missouri Basin. 
Certain fishes of present-day northern aflSnities may, in Pliocene 
time, have existed in the Hudson Bay and otlier Arctic drainages 
of Canada. 

Species that are in accord with both criteria mentioned above are 
Catostomus commersoni, Pimephales promelas, Moxostotna mac- 
rolepidotum and Hijbopsis gracilis. The three species Hsted first 
or their progenitors could have inhabited all three preglacial drain- 
ages treated here, in view of their present wide ranges. Hybopsis 
gracilis may have inhabited only the Preglacial Missouri and the 
Preglacial Plains Stream drainages. 

Other species, occurring at present in the Missouri River System, 
meeting the second criterion above, and considered Hkely in- 
habitants of a Preglacial Missouri System are: Hiodon alosoides, 
Rhinichthys cataractae, Notropis atherinoides, Percopsis omiscomay- 
cus, Culaea inconstans, Notropis heterolepis and Lota lota. 

The virtual restriction of Scaphirhynchus album, Hybopsis geUda, 
and Hybopsis meeki to the Missouri River System suggests an origin 
for these species in the Preglacial Missouri System. 

MISSISSIPPI COMPONENT 

Probably the easternmost part of the Kansas River System, east 
of the FHnt HiUs, long flowed to the east (Frye and Leonard, 
1952:185) and was a part of the Preglacial Teays/Mississippi 
System (Fig. 3), whereas the western part of the system flowed 
south, west of the Flint Hills, through the time of Kansan glacia- 
tion. Because the Kansan glacier overrode most of the present 
Kansas River Basin east of the Flint Hills, repopulation from the 



76 University of Kansas Publs., Mus. Nat. Hist. 

east must have taken place during glacial retreat or thereafter. 
After this retreat, capture of streams progressing farther westward, 
incorporated several tributaries of tlie Plains Stream into the Kan- 
sas River system, culminating in a drainage pattern closely re- 
sembling that of the present (Frye and Leonard, 1952:193-196). 

Downcutting westward in mid-Pleistocene time has been docu- 
mented for the Kansas River (Frye and Leonard, 1952:192ff.) 
at a time (Illinoian and/or during the preceding or succeeding 
interglacial ages) when the Ancestral Plains Stream, flowing south 
through McPherson Channel in central Kansas, was aggrading 
its channel. Ultimately the south-flowing stream proved unable 
to hold its own against the vigorous invader from the east. Frye 
and Leonard (1952:193) attributed this piracy mainly to isostatic 
readjustments. 

Quinn's (1958:42) evidence, although scant, indicated a similar 
westward cutting through an Ozark/Ouachita Highland by a tribu- 
tary of the Mississippi, the Ancestral Arkansas River, to capture 
another segment of the Plains Stream System. 

In regard to deflection eastward of the Platte, Lueninghoener 
(1947:60) wrote, "It seems reasonable to visualize the initial erosion 
of the Platte River Valley to have taken place in Yarmouth time 
by a Missouri River tributary eroding headwardly across the till 
border. Headward erosion appears to have progressed until cap- 
ture of some of the drainages west of the till border took place." 
Lugn (1935:153) wrote, "A long time must have elapsed before 
the drainage west of the till border . . . was captured and 
diverted through the new post-Loveland Platte River." 

All these instances suggest that downcutting westward by streams 
of the Mississippi System occurred in the mid-Pleistocene, sub- 
sequent to Kansan glaciation, in Yarmouthian or Illinoian time. 
This rough correspondence in time of deflection of several stream 
systems may be fortuitous, may be related to isostatic changes in 
the region, or may reflect a widespread pattern of climatic change. 
A primarily climatic mechanism whereby a south-flowing Plains 
Stream might have been captured, segment by segment, by east- 
flowing tributaries of the Mississippi System is suggested by reports 
(Frye and Leonard, 1952:165; 1957:29; 1963:19) of the onset of 
widespread aridit>^ in the High Plains, especially to the southwest, 
in Illinoian and/or Yarmouthian time. These writers (1952:165) 
stated, "It is difiicult to escape the conclusion that a profound 
change in ecological conditions in the Great Plains occurred during 
or at the close of the Yarmouthian interglacial interval, or at the 



Fishes of Kansas River System 77 

beginning of the Illinoian cycle of erosion and deposition. Whole- 
sale extinction of great populations of branchiate and other 
gastropods adapted to life in permanent water, which thrived in 
western Kansas in late Kansan and early Yarmoutliian time, is in- 
dicative of a less humid environment, or at least of a marked 
decline in tlie prevalence of permanent ponds and lakes of clear 
water in the Great Plains region." These authors fiirther stated (p. 
165) ". . . the aquatic gastropods in Illinoian deposits indicate 
an environment of ephemeral ponds and silt-laden streams." The 
effect of such a climatic change on streams of the High Plains is 
further suggested by tlieir description (Frye and Leonard, 1957:29) 
of conditions in the High Plains of Texas in Yarmoutliian and IIH- 
noian time. They wrote tliat streams were less competent in these 
Ages than in either the Kansan or Early Wisconsinan and that there 
may have been reduction of topographic relief due to the choking 
of minor valleys with sediments that streams were not able to carry 
away. The south-flowing iVncestral Plains Sti-eam, entering and at- 
tempting to cross an arid or semi-arid region such as the one just 
described, might, in accordance with the principles discussed by 
Quinn (1957:156), have deposited its load and aggraded its chan- 
nel as its waters evaporated. Such a choked, meandering stream 
might have been captured by tributaries of the Mississippi as they, 
debouching in an area of higher rainfall, cut westward relatively 
rapidly. 

The entire Kansas River System, then, eventually had its outlet 
into the Mississippi River. This provided access to the entire 
newly-integrated system of an eastern fauna. An earlier faunal 
contribution (discussed above) to the Ancestral Plains Stream 
from the Ancestral Teays/Mississippi might have been incorporated 
from tributaries of the Mississippi (for example, the Ancestral Des 
Moines River) that were deflected southward due to encroaching 
glaciers of the Nebraskan and Kansan advances. 

Many species, typical of the larger rivers of the Mississippi 
Valley, must have made their way up the major watercourses of 
the Missouri System, including the lower part of the Kansas River 
and its larger tributaries. Evidence as to wliich of the big-river 
fishes first reached the Kansas River Basin from this source is scant. 
The former distributions and area of origin are difficult to judge 
for species that now inhabit the mainstreams of large rivers, be- 
cause: (1) after confluence of formerly disjunct drainages, barriers 
to dispersal of such fishes seem nonexistent; (2) thus, formerly al- 
lopatric species may soon occupy almost wholly sympatric ranges; 



78 Unr'eesity of Kansas Publs., Mus. Nat. Hist. 

(3) mingling of stocks may be so nearly complete, due to the 
relatively large size and high vagility of some big-river species, that 
any racial diflFerences that may have existed soon disappear. Thus, 
less insight into their geographic history seems possible than in 
the case of upland species that inhabit peripheral waters of a major 
drainage basin. Consequently, big-river fishes probably have a 
lesser tendency and opportunity for speciation than do small-stream 
fishes. 

Underbill and Merrell (1959:140) theorized that intrabasinal 
variation in Notropis dorsalis is partly attributable to preference 
of this species for small streams where populations become isolated 
to some extent; the larger, more turbid intervening streams are 
thought to act as a barrier between populations in the tributaries. 
Neighboring, but unconnected stream systems may have many 
"large-river-species" in common. Thus, some species characteristic 
of large rivers, entering the Kansas River System from the east after 
retreat of the Kansan Glacier and downcutting westward into the 
Plains Stream System, probably encountered and mingled with 
populations of the same or closely related kinds that already in- 
habited the Plains Stream. C. L. Smith (1962) reported genera 
typical of larger rivers in some Pliocene faunas found in the High 
Plains {Lepisosteus, Ictiobus, Ictalurus, Aplodinotus? and Pomoxis). 
Some of these as well as other genera typical of larger waters have 
also been reported by C. L. Smith (1954, 1958) and G. R. Smith 
(1963) from beds of mid-Pleistocene age — all of which suggests a 
long sojourn in the central United States for many big-river fishes. 

In the case of several small (and possibly less vagile) cyprinids 
that occupy large rivers, mingling has not so completely erased 
differences between eastern and western populations. This is 
especially true of Notropis stramineus, which occurs in small as 
well as in large streams. In Notropis atherinoides, Notropis blen- 
nius, and Hybopsis aestivalis the evidence of mingling with eastern 
stocks seems stronger in the Kansas River Basin than in the Arkansas 
River Basin and other stream systems to the south. 

Species of the Kansas River Basin clearly tolerant of larger rivers 

and, in many cases, restricted to them and seeming to have had 

an eastern, or a mixed origin include: 

Ichthyomyzon castaneus Cycleptus elongatus 

Scaphirhynchus platorynchus Ictiobus cyprinellus 

Polyodon spathula Ictiobus bubalus 

Lepisosteus platostomus Ictiobus niger 

Lepisosteus osseus Ictalurus punctatus 

Dorosoma cepedianum Pylodictis olivaris 



Fishes of Kansas River System 79 

Hybopsis storeriana Anguilla rostrata 

Notropis atherinoides Pomoxis annularis 

Notropis blennius Stizostedion canadense 

Notropis buchanani Stizostedion vitreum 

Notropis stramineiis Aplodinotus grunniens 

Notropis shumardi Roccus chrysops 

Hybognathus nuchalis 

Bailey and Allum (1962:121) described a category of fishes, 
which, hke that above, probably entered South Dakota from the 
southeast by way of the Missouri River but which consisted of spe- 
cies more typical of upland creeks than of large and often turbid 
rivers. Species, so characterizable, also undoubtedly entered the 
Kansas River Basin from the east. Conditions for invasion by 
kinds that prefer clear water of upland tributaries likely were 
favorable in the interval following retreat of the Kansan glacier and 
before extensive downcutting westward had taken place (while 
the river headed east of the Flint Hills). The lower mainstream 
presumably was clearer than at present and probably possessed 
rubbly and gravelly bottoms with frequent riffles, as do the present 
Neosho and Verdigris rivers that head mainly in the Flint Hills. 
As discussed by Bailey and Allum ( 1962:122), upland fishes possibly 
venture into large rivers when they are clear and cool, as in winter. 
Some species may, even under present conditions, ascend stepwise 
from one year-round habitable tributary to the next by a process 
here termed "tributary-hopping." 

Especially important as a source-area for upland fishes was 
the rich faunal area of the Ozark Plateau, the northwestern limits 
of which are within 100 miles of the mouth of the Kansas River. 
Several upland species of Ozarkian aflBnities are, at present, re- 
stricted to the clearest and least-modified streams of both the 
Kansas and Arkansas river systems. 

Like the species typical of large rivers, some species that are 
typical of upland streams may have met and mingled with closely 
related populations already inhabiting the western part of the 
Kansas River Basin. (Species thought certainly to have inhabited 
the Ancestral Plains Stream are indicated by asterisks in the list 
below.) The populations of these species now present in the 
Kansas River possibly were derived from all three preglacial stream 
systems (Fig. 3). Following is a list of kinds that now occupy 
upland tributaries in the Kansas River Basin, and that probably 
entered the Basin from the east. In the case of Moxostoma mac- 
rolepidotum and Etheostoina spectabile the eastern influence is 
detectable only through the presence of intergrading subspecies. 



80 University of Kansas Publs., Mus. Nat. Hist. 

Carpiodes velifer * Semotilus atromoculatus 

*• Catostomus commersoni * Ictalurus melas 
Moxostoma ertjthrurum Ictalurus natalis 

Moxostoma macroJepidotum Noturus exilis 

Chrosomus erythrogaster Noturus flavus 

Hybopsis higuttata Etheostoma nigrum nigrum 

Notropis cornutus Percina caprodes 

Notropis dorsalis Percina maculata 

Notemigonus crysoleucas Micropterus salmoides 

Notropis rubellus Lepomis macrochirus 

Notropis umhratilis Lepomis megalotis 

Pimephales nototus ** Etheostoma spectabile 

* Pimephales promelas 

Along with fishes, the aquatic snails Campeloma snbsoUdiim 
(Anthony) (Leonard, 1959:31), Pletirocera acuta Raf. (Goodrich, 
1939:4), and the crayfish Orconectes lutetis (Greaser) (Metcalf 
and Distler, 1961:354) probably entered the Basin from the east. 
The snails are, at present, presumably extinct in the Basin. 

Several of the species listed probably entered the Kansas River 
Basin solely from the east after retreat of the Kansan Glacier. In 
this category would seem to belong those species that presently 
occur in the Kansas River System but not in the Arkansas River 
System — thus suggesting arrival in the Kansas River Basin too 
late to utilize the McPherson Ghannel Gonnection to the Arkansas 
River. Although the Arkansas River also came ultimately to flow 
into the Mississippi River (in post-Pliocene, but pre-Sangamonian 
time according to Quinn, 1958:42), its connections were far south 
of those of the Kansas River and thus introduced into the Arkansas 
River System a somewhat different fauna with more southern 
affinities. Species or subspecies that occur in the Kansas River 
System but not in the middle and upper Arkansas River System 
are few in number and include Hijbognathus hankinsoni, Hybopsis 
aestivalis htjostoma, Notropis cornutus, Notropis dorsalis, Lota 
lota, Hybopsis gelida, Hybopsis meeki, Scaphirhynchus albiis, and 
Hybopsis gracilis: gracilis x gulonella (only H. g. gulonella occurs 
in the Arkansas System). H. g. gracilis, H. gelida, H. meeki and 
S. albus are not found anywhere outside the Missouri River System 
and that part of the Mississippi River near and downstream from 
the mouth of the Missouri. As suggested elsewhere, these seem 
to be old, endemic fishes of the Missouri River, derived from the 
preglacial north-flowing Missouri rather than from an eastern 
source. 

Diversion to the east also opened pathways of dispersal for 
fishes originating in the Great Plains, Kinds that have invaded 
eastward from a probable Plains source include Carpiodes carpio 



Fishes of Kansas River System 81 

carpio, Notropis I. luirensis, Notropis topeka, Phenacobius mirahilis, 
Hybognathus placitiis, Campostoma anomaluin pullttm, Fundulus 
kansae, Fundulus sciadlcus, Lepomis cyanellus, and Lepomis humi- 
lis. Gene flow from western populations is detectable in popula- 
tions of Notropis stramineus and Etheostoma spectahile east of 
the mouth of the Kansas River in Missouri. 

VARIATION FROxM NORTHEAST TO SOUTHWEST 

As the Preglacial (north-flowing) Missouri River System was 
deflected southward (possibly incorporating some western tribu- 
taries of the Teays/Mississippi System) into the Ancestral Plains 
Stream, faunal elements were probably concomitantly deflected, 
resulting in a mingling of stocks. At least one mingling of eastern 
and western stocks occurred after deflection of some sections of 
the Plains Stream eastward. Probably other deflections with faunal 
mixing took place in glaciated areas and glacial fringe areas of the 
central United States outside the region considered here. 

To what extent, if any, northern connections to Hudson Bay may 
have been renewed during interglacial stages is not known. How- 
ever, at the present time, approximately 11,000 years (Frye and 
Wfllman, 1960: Fig. 1) since the Valderan glacial advance, streams 
tributary to Hudson Bay already have penetrated as far south as 
South Dakota. Surely during interglacial stages stocks must have 
met and mingled as they repopulated large sections of deglaciating 
terrain in Canada and the northern United States. 

Throughout the Pleistocene there probably was little opportunity 
for extended periods of isolation — and, in consequence, little im- 
petus to speciation — in areas of plainlike topography that were 
repeatedly glaciated or that were immediately adjacent to glaciers. 
In such areas repeated glacial retreat and advance probably fur- 
thered renewed intermingling of stocks rather than isolation of 
stocks with possible subsequent speciation. On the other hand, in 
regions where glaciers abutted areas of broken topography (for 
example the northern Appalachian Mountains and Cumberland and 
Ozark Plateaus) isolating mechanisms might have been strength- 
ened by encroachment and retreat of glaciers. Probably also in 
nonglaciated areas in the southwestern United States Pleistocene 
climatic fluctuations enhanced the process of speciation, whereas 
in glaciated areas of the Great Plains and Interior Lowlands climatic 
fluctuations inhibited the process. Thus, as pointed out by Grinnell 
(1924:228) and Miller (1948:128), arid regions have produced, 



82 University of Kansas Publs., Mus. Nat. Hist. 

through a process of isolation, new kinds, both of terrestrial and of 
aquatic animals. On the other hand, Thienemaim (1950:390) 
wrote of a "Glaziale Mischfauna" produced by encroachment of 
glaciers on the plains of Central Europe noting (p. 336) tliat re- 
sistant remnants of the preglacial, middle European fauna became 
mixed with immigrants from the Alpine and Polar regions. 

Within the present Mississippi River System, southwestern vari- 
ants of some fishes have been given taxonomic recognition as sub- 
species or as species having sibling species to the northeast. In 
these fishes, characters vary in the same manner in a southwesterly 
direction. Some kinds of variation, together with examples, are 
listed below. ("Northeastern" here refers to populations from the 
Ohio River System and upper Mississippi River System. "South- 
western" refers to the upper Arkansas River system, the upper Pecos 
River Drainage, and the upper Red River of the South. Intervening 
waters show various kinds and degrees of intergradation. ) 

General Body Shape 

Northeastern representative of the fishes considered here tend 
to be fusiform in body outline. Dorsal and ventral profiles curve 
smoothly and slightly anteriorly and posteriorly from the point 
of greatest body depth near the midpoint of total body length. 
The body is slender, the ratio of greatest body depth and head 
depth to standard length being less than in southwestern fishes. 
The smooth, slight degree of curvature continues along dorsal and 
ventral head profiles resulting in an acute, tapering snout, terminat- 
ing approximately midway of the vertical distance between the 
top and the bottom of the head. The head is usually small in rela- 
tion to the entire body — frequently being shorter, narrower and 
shallower than in southwestern kinds. 

Southwestern fishes tend to be chubbier in body form, especially 
anteriorly. The point of greatest body depth is generally anterior 
to the mid-point of total length. From this point there is a striking 
decurvature of the anterior dorsal profile resulting in a 'Tiump- 
backed" appearance. The posterior dorsal profile descends more 
gradually than the anterior to the caudal peduncle, which is gen- 
erally only shghtly deeper than the peduncle of northeastern fishes. 
The total impression is of a chubby fish, thick-bodied anteriorly, 
and tapering to a slender caudal peduncle. The head is relatively 
larger in southwestern kinds, generally being wider, deeper, longer 
and more massive. This may be in keeping with Hubbs' (1940b: 



Fishes of Kansas River System 83 

199) observation that fish in warmer waters typically have deeper 
bodies and larger heads than those of colder waters. As pointed 
out elsewhere, however, higher altitudes may, in the western parts 
of most of these southwestern streams, produce cold water tempera- 
tures. In southwestern kinds the snout is blunter and the ventral 
contour of the head is less upcurved than in northeastern kinds. 
Lines connecting (a) the occiput, (b) the ventral surface of the 
head directly below the occiput and (c) the anteriormost terminus 
of the snout tend to form a right triangle in southwestern fishes, in 
northeastern fishes such lines tend to form an isosceles triangle. 
Olund and Cross (1961:333), in comparing heads of a southwest- 
ern subspecies, Hijhopsis gracilis gulonella, and a more northern 
subspecies, H. g. gracilis, found the former to be rounded anteriorly 
( decurved ) in side view and deeper, whereas the latter was wedge- 
shaped in profile and much shallower. Probably the most extreme 
example of pronounced anterior decurvature to the southwest is 
found in the chub Htjbopsis aestivalis as represented in the Arkansas 
and Red rivers {H. a. tetranemus and H. a. australis respectively). 
The minnow Phnephales promelas and several of the species of 
Notropis also exhibit anterior decurvature especially well (for ex- 
ample southwestern as contrasted with northeastern populations of 
N. hlennius and N. stramineus.) This condition has been pointed 
out by Hubbs and Bonham (1951:103-107) in N. hlennius. Hybo- 
gnathus nuchalis from the northeast and its sibling species Hybo- 
gnathiis placitus from the southwest show a similar difference in 
body shape. This variance is also observed where these two species 
are sympatric in the lower Kansas River Basin. 

As concerns other than cyprinid fishes similar tendencies in body 
shape were observed in the sucker Catostomtis commersoni. Hubbs 
and Black (1940:227) noted that in the southwestern subspecies of 
the carpsucker Carpiodes carpio — called elongatus — the anterior 
part of the back is more arched than in the northern subspecies — 
carpio. The darter Etheostoma spectabile comprises, among others, 
a plains subspecies (£. s. pulchellum) with an arched and decurved 
anterior profile, whereas the typical subspecies, ranging from the 
northern Ozarks eastward, exhibits a more fusiform body profile. 
The same differences in profile were noted between Etheostoma 
cragini of the central and western Arkansas River Basin and Etheo- 
stoma pallididorsum of the Caddo River Drainage in Arkansas 
(Distler and Metcalf, 1962: Table 2). 



84 University of Kansas Publs., Mus. Nat. Hist. 

Mouth 

In correlation with the decurved snout of southwestern repre- 
sentatives tlie mouth tends to be less nearly terminal and more 
nearly horizontal, and that of northeastern fishes in more nearly 
terminal and is oblique. This is especially true of Pimephales 
promelas from northeast to southwest (Taylor, 1954:42; Hubbs and 
Ortenburger, 1929a: 38). In the species Notropis blennius north- 
eastern populations characteristically possess an oblique, terminal 
mouth, whereas southwestern populations tend to have a more 
inferior, horizontal mouth. Hybopsis gracilis, Rhinichthys catar- 
actae, Notropis stramineus and Campostoma anomalum, on the 
other hand, show little variation as regards this character, having 
inferior, horizontal mouths throughout their range. 

Eye 

Hubbs (1940b: 202) and Bailey (1956:333) pointed out that eye 
size tends to be smaller in plains representatives of some species 
of fish. Moore ( 1950 ) noted this in several members of the genus 
Hybopsis that inhabit plains streams. The reason for the smaller 
eye of Plains minnows is not entirely clear. Moore (1950:92) sug- 
gested that in some species smallness was related to decreased 
light in turbid waters of Plains streams and to the greater develop- 
ment of cutaneous sensoiy structures. That there is a selective 
value in some degree of reduction in the amount of eye surface 
exposed in sandy streams of the Great Plains is suggested by study 
of populations of Semotilus atromaculatus from shallow, sandy 
streams of the western Platte and Kansas river systems. In these 
fish there is, in addition to a smaller eye than in eastern populations 
(Table 8), a partial overgrowth of skin around the periphery of 
the eye, further decreasing the surface area exposed. Notropis 
atherinoides from the Kansas River also exhibits peripheral over- 
growth of skin around the eye and partial overgrowth of skin was 
pointed out by Moore (1950:92) for the eye of the chub Hybopsis 
meeki. 

Scales 

Many species of fishes have smaller scales in the Missouri River 
and other Plains streams than in the Ohio and upper Mississippi 
basins (see Bailey, 1956:333). For the sand shiner, Notropis stramin- 
eus, Bailey and Allum (1962: Table 5) presented data showing 
higher counts of body-circumference scales toward the west. Sev- 



Fishes of Kansas River System 85 

eral compaiisons reported on herein show a comparable tendency. 
In most species inhabiting tlie Great Plains sections of the Platte, 
Kansas, and Arkansas rivers, further smoothing of the external 
surface seems to be achieved by embedding of scales, especially 
the ventral and predorsal scales. In Notropis dorsalis of the west- 
ern Platte (piptolcpis of some authors) and in occasional speci- 
mens of Notropis stramineus, reduction in scale size and embed- 
ding have resulted in a predorsum naked in appearance. 

Adaptive Value 

A number of features have been noted that, to a greater or 
lesser degree, characterize southwestern races of some wide-rang- 
ing fishes. These have included decurved anterior dorsal profile, 
inferior, horizontal mouth and small eyes and scales. In some 
western fishes of turbid rivers there has been a marked develop- 
ment of cutaneous sense organs on the lower surface of the body 
and of barbels around the mouth. These features last-mentioned 
are probably adaptive and may (Moore, 1950:91-92) afford the 
fish greater cognizance of the substrate. Additional adaptive values 
were suggested by Hubbs and Walker (1942:101, 102), who in 
regard to certain psammophilous shiners wrote: "In correlation 
with their life on the open bottom in the current, they have terete 
bodies, with the dorsal contour more strongly arched than the 
ventral. The pectoral fins are large and horizontally expanded. 
The snout is long and decui"V'ed, and the large mouth in inferior. 
These morphological features adapt the fish to swimming about 
and feeding over the open bottom, in the current. The form of the 
head, body and fins is such as to increase the downward compo- 
nent of the force of the current, so as to help hold the fish against 
the bottom." Hubbs (1940:200) suggested that smaller scales 
provide a smoother surface that might be advantageous in facing 
the stress of water currents. 

Variation, Its Possible Causes 
and Its Taxonomic Interpretation 

The interpretation of the variation existing from northeast to 
southwest in terms of taxonomic treatment has been diverse. 

Southwestern or western (Great Plains) populations of the sev- 
eral species involved have been considered variously as (1) spe- 
cifically distinct, (2) subspecifically distinct or (3) not taxonom- 
ically distinct — from eastern or northeastern populations of the 



86 University of Kansas Publs., Mus. Nat. Hist. 

Mississippi Valley. In the course of time some kinds have been 
placed by various workers in more than one of the categories just 
mentioned. 

This lack of consistency in taxonomic treatment suggests the 
complex nature of the problems that these fishes present and in- 
timates that in some cases differences are more easily observed than 
in others. Despite this complexity there is among the various 
kinds of fishes concerned some concordance ( as pointed out above ) 
in regard to characters involved and in the clinal expression of 
these characters from east to west or northeast to southwest. Bailey 
(1956:334) suggested this in writing, "It is of interest to note that 
where . . . Mississippi basin fishes have intimately related 
species or subspecies on the Great Plains, the same character differ- 
ences are usually involved." Bailey specifically mentioned the 
smaller eye and greater number of circumferential body scales 
found in populations from the Great Plains. 

The problem arises whether this difference is purely a phenotypic 
response made during ontogeny to an environmental stimulus or is 
the expression of a genotype of wide occurrence in streams of the 
areas involved. Studies by several investigators have demonstrated 
the modifiability of the phenotype of some fishes experimentally by 
certain environmental stimuli. Bailey and Gosline (1955), Need- 
ham and Gard (1959), Underbill and Merrell (1959), Olund and 
Cross (1961), Strawn (1961), and Barlow (1961) have listed many 
of these studies and summarized important aspects applicable here. 
Several authors (Bailey, 1956:333-334; Bailey, Winn and Smith, 
1954:149, and Strawn, 1961:155) have suggested that some pheno- 
typic variation observed in natural populations may also be a direct 
result of environmental stimuli such as temperature, chemical con- 
tent of water, light or diet. On the other hand, Lagler and Bailey 
(1947:58) provided experimental evidence for the genetic fixity of 
degree of squamation in two subspecies of Etheostoma nigrum. 
Heuts (1947:100-102) found fresh and saltwater races of Gastero- 
steiis aculeatus to be genetically distinct. D. A. Distler (personal 
communication ) reared stocks of four geographic races of the darter 
Etheostoma spectahile for several generations under laboratory con- 
ditions as nearly identical as possible. No phenotypic differences 
were observed that would have made identification to subspecies 
any more difficult for F^ or F2 progeny than for the respective par- 
ental stocks. 



Fishes of Kansas River System 87 

In characters of Plains fishes that are most Hkely of adaptive value 
it seems unwarranted to attribute great significance to direct en- 
vironmental stimulus. There seems to be no reason to think that a 
character of advantage to the organism would necessarily be con- 
sistently produced by direct stimulus from the environment during 
an early developmental stage. Such an advantageous response to 
environmental stimulus, where and if it occurs, probably is fortui- 
tous. If advantageous modifications could be attained simply by 
direct environmental stimulus there would be little need for geno- 
typic change (allowing for speciation) to take place. 

Hubbs (1943:118) noted that a majority of the taxonomic char- 
acters employed in racial analysis of fishes behaved according to a 
Galtonian type of inheritance — that is to say, in a manner best 
explained as being controlled by multiple factors. Underbill and 
Merrell (1959:144-145) pointed out that certain meristic characters 
such as anal rays may be remarkably stable within a population, 
whereas other characters may exhibit much variation from popula- 
tion to population. They suggested that these variable characters 
might be controlled by multiple genes. Needham and Gard (1959: 
76) postulated that multiple genes may control the migratory in- 
stinct in the rainbow trout, which shows various gradations of de- 
velopment. The broad "normal curves" obtained from analysis of 
various body proportions and of some meristic characters of Plains 
fishes suggests control by multiple genes. If control of such char- 
acters is rather by direct environmental stimulus the results seem, 
in any event, to be indistinguishable. 

Underbill and Merrell (1959:145) speculated that genotype may 
simply determine the upper and lower limits of variation in regard 
to highly variable meristic characters. They further suggested that 
environment "operates on the existing genotypes, modifying their 
expression within their reaction ranges and also selecting the better 
adapted genotypes." Needham and Gard (1959:75) termed such 
environmental eflFects as "direct" and "selective." Both (a) the 
ability of the organism to adapt to the environment and (b) its 
limits of modifiability during development would seem to be ulti- 
mately under genie control. 

The relationships discussed in the preceding paragraphs might be 
summarized thus: the genotype attained through natural selection 
produces a phenotype that is adaptive and, in addition, determines 
the limits to which this phenotype can be modified directly by the 



5—8449 



88 University of Kansas Publs., Mus. Nat. Hist. 

environment. Direct stimulus seemingly could produce a pheno- 
type that is adaptive (and thus additive to the process of natural 
selection), neutral, or even non-adaptive. 

Thus, evidence seems great as pointed out by Miller (1948:148, 
1961:555), Bailey and Gosline (1955:24), Needham and Card 
(1959:73, 77), Underbill and Merrell (1959:145), Olund and Cross 
(1961:342) and Strawn (1961:155) that the phenotype observed 
(and which must be dealt with taxonomically) is the product both 
of heredity and of direct environmental stimulus. Strawn (1961: 
155) termed this relationship the "product of a given environment 
interacting on the genotype." 

For fishes of the Great Plains such interacting forces might be the 
following: 

(1) Natural selection for certain genotypes. This is controlled by the en- 
vironment that is characteristic of Plains Streams (to what extent this 
environment may have varied in the past is not fully known, although 
some inferences have been presented in preceding sections). 

(2) Isolation. Drainages of the Great Plains have been variously isolated 
from northern and eastern drainages in Preglacial and Pleistocene time. 
This isolation would reinforce the selective mechanisms mentioned 
imder ( 1 ) . 

(3) Mingling of stocks. This has been due to drainage changes of the 
Pleistocene alternating with or subsequent to (2) above. 

(4) Direct environmental influence on phenotypic expression. 

Items (1), (2), and (4), above, would, in most instances, tend 
to yield a more distinctive kind of fish toward the southwest. Item 
(3) would, in cases where interfertility was unimpaired, tend to 
produce a more homogeneous population. 

The interrelations of isolation and mingling have been discussed 
by Hubbs (1943:116-117), who states "It will commonly be difficult 
or impossible to determine whether the intermediate characters of 
a given population date back to the initial divergence of the two 
types, or are due to the secondary or subsequent meeting and inter- 
breeding of populations, or of entire forms, which had previously 
been isolated for some time." 

In at least one case — Hijbognathus placitus and Hyhognathus 
nuchalis — the processes of difi^erentiation (just described) between 
stocks inhabiting a western. Plains Drainage and a more eastern 
Teays/Mississippi Drainage seem to have progressed far enough to 
produce different, although closely related, species. 

In other instances differentiation has not produced species but 
has resulted in well-differentiated variants that have been recog- 
nized as subspecies {Notropis s. straminetis and missuriensis; 
Etheostoma s. s^pectabile and pulchellum; Hybopsis aestivalis sspp. ) 



Fishes of Kansas River System 89 

In still other instances differentiation may have been less or sub- 
sequent mingling may have obscured differences that once were 
better defined. Such kinds vary over a broad zone of clinal inter- 
gradation. Such intergradation is seen to various degrees and in 
varying patterns between Plains and eastern populations of Semo- 
tihis atromaculatus, Hybopsis aestivalis htjostoma, Notropis atherin- 
oides, Notropis blennius, Pimephales promelas, Campostoma 
anomalmn pulhnn, Carpiodes cypriniis, Catostomtis commersoni 
and IctaJurus melas. 

RECORDS AND COLLECTIONS OF FISHES 

An early summary of collections from the Missouri River Basin 
was compiled by Evermann and Cox (1896). The status of 
ichthyological collecting in Kansas prior to 1946 was summarized 
by Breukelman (1946). His account contains many references to 
the Kansas River Basin. Minckley (1959) listed collections that 
had been made in the Blue River Basin prior to his work in that 
stream system. 

The earliest reports that are possibly attributable to the Kansas 
River Basin all seem to deal with catfishes. Thus, Abbott (1861: 
568-569 ) described two new species of "Pimelodus" ( both probably 
Ictalurus punctatus according to Evermann and Cox, 1896:354) 
from "Fort RHey, Kansas" and Gill (1862:43, 44; 1876:417-419, PI. 
6) named and discussed Ictalurus simpsonii (also identifiable as 
I. punctatus) noting (1862:44) that "Two specimens were obtained 
by Dr. Suckley in the 'Big Sandy River' of Kansas." The reports of 
GiU noted above pertain to fishes collected on the expedition of 
James Hervey Simpson in 1858 and 1859. I find no reference else- 
where in the report of Simpson (1876) confirming the presence of 
George Suckley on this expedition. According to a chart of mag- 
netic dip (Simpson, 1876:227), some members of the expedition 
crossed Big Sandy River ( also known as Big Sandy Creek or Sandy 
Creek), a tributary of the Little Blue River in southern Nebraska, 
on June 12, 1858, and, evidently, again in early October, 1859. 

Species named by Abbott (1861:568-569; 1861:473-474), and by 
Gill (1864:151) and the species reported by Cope (1864, 1865, 
1871 ) were from collections made by Dr. W. A. Hammond. Ham- 
mond was a surgeon assigned to an expedition under the command 
of Lt. F. T. Bryan to explore a route from Fort Riley, Kansas, to 
Bridgers Pass, Wyoming. The itinerary of the expedition is dis- 
cussed further in the chapter concerning species of improbable oc- 
currence in the Kansas River Basin. 



90 University of Kansas Publs., Mus. Nat, Hist. 

In 1875 F, H. Snow published a list of the more common fishes 
found in the Kansas River at Lawrence, Kansas. 

In the period 1880-1900 numerous collections were made and 
recorded in the literature of that time. Serving as a basis for knowl- 
edge of fishes of the Basin were the reports of Gilbert ( 1884, 1885a, 
1885b, 1886, 1889) based on material sent him by F. W. Cragin of 
Washburn University, Topeka, Kansas. A few (mostly types) of 
the specimens collected by Cragin and his co-workers are still 
housed at the United States National Museum. However, the part 
of the collection of Cragin retained at Washburn University was 
mostly destroyed by fire. 

In 1885 Cragin and I. D. Graham ( the latter at Kansas State Col- 
lege, Manhattan) issued (separately) the first and, to date, only 
lists of the fishes known to occur in Kansas. In addition Cragin 
published a note concerning lampreys (1885a) and Graham 
(1885b) listed the fishes housed in the museum of Kansas State 
College (now Kansas State University). Some of the specimens 
acquired by Graham are still preserved in the holdings of Kansas 
State University (Breukelman, 1946:54; Jennings, 1942). 

As mentioned heretofore, O. P. Hay investigated the fish fauna 
of northwestern Kansas and reported thereon in 1887. Part of Hay's 
collections are extant at the United States National Museum. 

In 1891 S. E. Meek (1894) collected in the Blue River at Crete, 
Saline County, Nebraska. Evermann and Cox (1896) reported 
collections made in 1892 and 1893 in the upper Big Blue River 
Basin in Nebraska. 

The State Biological Survey of Kansas actively collected fishes, 
many of them from the Kansas River Basin, at various times since 
1900 but especially in the years 1910-1912 (Breukelman, 1946:55) 
and from 1950 to the present. Many of these collections are pre- 
served in the Museum of Natural History, University of Kansas, and 
the older collections were reported by Breukelman ( 1940b ) . Breu- 
kelman (1946:55) described the work of Minna E. Jewell, who in 
the 1920's collected extensively in Kansas. The collection of Jewell 
is now housed in the Museum of Zoology, University of Michigan. 
Also housed at that institution are the extensive collections made 
by Raymond E. Johnson in Nebraska from 1939 to 1941, part of 
which are from the Kansas River System. Many smaller collections 
from the Kansas River Basin are also housed at UMMZ, notably 
those made by W. Ralph Taylor, John Breukelman, Claude Hib- 
bard and C. W. Greaser. 



Fishes of Kansas River System 91 

Aside from the collections already mentioned there are a number 
of smaller collections taken at various times and places from the 
Kansas River Basin in the holdings of the United States National 
Museum. Among these are a few specimens taken by F. F. 
Crevecoeur, who made early observations on the fishes in the Ver- 
million River near Onaga, Kansas (1903, 1908). 

At the Museum of Comparative Zoology, Harvard, a number of 
specimens are preserved that were taken in the 1800's near Hays 
and Topeka, Kansas. Some of these collections are credited to "R. 
Bliss" or "Dr. L. Watson" of Ellis, Kansas ( See Evermann and Cox, 
1896:360). 

At the Academy of Natural Sciences of Philadelphia there are a 
few old specimens, mostly types, and a number of specimens col- 
lected in the Kansas River near Topeka, Kansas, in 1958 in a hm- 
nological survey under the supervision of Ruth Patrick. 

W. L. Minckley collected in the Big Blue River System of Kansas 
in 1957 and 1958 and J. E. Deacon and A. L. Metcalf made collec- 
tions in the Wakarusa River Basin in 1959. Frank B. Cross has 
made many collections in the Kansas River Basin assisted by various 
students among whom Bernard C. Nelson should especially be 
noted. My own collections have been made at various times be- 
tween 1956 and 1963. The collections mentioned in this paragraph 
are housed in the Museum of Natural History, University of Kansas. 

Species of fishes that have been described from the Kansas River 
Basin are listed in Table 6. As indicated in the table it is likely 
tliat some of the type localities are in error. 

For museums from which material from the Kansas River Basin 

was studied the following abbre\'iations are used: 

ANSP — Academy of Natural Sciences of Philadelphia. 

KU — Museum of Natural History, The University of Kansas, Lawrence. 

KSU — Kansas State University, Manhattan. 

MCZ — Museum of Comparative Zoology, Harvard. 

OAM — Oklahoma State University, Stillwater. 

UMMZ — University of Michigan, Museum of Zoology. 

USNM — United States National Museum. 

ACCOUNTS OF SPECIES 

Ichthyomyzon castaneus Girard 

Four specimens of the "chestnut lamprey" were reported from the 
mouth of Mill Creek by Cragin (1885a: 99). His description of the 
specimens, specifying that all possessed 3 maxillary (supraoral). 



92 



University of Kansas Publs., Mus. Nat. Hist. 



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Fishes of Kansas River System 



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94 University of Kansas Publs,, Mus. Nat. Hist. 

cusps strongly suggests that the specimens were, in fact, 7. castaneus. 
Hubbs and Trautman ( 1937 ) listed no further occurrences of chest- 
nut lampreys in the Kansas River System. Bailey (1959b: 163) re- 
ported I. unicuspis Hubbs and Trautman from the Missouri River 
in South Dakota and suggested that 7. unicuspis might range farther 
upstream in the Missouri System than did 7. castaneus. Cross and 
Metcalf (1963:187) hsted new records of 7, castaneus from the Mis- 
souri River near St. Joseph, Missouri, and in North Dakota and of 
7. unicuspis from Omaha, Nebraska. It seems likely, then, that both 
species formerly penetrated far up the Missouri River and its main 
tributaries; as pointed out by Bailey (1959b: 163), there may have 
been a general pattern of extirpation of lampreys in the region in 
recent years. Cross and Metcalf (1963:187) noted a report by an 
angler in 1960 of a lamprey from the Kansas River near its mouth. 
To my knowledge 7. unicuspis has never been taken from the Kan- 
sas River Basin. 

Scaphirhynchus platorynchus (Rafinesque) 

The shovel-nosed sturgeon is taken frequently in the lower part 
of the Kansas River and its largest tributaries (Map 1). Western- 
most records are from the Republican River, including two early 
collections and one recent collection as follows: (1) USNM 3246 
taken by W. A. Hammond, (2) USNM 3294 taken by Lt. F. T. 
Bryan on June 23, 1856, and (3) KU 4065 taken by F. B. Cross and 
Bernard Nelson at Concordia, Cloud County, Kansas, on June 25, 
1958. (Tlie earlier records have not been included in Map 1 due 
to the lack of specific locality data. ) 

The taxonomic status and some aspects of the ecology of this and 
the related species S. alhus have been discussed by Bailey and Cross 
(1954). 

Scaphirhynchus albus (Forbes and Richardson) 

The pallid sturgeon has been taken only in the lower Kansas 
River. It is much less abundant than S. platorynchus, which occurs 
in the same waters. According to Bailey and Cross (1954:201) 
eight percent of the specimens of Scaphirhynchus observed from 
near Lawrence, Kansas, were S. alhus. However, since 1952, a year 
of unusually high water and flooding, no specimens of albus have 
been taken at Lawrence. 

Bailey and Cross (1954:174) contended that S. albus and S. 
platorynchus are closely related. This being the case the two kinds 



Fishes of Kansas River System 95 

probably diverged from an ancestral stock at a time v^^hen they 
were not sympatric as at present. Of the three preglacial drain- 
ages mentioned herein (Fig. 3) it seems that the widespread S, 
platonjnchus inhabited the Preglacial Teays/Mississippi and pos- 
sibly the Preglacial Plains Stream and some other southwestern 
drainages. The most likely area of origin for S. alhus, on the other 
hand, seems to be the Preglacial Hudson Bay System, whence it 
invaded the present Missouri 'Mississippi System by deflection 
southward of north-flowing streams. 

Polyodon spathula (Walbaum) 

The paddlefish seems restricted to the lower part of the Kansas 
River proper. Snow (1875:140) wrote "Previous to 1874 only two 
specimens of this curious fish had ever been taken at Lawrence, but 
since the building of the dam several have been taken, varying in 
weight from 1 lb. to 20 lbs., and in length from 2 to 5 feet." The 
paddlefish still occurs in the vicinity of Lawrence, Kansas (Cross 
and Hastings, 1956:86) but its rarity is such that a paddlefish 
snagged by an angler in 1963 received sj^ecial attention in the local 
newspaper as an oddity. The westernmost records are from the 
vicinity of Manhattan, Kansas (Graham, 1885a:70; Jennings, 1942; 
364). I have examined one specimen (KSU 4960) taken in 1938 at 
the mouth of Wildcat Creek, Riley County, Kansas. 

^ Lepisosteus platostomus Rafinesque 

The short-nosed gar has been taken only from the Kansas River 
Mainstream, the lowermost part of some of the largest tributaries, 
and from oxbow lakes along the Kansas River (Map 2). Further 
discussion is in the account of L. osseus, below. 

Lepisosteus osseus Linnaeus 

The long-nosed gar is common in the larger, downstream parts 
of the Kansas River and in some of its major tributaries (Map 3). 
The long-nosed gar seems to be more common than the short-nosed 
gar and to ascend smaller tributaries tlian does the latter. Minckley 
( 1949:415) reported L. osseus from 14 stations in the Big Blue River 
System, L. platostomus from only two stations. In the winter of 
1956-57, I observed large numbers of dead long-nosed gars and a 
few short-nosed gars along the Kansas River at Lawrence. 

There is some indication that this species once ranged farther 



96 University of Kansas Publs., Mus. Nat. Hist. 

west in the Kansas River Basin than at present. Hay (1887:247) 
reported it from Beloit on the Solomon River. This record is rep- 
resented by a juvenile (USNM 37959), 82.5 mm in total length and 
17.4 mm in snout-length. An entry (USNM 3244) of February, 
1861, in the catalogue of the Division of Fishes, United States Na- 
tional Museum, lists L. osseiis from the Republican River with Wm. 
S, Wood as collector. Attempts to locate this specimen were un- 
successful. 

Both species of gar reported here, being characteristically species 
of big rivers, enter the Kansas River Basin along the main channels 
from the east. 

Dorosoma cepedianum (LeSueur) 

The gizzard shad is common in the lower part of the Kansas 
River, in its major tributaries, and in reservoirs in the western part 
of the Basin (Map 4). It is another of a large number of fishes, 
typical of big rivers, that ascend the Kansas River from the east. 

Salmo trutta Linnaeus and Salmo gairdneri Richardson 

The brown trout and the rainbow trout have been stocked in a 
few streams and springs in the Kansas River System in Nebraska, 
Colorado, and Kansas. Rainbow trout are placed periodically in 
Elm Creek, Webster County, Nebraska; Rock Creek, Dundy County, 
Nebraska, and in several streams in the vicinity of Wray, Yuma 
County, Colorado. According to members that I have contacted 
of the Colorado Game and Fish Department and of the Game, For- 
estation and Parks Commission, State of Nebraska, rainbow trout 
do not reproduce in these streams. Brown trout have been stocked 
in some tributaries of the North Fork Republican River in Yuma 
County, Colorado, and are said to reproduce there. The streams 
mentioned above are cool, clear brooks deriving their water from 
the area of Pleistocene sands and gravels or from the base of the 
Ogallala Formation where it contacts underlying Cretaceous beds. 

Hiodon alosoides (Rafinesque) 

The goldeye has been taken at numerous localities in the lower 
Kansas River (Map 5) and in its larger tributaries. Hay (1887: 
250) reported it as far upstream as WaKeeney on the Saline River 
and Concordia on the Republican River (I have checked the iden- 
tity of the Concordia specimen — USNM 37932). Another specimen 



Fishes of Kansas River System 97 

from the Republican River (USNM 3548) bears the data "Repub- 
Hcan — Wood," There are reports by fishermen as far west as Scan- 
dia, Republic County, Kansas, on the Republican River, and Salina, 
Kansas, on the Smoky Hill River. Trautman (1957:173, 174) re- 
marked that: 

"The Goldeye appears to be far more tolerant of turbid water than is the 
Mooneye Hiodan tergisus. Tolerance to turbidity is indicated by its presence, 
often in large numbers, in the muddy streams of the Missouri system and those 
of the province of Manitoba. It is therefore possible that the clear-water in- 
habiting Mooneye was fonnerly more numerous in the upper Ohio Drainage 
than was the Goldeye, but that since the waters of tliis drainage have become 
more turbid, the turbid-tolerant Goldeye has become the most numerous." 

This tolerance of turbidity and the present distribution of H. 
alosoides both suggest a western or Plains-origin. Perhaps H. 
alosoides, which also possesses northern affinities, originally in- 
habited the Preglacial, north-flowing Missouri River and became a 
part of the fauna of the Ancestral Plains Stream after deflection of 
the Ancestral Missouri southward into the Ancestral Plains Stream 
Drainage at the onset of glaciation. Subsequently, it has extended 
its range eastward. Hiodon tergisus, on the other hand, may have 
evolved in the Preglacial Teays/Mississippi System. 

Esox lucius Linnaeus 

Although occurring farther north in the Missouri River Basin 
(Evermann and Cox, 1896:415; Bailey and Allum, 1962:33), tlie 
northern pike does not seem to have been native to the Kansas 
River System. Both Cragin (1885b:110) and Graham (1885a:78) 
noted that the northern pike had been introduced in Kansas. Lack 
of subsequent records indicates, however, that these early introduc- 
tions were unsuccessful. More recently northern pike have been 
planted in several reservoirs of Nebraska (I observed a large speci- 
men taken in Harlan County Reservoir), Kansas (Tuttle Creek and 
Webster reservoirs, R. E. Schoonover, personal communication) 
and in Bonny Reservoir in Yuma County, Colorado (McCarraher, 
1961:223). 

Cyprinus carpio Linnaeus 

Graham (1885a: 78) and Cragin (1885b: 109) listed carp as hav- 
ing been planted in Kansas, and Dyche (1914:125) noted that it 
was found in nearly all Kansas streams. At present it occurs 
throughout the Basin. It is less abundant in the smaller, clearer 



98 University of Kansas Publs., Mus. Nat. Hist. 

tributaries having high gradient than in rivers, but it is not com- 
pletely alien to small streams. In the headwaters of Medicine 
Creek at Wellfleet, Nebraska, one of the most turbid creeks in the 
Kansas River Basin (Secchi disc approximately six inches), carp 
approximately one foot long were observed in a brook eight to 15 
feet wide and rarely as deep as three feet. 

Carassius auratus (Linnaeus) 

The goldfish has been reported from a few, scattered localities in 
the Kansas River Basin; it does not seem to have become common. 

Notemigonus crysoleucas (Mitchill) 

The golden shiner has been taken in the extreme eastern and 
extreme western sections of the Kansas River Basin (Map 6) but 
there are no records from the central part. Minckley ( 1959 ) failed 
to take the golden shiner in his extensive survey of the Blue River 
Basin in Kansas. Several of the western records are in or near im- 
poundments. That N. crysoleucas may not be native to these west- 
em waters is given credence by Breukelman (1940:370): "The 
only specimens on record are two series we obtained on the 1938 
field trip; one from Beaver creek below the Lake Atwood dam and 
the other from the Saline river below the Sheridan county dam. At 
both places fishermen reported the species common. Since this is 
an easily recognized fish it was probably not present in any consid- 
erable numbers until recently; the earlier collectors would not all 
have missed it." 

There are no verifiable records of N. crysoleucas from the Kansas 
River Basin prior to the 1930's, although Graham (1885a: 74) listed 
it from "Eastern Kansas." Probably this cyprinid is native to the 
extreme eastern part of the basin but it has not penetrated far 
westward. 

Most records from the extreme eastern section of the basin are 
from creeks. Deacon and Metcalf (1961:316), in a survey of the 
Wakarusa River System, reported the species from 5 stations — all 
on tributaries of the Wakarusa River Mainstream. 

Semotilus atromaculatus (Mitchill) 

Habitat and Distribution. — The creek chub is common in the 
Kansas River Basin except in the drainage of the upper Big Blue 
River (Map 7), an area of especially silty streams with muddy 



Fishes of Kansas River System 



99 



Table 7. — Ranges and Means of Lateral-line Scales for Some Speci- 
mens OF Semotilus atromaculatus. Arrangement of Localities Is North 
TO South by Stream System and East to West Within Each Stream 

System. 



Locality 



Niobrara River Drainage: 

Nebraska, Antelope County, Verdigre Creek 
(KU 7170 and KU 7227) 



Platte River Drainage: 

Nebraska, Dixon County, Logan Creek 
(KU7156) 



Nebraska, Keith County, North Platte River at 
Keystone (KU 4851) 



Colorado, Douglas County, Tributarv of Cherry 
Creek (KU 5518) ": 



Colorado, Larimer County, Big Thompson River 
(KU4774) 

Colorado, Larimer County, Tributary of Cache 
la Poudre River at Liver more (9 September 
1959) (KU 4804) 



(2 September 1960) (KU 5530). 



Wyoming, Albany County, Laramie River, 17 
miles north Laramie (KU 4812) 



Lower Missouri River Drainage: 

Missouri, Moniteau Countv, Moniteau Creek 
(KU6490) '. 



Missouri, Morgan County, Richland Creek 
(KU6458) 



Kansas River Drainage: 

Kansas, Douglas County, Tributary of Kansas 
River (KU 1290) 



Kansas, Shawnee County, Shunganunga Creek 
(KU 1954) 



Kansas, Riley Countv, Deep Creek 
(14 December 1957) (KU 3869). 

(7 September 1961) (KU 7219). . 



Nebraska, Webster County, Elm Creek 

(Republican River Drainage) (KU 7052). 



Kansas, Scott County, Timber Creek (KU 4012) 

Kansas, Wallace County, Rose Creek 

(Smoky Hill River Drainage) (KU 4023 and 
KU 7207) 



Range 


Mean 


55-63 


59.5 


52-63 


58.3 


57-65 


60.4 


55-67 


60.6 


57-62 


58.9 


56-67 


62.2 


56-65 


60.1 


55-63 


59.1 


55-64 


59.7 


56-65 


60.0 


54-61 


57.6 


56-65 


60.0 


54-65 


59.4 


56-67 


60.6 


55-67 


61.0 


53-63 


57.3 


54-63 


58.7 



No. 



10 

16 
10 
25 
10 

10 
15 

20 

16 

12 

11 
10 

14 

18 

16 
9 

14 



100 University of Kansas Publs., Mus. Nat, Hist. 

Table 7. — Concluded. 



Locality 



Range 



Mean 



No. 



Nebraska, Hitchcock County, Frenchman Creek 
at Enders Dam (KU 4855) 

Arkansas River Drainage: 

Missouri, Ripley County, Tributary of Eleven 
Point River (KU 6583) 

Arkansas, Franklin County, Spirit Creek 

(KU6293) 

Arkansas, Benton County, McKisic Creek 

(Elk River Drainage) (KU 4557) 

Oklahoma, Mayes County, Tributary of Neosho 
River (KU 6899) 

Kansas, Upper Neosho Drainage (Composite). . . 

New Mexico, Colfax County, Cimarron River at 
Springer (KU 4218) 

New Mexico, Mora County, Mora River at 
Valmora (KU 4234) 

Red River Drainage: 

Arkansas, Montgomery County, Caddo River 
(KU6159) 

Arkansas, Sevier County, Winters Creek 

(KU6108) 

Trinity River Drainage: 

Texas, Nacogdoches County (Composite) 



56-65 

55-61 

53-60 

54-61 

56-60 
57-63 

52-58 

51-59 

51-55 
51-58 
51-55 



59.5 

58.1 
56.1 
57.6 

58.0 

58.7 

55.0 
53.8 

52.7 
53.6 
52.4 



20 

10 
15 
14 

10 

7 

10 

18 

10 
10 

7 



bottoms. As suggested by its common name, this species prefers 
small^ upland streams. Only two records are known to me from 
the lower 100 miles of the Kansas River Mainstream, although the 
species abounds in creeks on both sides of the mainstream. 

Variation. — A check of some meristic characters and body- 
proportions of specimens from several localities revealed only slight 
variation. West of the Mississippi, scale-size seems to increase 
from north to south. A similar southward increase in scale-size 
was noted in eastern populations by Bailey, Winn and Smith 
(1954:124). Some counts of lateral-line scale are in Table 7. 
Ventral scales tend to decrease in size and to become embedded 
in a thick epidermis in some western tributaries of the upper Platte 
and Kansas river systems. 



Fishes of Kansas River System 



101 



Eye-size diminishes westward, as in several other species of wide 
occurrence in the Great Plains (Table 8). In western streams of 
the Platte and Kansas systems the skin surrounding the eye thickens 
and extends inward slightly, sheathing the peripheral surface of 
the eye. Two mechanisms seem at work here to decrease the 
amount of eye-tissue exposed to the environment, suggesting that 
this reduction has adaptive value to western representatives of this 
species. 

Populations from the Kansas, Arkansas, and Red River systems 
differ subtly from those of the Platte River in possessing (1) 
deeper bodies, shghtly tliicker anteriorly, (2) longer and deeper 
heads and (3) a more decurved anterior dorsal profile. Popu- 
lations from tlie upper Canadian and Pecos rivers in northeastern 
New Mexico are exceptional in possessing a smaller head. 

Zoogeography. — This species has been reported as a mid-Pleisto- 
cene fossil from southwestern Kansas and nortliwestern Oklahoma 
(C. L. Smith, 1958:178; G. R. Smith, 1963:279), and it now occurs 
as a rehct in tlie upper Canadian and Pecos river systems. Probably 
it occurred in the drainage of the Ancestral Plains Stream. How- 



Table 8. — Corneal Width (Expressed in Thousandths of Standard 
Length) of Some Specimens of Semotilus atromaculatus (40-50 mm in 
Standard Length). Localities Are Arranged Generally in an East- 
West Manner. 



Locality- 


Range 


Mean 


No. 


Kentucky, Bullitt County, Rolling Fork River 

Drainage (KU 4105 and 5707) 


69-76 


73.1 


10 


Wisconsin, Iowa County, Tributary to Trout Creek 
(KU5381) 


73-85 


76.5 


26 


Arkansas, Montgomery County, Caddo River 

(Ouachita River Drainage) (KU 6915) 


71-81 


74 8 


10 


Missouri, Morgan County, Richland Creek 

(Missouri River Drainage) (KU 6458) 


67-78 


72 9 


7 


Kansas, Shawnee County, Shunganunga Creek 

(Kansas River Drainage) (KU 1954) 


64-71 


67.5 


10 


Nebraska, Webster County, Elm Creek 

(Republican River Drainage) (KU 7052) 


62-71 


65.6 


12 


Wyoming, Albany County, Laramie River 

(North Platte River Drainage) (KU 4812) 


60-67 


63.2 


12 


New Mexico, Colfax County, Cimarron River at 
Springer (KU 4218) 


60-70 


62.9 


14 



102 University of Kansas Publs., Mus. Nat. Hist. 

ever, diversion of the Kansas River eastward placed any popula- 
tions previously inhabiting that basin in contact with populations to 
the east, in the Mississippi Valley. Presently, populations in the 
Kansas River resemble those to the east, and characters that do vary 
show smoothly clinal change from east to west or northeast to 
southwest. The diflFerences that I found in western creek chubs 
(smaller eye, smaller scales and a chubbier body) seem too slight 
to merit taxonomic recognition. 

Chrosomus erythrogaster (Rafinesque) 

Habitat and Distribution. — At present the southern red-bellied 
dace is found in the Kansas River Basin ( Map 8 ) only in streams of 
the Flint Hills Area, where it is restricted to headwaters of creeks, 
usually near springs. Within the limited confines of this habitat 
it is often abundant. 

Evidence of a former, more widespread distribution is provided 
by the record of Hay (1887:249) from Lenora, Norton County, 
Kansas. I have examined a specimen (USNM 38241), collected 
by Hay at this locality, that has the small scales and looped intestine 
characteristic of C. enjthrogaster and possesses 8 anal rays. Al- 
though suffering much depigmentation, there are still discernible 
two dark lateral bands as in Chrosomus eos Cope and C. erythro- 
gaster. A few scattered melanophores dorsal to the upper band 
suggest the presence, originally, of spots in that region. The mouth 
is long and subhorizontal as in C. erythrogaster, not short and 
strongly oblique as in C. eos. 

Chrosomus erythrogaster seems to be one of a number of species 
that has suffered restriction of range in the Kansas River Basin 
since settlement of the land and subsequent modification of the 
watershed. 

Zoogeography. — To my knowledge the southern red-bellied dace 
does not occur north of the Kansas River Basin in the Missouri 
River System. In the Platte and Niobrara drainages both C. eos 
Cope and C. neogaeus (Cope) are found. To the south in parts 
of the Arkansas River and Red River drainages, C. erythrogaster 
prevails. C. erythrogaster is also common in the Ozark Region, 
including Ozarkian tributaries of the lower Missouri River. It 
seems probable, then, that C. erythrogaster entered the Kansas 



Fishes of Kansas River System 103 

River Basin from the Ozark Region. Probably it has occupied 
the eastern part of the Basin for a long period of time, as streams 
east of the Flint Hills have long flowed eastward toward the Ozark 
Region. 

Hybopsis gracilis (Richardson) 

Habitat and Distribution. — The flat-headed chub has been taken 
only rarely in the Kansas River Basin. All collections are from 
streams with sandy bottoms. Olund and Cross (1961:334-336) 
discussed the habitat of the species in the Great Plains. H. gracilis 
has an unexplained disjunction in its distribution in the Kansas 
River Basin, having been taken in the lowermost part of the main- 
stream and again, after a hiatus of approximately 200 miles, in the 
upper Republican River ( Map 9 ) . There are no records to suggest 
a former, more widespread distribution. 

Zoogeography. — Olund and Cross (1961) recognized two sub- 
species of Hybopsis gracilis. They mapped the typical subspecies 
(PI. 21) as occurring in the Mississippi River, below the mouth 
of the Missouri River, in the Missouri River and some of its 
northwestern tributaries and in the basins of the Mackenzie and 
Saskatchewan rivers of Canada. The other subspecies, H. g. 
gulonella, was found to occur in some western tributaries of the 
Missouri River and in the upper Arkansas, Pecos and Rio Grande 
drainages. Many populations from the Missouri River Basin, in- 
cluding those from the Kansas River Basin, were considered in- 
tergrades between the two subspecies. 

Bailey and Allum (1962:44) suggested that differences in water 
temperature between the Missouri River and its western tributaries 
produce, through phenotypic modification, the morphological dif- 
ferences that were observed by Olund and Cross and thought by 
the latter authors to deserve taxonomic recognition. Bailey and 
Allum (1962:44) noted that the upper Missouri River is fed by 
mountain tributaries and remains relatively cool throughout the 
summer. It should be pointed out, however, that the North Platte 
River, in which H. g. gulonella occurs, \s also fed by tributaries 
carrying melt-waters from mountains in northern Colorado and 
southern Wyoming. Olund and Cross (1961:333) found popula- 
tions of H. g. gulonella at the uppermost stations sampled on the 



6—8449 



104 University of Kansas Publs., Mus. Nat. Hist. 

North Platte River where summer temperatures of water presum- 
ably are colder, whereas downstream where waters become warmer 
there was evidence of intergradation with H. g. gracilis. This 
suggests a zone of intergradation of difiFerent kinds of fishes rather 
than a purely phenotypic response. The upper Arkansas River is 
also fed by mountain streams carrying meltwaters tliroughout the 
summer from the massive Sawatch Range and other mountain 
ranges of Central Colorado. Specimens collected from the upper 
Arkansas River also have the phenotype of H. g. gulonella. Flat- 
headed Chubs were taken at Florence, Fremont County, Colorado, 
on September 7, 1959, in water at 65° F. It can be argued, of 
course, that it is not known whether these fish undergo their early 
development in these waters. However, the upper Arkansas River 
has few tributaries and most of these also drain mountainous areas. 
The present distribution of H. g. gulonella suggests that it dif- 
ferentiated in the Ancestral Plains Stream. If so, a question remains 
concerning when and where H. g. gracilis was sufficiently separated 
from the stock of incipient H. g. gulonella to diverge racially from 
it. Possibly H. g. gracilis inhabited the Preglacial Missouri River 
described by Meneley, Christiansen and Kupsch (1957). This 
stream possessed, as headwaters, the present-day upper tributaries 
of the Missouri River but flowed northeastward into Hudson Bay. 
Perhaps with the advent of glaciation and diversion of the upper 
Missouri southward into the Ancestral Plains Stream, populations 
of H. g. gracilis commingled with H. g. gulonello, the latter kind 
having held forth in that segment of the Ancestral Plains Stream 
south of the divide between Hudson Bay and the Gulf of Mexico 
that lay somewhere in the vicinity of present-day South Dakota 
(Flint, 1955:148). Such a northern origin might account for the 
preference of H. g. gracilis for the cooler waters of the Missouri 
River over the adjacent downstream parts of its western tributaries 
and for its reinvasion of the Saskatchewan and Mackenzie drainages 
of Canada since the last glaciation there. The hypothesis of an 
origin in the ancestral north-flowing Missouri System might explain 
the absence of H. g. gracilis from the upper Mississippi and Ohio 
rivers and in the Red River of the North. A cold-tolerant fish in- 
habiting an eastern relicitium in the central Mississippi Valley would 
be expected to have ascended the Mississippi and to have entered 
the Red River through the well-documented connection to Lake 
Agassiz offered by the River Warren (Underbill, 1957:7). 



Fishes of Kansas Rutr System 



105 



Hybopsis biguttata (Kirdand) 

Habitat and Distribution. — At present the horny-headed chub 
seems to have one of the most restricted ranges of any fish found 
in the Kansas River Basin (Map 10). It has, in recent years, been 
taken only in Mill Creek, Wabaimsee County, Kansas (KU 2968). 
There are a number of earlier records that indicate that the species 
was formerly much more widespread in the Basin. Specimens that 
I have examined are listed in Table 9. 

In discussing the status of this species in northwestern Kansas, 
Breukelman (1940:370) noted that, "Careful seining of Hay's locali- 
ties at the same time of the year in 1938 failed to reveal a single 
specimen. Questioning of fishermen indicated that if present in 
northwestern Kansas, the species is very rare. In view of the fact 
that large specimens of this clearwater species are not apt to be 
misidentified, a change in the fauna of the area seems to have 
occurred." In the case of Rock Creek, Douglas County, the species 
seems to have disappeared since 1941 (Table 9) as frequent col- 
lections made since 1950 have failed to reveal it there. 

Elsewhere in the Missouri River System H. biguttata is known 
at present only from a few rather widely separated areas : ( 1 ) the 
Ozark Region of Missouri, (2) a few creeks in eastern Wyoming 



Table 9. — Early Records of Hybopsis biguttata (Kirtland) From the 

Kansas River Basin. 



Museum number 


Locality 


Date collected 


USNM3552 


"Republican" 

Fort Hays, Kansas 

Fort Hays, Kansas 

Fort Hays, Kansas 

Saline River, 

Trego County, Kansas 

Rock Creek, 

Douglas County Kansas 

Rock Creek, 

Douglas County, Kansas 

Rock Creek, 

Douglas County, Kansas 


probably 1857 
1871 


MCZ 18480 


MCZ 18483 


1871 


UMMZ 86429 


1871 


KU8188 




KU 403, 416, 420, 423-429, 
587, 593 


1898 


KU389, 391, 392 




UMMZ 63170 


1899 




1924 



106 University of Kansas Publs., Mus. Nat. Hist. 

(Simon, 1946:71 and KU 4813), and (3) the area drained by the 
Big Sioux and Floyd rivers in Iowa (Cleary, 1956: Map 37). Older 
records (Hay, 1887:250, 252; Evermann and Cox, 1896:410; Ellis, 
1914:62) indicate that the species was more widespread in the 
Missouri River Basin in the past. 

Disjunctions in range that now exist, with preservation of only a 
few semi-relict populations in especially favorable streams, suggest 
a process of extirpation over much of the Missouri River Basin. A 
similiar pattern was observed by Trautman (1957:294), who traced 
the decrease in abundance of the horny-headed chub in several 
Ohio streams. 

Greene (1935:75) described the preferred habitat of the homy- 
headed chub as ". . . clear water, moderate current with some 
gravelly riffles and a moderate amount of aquatic vegetation." Some 
or all of the requisites specified by Greene are thought to have dis- 
appeared from much of the basin of the Kansas River during the 
years since settlement began. Water in streams has, in many 
parts of the basin, decreased in clarity and in constancy and volume 
of flow. Decrease in constancy of flow may have been deleterious 
to growth of aquatic plants. Siltation due to cultivation may have 
increased fine sediment on stream-bottoms at the expense of coarser 
sediments, especially in the central part of the basin. Greene 
(1935:76) and Lachner (1952:435) pointed out the need of H. 
biguttata for gravelly areas at the time of spawning; gravel is needed 
to build the "nest" characteristic of the species. 

Zoogeography. — As pointed out by Bailey and AUum (1962:121), 
H. biguttata is not a typical inhabitant of large, turbid streams. 
Probably it made its way up the Missouri River System from the 
Ozark Region at times when conditions were less inimical, or by a 
process of "tributary-hopping." One now sees this process some- 
what in reverse, with extirpation from all but a few of the tributaries 
in which the species had established populations prior to settlement. 

Hybopsis storeriana (Kirtland) 

The silver chub is taken frequently in the lower mainstream of 
the Kansas River but is rare upstream (Map 11). The westernmost 
record is UMMZ 134348 — Republican River, IK miles east of Benkel- 
man, Dundy County, Nebraska, July 24, 1939 (one specimen). 
Other marginal records to the west include (1) KU 439 — Repub- 



Fishes of Kansas River System 107 

lican River, Clay Co., Kansas, 1910, (2) KU 8129— Kansas River at 
Junction City, Geary Co., Kansas, May 9, 1964, (3) UMMZ 122191 
—Wildcat Creek, Riley Co., Kansas, September 7, 1923 and (4) 
KU 3810— Blue River, 1 mile south of Rocky Ford Dam, Riley Co., 
Kansas, April 6, 1957. Hay (1887:250) reported, with some reserva- 
tion, H. storeriana from WaKeeney, Trego County, Kansas. Breukel- 
man (1940:370) questioned the validity of this identification. I 
was unable to find specimens of H. storeriana among Hay's collec- 
tions from WaKeeney at the U. S. National Museum. There are, 
ho\\'e\'er, 3 specimens (USNM 37938) referable to this species 
bearing the locality data "Wallace, Kansas." Hay (1887:250) men- 
tioned that the specimens from WaKeeney had lost their scales and 
were otherwise in poor condition. This characterization applies to 
USNM 37938 and it seems possible that the locality data are in error. 
(I have not included H. storeriana in Table 5, which Hsts species 
taken by Hay at Wallace. ) Perhaps the failure to take H. storeriana 
frequently in upper tributaries of the Kansas River System in recent 
years is related to a decreased volume of flow in those streams. In 
this respect H. storeriana is like Hiodon alosoides and Lepisosteus^ 
osseus. Several writers have pointed out the proclivity of the silver 
chub for large streams. Forbes and Richardson (1909:167) noted 
that it was a fish of larger streams and lowland lakes. Greene 
(1935:79) characterized it as typical of "large-water fishes of south- 
ern range." Greene's comment seems pertinent to this species in 
the Kansas River Basin as it seems to have penetrated from the east 
along the larger streams, 

Hybopsis meeki Jordan and Evermann 

Habitat and Distribution. — The sickle-finned chub has been taken- 
only in the lower mainstream of the Kansas River and seems rare 
there (Map 12). Three records, consisting of one specimen each, 
are known to me: KU 1842 and KU 2172 taken on October 24, 1951, 
and April 5, 1952, respectively, below the dam at Lawrence, Douglas 
County, Kansas, and one specimen taken on April 20, 1962, at 
Eudora, Douglas County, Kansas, but not preserved. 

Zoogeography. — Bailey and Allum (1962:48, Fig. 5) have 
mapped known records of the sickle-finned chub and pointed out 
its virtual restriction to the Missouri River and that part of the 
Mississippi River downstream from the mouth of the Missouri. 



108 University of Kansas Publs., Mus. Nat. Hist. 

Such a distribution suggests an origin in the ancestral, north-flowing 
Missouri River. Possibly a stock ancestral to this species and to 
Hybopsis storeriana gave rise to the former in the ancestral, north- 
flowing Missouri and to the latter in the Teays/Mississippi System 
in pre-Pleistocene time. From the upper Mississippi H. storeriana 
penetrated northward soon enough after the last glacial retreat to 
enter the Red River (Underbill, 1957:29); whereas, in the upper 
Missouri River it has penetrated only as far northward as south- 
eastern South Dakota according to Bailey and Allum (1962:46). 

Hybopsis aestivalis hyostoma (Gilbert) 

Habitat and Distribution. — Most records of the speckled chub 
are from the sandy mainstream of the Kansas, Big Blue, and Re- 
publican rivers (Map 13). Available records suggest that it ascends 
far upstream in the Republican River but not in the Smoky Hill 
River. The same is true of Hybopsis gelida and Hybopsis gracilis. 
The reason for this is not clear. 

In the summer of 1961, H. a. hyostoma was abundant in the 
Kansas River below Bowersock Dam at Lawrence, Douglas County. 
Abundant populations are also indicated by a collection of 122 
specimens (UMMZ 135100) by R. E. Jolinson on July 17, 1940, in 
the RepubHcan River, one mile south of Indianola, Red Willow 
County, Nebraska. This collection contained females distended 
with eggs as did another collection taken on the following day in 
Blackwood Creek, Hitchcock County, Nebraska. 

Variation and Zoogeography. — As pointed out by Greene (1935: 
79) H. aestivalis seems to have southern affinities, judging from its 
occurrence in Gulf Coast drainages from the Rio Grande to the 
Choctawhatchee River in Florida (Yerger and Suttkus, 1962:327). 
Northward, the species is a hesitant invader. Bailey and Allum 
noted (1962:108) that the northernmost record in the upper Mis- 
sissippi Drainage is in Vernon County, Wisconsin. (Another pe- 
ripheral record in this area is USNM 118910 taken by Surber at 
Brownsville, Houston County, Minnesota in 1928.) Bailey and 
Allum (1962:108) further stated that there are no records of H. 
aestivalis above the Platte River Drainage in the Missouri River 
Basin. In these preferences H. aestivalis exhibits a tendency op- 
posite to that of H. gracilis and H. gelida, which flourish northward 
and at higher elevations. 



Fishes of Kansas Ri\'er System 109 

Hybopsis aestivalis, as now recognized, comprises two diverse 
kinds as regards number of barbels. One, a widespread, two- 
barbeled kind, is found in much of the Mississippi River System and 
in some Gulf Coast drainages. The other, a four-barbeled kind, is 
found in the Arkansas and Red rivers in the west and in the Es- 
cambia and Choctawhatchce rivers in the southeast (Yerger and 
Suttkus, 1962:327). To my knowledge there has been no adequate 
demonstration of intergradation between these kinds. The problem 
of relationship of the populations of H. aestivalis occupying the vari- 
ous streams of the Gulf Coast and southwestern tributaries of the 
Mississippi River is complex and in need of study. 

Within the range of the widespread two-barbeled chub (to 
which populations from the Kansas River System appertain) there 
is variation from east to west; smaller eyes, smaller scales, a more 
naked venter, longer barbels, and smaller melanophores character- 
ize western populations. Some writers have considered this varia- 
tion deserving of taxonomic recognition. Breukelman (1940:380) 
employed the name "Extrarius aestivalis: sesquialis x tetranemiis" 
for speckled chubs from the Kansas River System. Breukelman 
seems to have appreciated the fact that in the characters mentioned 
above, which distinguish Missouri River populations from those of 
the Ohio River, that the former parallel to a slight extent those 
found in the Arkansas and Red rivers. Notwithstanding this paral- 
lelism, the difference in number of barbels, a much smaller eye 
and a more extreme decurvature of the snout readily distinguish 
populations in the Arkansas and Red rivers from Missouri River 
representatives. 

As this fish inhabits the lai-ger rivers of the Mississippi Valley, 
gene-flow must be considerable between populations. Some speci- 
mens that I have observed from the Mississippi River seem interme- 
diate between those found in the Ohio and Missouri Rivers whereas 
other specimens closely resemble one or the other. The Mississippi 
River probably acts as a "mixing bowl" for this and other large 
river species, receiving gene flow from its major tributaries, mixing 
these strains and probably providing gene flow back towards tribu- 
taries as well, thereby slowing any tendency toward speciation. As 
the zone of intergradation between extremes of the two-barbeled 
fraction of H. aestivalis in the Mississippi River System is wide and 
ill-defined it seems best to consider it one taxon in accordance with 



110 University of Kansas Publs., Mus. Nat. Hist. 

the principles suggested by Bailey, Winn and Smith (1954:148-149). 
In this case, however, retention of a trinomial is necessary at least 
until such time as the status of the four-barbeled kinds and other 
named subspecies of the Southwest is elucidated. As Hybopsis 
aestivalis hyostoma (Gilbert) is the oldest available name for the 
two-barbeled form of the Mississippi Basin, it is here employed to 
include populations of the Kansas River. 

As mentioned above, Breukelman (1940:380) employed the name 
"sesquialis" in referring to Kansas River populations of this species. 
However, "sesquialis" is a nomen nudum. An earlier and seemingly 
available name, applicable to populations of the upper Missouri 
Basin, is Hybopsis montanus Meek, Although this name was placed 
in the synonymy of Notropis stramineus by Bailey and Allum 
(1962:68), Meek's description (1884:526) of a barbeled minnow 
seems to apply to H. aestivalis and he allied H. montana with H. 
aestivalis in this description. Three specimens among the types in 
the national collection, labeled H. montanus and bearing the num- 
ber indicated by Meek in the original description (USNM 36882), 
are H. aestivalis. There are two labels in the jar, bearing the in- 
scriptions (1) "Hybopsis montanus, E. Slope Rocky Mts., F. V. 
Hayden" and (2) "Hybopsis montanus, Jordan and Meek (Type) 
Upper Missouri Region. Dr. F. V. Hayden." In number of barbels, 
eye-size, scalation, head-shape, and body proportions, these speci- 
mens are like H. aestivalis from the Missouri River Basin. 

Hybopsis gelida (Girard) 

Records of the sturgeon chub in the Kansas River System were 
listed by Cross (1953:90) and Bailey and Allum (1962:46, 47). The 
species has been taken on numerous other occasions in the Kansas 
River near Lawrence and Eudora, Kansas (Map 14). Two recent 
additional records are: (1) KU 8111 — Kansas River at Willard, 
Shawnee Co., Kansas, May 9, 1964 (1 specimen) and (2) KU 8124— 
Smoky Hill River at U. S. Highway 77, south of Junction City, 
Geary Co., Kansas, May 9, 1964 (3 specimens). These last- 
enumerated collections bridge, to some extent, a hiatus that pre- 
viously existed between downstream records in eastern Kansas and 
records in the upper Republican River in southwestern Nebraska. 

Cross (1959:90) noted that H. gelida was locally abundant in 
the Kansas River near Lawrence, 42 specimens having been secured 
there between October, 1951, and June, 1952 (years of heavy rain- 



Fishes of Kansas River System 111 

fall). UMMZ 135099 also consists of a large series (54 specimens) 
taken in the Republican River, one mile south of Indianola, by 
R. E. Johnson on July 17, 1940. The present distribution of H. 
gelida almost exclusively in the Missouri River System suggests 
that it may have evolved in the Preglacial north-flowing Missouri 
System. Possibly the same ancestral stock gave rise in the Teays/ 
Mississippi and Preglacial Plains System to H. aestivalis. 

Phenacobius mirabilis (Girard) 

Distribution ami Habitat. — The sucker-mouthed minnow occurs 
widely in the Kansas River Basin and in habitats ranging from the 
Kansas River Mainstream to small, upland tributaries (Map 15). 
The largest concentrations were found in tributaries of the Wakarusa 
and Delaware rivers in the eastern part of the basin. 

Zoogeography. — Knowledge concerning the present and past 
distribution of P. mirabilis suggests that it evolved in the Ancestral 
Plains Stream. The species is widely distributed and common in 
the streams that probably comprised this drainage. 

Minckley and Craddock (1962:374) suggested a possible western 
origin for this species, stating, "Of the five species of Phenacobius, 
one, P. mirabilis, may have arisen from stock isolated west of the 
St. Louis Embayment and/or the line of Pleistocene glaciation 
(possibly in the Ozarldan Region). This form became adapted for 
Hfe in the sandy, variable streams of the Plains . . ." The An- 
cestral Plains Stream probably was the western drainage involved. 

P. mirabilis has invaded some waters peripheral to the drainages 
that were, in part, contained in the Ancestral Plains Stream, but its 
penetration has been slight in some directions and probably of a 
recent nature in others. 

Presumably, after western drainages were diverted into the 
middle reaches of the Mississippi System, this fish followed the 
Missouri River downstream. In the Mississippi River it penetrated 
northward to southwestern Wisconsin where Greene (1935:115) 
characterized it as a fish of southern origin, and to southeastern 
Minnesota (Underbill, 1957:15). Eastward from the Mississippi 
its spread seems to have occurred partly in recent time and to have 
been rapid. Forbes and Richardson (1909:159) noted, "This little 
fish is, in Illinois, upon the eastern border of its range." Trautman 
(1957:323) stated, "Before 1800 this plains-inhabiting species pre- 
sumably ranged only as far eastward as the Mississippi River, except 
perhaps for outlying and/or existing relict, eastern populations." 



112 University of Kansas Publs., Mus. Nat. Hist. 

Trautman traced the invasion of this species eastward into and 
across parts of Ohio, correlating the extension of range with the 
spread of land-use practices that resulted in an increase in turbid, 
prairie-type streams with silty bottoms. Zahuranec (1962:843) 
recorded an additional eastward penetration of this species in Ohio, 
which he also correlated with increased siltation of streams. 

Northward from the Platte River in the Missouri River Basin, P. 
mirabilis has not penetrated far. According to Bailey and Allum 
(1962:54), it has not been reported north of the Platte River in 
Nebraska and only once in South Dakota. However, Cleary ( 1956, 
Map 40) reported it from several tributaries of the Missouri River 
in northeastern Iowa. 

Southward in the Great Plains Region P. mirahilis is found in 
the Red River System (Lewis and Dalquest, 1957:42; Riggs and 
Bonn, 1959:162), and rarely in some coastal drainages of Texas 
according to Hubbs and Herzog (1955:69-71). Probably, part of 
the Ancestral Red River was incorporated in the Ancestral Plains 
Stream, and possibly one or more of the coastal streams of Texas 
had connections to this system. The evidence of Quinn (1957:160) 
for widespread alluviation in Sangamonian time in the lower parts 
of Texas coastal streams such as the Brazos and Colorado and 
eventual transgression of interstream divides might, in addition to 
extensive flooding (Hubbs and Herzog, 1955:70), have led to major 
interchanges of faunas in streams of this region. Such interchange 
may account for the infrequency, noted by Clark Hubbs (1957: 
102), with which species-ranges correlate with present stream- 
systems in Texas. 

Notropis atherinoides Rafinesque 

Habitat and Distribution. — In the Kansas River Basin the 
emerald shiner is clearly a species of the larger rivers. All collec- 
tions in the Basin are from the mainstream of the Kansas and 
Republican rivers or from the lowermost parts of the largest tribu- 
taries of these streams (Map 16). Although seemingly tolerant of 
turbid streams with sandy bottoms N, atherinoides does not pene- 
trate far west in the Basin, even though sandy streams are common 
westward. 

Variation. — Recently Bailey and Allum (1962:57) placed No- 
tropis percobromus (Cope) in the synonymy of N. atherinoides. 
Previously, N. percobromus had been considered (Hubbs, 1945: 



Fishes of Kansas River System 113 

16-17) a related but distinct species inhabiting the Great Plains. 
Bailey and Allum (1962:57) found that "Plains collections average 
somewhat chubbier in body form, as measured by head depth, 
body depth, or peduncle depth, and the head is usually a bit 
longer . . . But these and other characters vary with locality 
and are not geographically consistent." Variability in body propor- 
tions was observed also in series examined from the Kansas River 
Basin. For example, means of body depth (taken at dorsal in- 
sertion) divided by standard length varied from 205 to 229 for 
collections that I examined from the Kansas River Basin. Three 
collections made in the Kansas River in Douglas County, Kansas, 
yielded means as follows: 205 (November, 1957); 207 (October, 
1951); and 212 (April, 1961). This same proportion also showed 
much variation in three series of ten specimens each from the 
Arkansas River Basin: 201 (September, 1951, North Canadian 
River, Beaver County, Oklahoma); 214 (April, 1953, Spring River, 
Cherokee County, Kansas); and 225 (June, 1958, Medicine River, 
Barber County, Kansas). Despite this variability, which is prob- 
ably due in part to seasonal nutritional or reproductive fluctuations, 
populations in the Great Plains usually have mean \'alues greater 
than 200; populations east of the Mississippi yield mean values less 
than 200 for the same character. Measurements of distance across 
the exposed part of the eye revealed less variability than in the case 
of body depth. Western populations seem consistently to have a 
smaller exposed surface. However, this smallness is achieved dif- 
ferently in populations in the Kansas River System than in those 
in the western part of the Arkansas River System. In the latter the 
cornea is actually small, but in the former the cornea is larger, ap- 
proaching the size found in eastern populations. The periphery of 
the cornea is covered by an ingrowth of thin skin. Although dif- 
ferent mechanisms are at work, approximately the same area of 
the cornea is exposed in both cases. Distance across the exposed 
surface of the eye divided by standard length yielded means of 74 
to 77 for six lots from the Kansas River System and 73-75 for three 
lots from the Arkansas River System in Kansas and western 
Oklahoma. 

Zoogeography. — Possibly a member of the Notropis atherinoides 
complex occupied the Ancestral Plains Drainage. This seems likely 
since related kinds are found in the Brazos and Rio Grande ( Bailey 
and Allum, 1962:60), suggesting an early penetration south westward 



114 University of Kansas Publs., Mus. Nat. Hist, 

by members of the group. Also the weak cHnal differences seen in 
Plains populations may be a heritage from a slightly differentiated 
stock of the Ancestral Plains Drainage and not due entirely to 
ontogenetic responses to environmental stimuli as suggested by 
Bailey and AUum (1962:60). 

Notropis rubellus (Agassiz) 

Habitat and Distribution. — In the Kansas River Basin the rosy- 
faced shiner seems to be almost confined to Mill Creek in Wa- 
baunsee County, Kansas, where it is common (Map 17). This is a 
clear, upland stream having rubbly bottoms and frequent riflfles. 
Such a restricted range might provide grounds for suspecting trans- 
fer of this species through Lake Wabaunsee on the headwaters of 
Mill Creek from the Neosho or Marais des Cygnes rivers to the 
south. However, Gilbert (1886:208) reported "Notropis rubrifrons" 
from a tributary of Mill Creek and from Blacksmith Creek immedi- 
ately east of Mill Creek many years before construction of Lake 
Wabaunsee. Gilbert specified that "The males were flushed with 
brick-red on head and bases of fins." This coloration together with 
the fact that the specimens came from small, upland creeks make 
it probable that Gilbert's specimens were actually N. rubellus and 
not the related species of larger streams, N. atherinoides. The only 
other record of N. rubellus from the Kansas River Basin is that of 
Minckley (1959:420) based on one specimen (KU 4195) taken in 
a small tributary of the Big Blue River. 

Proportional measurements made on specimens from Mill Creek, 
from the Neosho River System and from the Missouri River System 
( Osage River in Kansas and Richland Creek, Morgan County, Mis- 
souri) suggested that affinities of the specimens from Mill Creek 
were with those from tributaries of Missouri River (Table 10). 
These populations had deeper bodies and a larger head and eye 
than did specimens from the Neosho River. 

Zoogeography. — Notropis rubellus is unknown in the Missouri 
River Basin north of the Kansas River (Bailey and Allum, 1962:60) 
and, therefore, probably did not disperse downstream from the 
north. Ingress to the System was probably from the northern 
Ozarks to the east. The species may long have inhabited the 
eastern part of the Basin as this part has consistendy drained east- 
ward into the Mississippi System. The population in Mill Creek 
seems semi-relict; possibly the species is on its way to extirpation 
in the Basin. 



Fishes of Kansas River System 



115 



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116 University of Kansas Publs., Mus. Nat. Hist. 

Notropis umbratilis (Girard) 

Distribution and Habitat. — All collections of the red-finned shiner 
are from the eastern part of the Kansas River Basin, although none 
is from the Kansas River Mainstream (Map 18). Within this basin 
the species varies considerably in abundance. Deacon and Metcalf 
(1961:316) found it to be the most abundant species at several 
stations, especially in the lower and middle parts, on tributaries to 
the mainstream of the Wakarusa River, but Minckley (1959:420) 
found only one specimen in an intensive survey of the fishes of the 
Big Blue River System. The only record outside the eastern part 
of the Basin is from Lenora, Kansas (Hay, 1887:249), on the North 
Fork of the Solomon River. I have examined these specimens 
(USNM 38232) and find them to be N. umbratilis. The western- 
most records since that of Hay are: ( 1 ) KU 8133 — Kans., Dickinson 
Co. at Rock Springs 4-H Camp, May 9, 1964 (20 specimens), (2) 
KU 8211 — Kans., Dickinson Co., Lyon Cr., 3/2 mi. northeast of 
Herrington, April 29, 1962, and (3) KU 8225— Kans., Morris Co., 
Clarks Cr., 4 mi. south and 2 mi. west of Skiddy, April 29, 1962 
(1 specimen). 

Variation and Zoogeography. — Hubbs and Ortenburger (1929b: 
87) allocated specimens from the Arkansas and Red rivers in Ar- 
kansas and Oklahoma to Notropis umbratilis umbratilis. Males in 
breeding condition from the Kansas, Marais des Cygnes, and Ar- 
kansas river drainages in Kansas possess, in common, many small 
tubercles on the entire opercle, a character not shared by populations 
that I have examined from east of the Mississippi River nor by the 
specimen depicted by Trautman (1957: Fig. 79). Hubbs and Orten- 
burger (1929b: 87) noted that the tubercles of N. u. umbratilis 
"cover the entire head." However, it seems unwise to assign 
specimens from the Kansas River Basin to N. u. umbratilis or another 
subspecies pending further study of this difficult group, the com- 
plexities of which are in need of thorough investigation. 

Lack of penetration westward of this species and its widespread 
occurrence in the Ozark Region, suggest for western subspecies of 
N. umbratilis an area of origin in western tributaries of the Teays/ 
Mississippi System draining the Ozark and Ouachita highlands and 
adjoining areas to the south. Possibly the fish was not present in 
the Ancestral Plains or Ancestral Missouri River drainages. Mor- 
phological adaptations suggestive of existence in the Great Plains 
are not clearly observable, although there seems to be a tendency 
towards a relatively larger head in progressing westward in both 



Fishes of Kansas River System 117 

the Arkansas River and Missouri River drainages (Table 12, un- 
published, of doctoral dissertation by A. L. Metcalf, filed in Univ. 
Kansas Library, 1964). 

Notropis cornutus (Mitcliill) 

Habitat and Distribution. — Collections made in the 1800's suggest 
that the northern common shiner then occurred widely in the Kansas 
River Basin (Map 19). Presently it seems to be restricted to two 
separate areas: (1) in streams draining the Flint Hills and Osage 
Cuestas and ( 2 ) in the headwaters of Smoky Hill River and possibly 
in the upper Republican River System in western Nebraska. 
Whether the species occurs naturally in the upper Republican is 
doubtful, as the only record is of three specimens (UMMZ 134375) 
taken from Champion Lake, Chase County, Nebraska, by R. E. 
Johnson on July 25, 1939. Champion Lake is an impoundment on 
Frenchman Creek. 

In the Kansas River Basin, Notropis cornutus is almost restricted 
to clear, spring-fed creeks. Probably the siltation and other modi- 
fications that streams over much of the Basin have suffered due to 
agricultural practices have contributed to a reduction in range of 
this species. It may never have been common in the Glacial Till 
Plains and Loess Plains districts of the Basin. In the Stranger 
Creek System, for example, it is found only in areas where till has 
been removed and the underlying Oread Escarpment chiefly de- 
termines stream physiography. Cleary (1956: Map 44) indicated 
a similar scarcity of this species in the till plains of southeastern 
Iowa and Harlan and Speaker (1956:95) wrote, "This shiner has 
an ecological preference for clear water, and reaches its greatest 
abundance in the upstream tributaries to the major inland rivers. 
As collections progress dowTistream in the more turbid waters, the 
common shiner becomes reduced in numbers." 

Zoogeography. — This species may have originated in Pliocene 
time according to Gilbert (1961:189). N. cornutus must have been 
at some time and to some degree isolated from the closely related 
N. chrysocephalus in order for speciation to take place. Since much 
of the unglaciated part of the Teays/Mississippi Drainage is pres- 
ently occupied by the latter species, N. cornutus probably originated 
in a drainage to Hudson Bay or to the eastern, Atiantic Coast or 
both. That it did not originate in the Ancestral Plains Stream 
System is indicated by its limited penetration southwestward at the 
present time, and by the absence of fossils or historical records of 
this species in the southern Great Plains. 



118 University of Kansas Publs., Mus. Nat. Hist. 

Notropis shumardi (Girard) 

The synonymy, characters and distribution of the silver-banded 
shiner have recently been clarified and discussed by Gilbert and 
Bailey (1962). They map several localities of occurrence in the 
Missouri River from Missouri to South Dakota but none in the 
Kansas River System. I have seen only one specimen (MCZ 
31885) from the Kansas River. This specimen is 37.0 mm in 
standard length and bears only the data "Topeka, Kansas." This 
specimen possesses eight anal rays (the anal fin is well-preserved); 
pharyngeal teeth are 2:4-4:2; the origin of the dorsal fin is only 
slightly behind that of the pelvic fin. Although the date of capture 
is not specified it seems that this was in the 1800's for the following 
reason: (1) the fish appears to have been preserved for a long 
time and (2) fishes taken in 1871 at Hays, Kansas, bear MCZ lot 
numbers close to that (18435) originally borne by specimen under 
consideration, 

Notropis lutrensis lutrensis (Baird and Girard) 

Distribution and Habitat. — The red shiner is one of the two most 
ubiquitous fishes in the Kansas River Basin — the other being Pime- 
phales promelas ( Maps 20 and 27 ) . Almost every stream, large or 
small, from which collections exist has yielded this species. The 
only exceptions are headwater tributaries of the Big Blue River in 
Nebraska (Lincoln Creek and North Fork Blue River), which are 
exceedingly silty streams originating in an area mantled both by 
loess and glacial till (Fig. 2). 

Zoogeography and Variation. — The red shiner is a species of the 
southern and southwestern Great Plains in terms of its present 
distribution, its preferred habitat, and its probable area of origin. 
Some of the environmental conditions that N. I. lutrensis tolerates 
in streams of the Great Plains are turbidity of water, silty and 
sandy bottoms, fluctuations in water level, intermittency, and ex- 
tremes in water temperature (especially in shallow creeks of the 
High Plains). N. lutrensis is widespread in Atlantic Drainages of 
the southwestern United States and northern Mexico (Hubbs, 
1940a: 6-9; Koster, 1957:67; Meek, 1904:72) and, as pointed out by 
Gibbs (1957:192), a southwestern origin seems likely for the 
"lutrensis-ornatus" group. 

Presumably the Ancestral Plains Stream early received popula- 



Fishes of Kansas River System 119 

tions of N. lutrensis from the south. The species presently abounds 
in streams of the Great Plains as far north as the Platte System. 
In the Niobrara River it seems less common and it has not been 
taken in the western tributaries of the Missouri River in South 
Dakota, according to Bailey and Allum (1962:63). (The Pre- 
glacial Plains Stream System ( Fig. 3 ) possibly extended no farther 
to the north than the present Platte Drainage. ) From the hypothe- 
sized southwestern center of origin this species has not pene- 
trated far eastward. In Iowa (Cleary, Map 49) it occurs widely 
in the southwestern part of the state but becomes rare northeast- 
ward. N. lutrensis was not found by Underbill (1957:18) in tribu- 
taries of the Mississippi River in Minnesota nor was it recorded from 
Wisconsin by Green ( 1935 ) . In Illinois the species was interpreted 
by Forbes and Richardson (1909:144) as a "western" species. In 
Mississippi Cook (1958:112) reported it only from the Mississippi 
River Drainage. 

Counts were made of the number of circumferential scales on 
series of N. I. lutrensis (10 per series) from the Platte, Kansas, 
and Arkansas river systems. The range of means for six series 
from the Platte was 25.9-26.7; for 14 series from the Kansas System 
26.0-27.6 and for nine series from the Arkansas River in Kansas, 
western Oklahoma, and eastern Colorado 26.2-28.6. These figures 
suggest a slight increase in number of circumferential scales south- 
ward in the Great Plains. There were no discernible trends from 
east to west, however, in the streams considered. In the Kansas 
River Basin from east to west the following means were obtained: 

26.4 — 26.9 — 26.0 — 27.5 — 26.3 — 26.8 — 26.9 — 27.4 — 27.6 — 

26.5 — 26.9 — 27.3 — 26.3 — 27.5. 

Notropis blennius (Girard) 

Distribution and Habitat. — The distribution of tlie river shiner 
in the Kansas River System is disjunct (Map 21). It has been 
taken (1) in the mainstream of the Kansas River near Lawrence, 
Kansas, and (2) in the upper reaches of the Republican and 
Smoky Hill river systems in western Kansas and Nebraska. The 
species has been taken on several occasions in the Missouri River 
in Kansas and might, therefore, be expected to enter the lower part 
of the Kansas River. The distributional pattern of the upstream 
populations in certain shallow, clear streams resembles greatly 



7—8449 



120 University of Kansas Publs., Mus. Nat. Hist. 

that of Htjbognathus Jiankinsoni. Notropis blennius in the Mis- 
souri River System does not, however, possess the northern aflBni- 
ties of H. hankinsoni and, according to Bailey and Allum (1962:63), 
it is doubtful whether the species occurs in the System north of the 
Platte River. 

The following specimens have been examined by me from the 
Kansas Basin (asterisk denotes presence of female distended by 
eggs in collection ) : 



UMMZ 134366 (1 
*UMMZ 135097 (2 



UMMZ 134345 (1) Nebraska, Dundy Co., Republican River, July 

24, 1939. 

Nebraska, Dundy Co., Rock Creek, July 24, 1939. 

Nebraska, Red Willow Co., Republican River, 
July 17, 1940. 

Nebraska, Dundy Co., Arikaree River, July 19, 
1940. 

Nebraska, Dundy Co., Arikaree River, July 19, 

1940. 

Kansas, Douglas Co., Kansas River at Lawrence, 

October 24, 1951. 

Nebraska, Dundy Co., Arikaree River, April 29, 

1947. 

Kansas, Logan Co., Smoky Hill River, June 19, 

1958. 

Kansas, Shawnee Co., Kansas River at Topeka. 



" UMMZ 135124 (6 

*UMMZ 135133 (1 

KU 1850 (1 

KU 3787 (2 

KU 4017 (1 

MCZ 31881 (2 

Zoogeography. — Recognition of two subspecies of Notropis blen- 
nius was proposed by Hubbs and Bonham (1951:103-107): N. b. 
jefunus, wide-ranging; N. b. blennius in a part of the Arkansas 
River System. Within the Arkansas River System, however, Hubbs 
and Bonham (p. 103) suggested the presence of N. b. jejunus in the 
Neosho (Grand) and Illinois Rivers in Oklahoma. Moreover, they 
found specimens (p. 103) "from Nebraska" to be intermediate be- 
tween the two subspecies. 

Knowledge concerning this species in the Great Plains is slight 
owing to the small number of specimens in collections. Analysis 
of four characters (Table 11) in which four western series were 
contrasted with three eastern series suggested, despite its cursory 
nature, that (1) eye size, although seemingly highly variable, may 
average slightly smaller in western populations, (2) the head is 
deeper and wider in the range of "N. b. blennius" in the Arkansas 
River System, and (3) southern and especially southwestern popu- 
lations are deeper-bodied than northeastern populations. 

Specimens from the Kansas River Basin seem intermediate be- 
tween southwestern and northeastern populations. In addition to 
the characters listed above the anterior dorsal profile exhibits less 



Fishes of Kansas River System 



121 



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122 University of Kansas Publs., Mus. Nat. Hist. 

decurvature than was observed in specimens from the Arkansas 
River System. The upper limb of the pharyngeal arch is broad, 
flattened, and smoothly curving anteriorly in specimens from the 
Kansas River. Specimens from Noble County, Oklahoma, have 
pharyngeal arches with the upper limb more rodlike and less 
smoothly curving anteriorly as described by Hubbs and Bonham 
(1951:104). 

It seems impossible in this survey, because of insufficient material, 
to elucidate the subspecific status of several populations of this 
species. 

It has been suggested that Notropis braytoni (Jordan and Ever- 
mann, 1896:264) and Notropis potteri (Hubbs and Bonham, 1951: 
102) are aUied to Notropis blennius. If such relationships are the 
fact this group of shiners seems marked by endemism in south- 
western drainages. Within such a general pattern of endemism 
toward the Southwest, speciation in the direction of "Notropis b. 
blennius" as defined by Hubbs and Bonham could have been going 
on in the Arkansas River or possibly in the Ancestral Plains Stream 
System of an earlier time. If in the latter, upon deflection of west- 
em drainages into the central Mississippi Valley an eastern repre- 
sentative of the species, originally inhabiting the Teays/Mississippi, 
probably extended its range westward, invading the domain of the 
southwestern representative. There it intergraded extensively leav- 
ing evidence of the southwestern representative mainly in the 
southwestern part of the former Plains Stream System. 

The pattern that seems to prevail here of a northern and eastern 
kind grading to a southwestern kind across the Kansas and Arkansas 
river systems suggests the pattern, discussed elsewhere, exhibited 
by Pimephales promelas, Catostomiis commersoni and Ictalurus 
melas. 

Notropis dorsalis (Agassiz) 

To my knowledge, the only record of the big-mouthed shiner 
from the Kansas River Basin is of one specimen (KU 3479) taken 
from the Kansas River, Douglas County, Kansas (Sec. 5, T. 13 S, 
R. 21 E) on April 20, 1955. 

The rarity of the species in the Kansas River Basin is difficult to 
understand because it is abundant in streams of southwestern Iowa, 
southeastern Nebraska and throughout most of the Platte River 
System. 

The specimen taken from the Kansas River has the predorsal area 



Fishes of Kansas Rivter System 123 

fully-scaled as do specimens from tributaries of the Missouri River 
in northeastern Kansas and from the Nemaha River System in 
southwestern Nebraska. 

Notropis buchanani Meek 

There are few records of the ghost shiner from the Kansas River 
Basin (Map 22). Minckley (1959:423), who first recorded the 
species from the Basin, caught two specimens (KU 3833), a female 
and a tuberculate male, along a sandbar in the lower part of the 
Big Blue River on August 14, 1957. Other specimens have been 
collected in the lower mainstream of Mill Creek in Wabaunsee 
County, Kansas. Deacon and Metcalf (1961:317) reported N. 
buchanani from the Wakarusa River, basing their record on two 
specimens collected by C. W. Creaser in 1924 (UMMZ 63107). 

The ghost shiner seems to be restricted to the eastern part of the 
Basin, where it is rare. Hubbs and Ortenburger (1929b: 68), Met- 
calf (1959:367), and Deacon (1961:382) discussed the predilection 
of this species for larger streams over smaller tributaries in Okla- 
homa and southeastern Kansas; Deacon found it to prefer relatively 
quiet water adjacent to the strong current of the mainstream. 
Seemingly it has entered the Kansas River Basin from the south- 
east. 

Notropis heterolepis Eigenmann and Eigenmann 

There are no records, known to me, of the black-nosed shiner 
from the Kansas River Basin. The only basis for its inclusion in the 
fauna is Hubbs' (1951a) interpretation of Notropis germanus Hay 
from the Smoky Hill River at Wallace as a hybrid between N. 
heterolepis and Hybognuthus hankinsoni. As pointed out by Hubbs 
(1951a:451), N. heterolepis probably once occurred in the Kansas 
River Basin. The species has been taken in recent years in Mis- 
soini, downstream from the mouth of the Kansas River, and it is 
known to have existed in the Missouri River Drainage of South 
Dakota (Bailey and AUum, 1962:70), Iowa (Harlan and Speaker, 
1956:98), Nebraska (Evermann and Cox, 1896:401) and Colorado 
(Ellis, 1914:49) in former times; thus it might easily have had 
access to the Kansas River Basin. Extirpation over much of the 
southern part of the former range of the species has been discussed 
by Hubbs (1951a:451), Cleary (1956:272), and Trautman (1957: 
388-390). As noted by Hubbs (1951a:451) the specimens (UMMZ 
167085, USNM 69382, 69383), reported by Jordan (1891:17, 18) as 



124 University of Kansas Publs., Mus. Nat. Hist. 

Notropis cayuga from the Arkansas River at Wichita, Kansas, are 
N. heterolepis. Thus, barring an error in locaHty data, the range of 
this shiner once extended south to the Arkansas River Drainage. 
If this be true the species probably gained access to the Arkansas 
Drainage from the north by way of its former connections with the 
Ancestral Plains Stream. 

Notropis stramineus missuriensis (Cope) 

Habitat and Distribution. — The sand shiner is one of the most 
abundant and widespread species of tlie Kansas River Basin (Map 
23 ) . It has been collected from approximately 200 localities in the 
Basin and is exceeded in its ubiquity only by Pimephales promelas 
and Notropis hitrensis. Although taken in many kinds of streams, 
large and small, it was most common over sandy bottoms. 

Variation and Zoogeography. — Bailey and Allum (1962:64-68) 
found two subspecies of the sand shiner in South Dakota — a small- 
scaled subspecies (N. s. missuriensis) generally west of the Mis- 
souri River and a large-scaled subspecies {N. s. stramineus) east 
of the Missouri. The two subspecies were found to intergrade over 
a narrow zone in that state. Southward in the Great Plains a similar 
situation seems to prevail (see Table 12, which has the same 
format as tliat used by Bailey and Allum, Table 15, for ease of 
comparison ) . 

Populations from the Kansas and Platte rivers seem assignable to 
Notropis s. missuriensis (circumferential scales averaging 26.3- 
29.4; predorsal scales averaging 16.0-18.5 in the western parts of 
the basins to 15.0-17.0 in the eastern parts). Although some 
populations of the Big Blue River System in Kansas were considered 
intergrades by Minckley (1959:423), intergradation in that stream 
and in other streams in the eastern part of the Kansas River System 
is slight and mosaic in nature. Intergrading populations inhabit 
other tributaries of the Missouri River south of the mouth of the 
Platte and eastward at least to and including the Chariton (Bailey 
and Allum, 1962:64) and Blackwater rivers in Missouri (circum- 
ferential scales averaging 24:9-25.7; predorsal scales 14.4-15.1. In 
the Moreau, Osage, and Gasconade systems and eastward in the 
Missouri River System N. s. stramineus occurs (circumferential 
scales averaging 23.0-24.5; predorsal scales 12.9-13.6). The 
geographical pattern of intergradation in these areas resembles that 
between subspecies of the orange-throated darter — Etheostoma s. 
spectabile and E. s. pulchellum. 



Fishes of Kansas River System 125 

In the Arkansas River System relationships similar to those noted 
above exist. West of the Verdigris River System, N. s. missuriensis 
occurs through Kansas, Oklahoma, Colorado and New Mexico ( cir- 
cumferential scales averaging 28.7-30.7; predorsal scales 16.0-17.5). 
In the Neosho River System of Kansas, however, intergrading 
populations are found (circumferential scales, 24.9-25.5; predorsal 
scales 13.4-14.2). The pattern of intergradation in northeastern 
Oklahoma and Arkansas has not been investigated by me. 

Sand shiners from the Red River (of the south) have been re- 
ferred to "Notropis d. deliciosus" by Hubbs and Ortenburger 
(1929a:28) and Hubbs and Lagler (1958:84). Specimens that I 
have examined from the Red River Drainage of eastern and central 
Oklahoma have the large scales of N. s. stramineus (circumferential 
scales averaging 23.8-24.4; predorsal scales 12.9-14.3); however, 
series from the North Fork Red River in southwestern Oklahoma 
give evidence of intergradation with N. s. missuriensis (circum- 
ferential scales averaging 25.5-26.2; predorsal scales 14.3-14.4). 

A few specimens examined from the Colorado River System of 
Texas and the Rio Grande System in Coahuila, Mexico, have the 
large scales of N, s. stramineus. Circumferential scales averaged 
23.6 and predorsal scales 13.5 for eight specimens from the Colorado 
Drainage. Circumferential scales averaged 23.7 and predorsal 
scales 14.2 for 23 specimens from the Rio Grande Drainage. 

The subspecies N. s. missuriensis seems especially tolerant of 
(adapted to) rivers of the Great Plains. Probably its squamation 
exemplifies such an adaptation. In addition to possessing smaller 
scales westward, occasional individuals or, in some collections, a 
large percentage of specimens, have embedded scales predorsally 
or lack scales on the nape. As N. s. missuriensis occurs in the Ar- 
kansas River and other Plains streams to the north, and as it varies 
little from stream to stream, it probably dispersed throughout this 
region when streams formed an integrated drainageway (Ancestral 
Plains Stream) (Figs. 3 and 4). The presence of intergradational 
populations in the western part of the Red River System suggests 
that it has had connections to the drainageway mentioned above, 
but that these connections were less intimate or less recent than 
those between streams to the north. Subsequent to deflection of 
the upper Kansas River eastward the plains subspecies penetrated 
slightly to the east in the lower Kansas and Platte river basins and 
adjacent parts of the Missouri River System where intergrades now 
occur. 



126 



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132 University of Kansas Publs., Mus. Nat. Hist. 

Notropis topeka (Gilbert) 

Distribution and Habitat.— The Topeka shiner seems, at the 
present time, to be almost restricted to streams draining the Fhnt 
Hills and the Oread Escarpment in the Kansas River Basin (Map 
24). The only record outside these areas since 1900 is a specimen 
(KU 3108) taken by A. B. Leonard and A. B. Williams from 
Cherry Creek, Cheyenne County, Kansas in 1947. Prior to 1900 
there were several records of the species from the western part of 
the Drainage (Gilbert, 1884, 1885a, 1885b, 1886, 1889; Cragin, 
1885b; Hay, 1887). 

The distribution, habitat and abundance of N. topeka in Kansas 
have been discussed by Minckley and Cross (1959). They found 
evidence that this species increased in abundance in streams of 
the Flint Hills during a period of drought (in 1956) marked by 
lowering of water level and intermittency. Conversely, they found 
that as years of greater rainfall returned (1957-58) that spawning 
by N. topeka seemed less successful than that of other species of 
minnows in the same streams. 

Minckley and Cross (1959:215-216) suggested that N. topeka 
may have been rare in streams of the Flint Hills prior to and at 
the time of setriement noting "In Kansas, it seems that the Topeka 
Shiner has almost disappeared from areas other than the Fhnt Hills 
but has increased in abundance in that area." 

The pattern of distribution and of extirpation of N. topeka re- 
sembles greatly that of Notropis corntittis, Chrosomm enjthrogaster, 
and Hybopsis biguttata, which are interpreted elsewhere as species 
that were once more widely distributed in the clearer streams of 
the Basin including the Fhnt Hills and Shawnee Escarpments area 
but are now almost restricted to these areas as they possess the 
best remaining habitat. 

Zoogeography. — The zoogeography of the Topeka Shiner is 
puzzling. Its absence east of the Mississippi River is notable. Its 
western distribution (Bailey and Allum, 1962: Fig. 6) suggests 
an origin in the Pleistocene south-draining master stream of the 
Great Plains that integrated at one time or another parts of the 
present Arkansas, Kansas, Platte and upper Missouri systems. The 
species still occurs in and reaches its greatest abundance in these 
systems. From such a center of origin I envisage a modest inva- 
sion by this headwater species eastward, as suggested by Bailey 
and Allum (1962:125), into the upper parts of the Des Moines, 
Iowa and Cedar rivers (Underbill, 1957:19; Cleary, 1956: Map 51; 



Fishes of Kansas Rtver System 133 

Bailey, 1959:120). Such an invasion almost certainly occurred 
by headwater transfer subsequent to tlie retreat of the Wisconsinan 
glacier (the distribution in Iowa of this fish is restricted to areas 
glaciated in the Wisconsinan Age). Thus the dispersal northeast- 
ward is of recent occurrence and might easily have been accom- 
plished by a fish with the strong preferences for shallow head- 
water streams suggested by Minckley and Cross (1959:211-212). 
"Tributary hopping" in an eastward direction may have taken 
place along the Missouri River in Missouri as populations are 
found in short tributaries on both sides of Missouri River as far 
east as Gasconade County (Bailey and Allum, 1962: Fig. 6; Fisher, 
1962:427). 

Bailey (1959a: 120) pointed out certain anatomical resemblances 
between N. topeka and Notropis anogenus Forbes. Possibly an 
ancestral stock gave rise to N, topeka in the Ancestral Plains Stream 
and to N. anogenus in a north-central drainage of tlie Mississippi 
Valley, such as the Driftless Area centered in Wisconsin. As pointed 
out by Bailey (1959a: 120) the two species were probably sympatric 
in southern Minnesota and north-central Iowa prior to settiement 
of the land. However, invasion of this glaciated area by these forms 
could only have occurred in post- Wisconsinan time. 

Hybognathus hankinsoni Hubbs 

Habitat and Distribution. — The brassy minnow seems rare in the 
Kansas River Basin (Map 25). I know of only ten records as 
indicated below. All are from cool creeks of the western or High 
Plains section of the Basin. These creeks are fed by springs issuing 
either from the contact of the Ogallala Formation and the under- 
lying, aquicluding Cretaceous rocks or from aquiferous Pleistocene 
gravels of the Holdrege or Grand Island formations or both. Else- 
where in Kansas this species is known only from one small but 
permanent, direct tributary of the Missouri River in Atchison 
County. 

UMMZ 66144 (5) Colorado, Yuma Co., Arikaree River at Beecher's 

Island, October 10, 1915. 
UMMZ 134328 (3) Nebraska, Lincoln Co., Medicine Creek, July 23, 

1939. 
UMMZ 134361 (7) Nebraska, Dundy Co., Rock Creek, July 24, 1939. 
UMMZ 160450 Kansas, Wallace Co., Rose Cr. and South Fork 

160462 Smoky Hill River, (2 mi. S, 2 mi. west of Wallace) 

July 26, 1950. 
USNM 38237 ( 1 ) Kansas, Wallace Co., Smoky Hill River at Wallace, 

July, 1885 (O. P. Hay). 



KU 2680 


(7) 


KU 3788 


(4) 


KU 4043 


(1) 


KU 4859 


(2) 


KU 7039 


(1) 



134 University of Kansas Publs., Mus. Nat. Hist. 

Nebraska, Dundy Co., Arikaree River, 3 miles W. 
Hajgler, November 1, 1952. 

Kansas, Sherman Co., North Branch, Smoky Hill 
River (Sec. 13, T. 10 S, R. 40 W), AprU 28, 1947. 
Kansas, Cheyenne Co., Republican River, (Sees. 
16 and 17, T. 4 S, R. 41 W) June 21, 1958. 
Nebraska, Hitchcock Co., Frenchman Creek, be- 
low Enders Reservoir, September 11, 1959. 
Nebraska, Franklin Co., Cottonwood Creek, 1 mile 
W. Bloomington, July 10, 1961. 

These few, isolated records suggest that H. hankinsoni, mainly 
northern in its present distribution, is a relict in the Kansas River 
Basin. That it occurred farther south in southwestern Kansas in 
Illinoian time has been pointed out by G. R. Smith (1963:279). 

Zoogeography. — This minnow's rarity in the Basin contrasts with 
its status in Minnesota, where Underbill (1957:20) judged it to be 
probably the most abundant minnow in the state. 

Much of the range presently occupied by H. hankinsoni (as 
mapped by Bailey, 1954:290) was glaciated once or two or more 
times in the Wisconsinan Glacial Stage. Bailey's (1954:291) specu- 
lation that the fish invaded these glaciated areas from refugia in the 
upper Missouri and upper Mississippi River systems seem logical. 
Its absence in the Ohio River Valley suggests that the species 
evolved in a drainage far from and/or not intimately connected to 
the Ohio. Its widespread distribution in the upper Missouri Basin 
and its occurrence in beds of Illinoian age in southwestern Kansas 
(G. R. Smith, 1963:279) suggest a possible origin in a western 
drainage such as the ancestral north-flowing Missouri or the An- 
cestral Plains Stream or both. 

Hybognathus placitus Girard 

Distribution and Habitat. — Most records for the plains minnow 
from the Kansas River Basin are unreliable due to confusion con- 
cerning the specific distinctness of H. placitus and H. n. nuchalis. 
H. placitus occurs widely in the Kansas River Basin (Al-Rawi and 
Cross, 1964:164,165); H. n. nuchalis has been found only in the 
lower mainstieam. Al-Rawi and Cross (1964:163) aptly character- 
ized the habitat of H. placitus as ". . . shallow, silty water ad- 
jacent to shifting sand-bottoms of the larger prairie streams that 
have low gradients." Trautman (1957:397) noted that in the Ohio 
River Bason H. n. nuchalis eschewed streams with silty bottoms 
and turbid water. 

Zoogeography. — Of the three major pre-glacial drainages de- 
scribed previously (Fig. 3) the ancestral Plains Stream probably 



Fishes of Kansas River System 135 

harbored H. phcitus and the ancestral Teays/Mississippi and pos- 
sibly some other streams of the Atlantic and Gulf coasts harbored 
H. n. nuchalis. This is suggested by the present range of H. placitus 
throughout most of the length and breadth of the Great Plains. 
With Pleistocene disruption and diversion of these drainages and 
incorporation of parts of both into the present Missouri River 
System both species had access to the Kansas River Basin. Even 
today the distribution and habitat preferences exhibited by the two 
species in the Kansas River System betoken a Plains origin and 
an eastern origin. 

Hybognathus nuchalis nuchalis Agassiz 

The silvery minnow occurs in the lower part of the Kansas River 
Mainstream. Probably it does not penetrate far to the west; how- 
ever, its distributional pattern is not clear, because of former con- 
fusion of this species with Hybognathus placitis. Habitat prefer- 
ences and zoogeography of the two species have been discussed 
above in the account of H. placittis. 

Pimephales notatus (Rafinesque) 

Habitat and Distribution. — The blunt-nosed minnow reaches its 

greatest abundance in the Kansas River Basin in clear creeks having 

high gradient and rubbly bottoms in the Shawnee Escarpments and 

Flint Hills. West and north of this area of concentration there are 

only scattered records and none at all from the entire Republican 

River System (Map 26). Specimens that I have seen, representing 

peripheral records, are indicated below together with collectors and 

year of collection. 

KU 3977 Kansas, Ellis Co., Ellis, Big Creek— F. B. Cross and Bernard 
Nelson, 1958. 

KU 3981 Kansas, Trego Co., Saline River — F. B. Cross and Bernard 
Nelson, 1958. 

UMMZ 160471 Kansas, Logan Co., 8)i mi. W. Russell Springs, S. Fk., 
Smoky Hill River— W. R. Taylor and H. Hildebrand, 1950. 

USNM 37939 Kansas, Norton Co., Lenora, N. Fk., Solomon River— O. P. 
Hay, 1886. 

USNM 37955 Kansas, Phillips Co., Kirwin, N. Fk., Solomon River— O. P. 
Hay, 1886. 

USNM 68214 Nebraska, Saline Co., Crete, Blue River— S. E. Meek, 1891. 

Zoogeography. — In western tributaries of the Missouri River, 

north of the Kansas River, P. notatus is recorded only from the 

Elkhom River (Meek, 1894:136— MCZ 31845). Its absence in so 

large a part of the Missouri Basin reflects scarcity of habitat suitable 

8—8449 



136 University of Kansas Publs., Mus. Nat. Hist. 

for blunt-nosed minnows. P. notatus probably has little ability to 
penehate the many miles of monotonous, sandy mainstream that 
intervene between rubble-bottomed tributaries. 

The blunt-nosed minnow probably entered the Kansas River 
Basin from the southeast; the species is common in the Ozark 
Region and eastward. Bailey and Allum (1962:76) speculated that 
populations in Missouri River tributaries in eastern South Dakota 
entered that area from the upper Mississippi River System. 

Pimephales promelas Rafinesque 

Habitat and Distribution. — The fat-headed minnow is the most 
widespread species occurring in the Kansas River Basin. It has been 
taken at approximately 240 localities in all kinds of streams (Map 
27 ) . A number of records prior to 1900 from both the eastern and 
western parts of the Basin are available. 

Northward from the Kansas River in the Missouri River Basin 
P. promelas is also abundant. Bailey and Allum (1962:77) found 
it to be "probably the most ubiquitous fish in South Dakota." South- 
ward from the Kansas River in the Arkansas River System the 
species seems generally less abundant. 

Variation and Zoogeography. — Hubbs and Ortenburger (1929a: 
38) and Taylor (1954:42) noted differences between populations of 
fat-headed minnows from north-central and southwestern (Okla- 
homa and Texas) states. These differences prevail from north to 
south in the Great Plains. Populations from the Platte River Drain- 
age northward differ from populations in the Arkansas River and 
southward into Mexico, as pointed out by Hubbs and Ortenburger 
(1929a: 38); populations from the Kansas River Basin are inter- 
gradational between the two. Northern populations in the Great 
Plains (Platte and Nemaha rivers northward) have fewer pored 
scales in the lateral line and more head-tubercles ( especially on the 
chin) than do southern specimens. In addition, the northern rep- 
resentatives possess slightly fewer circumferential scales and have 
a slenderer body form, with less decurvature of the anterior profile 
and a less chubby head. 

The diversity between southwestern and northern populations 
has been given taxonomic status by recognition of a southwestern 
representative called Pimephales promelas confertus. However, I 
follow Taylor (1954) and Bailey (1956:328) in utilizing only the 
binomial. The geographic variation of P. promelas is currently 
being studied by John Vandermeer, University of Kansas. 



Fishes of Kansas Rin'er System 137 

The differences between representatives of the species from the 
southern and northern Great Plains might be attributed to one or 
more of the following: 1) Direct effect of environment in produc- 
ing different phenotypes; 2) Influence of a stock lingering in the 
upper Missouri Basin from the fauna of the preglacial north-flowing 
Missouri River; or 3) Invasion of a stock originating in the Teays/ 
Mississippi into the middle and upper Missouri River System. P. 
promelos, a widespread species (Canada to Mexico) seemingly 
tolerant of a wide range of temperatures and water conditions, 
possibly occupied all three preglacial drainages discussed herein 
(Fig. 3), and populations in the Missouri River Basin may be a 
melange, resulting from mixing of two or three stocks. 

Campostoma anomalum puUum (Agassiz) 

Habitat and Distribution. — The central stoneroller is distributed 
throughout the Kansas River Basin, but is rare or absent in areas 
where thick glacial till or loess is deposited and in other areas 
where siltation of streams is extreme (Map 28). Thus, it is absent 
in the Loess Hills ( Fig. 2 ) north of the Republican River in south- 
central Nebraska (Medicine Creek and Red Willow Creek) and in 
the upper part of the watershed of the Big Blue River in Nebraska. 
It is uncommon in the glacial till-plains in Kansas and Nebraska 
(in collections made by R. E. Johnson and by myself). In Iowa, 
Cleary (1956: Map 59) showed it to be almost absent from the 
till-plains south of the Wisconsinan end-moraines in southwestern 
Iowa. In Illinois, Forbes and Richardson (1909:111) found the 
stoneroller to be among those fishes that assiduously avoided till- 
plains of the area covered by glaciers of Illinoian age. The species 
is most common in clear streams with bottoms of rubble, gravel or 
coarse sand. It prefers smaller creeks in the Kansas River Basin, 
often ascending to their ultimate headwaters. In the western part 
of the Basin it is most common in the area fed by springs issuing 
from Pleistocene gravel deposits or from the contact of the Ogallala 
Formation with underlying beds of Cretaceous age. 

Variation and Zoogeography. — Stonerollers from parts of the 
Kansas River Basin have been assigned to the subspecies plumbeum 
by Breukelman (1940a:371) and by Minckley (1959:426), but the 
distinguishing characters of plumbeum were not well defined by 
anyone including Girard (1856:178; 1858:228,229) who proposed 
the name "Dionda plumhea" and it remained for Hubbs and 
Ortenburger (1929b: 95) to show that D. plumbea, together with 



138 University of Kansas Publs., Mus. Nat. Hist. 

Dionda spadicea Girard and Dionda grisea Girard, was conspecific 
with Camposfoma anomalum (Rafinesque). Hubbs and Orten- 
burger (1929b: 95) did not specifically propose use of the trinomial. 

Subspecific allocations in Campostoma anomalum have been 
made chiefly on the basis of the number of scales in the lateral line 
and around the body. Counts of lateral line and circumferential 
scales were made (unpublished tables 15 and 16 of doctoral dis- 
sertation by A. L. Metcalf, filed in Univ. Kansas Library, 1964) on 
40 series of stonerollers from the Great Plains and on the types of 
Chondrostoma pullum Agassiz (UMMZ 86455). Specimens (PAS 
5066) labeled "Cotype of Dionda plumbea Girard. Headwaters of 
the Canadian River (Llano Estacado). H. B. MoUhausen. Smiths. 
Inst. (2981) 5066" are too poorly preserved to allow reliable counts 
to be made. 

Although populations from the Kansas River Basin and from 
some other parts of the Great Plains possess, on the average, higher 
circumferential scale-counts than those reported for some popula- 
tions of C. a. ptdlum to the east and north, the differences seem 
gradational from one area to the other. Following the reasoning 
of Bailey, Winn and Smith (1954:148-149) I think it judicious not 
to recognize subspecies of fishes where differences are clinal and 
of slight degree from one geographic area to another. In this case 
the cline involving squamation is poorly defined from east to west 
and in some places seems to be disrupted or even reversed. In the 
Arkansas River Basin, however, there is a recognizable cline toward 
decreasing size of scale from east to west. Confounding the ex- 
pression of an east-west cline may be a north-south cline. Speci- 
mens from the upper Platte River have consistently lower scale- 
counts than do specimens from the adjacent upper Arkansas River 
in Colorado. 

Subspecific partitioning of populations of the central stoneroller 
into pullum and plwnbeum seems to me to obscure the basic unity 
of morphology which distinguishes this wide-ranging land and 
which sets it off from the coarse-scaled stoneroller (C anomalum 
oligolepis). In the case of C. a. pullum and C. a. plumbeum the 
degree of difference is slight and involves, so far as has been 
pointed out, only clinal differences in scale size. In C. a. pullum 
as compared with coarse-scaled stonerollers there are trenchant 
differences not only in scalation but also in body-proportions, shape 
of mouth, tubercle-patterns and breeding coloration. 



Fishes of Kansas Riveb System 139 

Greene (1935:132) suggested that C. a. oligolepis had its origin 
in the Driftless Area, centered in Wisconsin, whereas C. a. pullum 
originated in the Mississippi River Valley to the south of the 
Driftless Area. However, the subsequent discovery that C. a, 
oligolepis also occurs in the Ozark Region and the fact that much 
of the Ohio River Basin is populated by a related, coarse-scaled 
stoneroller (C. a. anomalum) makes Greene's hypothesis untenable 
and argues for an original (possibly preglacial) habitation by 
coarse-scaled stonerollers of the Teays/Mississippi System. An- 
cestral stock of the central stoneroller must have originated west, 
south or southwest of the main part of the Teays/Mississippi Basin. 
That fine-scaled members of the genus have long inhabited south- 
western drainages is suggested by the presence of C. ornatum, ex- 
treme in regard to smallness of scales, in a part of the Rio Grande 
Drainage and in some other drainages of northern Mexico. Areas 
of origin for stock of the central stoneroller might have included 
one or more of the following: (1) Ancestral Plains Drainage, (2) 
Ouachita Mountains, and (3) coastal streams of Texas and Louisi- 
ana. That the Ancestral Plains Drainage was among areas early 
inhabited by the central stoneroller is suggested by fossils that 
possess, according to G. R. Smith (1963:279), dentaries that "are 
referable to the narrow-jawed subspecies, C. a. pullum (Agassiz) 
of the Mississippi Valley or C. a. plumbeum (Girard), of the High 
Plains, on the basis of the narrow gnathic ramus . . ." The 
penetration of the central stoneroller eastward into certain Atlantic 
Coast-drainages of New York and into the upper Allegheny River 
(Ross, 1958:3) suggests that its invasion northeastward has long 
been in progress, possibly dating from one or more Pleistocene 
interglacial stages. 

Cycleptus elongatus (LeSueur) 

I have seen no specimens of the blue sucker from the Kansas 
River Basin other than KSU 2917, collected by I. D. Graham from 
the Blue River, probably in the 1880's and discussed by Minckley 
( 1959:416 ) . C. elongatus was listed by Graham ( 1885a:72; 1885b ) 
from "Kansas River." Cragin (1885b: 107) specified "Neither rare 
nor abundant in Kansas R. I have known of a fisherman taking 
twenty specimens or thereabout in a trip from Manhattan to 
Topeka." As misidentification of this distinctive fish is unlikely it 
must be assumed that the absence of more recent records than those 
just quoted indicates a marked depletion in numbers of this species 



140 University of Kansas Publs., Mus. Nat. Hist. 

in the Kansas River. However, as it penetrates as far north in the 
Missouri River as Iowa (Harlan and Speaker, 1956:71) and South 
Dakota (Bailey and Allum, 1962:78) it doubtless still enters the 
lowermost Kansas River on occasion. 

Ictiobus cyprinellus (Valenciennes) 

Distribution and Habitat. — The big-mouthed buffalo is abundant 
in the lower mainstream of the Kansas River and in the downistream 
parts of major tributaries such as the Wakarusa River ( Deacon and 
Metcalf, 1961:315). It seems to range farther upstream than the 
other species of Ictiobus (Maps 29 and 30). 

Ictiobus bubalus (Rafinesque) 

The small-mouthed buffalo occurs commonly in the larger streams 
of the lower part of the Kansas River Basin (Map 30). Many of 
the specimens examined by me are young fish and are diflBcult to 
separate from Ictiobus niger. For specimens more than 20 cm in 
standard length, values for height of anteriormost dorsal ray divided 
by head length ranged from .85 to 1.07 (see Trautman, 1957:226). 

Ictiobus niger (Rafinesque) 

The black buffalo seems rare in the Kansas River Basin, although 
its rarity may reflect only inadequate sampling of the larger waters. 
It is much less common in collections than either I. bubalus or /, 
cyprinellus. I have examined only four specimens, all mature, from 
the Basin in Kansas as follows: KU 3134 and 3135 from Douglas 
County at Lakeview, an oxbow lake south of the Kansas River; 
KU 3912 from the Big Blue River, Riley County, and KU 4373 
from the Little Blue River, Marshall County. Values for length of 
anteriormost dorsal ray divided by head length ranged from .64 
to .71 for these specimens. 

Carpiodes cyprinus (LeSueur) 

Habitat and Distribution. — Records of the quillback from the 
Kansas River Basin are two few to allow much insight regarding its 
choice of habitat; however a preference may exist for the shallow, 
sandy streams of the central and western parts of the Basin. Speci- 
mens that I have seen are listed below: 

UMMZ 135904 (3) Nebraska, Red Willow Co., Republican River, 1 

mi. S. Indianola, July 17, 1940. 



UMMZ 135121 


(1) 


KU 4180 


(1) 


KU 4864 


(1) 


KU 7136 


(1) 


PAS 89957 


(2) 



Fishes of Kansas River System 141 

Nebraska, Dundy Co., Arikaree River, July 19, 
1940. 

Kansas, RUey Co., Big Blue River, June 20, 1940. 
Nebraska, Furnas Co., Republican River at Cam- 
bridge, September 12, 1959. 

Nebraska, Dundy Co., South Republican River, 
August 23, 1961. 

Kansas, Jefferson Co., Kansas River below Tecum- . 
seh, September, 1958. 

Although neither Carpiodes cyprinus nor Carpiodes forbesi (see 
below) was mapped by Trautman (1956:Map 1) south of the Mis- 
souri River System, west of the Mississippi River, recent collections 
show it to be present in the Arkansas River System in Kansas ( KU 
8164 from Sedgwick Co. and KU 8241 from Sumner Co. ) and pos- 
sibly in the Colorado River System of Texas (OAM 5440 from the 
vicinity of San Angelo). 

Variation and Zoogeography. — This carpsucker has, in several 
instances prior to the recent analysis of Railey and Allum (1962:81) 
been recorded as Carpiodes forbesi Hubbs when mentioned from 
waters of the Great Plains. Bailey and Allum (1962:81) placed 
C. forbesi in the synonymy of C. cyprinus, noting that ". . . the 
slender fish with a low dorsal are likely the product of their en- 
vironment." Although some enviionmental modification of the fish 
in the manner suggested by Bailey and Allum seems possible, the 
generally distinctive phenotype of C. cyprinus in the Great Plains 
probably is due in part to a different genetic heritage. 

A fish resembling present-day plains populations of C. cyrinus 
may have originated in the Ancestral Plains Drainage. Subsequent 
to diversion of segments of the Ancestral Plains Drainage eastward, 
western populations may have mingled with closely related, but 
slightly different, congeners to the east. The result could reason- 
ably be a wide zone of inter gradation in the western Interior Low- 
lands, which might account for some of the difficulties in identifying 
specimens of this complex. Such a zone is suggested by the dis- 
cernment of phenotypes of both "cyprinus" and "forbesi" in the Big 
Sioux River System in South Dakota and Iowa (Cleary, 1956, 
Maps 20 and 21; Bailey and Allum, 1962:81). Height of the dorsal 
fin in specimens from the Arkansas River System of Kansas varies 
to a degree that suggests a mosaic kind of intergradation. 

Possibly western populations of C. cyprinus should be accorded 
subspecific status but until more specimens are available, allowing 
better insight into patterns of intergradation, this seems premature. 



142 University of Kansas Publs., Mus. Nat. Hist. 

In any event the name "forbesi" is probably unavailable as the 
type locality in the Illinois River (Hubbs, 1930:13) was considered 
by Trautman (1956:39) as within the range of another subspecies — 
Carpiodes cyprimis hinei Trautman. 

Carpiodes velifer (Rafinesque) 

The high-finned carpsucker probably has been extirpated in the 
Kansas River Basin (Map 31); none is known to have been ob- 
tained since the 1920's, Records from the Basin were summarized 
by Minckley (1959:417-418). I have seen additional specimens 
( USNM 3549 ) attributed to the Republican River and collected by 
W. A. Hammond in the 1850's. Hay's (1887:247) record of "Ictio- 
bus velifer" from Beloit, Kansas, probably is based on USNM 
37936, which is instead Carpiodes carpio carpio. The pattern of 
extirpation for C. velifer resembles that for other species known to 
prefer the clearer waters seemingly more common prior to man's 
modifaction of the Kansas River Watershed. 

Carpiodes carpio carpio (Rafinesque) 

The river carpsucker is, by far, the most common of the deep- 
bodied suckers in the Kansas River Basin. It is found throughout 
the Basin from the larger waters of the east to shallow, sandy tribu- 
taries of the west (Map 32). It is rare in rubble-bottomed, upland 
creeks in the eastern part of the Basin. The western and south- 
western inclination of the range of this species suggests an origin 
in the Ancestral Plains Stream and streams to the southwest, whence 
it penetrated eastward to a limited extent. The related species, 
Carpiodes velifer, on the other hand, might reasonably have orig- 
inated in the Ancient Teays/Mississippi System. 

Moxostoma erythmrum (Rafinesque) 

Habitat and Distribution. — In the Kansas River Basin the golden 
redhorse has been taken only in Mill Creek, Wabaunsee County, 
Kansas (Map 33). It inhabits the larger pools of this clear, rubble- 
bottomed stream of the Flint Hills Uplands. 

Zoogeography. — M. ert/thrurum is one of several fishes that are 
restricted to Mill Creek or to Mill Creek and a few nearby streams 
of the Flint Hills Area. The isolated occurrence of M> erythrurum 
is interpreted as relict; Mill Creek preserves habitat especially 



Fishes of Kansas River System 143 

favorable for fishes of clear upland streams. Elsewhere in the 
Missouri River System this species is presently found in Missouri 
in streams in or near the Ozark Uplift, in the Rock River, Iowa 
(Cleary, 1956: May 24) and in the lower Platte River in Nebraska 
(Larry A. Morris, unpublished master's dissertation. University of 
Missouri, 1960:28). Cope's (1871) types of "Ptychostomiis bucco" 
from "St. Joseph, Missouri" (PAS 6961-64) are golden redhorse. 

Moxostoma carinatum (Cope) 

The only record of the river redhorse from the Kansas River 
System is based on an adult (KSU 2732) listed by Jennings 
(1942:364) as Moxostoma aureolum and reidentified by Frank B. 
Cross as M. carinatum. Jennings (1952:364) noted that the speci- 
men was from the Wakarusa River, obtained by I. D. Graham, pre- 
sumably in 1886 or a few years earlier. 

Moxostoma macrolepidotum (LeSueur) 

Habitat and Distribution. — The northern redhorse seems to in- 
habit most commonly the medium-sized streams of the eastern 
part of the Kansas River System (Map 34). In addition to the 
records presented by Minckley and Cross (1960) I have examined 
two specimens that provide the westernmost records from the 
basin, namely USNM 37930 (reported by Hay, 1887:247) from Be- 
loit, Mitchell County, Kansas, in the Solomon River, and UMMZ 
134994, obtained on July 8, 1940, by R. E. Johnson in the Litde 
Blue River one mile southeast of Oak, Nuckolls County, Nebraska. 

Zoogeography. — Minkley and Cross (1960) in assessing the 
taxonomic status of the northern redhorse in the Kansas River 
Basin considered specimens they examined to be intergrades be- 
tween the subspecies M. a. aureolum (LeSueur) and M. a. pisolab- 
rum. Trautman and Martin. Hubbs and Lagler (1958:65) indi- 
cated that the name aureolum is unavailable in the genus Moxos- 
toma. The name "haydenf (Teretulus haydeni Girard) is avail- 
able for populations of this species from the upper Missouri River 
according to Robins and Raney (1957:154), should these popula- 
tions eventually be shown to differ subspecifically from the eastern 
m. macrolepidotum. 

M. m. macrolepidotum is found in the upper Missouri River Basin 
and elsewhere to the north and east, whereas M. m. pisolahrum has 
its center of distribution in the Ozark Region. It seems, then, that 



144 University of Kansas Publs., Mus. Nat. Hist. 

gene flow responsible for the element of M. m, macrolepidotum 
discerned in populations of the Kansas River Basin must have come 
from the north, and gene flow responsible for the element of M. m. 
pisolabriun from the southeast. A similar zone of intergradation 
between these subspecies was indicated for northeastern Missouri 
by Trautman and Martin (1951:8, Map 1). The present distribu- 
tion of M. m. macrolepidotum (Trautman, 1957, Map 46; Minckley 
and Cross, 1960:35) over much of Canada indicates a fish tolerant 
of cold waters. M. m. macrolepidotum, may have inhabited the 
upper Missouri System for a long period, perhaps in the preglacial 
Hudson Bay Drainage, and perhaps came into contact with M. m. 
pisolahriim only after deflection of the Missouri Drainage southward 
and eastward in one or more glacial advances. (It is not known 
whether the Missouri System after an initial deflection southward 
reacquired a northward direction of flow in any interglacial stage. ) 
Either M. m. macrolepidotum endured the Wisconsinan glacial stage 
in the Upper Missouri Basin, or gained access to the Upper Missouri 
directly from the east through headwater connections with the Red 
or Minnesota rivers, such as were hypothesized by Bailey and Allum 
(1962:114) for some species inhabiting South Dakota. One of the 
above alternatives seems to apply because it is unlikely that M. m. 
macrolcpidohnn could, in postglacial time, have ascended the Mis- 
souri River System, passed genetically unaltered through the range 
of M. m. pisolahrum inhabiting the lower Basin, and then invaded 
and flourished in much of the upper Missouri River Basin. 

Catostomus commersoni (Lacepede) 

Habitat and Distribution. — The white sucker is found in smaller, 
clearer streams throughout much of the Kansas River Basin (Map 
35). The distribution of this slender-bodied sucker complements 
that of the deeper-bodied sucker Carpiodes carpio carpio. 

Variation and Zoogeography. — Evermann and Cox (1896:394) 
showed that C. commersoni in the Upper Missouri River Basin 
( Platte River northward ) have larger scales, smaller eyes and more 
rows of papillae on the upper lip than do specimens from Ohio and 
Pennsylvania. They noted that the name Catostomus sucklii Girard 
is available for populations in the Upper Missouri Basin but added 
( p. 394 ) , "We doubt, however, if sucklii should be recognized even 
as a subspecies . . ." Subsequently Ellis (1914:22-25) assigned 
white suckers from the Kansas River Basin in Colorado to C. c. 



Fishes of Kansas River System 145 

siicklii and Breukelman (1940:369), referring to suckers from the 
Kansas River Basin in northwestern Kansas, wrote, "The specimens 
we took are referable to subspecies mcklii, and the records probably 
apply to this form. Typical commersonnii is not known to occur 
farther west than the eastern third of Kansas, but the subspecies 
of this species have not been critically studied." Bailey and Allum 
(1962:85) did not partition Catostoinus commersoni into sub- 
species. 

My counts indicate ( unpublished table 17 of doctoral dissertation 
by A. L. Metcalf, filed in Univ. Kansas Library, 1964) that white 
suckers from the Kansas, Arkansas, Canadian, and Pecos rivers are 
smaller-scaled than populations from the Platte System northward 
in the Great Plains. The mean number of lateral-line scales in 
specimens from these stream systems are as follows (listed east to 
west within system ) : Niobrara River, 59.6; western half of Platte 
River, 58.4, 59.7, 59.9; Marais des Cygnes River, Kansas, 67.5; 
xMoniteau Creek, Missouri, 66.3; Kansas River, 65.3, 66.4, 65.8, 61.0; 
Arkansas River, Colorado, 63.8, 63.5, 62.7; Mora River, New Mexico, 
65.6; Pecos River, New Mexico, 65.7. The mean of 61.0 from 
the Kansas River System was obtained from specimens taken below 
Enders Dam on Frenchman Creek in Hitchcock County, Nebraska. 
Introductions from the Platte River Drainage to the north may 
have taken place at this locality (see account of Fundidiis sciadicus). 
Scale-counts in populations from the southern Great Plains closely 
resemble those from the lower Missouri River in Missouri and 
eastward in the Mississippi Valley. 

Evermann and Cox (1896:393) noted that the number of dorsal 
rays in populations from the upper Missouri Basin is highly variable, 
but they stipulated no differences between these populations and 
those elsewhere. Bailey and Allum (1962:85) found that specimens 
from western South Dakota have slighdy fewer dorsal rays than 
those from the eastern part of that state. I observed little difference 
in dorsal ray-counts of specimens from the Platte, Kansas, Arkansas 
and Pecos river systems, the number usually ranging from 11 to 13 
with the average number approximately 12. 

Evermann and Cox (1896:394) found, on the average, three or 
four rows of papillae on the upper lip of white suckers from the 
upper Missouri Basin and two or three rows on specimens from Oliio 
and Pennsylvania. ( Rows of papillae are difficult to count because 
of their irregularity and the minuteness of some marginal papillae, 



146 University of Kansas Publs., Mus. Nat. Hist. 

especially on small fish.) The average number of rows on speci- 
mens seen by me are as follows (series contain 7-20 specimens): 

Ohio, Cuyahoga Co., Lake Erie Dr 3.5 

Wisconsin, Iowa Co., Fox River Dr 3.7 

Missouri, Moniteau Co., Moniteau Creek, Missouri River Dr 3.4 

Kansas, Riley Co., Kansas River Dr 3.8 

Kansas, Upper Smoky Hill River Dr 4.2 

Nebraska, Antelope Co., Verdigre Creek, Niobrara River Dr 4.3 

Wyoming, Platte Co., Laramie River 4.6 

Colorado, Custer Co., Arkansas River Dr 4.7 

New Mexico, Mora Co., Mora River, Canadian River Dr 4.5 

New Mexico, San Miguel Co., Pecos Dr 4.6 

The number of rows of papillae increases westward and in this 
respect specimens from the Kansas River Basin are intermediate 
betAveen those of the lower Missouri River Basin and those of 
western drainages such as the Arkansas, Pecos and upper Platte 
rivers. Specimens from the Smoky Hill River System resemble 
western populations. 

Southwestern populations of C. commersoni in the Great Plains 
seem chubbier anteriorly, those from the Platte System northward 
more slender; but, 22 proportional measurements made on young 
fish 50-80 mm in standard length failed to reveal outstanding differ- 
ences. Seemingly the differences become more pronounced in older 
fish. 

Examination of a few series of white suckers suggests that Ever- 
mann and Cox (1896:394) rightly ascribed relative smallness of 
eye to populations from the Great Plains. Means obtained for 
distance across exposed surface of the eye expressed as thousandths 
of standard length were as follow: Wisconsin, Iowa Co., Fox River 
Drainage (KU 5379), 64.7; Kansas, Kansas River (Composite), 
60.1; Nebraska, Garden Co., North Platte River (KU 4840), 58.8; 
Colorado, Weld Co., South Platte River (KU 4799), 56.6; New 
Mexico, San Miguel Co., Pecos River (KU 4268), 57.0. 

In summary, there is a trend in the western part of the Mississippi 
River System towards a smaller eye and more papillose lips from 
east to west; towards smaller scales from north to south, and possibly 
towards a chubbier body-form from northeast to southwest. Per- 
haps these trends have zoogeographic significance. The Ancestral 
Plains Stream System probably was inhabited by Catostomus com- 
mersoni since (1) fossil remains of it have been reported from 
mid-Pleistocene (Illinoian?) beds of northwestern Oklahoma and 



Fishes of Kansas River System 147 

southwestern Kansas (C. L. Smith, 1954:283; 1958:177, 179, and 
G. R. Smith, 1963:281), and (2) the species still occurs in the 
western (Arkansas and Canadian rivers) and northern (Kansas 
River northward) parts of this hypothetical early drainage. It 
seems reasonable to suppose that the ancient Hudson Bay drainage 
also harbored C. commersoni because its present range extends far 
to the north in Canada and to high altitudes in the Rocky Moun- 
tains. If, as suggested by Greene (1935:59), this cold-tolerant fish 
followed glacial fronts northward in association with coregonids 
and salmonids it may have inhabited the northern part of the 
Great Plains ( Dakotas, Montana, and Wyoming ) throughout much 
or all of the Pleistocene. The widespread occurrence of this species 
in the eastern United States, including some Atlantic coastal drain- 
ages, strongly suggests that it also inhabited the preglacial Teays/ 
Mississippi System. 

If three geographically-separated representatives formerly existed, 
the representative of the Ancestral Plains Stream Drainage may 
have been a chubby fish with many lip-papillae, and small eyes and 
scales. Fish from the western part of the Hudson Bay Drainage 
may have resembled the representative from the Plains except for 
slenderer body and larger scales. Populations from the Teays/ 
Mississippi may have possessed a large eye, slender body, small 
scales, and lips with few papillae. Assuming that such distinctive- 
ness did exist, it has been blurred subsequently in the Mississippi 
River System by Pleistocene minghng of stocks caused by advances 
of glacial ice, compression of ranges and changes in stream connec- 
tions. Such blurring seems especially detectable in the Kansas 
River Basin, which is occupied by a fish most nearly resembHng 
populations of the Missouri/Mississippi System to the east but 
sharing a few characters with populations to the north and 
southwest. 

Ictalurus punctatus (Rafinesque) 

The channel catfish is widespread in the Kansas River Basin 
(Map 36) and has long inhabited streams of the Great Plains as 
shown by Pliocene and Pleistocene fossil remains (C. L. Smith, 
1954:285; 1958:179; 1962:507 and G. R. Smith, 1963:281). Like 
several other catfishes of the Mississippi Basin, 7. punctatus is tol- 
erant of large, turbid rivers, readily disperses therein, and therefore 
is of little significance in elucidating zoogeographic patterns. 



148 University of Kansas Publs., Mus. Nat. Hist. 

Ictalurus furcatus (LeSueur) 

The only specimen of blue catfish that I have examined from 
the Kansas River Basin is KU 7488 (head only) taken in the Kansas 
River at Lawrence, Douglas County, Kansas, in July, 1942. Both 
Snow (1875:141) and Dyche (1914:76) wrote of large blue catfish 
taken at Lawrence. The species is known from the Missouri River 
as far upstream as Iowa (Harlan and Speaker, 1956:108) and South 
Dakota (Bailey and Allum, 1962:90). 

Ictalurus natalis (LeSueur) 

Habitat and Distribution. — The yellow bullhead is common in 
small streams of the Basin from the Flint Hills eastward ( Map 37 ) . 
Minckley (1959:427) observed it to inhabit the mud-bottomed 
streams and upland, gravelly creeks, of the Big Blue River System 
in Kansas. 

Westward from the Flint Hills this species is rare in collections. 
Early collectors failed to record it from the western part of the 
Basin. Possibly its occurrence there in more recent years (first 
recorded from the Saline River by Breukelman, 1940:372) is due 
to introduction by man. Peripheral records in addition to those 
of Breukelman are as follows: KU 7203— Smoky Hill River, 2.5 
mi. SW Wallace, Wallace Co., Kansas, August 24, 1961 (12); KU 
7011— Foster Creek, 5.5 mi. NW Orleans, Harlan Co., Nebraska, 
July 18, 1961 (5); KU 4051— Sappa Creek, Decatur Co., Kansas, 
sec. 29, T. 2 S, R. 28 W, June 23, 1958 (1). 

Zoogeography. — Ictalurus natalis is widespread in the eastern 
United States (Hubbs and Lagler, 1958:90; Trautman, 1957: Map 
106). Populations in the Kansas River System are on the north- 
western periphery of the range of the species and probably entered 
the Basin from the southeast. 

Ictalurus melas (Rafinesque) 

Habitat and Distribution. — The black bullhead is the commonest 
of the catfishes inhabiting the Kansas River Basin (Map 38). It 
seems to tolerate a wide variety of habitats, although its distribu- 
tion is somewhat complementary to that of the larger catfishes, 
Ictalurus punctatus and Plyodictis olivaris — the latter species being 
more common in the larger streams of the Basin. /. melas and 
Pimephales promelas seem to be the only species that presently 



Fishes of Kansas Ri\ter System 149 

thrive in the highly turbid, mud-bottomed, low gradient streams of 
the upper Big Blue River in Nebraska (Lincoln Creek, Beaver 
Creek and North Fork Big Blue River). 

Variation and Zoogeography. — Northern ( I. m. melas ) and south- 
em ( 7. m. catulus ) subspecies have been recognized by some workers 
(Ortenburger and Hubbs, 1926:134; Hubbs and Ortenburger, 
1929a .-39; Hubbs and Lagler, 1958:90) but not by others (Traut- 
man, 1957:427; Bailey, 1956:328; Bailey and Allum, 1962:88). 
Hubbs (1940b: 209-210; 1945:19) noted that northern populations 
possess a heavier body, smaller spines and shorter anal fin than 
southern representatives. Anal fin-rays in I. m. catulus have been 
enumerated as 19-2.3 (Ortenburger and Hubbs, 1926:134); for 
7. m. melas as 17-21 (Hubbs and Lagler, 1958:89). The number of 
anal rays in specimens examined by me from the Kansas River 
System seems to be intermediate between that of specimens from 
the Arkansas River System and some from tributaries of the Missouri 
River north of the Kansas River in Kansas. Means and ranges for 
13 specimens from the area last named are 19.2 (18-20); for 37 
specimens from the Kansas River System 20.9 (19-24) and for 20 
specimens from the Arkansas River in Kansas 22.0 (21-24). The 
average number of anal rays in specimens from the eastern part of 
the Kansas River Basin is 20.8 and in 9 specimens from Hackberry 
Creek in the Smoky Hill Drainage is 21.2. 

Fossils (C. L. Smith, 1945:285; 1958:178 and G. R. Smith, 1963: 
281) show that 7. melas occupied a part of the Great Plains in 
southwestern Kansas and northwestern Oklahoma in the Middle 
Pleistocene (Illinoian?). This fact and the ubiquity of the species 
in the Great Plains at present, suggest that it inhabited the Ancestral 
Plains Stream Drainage. Possibly the species inhabited also the 
Hudson Bay Drainage. Initial separation with subsequent mingling 
of faunas from these two drainages may have given rise to a situa- 
tion in which slightly different kinds inhabit northern and south- 
western segments of the western part of the Mississippi River 
System; these kinds are connected by a wide zone of intergrading 
populations. 

Ictalurus nebulosus (LeSueur) 

Cragin (1885b: 107) listed the brown bullhead from "Various 
streams about Topeka. Also Lawrence (Snow) and Ottawa 
(Wheeler)." Graham hsted it ( 1885a :71) as "Plentifur in Kansas 



150 University of Kansas Publs., Mus. Nat. Hist. 

and specified ( 1885b ) its presence in the Kansas River. The "bull- 
heads" seem to have been confused by workers of the last century. 
Later workers have failed to find this species in the Kansas River 
Basin, However, in recent years it has been introduced into farm 
ponds in the Basin. 

Noturus flavus Rafinesque 

Habitat and Distribution. — Records of the stonecat are scattered 
throughout much of the Kansas River Basin ( Map 39 ) . There are 
both older (1800's) and more recent records from most parts of 
the Basin, indicating that N. flavus is native to the Basin and that 
it has withstood modifications of stream-habitats that seem to have 
taken place. 

Nottirus flavus occurs in streams with either sandy or rubbly 
bottoms. As pointed out by Minckley (1959:428), IV. flavus occurs 
in the Blue River along sand flats where no rubble cover is avail- 
able. In nearby rubbly riffle-habitats of Flint Hills streams ( Camp 
Creek in Pottawatomie County and Clark Creek in Morris County ) 
I found large concentrations of this species in April, 1962. In the 
western part of the Basin the stonecat was usually found under 
large flat stones that occasionally strewed the predominantly sandy 
streambeds. 

Zoogeography. — Noturus flavus is widespread in the upper Mis- 
sissippi River Valley (Hubbs and Lagler, 1958:90) including most 
parts of the Missouri River System. As suggested by Bailey and 
Allum (1962:121) it is one of a large number of species that could 
employ the Missouri River mainstream as an avenue of dispersal. 

Noturus exilis Nelson 

Habitat and Distribution. — The slender madtom has a restricted 
distribution in the Kansas River Basin ( Map 40 ) , having been taken 
only in the Mill Creek System in Wabaunsee County, Kansas ( KU 
4672, KU 8117) and in Washington Creek, a tributary of the Wa- 
karusa River, Douglas County, Kansas ( KU 4675 ) . In these streams 
it is restricted to riffle habitats of the headwater creeks typical of 
the uplands of the Flint Hills and Shawnee Escarpments. In the 
Mill Creek System it has been taken at numerous stations in head- 
water tributaries. In Washington Creek, however, it has been 
found only in a short section of permanently-flowing stream down- 
stream from Lone Star Lake, an impoundment of approximately 



Fishes of Kansas River System 151 

200 acres completed in 1939. No specimens were taken in the 
lowermost, mud-bottomed sections of either Mill Creek or Wash- 
ington Creek. 

Zoogeography. — Notnrus exiUs is a representative in the Kansas 
River Basin of a clear-water fauna of Ozarldan affinities. Probably 
it entered the Basin from the east at a time when streams were 
less inimical to it than at present. West of the Osage River in the 
Missouri River System it has persisted only in favorable habitat, 
as in the streams noted above and in Moniteau Creek, Moniteau 
County, Missouri (KU 6477), Moreau Creek, Morgan County, Mis- 
souri (KU 6470), and Indian Creek, a tributary of Blue River 
(entering the Missouri River in Jackson County, Missouri), Johnson 
County, Kansas (KU 4335). 

Pylodictis oHvaris (Rafinesque) 

The biology of the flat-headed catfish in Kansas, including the 
Big Blue River System, was studied by Minckley and Deacon 
( 1959 ) . They noted that in Kansas this species is present in major 
rivers and impoundments and usually is absent from intermittent 
streams (Map 41). P. olivaris has been reported as far west as 
Gove County, Kansas (KU 2073, from Hackberry Creek). It is of 
little significance zoogeographically here, because it is widespread 
in larger streams of the Mississippi River Basin and some other Gulf 
drainages (Moore, 1957:142). 

Anguilla rostrata (LeSueur) 

In recent years the American eel has been taken only in the 
lower part of the Kansas River Mainstream (Map 42). Formerly, 
as indicated by Breukelman (1940:372), this species was found 
upstream as far as Big Creek (probably in Ellis County, Kansas) 
and possibly to Beaver Creek in Rawlins County, Kansas. 

Fundulus kansae Carman 

Habitat and Distribution. — The plains killifish seems pre-emi- 
nently adapted to life in shallow, sandy streams of the Great Plains. 
It reverses the distributional pattern exhibited by most fishes of the 
Kansas River Basin in that it is most abundant in the western rather 
than in the eastern half of the Basin (Map 43). Eastward it seems 
to be restricted to the mainstream of the Kansas River. To the 

9—8449 



152 University of Kansas Publs., Mus. Nat. Hist. 

northeast, Fiinchdus kansae has been recorded in Clay and Howard 
counties, Missouri (Miller, 1955:11) and from the Nemaha River 
Drainage of southeastern Nebraska ( KU 7152, 8 mi. south of Syra- 
cuse in Muddy Creek). In the Missouri River mainstream it has 
been taken as far downstream as New Haven, Franklin Co., Missouri 
(Fisher, 1962:428). 

Zoogeography. — Fundulus kansae probably originated in the An- 
cient Plains Stream. Its present range coincides well with tlie 
supposed limits of such a stream system (Fig. 3). To the east a 
few records are known beyond such limits (see above) possibly 
representing stragglers from the west. To the nortli this species is 
thought to have occurred naturally only as far as the Platte or Nio- 
brara systems (Miller, 1955: 11, 12). Recent records in South 
Dakota are interpreted by Miller (1955) as introductions by man. 
Introductions may account also for the presence of the species in 
the Big Horn River in Wyoming (KU 6869, taken by Charles A. 
Long on June 22, 1961, one mile west of Greybull). 

To the south either F. kansae or F. zebriniis Jordan and Gilbert 
(a possibly distinct, but closely related species) has been reported 
from the Arkansas, Red, Brazos, Colorado and Pecos drainages. 
This distributional pattern suggests some interchange of fauna! 
elements of these streams in the past. The incorporation of parts 
of the Platte, Kansas and Arkansas rivers into the Ancient Plains 
Stream has been described elsewhere and the possible inclusion 
of, or interchange with, parts of the Red, Brazos and Colorado 
rivers has been suggested. Koster (1957:1) postulated a connec- 
tion of the Upper Canadian and Upper Pecos systems in New 
Mexico. 

Fundulus sciadicus Cope 

Habitat and Distribution. — To my knowledge the only records 
of the plains topminnow from the Kansas River Basin are provided 
by Cope (1865:78,85) and by a series (KU 4856) taken below 
Enders Reservoir, Hitchcock County, Nebraska, on September 11, 
1959. In his original description Cope lists the species "from the 
Nebraska or Platte River," whereas later in the same paper he 
includes it in a list of fishes "from the Platte River, near Fort Riley." 
Cope's material probably came from the Platte Drainage where 
the species occurs naturally. R. E. Johnson (unpublished disserta- 
tion, University of Michigan, 1942) in his intensive survey of the 
waters of Nebraska did not find F. sciadicus in the Kansas River 



Fishes of Kansas River System 153 

System in that state. I suspect, therefore, that the species has been 
introduced into the RepubHcan River Drainage, adjacent to the 
Platte, in recent years. Several reservoirs, intensively fished, are 
located in the upper Republican Drainage. 

In addition to its occurrence in the Platte River to the north, F. 
sciadicus is found in some tributaries of the Missouri River down- 
stream from the Kansas River in central Missouri ( KU 6451, 6454, 
6494). 

Probably the species was formerly more widespread in the Great 
Plains. Bailey and Allum (1962:93) noted that it "was formerly 
rather common in clear creeks in southern South Dakota, but ap- 
pears now to be uncommon." Bailey (1956:157) suggested that it 
might be on its way to extinction in Iowa. In Nebraska it occurs 
more widely but this may be due, as suggested above, to artificial 
transplanting. I have collected F. sciadicus (KU 7116) in associa- 
tion with Chrosomus eos (KU 7118) and Chrosomus neogaeus (KU 
7119) in the sandhills region of Nebraska. The Chrosomus spp. 
occurring in this region have been considered relicts (Hubbs and 
Lagler, 1958:80; Bailey and Allum, 1962:40-42). 

Zoogeography. — Zoogeographically F. sciadicus is puzzling. Its 
restriction to waters west of the Mississippi strongly suggests an 
origin in the Ancient Plains or Ancestral north-flowing Missouri 
drainages. However, its current absence over much of the area 
thought to have been drained by these systems is hard to explain, 
other than by ^videspread extirpation. Its occurrence in tributaries 
of the Neosho River System in southeastern Kansas, southwestern 
Missouri and northeastern Oklahoma is also puzzling. These 
populations may be ( 1 ) relicts of the fauna of the Ancestral Plains 
Stream (this stream-system may have included parts of the an- 
cestral Neosho drainage — see Quinn, 1958:41,42, Fig. 1), or (2) 
due to headwater-transfer to the Neosho River Drainage (Spring 
River) from southern tributaries of the Missouri River. 

Lota lota (Linnaeus) 

From the Kansas River I have seen only one specimen ( KU 7487 ) 
of the burbot. It bears the data "Kaw River, Lawrence, Kansas, 
1906." Gilbert (1886:210) reported the burbot from the Missouri 
River at Leavenworth, Kansas, and Fisher (1962:428) took one 
specimen at St. Joseph, Missouri. An additional specimen from the 
Missouri River is KU 3162, taken eight miles south of Atchison, 



154 University of Kansas Publs., Mus. Nat. Hist, 

Kansas, on April 3, 1953. Occurrence of the burbot in this part 
of the Missouri River not far from the mouth of the Kansas River 
suggests that the species may still enter the lowermost mainstream 
on occasion. 

Lota lota, a holarctic species v^^idespread in Canada and northern 
United States, might have gained access to the Missouri River Basin 
either from the north (preglacial Missouri) or the east or from 
both directions. 

Roccus chrysops (Rafinesque) 

The white bass is most abundant in ( 1 ) the lower Kansas River 
Mainstream, (2) impoundments of the Basin, and (3) streams near 
such impoundments (Map 44). Minckley (1959:428) assumed 
that the white bass was indigenous to Kansas on the basis of reports 
of Cragin (1885b: 111) and Graham (1885a:77; 1885b). Cragin 
attributed R. chrysops to Mill Creek in Shawnee Co., Kansas and 
to Eureka Lake ( Eureka Lake was an oxbow lake along the Kansas 
River immediately west of Manhattan, Riley County, Kansas). 
From this same area Minckley (1959:428) recorded R. chrysops 
caught in 1957 and 1958. Snow (1875:140) reported "Labrax 
chrysops" from the Kansas River at Lawrence. I found young white 
bass to be common in the Kansas River at Lawrence and at Eudora, 
Kansas, in the summer of 1961. 

Micropterus dolomieui Lacepede 

and 

Micropterus punctulatus Rafinesque 

According to Mr. Roy Schoonover, Chief, Fisheries Division, 
Kansas Forestry, Fish and Game Commission (personal communi- 
cation) a few small-mouthed bass and spotted bass have been 
planted in Cedar Bluff Reservoir, Trego County, Kansas. 

Micropterus salmoides salmoides (Lacepede) 

Records for the large-mouthed bass are scattered through much 
of the Kansas River Basin. Probably it is native in the Basin as 
Graham (1885a: 76) wrote that it was "Reported from branches of 
Kansas river, below dam at Lawrence." Cragin (1885b:110) listed 
the species from Soldier Creek, Shawnee Co., Kansas, and Dyche 
(1914:45) wrote "The Black bass is one of the original native 



Fishes of Kansas RmER System 155 

Kansas fishes, and at one time, in the early history of the country 
when settlements were few and far between, it was rather common 
in the smaller and the middle-sized streams of the state." In addi- 
tion this species of bass has been widely stocked in farm-ponds and 
other impoundments in Kansas (Minckley, 1959:429; Hastings and 
Cross, 1962:15). 

Chaenobryttus gulosus (Cuvier) 

I have seen no specimens of the warmouth from the Kansas River 
Basin other than those from impoundments. However, Graham 
(1885a: 75) listed the warmouth from the Kansas River, which 
suggests that the species may have occurred natively in streams of 
the Basin. 

Lepomis cyanellus Rafinesque 

Habitat and Distribution. — The green sunfish occiu-s widely in 
the Kansas River Basin, both in streams and impoundments (Map 
45). It is most abundant in lentic situations. 

Zoogeography. — Branson and Moore (1962:90,91) presented 
evidence indicating that L. cyanellus may be the most primitive 
member of its genus. That the species is a venerable one receives 
substantiation from C. L. Smith's (1962:516) report of fossil material 
of late Phocene age referable to this species. C. L. Smith (1954: 
286; 1958:179) and G. R. Smith (1963:282) also found material of 
Pleistocene (Illinoian?) age resembling L. cyanellus in southwestern 
Kansas and northwestern Oklahoma. Clearly, then, L. cyanellus 
or its progenitors occupied the Great Plains at least since Late 
Pliocene time, suggesting adaptibility to, and tolerance of, condi- 
tions of this region. Its tolerance of adverse conditions such as 
oxygen depletion, stagnation and partial desiccation of creeks and 
ponds was pointed out by Gerking (1945:99), Lewis and Dalquest 
(1957:46) and Metcalf (1959:392). 

The absence of L. cyanellus from Atlantic coastal drainages 
(Trautman, Map 133) suggests that its origin has been western 
or southwestern. From such a center it seems to have penetrated 
as far east in drainages of the Gulf Coast as the Alabama River 
and possibly the Escambia River (Bailey, Wiim and Smith, 1954: 
137) and up the Mississippi Valley where habitat is favorable to it. 
That this habitat is suggestive of Plains conditions is indicated by 
Forbes and Richardson (1909:249), who wTOte that the green 



156 University of Kansas Publs., Mus. Nat. Hist. 

sunfish was "almost the sole sunfish product of the net in the prairie 
creeks." In Ohio Trautman (1957:501) found it in streams having 
low or moderate gradient and noted its tolerance of turbidity and 
siltation. 

Lepomis megalotis (Rafinesque) 

The long-eared sunfish is rare in the Kansas River Basin (Map 
46). I have examined only four collections: (1) KU 1953 — Rock 
Creek (tributary of Wakarusa River), Douglas County, Kansas, 
August 11, 1951 (1 specimen), (2) KU 6743— tliis and the two 
succeeding lots are from the Mill Creek Drainage, Wabaunsee 
County, Kansas. August 9, 1953 (2 specimens), (3) KU 7446— 
July 25, 1963 (10 specimens) and (4) KU 8116— May 9, 1964 (14 
specimens). The species has been taken in the Mill Creek System 
on other occasions. The record of Minckley (1956:353) from Deep 
Creek, Riley County, Kansas, is probably invalid, as it was based 
on a field identification made by me that I suspect was in error. 

In the absence of early records it is not entirely clear that this 
species is native to the Basin. However, its association with other 
members of the "Mill Creek Fauna" (discussed elsewhere) that 
have eastern affinities suggests that it reached the Basin from the 
east at a time when streams may have been clearer, and has per- 
sisted only in the habitat most favorable to it in the Flint Hills and 
Shawnee Escarpments. 

Specimens from the Kansas River System possess a red predorsum 
(unpigmented in alcoholic specimens). 

Lepomis humilis (Girard) 

Habitat and Distribution. — The orange-spotted sunfish occurs 
widely in the Kansas River System (Map 47). It is almost as 
common as the green sunfish; the two are often taken at the same 
stations. Like that species it is most common in pools where 
aquatic vegetation provides some cover. In Deep Creek, Riley 
County, Kansas, Minckley (1956:355) found L. humilis "in water 
less than four feet deep, and in or near loose rock, algae, brush, or 
other cover." 

Zoogeography. — Like Lepomis cyanellus, L. humilis is primarily 
a Plains fish. Branson and Moore (1962:93) noted that "Its range 
indicates a preference for turbid waters . . ," Where it occurs 
outside the Great Plains, L. humilis seems to choose ( or is relegated 
by competition to) stream-habitats most nearly resembling those 



Fishes of Kansas River System 157 

of the Plains. In some instances a correlation between increase of 
such habitat in connection with agricultural modification of the 
watershed and the extension of range of L. humilis has been sug- 
gested. Thus, in Illinois, Forbes and Richardson (1909:256) found 
L. humilis to be most abundant in the prairie region of that state. 
Gerking (1945:101) found it much more abundant in silty, than in 
clear, streams, in Indiana. In Ohio, Trautman (1957:506-508) 
documented the in\'asion of this species from the west into the 
central part of the state. He correlated this invasion with an in- 
crease in stream-turbidity and increased siltation of stream bottoms. 
Possibly L. humilis had its origin in the Ancestral Plains Stream 
System. From this postulated center it has made modest invasions 
to the southeast in Gulf Coastal drainages as far as the Pearl 
River (according to Cook, 1959:182) and northeastward as noted 
above. 

Lepomis macrochirus Rafinesque 

The bluegill has been stocked in farm ponds (Hastings and 
Cross, 1962:15) and other impoundments in the Kansas River 
Basin. Its ubiquity and tolerance of a wide variety of habitats 
in the Basin suggest that it may be native, as does the allusion to 
this species by Graham (1885a: 76) and possibly by Snow (1875: 
140 — as Pomotis luna Girard). 

Ambloplites rupestris (Rafinesque) 

The record of Snow (1875:140) seems to be the only evidence 
that the rock bass occurred natively in the Kansas River Basin. 
However, Snow did not list Lepomis cijanellus from the Kansas 
River at Lawrence, a species that would have been expected there; 
possibly Snow confused these two species. The rock bass is found 
eastward in the Missouri River Drainage in the Ozark Region and 
may formerly have existed farther west. 

I have examined one specimen (UMMZ 134355) taken from a 
small creek below a fish hatchery 1/2 miles east of Benkelman, 
Dundy County, Nebraska, on July 24, 1939, by R. E. Johnson. 
This record is almost surely the result of introduction by man. 

Pomoxis annularis Rafinesque 

The white crappie has been taken at scattered localities through- 
out most of the Kansas River Basin. Several older records ( Snow, 
1875:140 — as Pomoxys hexacanthus; Graham, 1885a :75; Cragin, 



158 University of Kansas Publs., Mus. Nat. Hist. 

1885b: 110; Jennings, 1942:366) probably are referable to this 
species, and the statements of Dyche (1914:54-57) noted below 
strongly indicate that this species is native to the Basin. Dyche 
wrote (p. 54), "The Crappie is a native Kansas fish. As early as 
1867, '68, '69 and '70, the writer caught strings of them in the 
Wakarusa river and its tributaries. In 1871 and '72 he took them 
in Mill creek, Mission creek . . ." and (p. 56), "The crappies 
that the author caught in Kansas some forty years ago were, as 
he remembers them, of the white variety. All specimens collected 
some thirty years ago and preserved for the State University 
Museum are of the light-colored or white species." In addition the 
white crappie was planted early in the waters of Kansas according 
to Graham (1885a: 78) and in Nebraska, South Dakota and Kansas 
according to Evermann and Cox (1896:417). 

Pomoxis nigromaculatus (LeSueur) 

Records of the black crappie are scattered through much of the 
Kansas River Basin. It is not so abundant nor so widespread as 
the white crappie. That the black crappie, unhke the white crap- 
pie, may be an introduced species is suggested by the lack of older 
records, by Dyche's remarks noted above under the account of P. 
annularis, and by his statement (1914:57): "The first Black crap- 
pies that the author remembers having seen in Kansas were taken 
at Lake View, being propagated from stock that the United States 
Fish Commission car planted there nearly twenty years ago." 
( Lake View is an oxbow lake in Douglas County west of Lawrence. ) 
Graham (1885a) did not list this fish in the fauna of Kansas nor 
did Cragin (1885b: 110) list it in the Kansas River. 

Stizostedion vitreum vitreum (Mitchill) 

The walleye was, according to Graham (1885a: 78), stocked in 
the waters of Kansas by the 1880's. In recent years it has been 
stocked in several reservoirs in the Basin. 

There is also some indication (Cragin, 1885b: 11; Graham, 1885a: 
77, 1885b) that the species was native to the Basin. Three speci- 
mens (USNM 31464) in the National Collection, catalogued in 
1882, bear the following data: "Stizostedhwi vitreum var. 50/7710- 
neum. Valley Falls, Kansas, July 7, J. L. Whitman." The fish are in 
a good state of preservation. Characters are as follows: soft dorsal 
rays: 21, 21, 22; pyloric caeca: 4, unknown, 3; black spot at 



Fishes of Kansas River System 159 

posterior end of spinous dorsal fin in all; cheeks sparsely scaled in 
all. S. V. vitreum is wide-ranging, seemingly highly vagile, and 
di£Bcult to appraise zoogeographically. 

Stizostedion canadense (Smith) 

The sauger was recorded from the eastern part of the Kansas 
River Basin by several early workers (Cragin, 1885b: 111; Graham, 
1885a: 77, 1885b; and possibly by Snow, 1875:140, as Lucioperca 
americana). Specimens are still taken occasionally below Bower- 
sock Dam in the Kansas River at Lawrence. 

Perca flavescens (Mitchill) 

The yellow perch is not known to have been native to the Kansas 
River Basin but it has been introduced into several impoundments. 

According to Gile (1885), Graham (1885a:78) and Cragin 
(1885b: 111) this species was brought into Kansas in the 1800's. 
Ellis (1914:105) noted that P. jlavescens had been introduced into 
Colorado, Montana and the Pacific States, listing several specimens 
from Colorado, some taken as early as 1900. Simon (1946:98) 
mentioned introduction of this species into Wyoming waters. 

Bailey and Allum (1962:102) suggested that P. flavescens gained 
access to the middle Missouri River System in post-glacial time. 
However, the records of C. L. Smith (1954:286; 1958:178) and 
G. R. Smith (1963:283) of fossil material referred to this or a 
closely related kind from beds of Pleistocene age in southwestern 
Kansas and northwestern Oklahoma indicate that it ranged far 
to the south in the Great Plains in Illinoian (?) time. That sub- 
sequendy it was extirpated in the western Plains suggests occurrence 
of a period or periods of marked aridity there, possibly in Sanga- 
monian or in post-glacial time. 

Percina caprodes (Rafinesque) 

The logperch has been taken only in the Flint Hills and Shawnee 
Escarpments, in upland creeks with clear water and bottoms pre- 
dominantly of rubble and gravel (Map 48). The westernmost 
specimens that I have seen are from Riley County, Kansas (UMMZ 
122045, collected by Minna Jewell in September, 1923, from Deep 
Creek; KSU 3401, 3417, and 2999, taken in the 1880's from Wildcat 
Creek and KSU 25167, taken in 1958 from Eureka Lake, an oxbow 
two miles west of Manhattan). 



160 University of Kansas Publs., Mus. Nat, Hist. 

The logperch inhabits clear waters typically occupied by Kansas 
River fishes having Ozarkian afiinities. Logperch from the Kansas 
River System have been called P. c. carbonaria, a subspecies not 
known upstream from the Kansas River in the Missouri River Basin; 
another subspecies, P. c. semifasciata, is reported from eastern 
South Dakota by Bailey and Allum (1962:104). It is logical to 
suppose that Kansas River populations were derived from the south- 
east (probably from the Ozarkian Region). P. c. carbonaria is 
widespread in the lower Mississippi Valley and some drainages of 
the Gulf Coast (Bailey, Winn, and Smith, 1954:141; Cook, 1958:195). 

Percina maculata (Girard) 

To my knowledge, the first specimens of the black-sided darter 
to be found in the Kansas River Basin in almost 70 years were 
recently taken in the Mill Creek Drainage, Wabaunsee County, 
Kansas as follows: (1) KU 7447— sec. 11, T. 12 S, R. 10 E, taken 
on July 29, 1963, by Bill Cole and Verlyn Evert and (2) KU 8122— 
sec. 15, T. 12 S, R. 10 E, taken on May 9, 1964 by Frank B. Cross. 
Each of the collections contained only one specimen. There are, 
however, earlier records for this species. Gilbert wrote (1886:209) 
"Hadropterus aspro Cope and Jord. — Numerous specimens from 
Snokomo Cr., Wabaunsee Co. (Myers), this being its first definite 
record from Kansas." Jennings (1942:365) Hsted a specimen in the 
Museum of Kansas State University, Manhattan, taken in Wildcat 
Creek, Riley County, Kansas, in 1894. The record of Jennings 
probably refers to KSU 2884, a faded specimen collected in Sep- 
tember, 1894 from Wildcat Creek, that I have examined and found 
to be P. maculata. Two additional specimens (KSU 2742) with 
no data other than "I. D. Graham" are also extant in the collections 
of Kansas State University. 

The black-sided darter is one of a number of species that seems 
to have suffered restriction of range in the Kansas River Basin. The 
species has been reported from localities both downstream and up- 
stream from the mouth of the Kansas River in the Missouri River 
System. Perhaps it penetrated westward into the Kansas River 
Basin at a time when streams offered habitat more favorable to it 
than at present. This may have occurred during that time in the 
Pleistocene when the eastern part of the Kansas River headed east 
of or in the Flint Hills. 



Fishes of Kansas River System 161 

Etheostoma nigrum nigrum Rafinesque 

Early collections indicate that the jolrnny darter was widespread 
in small streams over much of the Kansas River Basin. Currently 
it is mainly restricted to some of the most favorable small-stream- 
habitat in the Flint Hills and Shawnee Escarpments (Map 49). 
Hay (1887:243, 249, 250) and Gilbert (1889:39) reported johnny 
darters from several western localities. I have been able to check 
the identity of Hay's specimens (three in number) from Beloit 
(USNM 37964). An isolated record from the Blue River System 
of Nebraska consists of one specimen (UMMZ 135368) taken by 
R. E. Johnson on the West Fork of the Big Blue River, three miles 
east of Stockham, Hamilton County, Nebraska, on July 7, 1940. 

The Kansas River Basin is on the western periphery of the 
range of this widespread species; it seems to have entered the Basin 
from the east. 

Etheostoma spectabile pulchellum Girard 

Habitat and Distribution. — The orange-throated darter is found 
in most parts of the Kansas River Basin (Map 50). It generally 
avoids the siltier streams, in the area mantled by glacial till and 
in the central and western parts of the Basin where agriculture is 
intensive. It is, therefore, an indicator-species of the better waters 
of the Basin and its extirpation from a stream is a sign of worsen- 
ing conditions (Deacon and Metcalf, 1961:320). 

Zoogeography. — Variation and systematics of this species are 
being studied by Mr. Donald A. Distler, University of Kansas. Most 
populations in the Kansas River Basin appertain to the subspecies 
E. s. ptilchelliim, although there is (Distler, personal communica- 
tion) progressive intergradation eastward in the Basin with the 
typical, eastern subspecies. The subspecies E. s. pulchellum in- 
habits the Great Plains from the Platte River Drainage southward 
into Texas. Probably this kind was dispersed by a common drain- 
age such as was provided by the Ancestral Plains Stream System. 

Aplodinotus grunniens Rafinesque 

The freshwater drum inhabits the larger streams and reservoirs, 
especially in the eastern part of the Kansas River Basin ( Map 51 ) . 
Westward the species becomes scarce. Judging from the localities 
represented (Map 51) this is one of many species of fishes that 
find large turbid rivers no barrier to dispersal in the Basin. 



162 University of Kansas Publs., Mus. Nat. Hist. 

SPECIES PROBABLY OCCURRING IN THE 
KANSAS RIVER BASIN 

These species have been reported in the literature, but supporting 
specimens are unknown: 

Acipenser fulvescens Rafinesque. — Snow (1875:140) reported 
"Acipenser maculosus" from the Kansas River, noting that the 
largest specimen taken weighed 26 pounds. Cragin (1885b: 106) 
and Graham (1858a: 70) listed "Acipenser rubicundus" from the 
Kansas River, Cragin indicating that this was on authority of Snow. 
Harlan and Speaker (1956:47) found the lake sturgeon to be con- 
fined to the Mississippi River in Iowa. Bailey and Allum (1962: 
108) failed to find the species in the Missouri River Drainage in 
South Dakota. They suggested that reports of large sturgeon from 
the upper Missouri River Drainage might be attributable to Sca- 
phirhynchus albtis, which is known to attain a large size. Bailey 
and Allum also stated (p. 108) "We know of no verified record 
of the lake sturgeon in the Missouri basin above the mouth of the 
Kansas River (Johnson, MS)." However, Johnson (unpublished 
dissertation, University of Michigan, 1942) recorded specimens of 
A. fulvescens from the lower Platte and Elkhom rivers in Nebraska. 
I know of no verifiable records from the Kansas River Basin. 

Amia calva Linnaeus. — Evidence is meager that the bowfin is 
native to the Kansas River Basin; indeed, Bailey and Allum (1962: 
30) doubted its occurrence anywhere in the Missouri River Basin. 
Subsequently, Fisher (1962:427) listed it from the lowermost part 
of the Missouri River. There is a record by Cope (1865:86) from 
"Platte River, near Fort Riley," based on collections of the expedi- 
tion of Lt. F. T. Bryan, discussed elsewhere. Graham (1885a: 71) 
listed this fish from "Branches of Missouri River" in Kansas. 

Alosa chrysochloris (Rafinesque). — There is little evidence that 
the skipjack herring ever occurred in the Kansas River System. 
Graham (1885a: 77) hsted it as abundant in the larger streams of 
Kansas; however, Graham did not list Dorosoma cepedianum, a 
species now abundant in these streams. I suspect that Graham 
confused the two species. Evermann and Cox (1896:413) listed 
no records from the Missouri River Basin other than those of 
Graham. Forbes and Richardson (1909:49) noted that A. chryso- 
chloris occurred in the "larger streams of Kansas"; possibly this 
information was also obtained from Graham. Harlan and Speaker 
(1956:59) did not find A. chrysochloris in the Missouri River 



Fishes of Kansas River System 163 

Drainage in Iowa, nor did Fisher (1962) list it from the lower 
Missouri River. However, Bailey and Allum (1962:31) reported 
one specimen from Fort Randall Dam on the Missouri River in 
South Dakota. 

Hiodon tergisus LeSueur. — The mooneve has been listed from 
Kansas by Cope (1865:85, "Fort Riley"), Graham (1885a: 74; 
1885b), Cragin (1885b: 109, "Kansas R. at Topeka"), and Evermann 
and Cox (1896:413). Bailey and Allum (1962:108) did not find 
any evidence justifying its inclusion in the faunal list of the upper 
Missouri River nor did Harlan and Speaker (1956:61) record it 
from the Missouri River Drainage of Iowa. However, Fisher ( 1962: 
427) listed it from several stations on the lower part of the Missouri 
River, as far upstream as Atchison County in northwestern Missouri, 
and Johnson (unpublished dissertation. University of Michigan, 
1942) reported one specimen from the lower Platte River in 
Nebraska. 

Notropis hudsonius hiidsonius (Clinton). — The spot-tailed sliiner 
was hsted by Cragin (1885b: 108) and Graham (1885b) from Wild- 
cat Creek (Riley County, Kansas) and by Graham (1885a: 73) from 
"Kansas River Branches." A specimen (KSU 2835) bearing the 
data "I. D. Graham, 1886" is extant in the collections of Kansas 
State University. This species is native to the Missouri River 
Basin in eastern South Dakota (Evermann and Cox, 1896:404, 
Bailey and Allum, 1962:62), northwestern Iowa (Cleary, 1956: 
Map 45) and southwestern Minnesota (Underbill, 1957:17, Map 
16), Formerly it may have occurred farther south in the Basin. 
This seems almost certain to have been the case during glacial 
stages of the Pleistocene. 

Hypentelium nigricans (LeSueur). — Graham (1885a: 72) listed 
the northern hog sucker from the Neosho and Kansas rivers. The 
species still occurs in the Neosho Drainage of Kansas. Possibly 
Graham erroneously ascribed specimens from that drainage to the 
Kansas River. However, H. nigricans occurrs in the Missouri River 
System as near to Kansas as Richland Creek, Morgan County, 
Missouri (KU 6889) and in the Osage River Drainage; it might 
formerly have ranged farther upstream. A specimen (KSU 2978) 
collected by I. D. Graham in 1886, but without further data, is 
preserved in the collections of Kansas State University. 

Percopsis omiscomaycus (Walbaum). — Recognition of the trout- 
perch in the fauna of the Kansas River Basin rests on Gill's (1864: 
151) description of "Percopsis hammondi" based on a specimen 



164 University of Kansas Publs., Mus. Nat. Hist. 

supposed to have been taken in Kansas by W. A. Hammond. Gill's 
account seems to be responsible for subsequent inclusion of the 
trout-perch in the ichthyofauna of Kansas (Cope, 1865:85; Graham, 
1885a:75; Cragin, 1885b: 109; Forbes and Richardson, 1909:226; 
Eddy and Surber, 1947:197; Moore, 1957:160 and Bailey and Allum, 
1962:94). If the specimen was actually obtained by W. A. Ham- 
mond it is likely, as pointed out elsewhere, that it came from the 
Kansas or Platte river systems. 

Whether or not Gill's specimens actually were from the Kansas 
River Basin it is reasonable to suppose that P. omiscomaycus in- 
habited the basin in historic time or in conjunction with glacial 
advances. Some other fishes with the northern aflSnities exhibited 
by P. omiscomaycus were shown by C. L. Smith (1954, 1958) to 
have existed even farther south than the present Kansas River 
Basin in Illinoian ( ? ) time. Furthermore, P. omiscomaycus ranges 
not far from the Kansas River Watershed even now, having been 
recorded in southwestern Iowa (Cleary, 1956: Map 71) and north- 
central Missouri (Hanson and Campbell, 1963:139). 

Etheostoma blennioides Rafinesque. — The sole record of the 
green-sided darter in the Kansas River Basin is that of Graham 
(1885a:76; 1885b) from Wildcat Creek, Riley County, Kansas (re- 
peated by Cragin, 1885b: 110). One specimen (KSU 2885), bear- 
ing only the data "I. D. Graham," is extant in the collections of 
Kansas State University. In regard to certain scale-counts (59 in 
lateral line and 19 around caudal peduncle) this specimen shows 
aflBnity to populations from the lower Missouri River System in 
Missouri but not to populations from the Arkansas River System 
in southeastern Kansas. As £, blennioides is common in part of 
the Missouri River Basin to the east and southeast of the Kansas 
River its past occurrence upstream in the Flint Hills section of the 
Kansas River System cannot be ruled out. As some other species 
with Ozarkian afiBnities have entered the Kansas River System it 
is conceivable that E. blennioides also accomplished such an in- 
vasion only to be extirpated subsequent to settlement of the Basin. 

SPECIES PROBABLY NOT OCCURRING IN THE 
KANSAS RIVER BASIN 

These species have been reported in the hterature, are not supported by 
specimens, and, as set forth below, are thought not to have occurred in the 
Kansas River Basin within historic time. 

Several of these doubtful records were introduced into the Hterature by 
Cope (1864, 1865, 1871), who based his reports on specimens collected by 



Fishes of Kansas River System 165 

Dr. W. A. Hammond. Hammond was a surgeon assigned to an expedition 
under the command of Lt. F. T. Bryan. The itinerary of this expedition was 
summarized by Olund and Cross (1961:331). The expedition traversed parts 
of the drainages of the Kansas, Platte, and Green (of Wyoming) rivers in a 
journey from Fort RUey, Kansas to Bridgers Pass, Wyoming, and back. The 
assortment of fishes Hsted by Cope ( 1864, 1865, 1871 ) from "Kansas" or from 
"Platte River near Fort Riley" seem not to have come from a single locaHty 
or even from a single stream system. Some entries in the catalogue of the 
Division of Mammals, United States National Museum, suggest how the designa- 
tion "Platte River near Fort Riley" might have originated — for example: 

3040 — Platte R. — 236 miles from Fort Riley — ^June 27 
3041— Platte R.— 248 miles from Fort Riley— June 29 
3042— Platte R.— 350 miles from Fort Riley— July 4 

Data entered in the catalogue and on labels of the Division of Fishes, 
United States National Museum, give the following locahties from which 
collections were made (attributed to the expedition of Bryan) and illustrate 
the variations that have crept into the records concerning these old collections: 

1. "Stranger's Creek, Kansas." 

2. "Fort Riley"; "Fort Riley, Kansas." 

3. "Republican R."; "Republican trib."; "Republican Fk." 

4. "Pole Creek"; "Pole Creek, Neb."; "Pole Creek, Wyoming"; "Pole Cr.— 
Ft. Riley"; "Pole Creek, Cheyenne Pass." 

5. "Bridger's Pass"; "Nr. Bridger's Pass"; "Wagon Road to Bridger's Pass." 

6. "Kansas." 

Entosphenus lamottenii (LeSueur). [As Ammocoetes niger Rafinesque by 
Cragin (1885b:106) and Graham (1885a:70).] Probably tliis record has 
resulted from misidentification. That Graham's specimen was not a brook 
lamprey is indicated by its being found attached to a "red-horse." Hubbs and 
Trautman (1937:73) placed these records in the synonymy of Ichthyomyzon 
castaneus. 

Lepisosteus spatula Lacepede. [As LitJwlepis tristoechus (Bloch and 
Schneider) by Cragin (1885b:106).] Possibly this record refers to a large 
specimen of Lepisosteus ossetis. The alligator gar, a southern species, probably 
approaches no nearer to the Kansas River Basin than the St. Louis area ( Forbes 
and Richardson, 1906:35). Fisher ( 1962) did not find it in the lower Missouri 
River. 

Salmo clarki Richardson. [As Trutta lewisi (Girard) by Cope (1865:85) 
and as Salmo (Salar) stomias Cope by Cope (1871).] These records were 
allocated to Salmo mykiss stomias by Everman and Cox (1896:356-357). I 
agree with Evemiann and Cox (1896:355) that ". . . die trout mentioned 
as Trutta leicisi probably came from some point in the headwaters of the 
South Platte rather than from Fort RUey." 

Umbra limi (Kirtland). To my knowledge, there is no evidence that the 
mudminnow has occurred, in historic time, in the Kansas River Basin despite 
inclusion of Kansas in the range of the species by some writers (Eddy and 
Surber, 1947:183; Moore, 1957:72). Probably it ranged farther south than 
at present during glacial stages of the Pleistocene. 

Gila robusta Baird and Girard. [As Gila affinis Abbott by Abbott 
(1861:474) and Cope (1865:85).] Abbott's 1861:474) description of G. 
afjinis (=^ G. robusta) list it from "Kansas." Evennann and Cox ( 1896:355) and 
Fowler (1925:397-398) pointed out that Gila robusta does not occur in the 
Missouri River Basin. It is, however, native to the upper Green River in 
Wyoming (Simon, 1946:79) in the area visited by the expedition of Lt. F. T. 
Bryan. An old label of red-bordered heavy paper, possibly the original label, 
with the type specimen (PAS 4196), states "Gila affinis (Abbott) — Dr. Ham- 



166 University of Kansas Publs., Mus. Nat. Hist. 

mond — Bridger Pass." Another label, newer in appearance, bears the data 
"TYPE of Gila affinis Abbott ("Kansas") Bridger Pass, Green River basin, 
Wyoming. Dr. Wm. A. Hammond." 

Rhinichthys cataractae ( Valenciermes ) . [As Rhinichthys maxillosus Cope 
by Cope (1864:278 and 1865:85).] Cope's (1864:278) description of R. 
maxillosus (= R. cataractae) lists it from "Kansas." His later account ( 1865:85) 
includes it in a list of fishes from "Platte River, near Fort Riley." Rhinichthyi 
cataractae occurs in the Platte Drainage but not in the Kansas River Drainage 
at the present time and there are no other records of the species from the 
Kansas River System. As the expedition under the command of F. T. Bryan 
traveled extensively in the Platte Drainage, it seems highly probable that 
"Rhinichythys maxillosus" was taken in that drainage. 

Minytrema melanops (Rafinesque). [Reported by Cragin (1885b: 108) 
from Mill Creek and Osage River.] To my knowledge there are no records 
of the spotted sucker from the Missouri River Basin other than those listed by 
Cragin. Ptychostoma haydeni Girard from the Upper Missouri River, although 
ascribed to M. melanops by some writers, has been shown to be Moxostoma 
macrolepidotum by Robins and Raney (1957:154). 

Erimyzon sucetta (Lacepede). [Listed by Cragin (1885b:108) from 
"Kansas R. (Snow)."] According to distributional maps prepared by the 
Missouri Conservation Commission (courtesy of Mr. William Pflieger) neither 
this species nor Erimyzon oblongus (Mitchill) occurs in the Missouri River 
Basin in Missouri; nor is either species known elsewhere in the Basin. 

Fundulus diaphanus (LeSueur). [Listed by Graham (1885a:75) from 
"Kansas River."] This record is far south and west of the present limits of the 
range of the banded killifish. Possibly Graham confused this Idllifish with F. 
kansae, a variable, sexually dimorphic species that does occur in the Kansas 
River. 

Culaea inconstans (Kirtland). [As Gasterosteus micropus Cope (1865:81, 
85) from "Platte River, near Fort Riley."] There are, to my knowledge, no 
original records of the brook stickleback from the Kansas River Basin except 
the description of G. micropus by Cope. As pointed out above, there is doubt 
whether Cope's specimens were collected in the Kansas or in the Platte river 
systems. As C. inconstans occurs in the Platte System it seems safer to suppose 
that Cope's material came from that system. Possibly the species existed far- 
ther south in the Plains during Pleistocene pluvial periods when its favored 
habitat of shallow ponds with aquatic vegetation might have been more wide- 
spread in this area of reatively low relief. 

Roccus mississippiensis (Jordan and Evermann). [Listed by Cragin, 
(1885b:lll) on authority of Snow but not reported by Snow (1875).] To my 
knowledge this is the sole record from the Kansas River Basin for the yellow 
bass. Probably it refers to Roccus chrysops, which seems to have been native 
to the Basin. R. mississippiensis has been widely introduced (Graham, 1885a: 
78) and its occurrence in the Basin would not be surprising. 

Ammocrypta pellucida (Baird). [Listed by Cragin (1885b: 110) and 
Graham (1885b) from Kansas River and Usted by Graham (1885a: 76) from 
". . . clear, sandy streams."] These records are far west of the range of 
A. pellucida (Moore, 1957:183; Linder, 1959:Fig. 1). Evermaim and Cox 
(1896) listed no additional records of any species of Ammocrypta from the 
Missouri River Basin, nor did Fisher (1962) find any in the lower Missouri 
River. Two specimens (KSU 3415) of Ammocrypta in the collections of 
Kansas State University bear only the data "L D, Graham." Although poorly 
preserved, these specimens seem assignable to A. pellucida (a few scales 
anteriorly on nape; opercle with broad, triangular spine; 6-8 scales below 
lateral line; five rows of scales from upper anterior comer of opercle to base 
of opercular spine; lateral-line scales 70 and 75; dorsal spines 10 and 11; anal 
spines and rays both I, 9). 



Fishes of Kansas River System 167 

Etheostoma cragini Gilbert. Regarding E. cragini, Gilbert (1887:63) wrote 
"A single specimen was also taken in Snokomo Creek, Wabaunsee County, 
Kansas." This is the sole record of this darter outside the Arkansas River 
Drainage. Gilbert's record seems in error, probably due to mixing of locality 
data or specimens from the Arkansas River System (reported in the same 
paper) with specimens from Snokomo Creek. 

SUMMARY 

Specimens of 71 species of fishes thought to be native were 
observed from the Kansas River System. The presence of seven 
species is known or believed to be entirely due to introduction by 
man. For six additional species evidence is inconclusive as regards 
native occurrence. There are records in the literature of eight 
species that may have inhabited the System in past, historic time 
but no longer occur there to the best of my knowledge. 

Several areas, all possessing streams characterized by relatively 
clear and cool water and by rubbly, gravelly or sandy bottoms, were 
found to support a relatively diverse fish fauna. These areas were 
(1) the Shawnee Escarpments of northeastern Kansas, (2) the 
Flint Hills of northern Kansas, (3) an area in the western part 
of the Basin draining escarpments where the Ogallala Formation 
makes contact with underlying, aquicluding Cretaceous beds, and 
(4) an area in southern Nebraska along the north side of the 
Republican River Valley draining thick aquiferous deposits of 
Pleistocene sands and gravels. Species found principally in one or 
more of these areas were Carpiodes velifer, Moxostoma erythrurum, 
Hijbopsis biguttata, Chrosomus erythrogaster, Hybognathus hankin- 
soni, Notropis rubellus, Notropis cornutus, Notropis topeka, Pime- 
phales notatus, Noturus exilis, Percina caprodes, Percina maculata, 
Etheostoma spectabile pulchellum, Ethostoma nigrum nigrum and 
Lepomis megalotis. 

At the opposite extreme were areas that supported few species of 
fish and in which streams were turbid, sometimes intermittent and 
possessed silty bottoms. Such areas included ( 1 ) an area mantled 
by glacial till in the northeastern part of the Basin; (2) an area 
(Loess Plains) mantled by thick deposits of loess in the north- 
western part of the Basin in Nebraska (on the eastern margin of 
this area was a terrain, mantled both by glacial till and and by loess, 
that yielded the lowest number of species of any area in the Basin ) ; 
(3) the "Dissected High Plains," an area in the central part of the 
Basin underlain by Cretaceous rocks of poor aquifying abihty; and 



10—8449 



168 University of Kansas Publs., Mus. Nat, Hist. 

(4) the Ogallala surface atop which all streams were observed to 
be highly intermittent, flowing only after rains. Ubiquitous species, 
especially characteristic of the areas, just mentioned, with the ex- 
ception of (4), which has no permanent fauna in streams were 
Notropis I. lutrensis, Notropis stramineus missuriensis, Pimephales 
promelas, Ictalurus melas and Lepomis cyanellus. 

Nineteenth-century accounts by visitors to the Kansas River Basin 
suggest that the larger streams fluctuated in amount of discharge 
and degree of turbidity then, as now. Tributaries, however, were 
probably clearer and more permanent than at present. The fauna 
both of the large and small streams seems to have changed to some 
extent. Early records from the lower mainstream indicate occur- 
rence of species that, at present, rarely if ever ascend the Kansas 
River, although some of them still occur in the Missouri River. In 
this connection it should be noted that pollution has been intense 
in the lowermost mainstream. A number of species typical of 
tributaries (including several listed in Paragraph 2 of this Sum- 
mary) seem to have been extirpated or their populations seem to 
have been decimated in much of the Basin, probably because of 
modifications of the watershed, especially agricultural modifications, 
that have resulted in increased intermittency, turbidity and siltation 
in these streams. 

The occurrence of some species is best explained in terms of 
preglacial drainage patterns, of which three systems seem significant. 
One, the Preglacial Plains System, flowed southward, west of the 
Flint Hills of Kansas, extending at least as far north as the Platte 
or Niobrara rivers and draining southward into the Gulf of Mexico 
either by way of an ancestral equivalent of the Red River or by 
way of some other outlet on the coast of Texas, This system long 
drained the western part of the Kansas River System. That part 
of the Kansas River System east of the Flint Hills, on the other 
hand, drained eastward into another major preglacial drainage — 
that of the Teays/Mississippi System. A third preglacial system, 
far removed from the Kansas River Basin, but probably having 
some effect on the ichthyofauna of the Missouri River System, con- 
sisted of streams (extending southward as far as central South 
Dakota) draining northward into Hudson Bay. 

From Pliocene fossil evidence and present distributions in the 
Great Plains it is hypothesized that species or subspecies that were 
contributed to the Kansas River System by the Preglacial Plains 
System (but not from other sources) were Hybopsis gracilis 



Fishes of Kansas River System 169 

gtilonella, Notropis I. lutrensis, Notropis topeka, Notropis stramineus 
mis^iriensis, Hybognathus placitus, Phenacobius mirabilis, Fundu- 
his kansae, Lepomis cyancllus, Lepomis humilis, and Etheostoma 
spectabile pttlcheUum. 

In the course of glacial advances, some of the western drainage 
of Hudson Bay was deflected southward into the Great Plains and 
is thought to have made its way southward more or less along the 
course of the Preglacial Plains Stream (hereafter termed Ancestral 
Plains Stream). Species or subspecies that may have been in the 
preglacial Hudson Bay Drainage and that were deflected into the 
Ancestral Plains Stream include Scaphirhynchus albus, Hybopsis 
meeki, Hybopsis gelida, Hybopsis gracilis gracilis, Hybognctus 
hankinsoni, and Moxostoma' m. macrolepidotiim. 

In epochs after retreat of the Kansan Glacier much downcutting 
westward by tributaries of the Mississippi River System caused 
several of those streams to reach the Ancestral Plains Stream, 
capture various segments of it, and thus fragment the System, The 
Kansas River had such an origin, with a tributary of the Missouri/ 
Mississippi System cutting through the Flint Hills and capturing 
streams to the west. Outflow of the Kansas River to the Mississippi 
gave ready access to the Basin for a large number of fishes that 
inhabited the Ancetral Teays/Mississippi System. 

Some species, especially those of larger rivers, probably reached 
the Kansas River Basin from two or three of the drainages noted 
above. Some species that have variational patterns, in the Great 
Plains and Interior Lowlands, suggestive of such a multiple origin 
are Hybopsis aestivalis, Semotilus atromaculatus, Notropis atheri- 
noides, Notropis blennius, Notropis stramineus, Pimephales prome- 
las, Campostoma anomalum, Carpiodes cyprinus, Catostomtis com- 
mersoni, Ictalurus melas, and Etheostoma spectabile. 

A mingling of eastern, western and northern stocks in the Great 
Plains and Interior Lowlands seems to have resulted from the 
disruptions and diversions of Pleistocene drainages, and from 
extirpation from, and reinvasion of, glaciated areas. Thus, differ- 
ences that might once have distinguished populations of the An- 
central Plains System from eastern or northern populations were, in 
many cases, obscured. In only a few cases do northeastern and 
southwestern congeners seem to have attained the status of species 
(for example, Hybognathus nuclialis and Hybognathus placitus). 
In other cases well-defined subspecies exist in the northeast and in 



170 University of Kansas Publs., Mus. Nat. Hist. 

the southwest (for example, Etheostoma s. spectabile and E. s. 
pulchellum; N. s. stramineus and N. s. missuriensis) . In an even 
larger number of cases differences are of lesser degree from south- 
west to northeast and a wide zone of intergradation exists between 
the extremes. 

ACKNOWLEDGMENTS 

For assistance in making collections I wish to thank Mr. Larry Witt, Dr. 
James Deacon, Prof. Frank B. Cross, Mr. Donald A. Distler, Mr. John Vander- 
meer and Mr. Martin Wiley. For courtesies extended in making specimens in 
their care available for study I am greatly indebted to Dr. Leonard P. Schultz 
and Dr. W. Ralph Taylor, United States National Museum; Prof. Reeve M. 
Bailey and Prof. Robert R. Miller, University of Michigan, Museum of Zoology; 
Dr. James Bohlke, The Academy of Natural Sciences of Philadelphia; Mrs. 
Myvanwy Dick, Museum of Comparative Zoology, Harvard University; and 
Prof. Donald J. Ameel, Kansas State University. For valuable criticisms and 
suggestions regarding the manuscript I acknowledge the help of Professors 
Frank B. Cross, A. Byron Leonard, Charles W. Pitrat, and E. Raymond Hall, 
The University of Kansas. Dr. W. Ralph Taylor, United States National Mu- 
seum, and Prof. Reeve M. Bailey, University of Michigan, Museum of Zoology, 
gave helpful suggestions. Mrs. A. C. Metcalf, Mr. William Pflieger, and Mr. 
Jon Barlow assisted in preparing the manuscript. I thank Mr. Tom Swearingen 
for advice concerning preparation of illustrations. 



Fishes of Kansas River System 



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Fishes of Kansas River System 177 

LITERATURE CITED 

Abbott, C. C. 

1861. Descriptions of four new species of North American Cyprinidae. 
Proc. Acad. Nat. Sci. Philadelphia, 1860:473-474. 

1861. Descriptions of two new species of Pimelodus, from Kansas. Proc. 
Acad. Nat. Sci. Philadelphia, 1860:568-569. 

Adams, F. G. 

1904. Reminiscences of Frederick Chouteau. Trans. Kansas State Hist. 
Soc, 8:423-434. 

Alden, W. C. 

1924. Physiographic development of the northern Great Plains. Geol. 
Soc. America Bull., 35:385-424, September 30. 

Al-Rawi, a. H. and Cross, F. B. 

1964. Variation in the plains minnow, Hybognathus placitus Girard. 
Trans. Kansas Acad. Sci., 67( 1):1.54-168, June 12. 

AUGHEY, S. 

1877. Catalogue of the land and fresh-water shells of Nebraska. Bull. 
U. S. Geol. and Geogr. Surv. Territories, 3(3):697-704, May 15. 

Bailey, R. M. 

1954. Distribution of the American cyprinid fish Hybognathus hankinsoni 
with comments on its original description. Copeia, 1954(4) :289- 
291, October 29. 

1956. A revised list of the fishes of Iowa, with keys for identification in 
Iowa Fish and Fishing. Iowa Conserv. Conmi., Des Moines, pp. 

327-377. 

1959a. Distribution of the American cyprinid fish Notropis anogenus. 
Copeia, 1959(2): 119-123, July 24. 

1959b. Parasitic lampreys (Ichthyomyzon) from the Missouri River, Mis- 
souri and South Dakota. Copeia, 1959(2) :162-163, July 24. 

Bailey, R. M. and Allum, M. O. 

1962. Fishes of South Dakota. Misc. Publ. Mus. Zool. Univ. Michigan, 
119:1-131, June 5. 

Bailey, R. M. and Cross, F. B. 

1954. River sturgeons of the American genus Scaphirhynchus: characters, 
distribution, and synonymy. Papers Michigan Acad. Sci., Arts 
and Letters, 39:169-208, May 5. 

Bailey, R. M. and Gosline, W. A. 

1955. Variation and systematic significance of vertebral counts in the 
American fishes of the family Percidae. Misc. Publ. Mus. Zool. 
Univ. Michigan, 93:1-44, September 28. 

Bailey, R. M., Winn, H. E. and Smith, C. L. 

1954. Fishes from the Escambia River, Alabama and Florida, with eco- 
logic and taxonomic notes. Proc. Acad. Nat. Sci. Philadelphia, 
106:109-164, December 17. 

Barlow, G. W. 

1961. Causes and significance of morphological variation in fishes. Syst. 
Zool., 10(3): 105-1 17. 

Bayne, C. K. 

1956. Geology and ground-water resources of Sheridan County, Kansas. 
Kansas Geol. Surv. Bull., 116:1-94, May 15. 



178 University of Kansas Publs., Mus. Nat. Hist. 

Bayne, C. K. and Fent, O. S. 

1963. The drainage history of the upper Kansas River Basin. Trans. 
Kansas Acad. Sci., 66 ( 3 ): 363-377, December 17. 

Bayne, C. K., Walters, K. L. and Plximmer, N. 

1959. Geology and ground-water resources of Cloud County, Kansas. 
Kansas Geol. Surv. Bull., 139:1-144, May. 

Berry, D, W. and Drumm, W. H. 

1952. Geology and groimd-water resources of Lincoln County, Kansas. 
Kansas Geol. Surv. Bull., 95:1-96, July. 

Bradley, E. and Johnson, C. R. 

1957. Geology and ground-water hydrology of the valleys of the Repub- 
lican and Frenchman rivers, Nebraska. U. S. Geol. Surv. Water- 
supply Paper, 1360-H:iv + 589-713. 

Bradley, E., Johnson, C. R. and Krieger, R. A. 

1957. Ground-water resources of the Ladder Creek area in Kansas. Kan- 
sas Geol. Surv. Bull., 126:1-94, December. 

Branson, B. A. 

1963. Additions to and distributional annotations on the Kansas gastropod 
fauna. Trans. Kansas Acad. Sci., 66(l):72-75, May 9. 

Branson, B. A. and Moore, G. A. 

1962. The lateralis components of the acoustico-lateraUs system in the 
svmfish family Centrarchidae. Copeia, 1962(1 ):1-108, April 11. 

Breukelman, J. 

1940a. The fishes of northwestern Kansas. Trans. Kansas Acad. Sci., 43: 
367-376. 

1940b. A collection of Kansas fishes in the State University Museum. 
Trans. Kansas Acad. Sci., 43:377-384. 

1946. A review of Kansas ichthyology. Trans. Kansas Acad. Sci., 49(1): 
51-70, August 6. 

Caldwell, M. B. 

1937. Exploring the Solomon River Valley in 1869. Kansas Hist. Quart., 
6(l):60-76. 

Call, R. E. 

1885. Contributions to a knowledge of the fresh-water Mollusca of Kan- 
sas. IL Fresh-water univalves. Bull. Washburn College Lab. 
Nat. Hist., l(2):51-54, February 15. 

1886. Fifth contribution to a knowledge of the fresh-water Mollusca of 
Kansas. Bull. Washburn College Lab. Nat. Hist, 1(6): 177-184, 
July. 

1887. Sixth contribution to a knowledge of the fresh-water Mollusca of 
Kansas. Bull. Washburn College Lab. Nat. Hist, 2(8): 11-25, No- 
vember 2. 

Carey, J. C. 

1954. Juniata: Gateway to Mid-Kansas. Kansas Hist. Quart., 21(2) i 
87-94. 

Chaney, R. W. and Eljas, M. K. 

1936. Late Tertiary floras from the High Plains. Carnegie Inst., Wash- 
ington Publ., 476:1-72, October. 

Cleary, R. E. 

1956. The distribution of the fishes of Iowa in Iowa fish and fishing. 
Iowa Conserv. Comm., Des Moines, pp. 267-324. 

Colby, C. C. and others 

1956. The Kansas Basin, pilot study of a watershed. Univ. Kansas Press,, 
Lawrence, x -\- 103 pp., September. 



Fishes of Kansas River System 179 

CoNDRA, G. E. and Reed, E. C. 

1950. Correlation of the Pleistocene deposits of Nebraska (Rev.). Ne- 
braska Geol. Surv. Bull., ISA: 1-74, March. 

1959. The geological section of Nebraska. Nebraska Geol. Surv. Bull., 
14A:viii + 82. 

Cook, F. A. 

1959. Freshwater fishes in Mississippi. Hederman Bros., Jackson, Mis- 
sissippi, 239 pp. 

Cope, E. D. 

1864. Partial catalogue of the cold-blooded Vertebrata of Michigan, Part 

I. Proc. Acad. Nat. Sci. Philadelphia, 1864:276-285. 

1865. Partial catalogue of the cold-blooded Vertebrata of Michigan, Part 

II. Proc. Acad. Nat. Sci. Philadelphia, 1865:78-88. 

1871. Recent reptiles and fishes in Report on the reptiles and fishes ob- 
tained by the naturahsts of the expedition. Preliminary Rept. Geol. 
Surv. of Wyoming and portions of contiguous territories, being a 
second aimual report of progress, 8:432-442. 

Cragin, F. W. 

1885a. Note on the chestnut lamprey. Bull. Washburn College Lab. Nat. 
Hist, 1(3):99-100, April4. 

1885b. Preliminary list of Kansas fishes. Bull. Washburn College Lab- 
Nat. Hist, 1(3): 105-111, May 2. 

Crevecoeur, F. F. 

1903. A new species of fish. Trans. Kansas Acad. Sci., 18:177-178. 

1908. A new species of Campostoma? Trans. Kansas Acad. Sci., 21: 
155-157. 

Cross, F. B. 

1953. Occurrence of the sturgeon chub, Hybopsis gelida (Girard) in Kan- 
sas. Trans. Kansas Acad. Sci., 56(1):90-91. 

Cross, F. B. and Hastings, C. E. 

1956. Ages and sizes of 29 flathead catfish from the Kansas River, 
Douglas County, Kansas. Trans. Kansas Acad. Sci., 59(l):85-86, 
April 12. 

Cross, F. B. and Metcalf, A. L. 

1963. Records of three lampreys (Ichthyomyzon) from the Missouri River 
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Dalquest, W. W. 

1964. Equus scotti from a high terrace near Childress, Texas. Texas Jour. 
Sci., 16(3):350-358, September. 

1965. New Pleistocene formation and local fatma from Hardeman County, 
Texas. Jour. Paleont, 39( 1 ): 63-79, January. 

Davis, S. N. 

1951. Studies of Pleistocene gravel lithologies in northeastern Kansas. 
Kansas Geol. Surv. Bull., 90(7): 173-192, September 30. 

De.\con, J. E. 

1961. Fish populations, following a drought, in the Neosho and Marais 
des Cygnes rivers of Kansas. Univ. Kansas Publ. Mus. Nat. Hist., 
13(9):359-427, August 11. 

Deacon, J. E. and Metcalf, A. L. 

1961. Fishes of the Wakarusa River in Kansas. Univ. Kansas Publ. Mus. 
Nat Hist, 13(6):309-322, February 10. 

Deevey, E. S. Jr. 

1949. Biogeography of the Pleistocene. Part I: Europe and North 
America. Geol. Soc. America Bull., 60:1315-1416, September. 



180 University of Kansas Publs., Mus. Nat. Hist. 

DiSTLER, D. A. and Metcalf, A. L. 

1962. Etiieostoma pallididorsum, a new percid fish from the Caddo River 
System of Arkansas. Copeia, 1962(3):556-561, September 28. 

Dyche, L. L. 

1914. Ponds, pond fish, and pond fish cuUure. Kansas State Dept. Fish 
and Game Bull., l:viii + 208 pp. 

Eddy, S. and Surber, T. 

1947. Northern fishes with special reference to the upper Mississippi Val- 
ley. (Second Ed.). University of Minnesota Press, Minneapolis, 
.\ii + 276 pp. 

Elus, M. M. 

1914. Fishes of Colorado. Univ. Colorado Studies, 11(1):1-136, March. 

Evermann, B. W. and Cox, U. O. 

1896. Report upon the fishes of the Missouri River Basin. Rept. Comm. 
Fish and Fisheries, 20:325-429, November 27. 

Fenneman, N. M. 

1931. Physiography of western United States. McGraw-Hill Book Co., 
New York, xiv -f 534 pp. 

FiSHEL, V. C, LoHMAN, S. W. and Stoltenberg, H. A. 

1948. Ground-water resources of Republic County and northern Cloud 
County, Kansas. Kansas Geol. Surv. Bull., 73:1-191, May. 

Fisher, H. J. 

1962. Some fishes of the lower Missouri River. Amer. Midi. Nat., 68(2): 
424-429, October. 

Fitch, H. S. and McGregor, R. L. 

1956. The forest habitat of the University of Kansas Natural History 
Reservation. Univ. Kansas Mus. Nat. Hist. Misc. Publ., 10(3): 
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Flint, R. F. 

1955. Pleistocene geology of eastern South Dakota. U. S. Geol. Surv. 
Prof. Paper, 262:vi + 173. 

1957. Glacial and Pleistocene geology. John Wiley and Sons, Inc., New 
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Forbes, S. A. and Richardson, R. E. 

1909. The fishes of Ilhnois. Illinois Nat. Hist. Surv. Div., cxxxii -f 357 pp. 

Fowler, H. W. 

1925. Notes on North American cyprinoid fishes. Proc. Acad. Nat. Sci. 
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Frye, J. C. 

1955. The erosional history of tlie Fhnt Hills. Trans. Kansas Acad. Sci., 
58(l):79-86, March 26. 

Frye, J. C. and Fent, O. S. 

1947. The late Pleistocene loesses of central Kansas. Kansas Geol. Surv. 
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Frye, J. C. and Leonard, A. B. 

1951. Stratigraphy of the late Pleistocene loesses of Kansas. Jour. Geol., 
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1952. Pleistocene geology of Kansas. Kansas Geol. Surv. Bull., 99:1-230, 
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1957. Studies of Cenozoic geology along eastern margin of Texas High 
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Texas Rept. Invest., 32:1-62, November. 



Fishes of Kansas River System 181 

1959. Correlation of the Ogallala Formation (Neogene) in western Texas 
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1963. Pleistocene geology of Red River Basin in Texas. Bur. Econ. Geo!., 
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Frye, J. C, Leonard, A. B. and Swineford, A. 

1956. Stratigraphy of the Ogallala Formation (Neogene) of northern 
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Frye, J. C. and Willman, H. B. 

1960. Classification of the Wisconsinan Stage in tlie Lake Michigan Gla- 
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Gerking, S. D. 

1945. The distribution of tlie fishes of Indiana. Invest. Indiana Lakes 
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GiBBS, R. H. 

1957. Cyprinid fishes of the subgenus Cyprinella of Notropis. I. Syste- 
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Gilbert, C. H. 

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1887. Descriptions of new and httle known etheostomids. Proc. U. S. 
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1889. Fourth series of notes on the fishes of Kansas. Bull. Washburn 
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Gilbert, C. R. 

1961. Hybridization versus intergradation : an inquiry into the relation- 
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Gilbert, C. R. and Bailey, R. M. 

1962. Synonymy, characters, and distribution of the American cyprinid 
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GiLE, W. S. 

1885. Fish culture. Kansas State Board Agric. 4th Bienn. Rept., 1883- 
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Gill, T. 

1862. Descriptions of new species of Pimelodinae, Proc. Boston Soc. Nat. 
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1864. Proceedings of June 7 (Untitled). Proc. Acad. Nat. Sci. Philadel- 
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1876. Report on ichthyology. In Report of explorations across the Great 
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GlRARD, C. 

1856. Researches upon the cyprinoid fishes inhabiting the fresh waters of 
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182 University of Kansas Publs., Mus. Nat. Hist. 

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Goodrich, C. 

1939. Pleuroceridae of the Mississippi River Basin exclusive of the Ohio 
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1885a. Preliminary list of Kansas fishes. Trans. Kansas Acad. Sci., 9:69-78. 
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Greene, A. R. 

1906. The Kansas River — its navigation. Trans. Kansas State Hist. Soc, 
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Greene, C. W. 

1935. The distribution of Wisconsin fishes. Wisconsin Conserv. Comm., 
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Grinnell, J. 

1924. Geography and evolution. Ecology, 5:225-229, July 29. 

Hall, E. R. 

1946. Mammals of Nevada. Univ. California Press, Berkeley, California, 
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1963. The effects of pool size and beaver activity on distribution and 
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Harlan, J. R. and Speaker, E. B. 

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Hay, O. p. 

1887. A contribution to the knowledge of the fishes of Kansas. Proc. 
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1960. Beaver in Kansas. Univ. Kansas Mus. Nat. Hist. Misc. Publ., 
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1947. Experimental studies on adaptive evolution in Gasterosteus actdea- 
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HiBBARD, C. W. 

1960. An interpretation of Pliocene and Pleistocene climates in North 
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Fishes of Kansas River System 183 

HiBBAKU, C. W., Fhye, J. C. and Leonaiid, A. B. 

1944. Reconnaissance of Pleistocene deposits in north-central Kansas. 
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HODSON, W. G. 

1959. Geology and ground-water resources of Mitchell County, Kansas. 
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1960. Geology and ground-water resources of Go\e Countv, Kansas. 
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HoRBERG, L. and Anderson, R. C. 

1956. Bedrock topography and Pleistocene glacial lobes in central United 
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1930. Materials for a revision of the catostomid fishes of eastern North 

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1942. An atherinid fish from the Pliocene of Oklahonui. Jour. Paleont., 
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HuBBS, C. L. and Black, J. D. 

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HuBBS, C. L. and Bonil^m, K. 

1951. New cyprinid fishes of the genus Notropis from Texas. Texas Jour. 
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1935. Two new subspecies of fishes from Wisconsin. Trans. Wisconsin 
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1958. Fishes of the Great Lakes Region. (Second Ed.). Bull. Cran- 
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HuBBS, C. L. and Ortenburger, A. I. 

1929a. Further notes on the fishes of Oklahoma with descriptions of new 

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IIuBBS, C. L. and TRAtrrMAN, M. B. 

1937. A revision of the lamprey genus Ichthyomtjzon. Misc. Publ. Mus. 
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HuBBS, C. L. and Walker, B. W. 

1942. Habitat and breeding behavior of the American cyprinid fish Notro- 
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1 1—8449 



184 Unr'ersity of Kansas Publs., Mus. Nat. Hist. 

HtTBBS, Clark 

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1955. The distribution of the suckermouth minnow, Phenacobius mirabilis, 
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Jennings, D. 

1942. Kansas fish in the Kansas State College Museum at Manhattan. 
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1958. Geology and ground-water resources of Logan Countv, Kansas. 
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1891. Report of explorations in Colorado and Utah during the summer of 
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1896. The fishes of North and Middle America. Bull. U. S. Nat. Mus., 
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Koster, W. J. 

1957. Guide to the fishes of New Mexico. Univ. New Mexico Press, Al- 
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Lachner, E. a. 

1952. Studies of the biology of the cyprinid fishes of the chub genus 
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L.A.GLER, K. F. and Bailey, R. M. 

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Langsdorf, E. 

1950. The first survey of the Kansas River. Kansas Hist. Quart., 18(2): 
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Latta, B. F. 

1949. Ground-water conditions in the Smoky Hill Valley in Saline, Dick- 
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Leonard, A. B. 

1959. Handbook of gastropods in Kansas. Univ. Kansas Mus. Nat. Hist. 
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1956. Cambiamenti climatici nei Great Plains. Quatemaria, 3:27-37. 

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1957. A fisheries survey of the Big Wichita River System and its impound- 
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Linder, a. D. 

1959. The American percid fishes Ammocrypta clara Jordan and Meek 
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Fishes of Kansas RnER System 185 

LoHAiAN, S. W. and Frye, J. C. 

1940. Geology and ground-water resources of the "Equus beds" area in 
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LUENIN'GHOENER, G. C. 

1947. The post-Kansan geologic Iiistory of the lower Platte Valley area. 
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1935. The Pleistocene geologv of Nebraska. Nebraska Geol. Surv. liuU., 
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1954. Additional studies of the Cenozoic of western Kansas. Kansas 
Geol. Surv. Bull., 109(4):49-64, May 15. 

Metcalf, a. L. 

1959. Fishes of Chautauqua, Cowlev and Elk counties, Kansas. Univ. 
Kansas Publ. Mus. Nat. Hi.st., 11(6) :345-400, May 6. 

Metcalf, A. L. and Distler, D. A. 

1961. New distributional records for two species of crayfish. Trans. 
Kansas Acad. Sci., 64(4) :353-356, January 5. 

Miller, R. R. 

1948. The cyprinodont fishes of the Death Valley System of eastern Cali- 
fornia and southwestern Nevada. Misc. Publ. Mus. Zool. Uni\-. 
Michigan, 68:1-155, April 20. 

1955. An annotated list of the American cyprinodontid fishes of tlie genus 
Fundulus, with the description of Fimduhis persimilis from Yu- 
catan. Occas. Papers Mus. Zool. Univ. Michigan, 568:1-25, 
August 1. 

1961. Speciation rates in some fresh-water fishes of western North 
America in Vertebrate Speciation, W. F. Blair, Editor. Uni\'. 
Texas Press, Austin, pp. 537-560. 

Minckley, W^. L. 

1956. A fish survey of the Pillsbury Crossing Area, Deep Creek, Riley 
County, Kansas. Trans. Kansas Acad. Sci., 59(3) :351-357, Octo- 
ber 31. 

1959. Fishes of the Big Blue River Basin, Kansas. Univ. Kansas Publ. 
Mus. Nat. Hist., 11(7) :401-442, May 8. 

Minckley, \\'. L. and Craddock, J. E. 

1962. A new species of Phenacobius (Cyprinidae) from the upper Ten- 
nessee River System. Copeia, 1962 (2):369-377, July 20. 



186 University of Kansas Publs., Mus. Nat. Hist. 

MiNCKLEY, W. L. and Cross, F. B. 

1959. Distribution, habitat and abundance of the Topeka Shiner Notropis 
topeka (Gilbert) in Kansas. Amer. Midi. Nat., 61( 1):210-217. 

1960. Taxonomic status of the Shorthead Redhorse, Moxostoma aureolitm 
(LeSueur) from the Kansas River Basin, Kansas. Trans., Kansas 
Acad. Sci., 63( l):3.5-.39, April 6. 

MiNCKLEY, W. L. and Deacon, J. E. 

1959. Biology of tlie flathead catfish in Kansas. Trans. Amer. Fish. Soc, 
88(4):344-355, October. 

Montgomery, Mrs. F. C. 

1928. Fort Wallace and its relation to the frontier. Coll. Kansas State 
Hist. Soc, 17:189-283. 

Moore, G. A. 

1950. The cutaneous sense organs of barbeled minnows adapted to life 

in the muddv waters of the Great Plains Region. Trans. Amer. 

Microscop. Soc, 69(l):69-95. 

1957. Fishes. (Part II in X'ertebrates of the United States by W. F. Blair 

and others). McGraw-Hill Book Co., Inc., New York, pp. 33-210. 

Moore, R. C. 

1949. Divisions of the Pennsylvanian System in Kansas. Kansas Geol. 
Sur\'. Bull., 83:1-203, November. 

Needham, p. R. and Gard, R. 

19.59. Rainbow trout in Mexico and California with notes on the cut- 
throat series. Univ. California Publ. Zool., 67(1):1-124, Septem- 
ber 25. 

O'Connor, H. G. 

1960. Ceologv and ground-water resources of Douglas County, Kansas. 
Kansas' Geol. Surv. Bull., 148:1-200, December. 

Olund, L. J. and Cross, F. B. 

1961. CJeographic variation in the North American cyprinid fish, Htjbopsis 
gracilis. Liniv. Kansas Publ. Mus. Nat. Hist., 13(7) :323-348, Feb- 
ruary 10. 

Osborn, H. F. 

1931. Cope: master naturalist. Princeton Uni\'. Press, Princeton, Ne^^' 
Jersey, xvi -f- 740 pp., April. 

Ortenburcer, a. 1. and Hubbs, C. L. 

1926. A report on the fishes of Oklahoma, with descriptions of new 
genera Proc. Oklahoma Acad. Sci., 6:123-141. 

Parker, H. N. 

1911. Quality of the water supplies of Kansas. U. S. Geol. Surv. Water- 
supply Paper, 273:1-375. 

Pi.ummer, N. and Romary, J. F. 

1942. Stratigraphv of the pre-Greenhorn Cretaceous beds of Kansas. 
Kansas Geol. Surv. Bull., 41:313-348, November 30. 

QUINN, J. H. 

1957. Paired river terraces and Pleistocene glaciation. Jour. Geol., 65(2) : 
149-166, March. 

1958. Plateau surfaces of the Ozarks. Proc. Arkansas Acad. Sci., 11: 
36-43. 

RiGGs, C. D. and Bonn, E. W. 

1959. An annotated list of the fishes of Lake Texoma, Oklahoma and 
Texas. Southwestern Nat., 4(4) :157-168, December 15. 



Fishes of Kansas Riv'er System 187 

Robins, C. R. and Rakey, E. C. 

1957. Distributional and nomcnclatorial notes on the suckers of the genus 
Moxostoma. Copeia, 1957 {2):154-155, July 15. 

Ross, R. D. 

1958. Races of the cyprinid fish Campostoma anomalum pullum ( Agassiz) 
in eastern United States. Virginia Agric. Expt. Sta. Tech. Bull., 
136:1-20, September. 

SCHOEWE, W. H. 

1949. The geography of Kansas. Piut II, Physical geography. Trans. 
Kansas Acad. Sci., 52(3):261-333, October 18. 

1951. The geography of Kansas. Part III, Hydrogeography. Trans. 
Kansas Acad. Sci., 54(3) :263-329, October 2. 

1952. Coal resources of the Cretaceous System (Dakota Formation) in 
central Kansas. Kansas Geol. Surv. Bull., 96:69-156, February 29. 

Simon, J. R. 

1946. Wyoming fishes. Wyoming Game and Fish Dept. Bull., 4:1-129. 

SiMPSOiV, J. H. 

1876. Report of explorations across the Great Basin of the territory of 
Utah for a direct wagon-route from C; nip Fl()\-d to Genoa, in 
Carson Valley, in 1859, by Captain J. H. Simpson. War Dept., 
Washington, 518 pp. 

Slaughter, B. H., Crook, W. W., Jr., Harris, R. K., Allen, D. C. and 

Seifert, M. 

1962. The Hill-Shuler local faunas of the upper Trinity River, Dallas and 
Denton counties, Texas. Bur. Econ. Geol. Univ. Texas Rept. In- 
vest., 48:viii+75, December. 

Smith, C. L. 

1954. Pleistocene fishes of the Berends Fauna of Beaver County, Okla- 
homa. Copeia, 1954(4):282-289, October 29. 

1958. Additional Pleistocene fishes from Kansas and Oklahoma. Copeia, 
1958(3) :176-180, August 28. 

1962. Some Pliocene fishes from Kansas, Oklahoma, and Nebraska. 
Copeia, 1962(3):505-520, September 28. 

Smith, G. R. 

1963. A late Illinoian fish fauna from southwestern Kansas and its climatic 
significance. Copeia, 1963(2):278-285, June 14. 

S mucker, S. M. 

1856. The life of Col. John Charles Fremont, and his narrative of ex- 
plorations and adventures, in Kansas, Nebraska, Oregon and Cali- 
fornia. Miller, Orton and Mulligan, New York and Auburn, 493 
pp. 

Snow, F. H. 

1875. The fishes of the Kansas River, as observed at Lawrence. Kansas 
State Board Agric, Annual Rept., 4:139-141. 

Strawn, K. 

1961. A comparison of meristic means and variances of wild and labora- 
tory-raised samples of the fishes Etlwostoma aruJiami and E. 
lepidum (Percidae). Texas Jour. Sci., 13(2):127-159, June. 

Swineford, a. and Frye, J. C. 

1951. Petrography of the Peoria Loess in Kansas. Jour. Geol., 59(4): 
306-322, July. 



1S8 University of Kansas Publs., Mus. Nat. Hist. 

S\\'iNEFORD, A., Leonard, A. B. and Frye, J. C. 

1958. Petrology of the Pliocene pisolitic limestone in the Great Plains. 
Kansas Geol. Surv. Bull., 130 (2):97-116, May 15. 

Taylor, W, R. 

1954. Records of fishes in the John N. Lowe collection from the Upper 
Peninsula of Michigan. Misc. Publ. Zool. Univ. Michigan, 87:1-50, 
November 9. 

Thienemann, a. 

1950. Verbreitungsgeschichte der Suesswassertierwelt Europas. E. Sch- 
weizerbart'sche Verlagsbuchhandlung, Stuttgart, xvi -j- 809 pp. 

Thwaites, R. G. 

1905a. Original journals of the Lewis and Clark expedition 1804-1806. 

Dodd, Mead and Co., New York, Vol. 5, ix -f 395 pp., March. 
1905b. Original journals of the Lewis and Clark expedition 1804-1806. 

Dodd, Mead and Co., New York, Vol. 6, ix + 280 pp., July. 

1905c. Account of an expedition from Pittsburgh to the Rocky Mountains 

performed in the years 1819, 1820. By order of the Hon. J. C. 

Calhoun, Secretary of War, under the command of Maj. S. H. 

Long, of the U.S. Top. Engineers. The Arthur H. Clark Co., 

Cleveland, Ohio, Vol. 14, 321 pp. 
1905d. Travels in the interior of North America by Maximilian, Prince of 

Wied. The Arthur II. Clark Co., Cleveland, Vol. 22, 393 pp. 
1906a. Travels in the far West 1836-1841. II. Letters and sketches, with 

a narrative of a year's residence among the Indian tribes of the 

Rocky Mountains. By Father Pierre Jean de Smet S. J. The Arthur 

H. Clark Co., Cleveland, Vol. 27, pp. 123-411. 

1906b. Journal of travels over the Rocky Mountains to the mouth of the 
Columbia River, made during the years 1845 and 1846. By Joel 
Palmer. The Arthur H. Clark Co., Cleveland, Vol. 30, 311 pp. 

Todd, J. E. 

1914. The Pleistocene history of the Missouri River. Science, n. s., 
39:263-274, February 20. 

1923. Is the channel of the Missouri River through North Dakota of 
Tertiary origin? Geol. Soc. America Bull., 34:469-494, Septem- 
ber 30. 

Trautman, M. B. 

1956. Carpiodes cyprinus hinei, a new subspecies of carpsucker from the 
Ohio and upper Mississippi river systems. Ohio Jour. Sci., 56:33-40, 
January 18. 

1957. The fishes of Ohio. Ohio State Universitv Press, Columbus, Ohio, 
xviii + 683 pp. 

Trautman, M. B. and Martin, R. G. 

1951. Moxostoma aureolum pisolabnim, a new subspecies of sucker from 
the Ozarkian streams of the Mississippi River System. Occas. 
Papers Mus. Zool. Univ. Michigan, 534:1-10, November 12. 

Underihll, J. C. 

1957. The distribution of Minnesota minnows and darters in relation to 
Pleistocene glaciation. Minnesota Mus. Nat. Hist. Occas. Pap., 
7:vi + 45. 

Underhill, J. C. and Merrell, D. J. 

1959. Intra-specific variation in the Bigmouth Shiner (Notropis dorsalis). 
Amer. Midi. Nat., 61( 1):133-147, January. 

\V alters, K. L. 

1956. Geology and ground-water resources of RawUns County, Kansas. 
Kansas Geol. Surv. Bull., 117:1-100, June. 



Fishes of Kansas Rh'er System 189 

Wedel, W. R. 

1936. An introduction to Pawnee archeology. Bur. Amer. Ethnology 
Bull., 112:1-122. 

1941. Environment and native subsistence economies in the central 
Great Plains. Smithson. Misc. Coll., 101 (3): 1-29, August 20. 

1946. The Kansa Indians. Trans. Kansas Acad. Sci., 49(l):l-35, 
August 6. 

1961. Prehistoric man on the Great Plains. Univ. Oklahoma Press, Nor- 
man, Oklahoma, xviii -|- 355 pp. 

Williams, A. B. 

1954. An explanation for the distribution of a North American crayfish. 
Ecology, 35(4):573-575. 

Yerger, R. W. and Sxjttkus, R. D. 

1962. Records of freshwater fishes in Florida. Tulane Stud. Zool., 
9(5):323-330, April 16. 

Zahuranec, B. J. 

1962. Range ex-tensions of some cyprinid fishes in southeastern Ohio. 
Copeia, 1962(4) :842-843, December 31. 

Transmitted May 25, 1965. 



D 
30-8449 



( Continued from inside of front cover ) 

10. A new genus of Pennsylvanian fish (Crossopterygii, Coelacanthiformes ) from 
Kansas. By Joan Echols. Pp. 475-501, 7 figiu-es. October 25, 1963. 

11. Observations on the Mississippi Kite in southwestern Kansas. By Henry S. 
Fitch. Pp. 503-519. October 25, 1963. 

12. Jaw musculature of the Mourning and White-winged doves. By Robert L. 
Merz. Pp. 521-551, 22 figures. October 25, 1963. 

13. Thoracic and coracoid arteries in two families of birds, Columbidae and 
Hirundinidae. By Marion Anne Jenkinson. Pp. 553-573, 7 figures. March 
2, 1964. 

14. The breeding birds of Kansas. By Richard F. Johnson. Pp. 575-655, 10 
figiu-es. May 18, 1964. 75 cents. 

15. The adductor muscles of the jaw in some primitive reptiles. By Richard C. 
Fox. Pp. 637-680, 11 figures in text. May 18, 1964. 

Index. Pp. 681-694. 

Vol. 13. 1. Five natural hybrid combinations in minnows ( Cvprinidae ) . 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, 

Mexico. By William E. Duellman. Pp. 19-72, plates 1-8, 3 figures in text. 

August 16, 1960. 50 cents. 
8. A new subspecies of the slider tiu-tle (Pseudemys scripta) from Coahuila, 

Mexico. By John M. Legler. Pp. 73-84, plates 9-12, 3 figures in text. 

August 16, 1960. 
"4. Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, plates 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. Bv James E. Deacon and Artie L. 
Metcalf. Pp. 309-322, 1 figure in text. February 10, 1961. 

7. Geographic variation in the North American cvprinid fish, Hybopsis gracilis. 
By Leonard J. Olund and Frank B. Cross. Pp. 323-348, plates 21-24, 2 
figures in text. February 10, 1961. 

8. Descriptions of two species of frogs, genus Ptychohyla; studies of Ameri- 
can hylid frogs, V. By William E. Duellman. Pp. 349-357, plate 25, 2 
figvures 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, plates 26-30, 
3 figures. August 11, 1961. 75 cents. 

10. Recent soft-sheUed turtles of North America (family Trionychidae ) . By 
Robert G. Webb. Pp. 429-611, plates 31-54, 24 figures in text. February 
16, 1962. $2.00. 

Index. Pp. 613-624. 

Vol. 14. 1. Neotropical bats from western Mexico.' 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. Miu-saloglu. Pp. 9-27, 1 figure in text. July 24, 1961. 

3. Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. 
Pp. 29-67, plates 1 and 2, 8 figures in text. July 24, 1961. 

4. A new subspecies of the black myotis (bat) from eastern Mexico. By E. 
RajTnond Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December 
29, 1961. 

5. North American yellow bats, "Dasypterus," and a list of the named kinds 
of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr. 
Pp. 73-98, 4 figures in text. December 29, 1961. 

6. Natural history of the brush mouse (Peromyscus boyKi) in Kansas with 
description of a new subspecies. By Charles A. Long. Pp. 99-111, 1 figiu-e 
in text. December 29, 1961. 

7. Taxonomic status of some mice of the Peromyscus boylii group in eastern 
Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 113- 
120,^1 figin-e in text. December 29, 1961. 

8. A new subspecies of ground squirrel ( Spermophilus spilosoma) from Ta- 
mauUpas, Mexico. By Ticvil Alvarez. Pp. 121-124. March 7, 1962. 

9. Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J. 
Knox Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in text. March 7, 
1962. 

10. A new doglike carnivore, genus Cynaretus, from the Clarendonian Pliocene, 
of Texas. By E. Raymond Hall and Walter W. Dalquest. Pp. 135-138, 
2 figures in text. April 30, 1962. 

11. A new subspecies of wood rat (Neotoma) from northeastern Mexico. By 
Ticul Alvarez. Pp. 139-143, April 30, 1962. 

12. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul 
Alvarez, and M. Raymond Lee. Pp. 145-159, 1 figure in text. May 18, 
1962. 

13. A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 
1 figure in text. May 21, 1962. 

*14. The mammals of Veracruz. Bv E. Raymond HaU and Walter W. Dalquest. 
Pp. 165-362, 2 figures. May 20. 1963. $2.00. 

(Continued on outside of back cover) 



^ (Continued from inside of back cover) 

15. The recent mammals of Tamaulipas, Mexico. By Ticul Alvarez. Pp. 363- 
473, 5 figures in text. May 20, 1963. $1.00. 

16. A new subspecies of the fruit-eating bat, Stumira ludovici, from western 
Mexico. By J. Knox Jones, Jr., and Gary L. Phillips. Pp. 475-481, 1 figure 
in text. March 2, 1964. 

17. Records of the fossil mammal Sinclairella, Family Apatemyidae, from the 
Chadronian and Orellan. By William A. Clemens. Pp. 483-491, 2 figures 
in text. March 2, 1964. 

18. The mammals of Wyoming. By Charles A. Long. Pp. 493-758, 82 figs. 
July 6, 196.5. $3.00. 

Index. Pp. 759-784. 

Vol. 15. 1. The amphibians and reptiles of Michoacan, Mexico. By William E, Duell- 
man. Pp. 1-148, plates 1-6, 11 figures in text. December 20, 1961. $1.50. 

2. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox 
Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962. 

3. A new species of frog (Genus Tomodactylus ) from western Mexico. By 
Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962. 

4. Type specimens of amphibians and reptiles in the Museum of Natural His- 
torv, the University of Kansas. By William E. DueUman and Barbara Berg. 
Pp. 183-204. October 26, 1962. 

5. Amphibians and Reptiles of the Rainforests of Southern El Peten, Guatemala. 
By William E. Duellman. Pp. 205-249, plates 7-10, 6 figures in text. Oc- 
tober 4, 1963. 

6. A revision of snakes of the genus Conophis (Family Colubridae, from Middle 
America). By John Wellman. Pp. 251-295, 9 figures in text. October 4, 
1963. 

7. A review of the Middle American tree frogs of the genus Ptychohyla. By 
William E. Duellman. Pp. 297-349, plates 11-18, 7 figures in text. October 
18, 1963. 50 cents. 

*8. Natural history of the racer Coluber constrictor. By Henry S. Fitch. Pp. 
351-468, plates 19-22, 20 figures in text. December 30, 1963. $1.00. 
9. A review of the frogs of the Hyla bistincta group. By William E. DueUman. 
Pp. 469-491, 4 figin-es in text. March 2, 1964. 

10. An ecological studv of the garter snake, Thamnophis sirtalis. By Henry S. 
Fitch. Pp. 493-564, plates 23-25, 14 figures in text. May 17, 1965. 

11. Breeding cycle in the ground skink, Lygosoma laterale. By Hemry S. Fitch 
and Harry W. Greene. Pp. 565-575, 3 figiwes in text. May 17, 1965. 

12. Amphibians and reptiles from the Yucatan Peninsula, Mexico. By William 
E. DueUman. Pp. 577-614, 1 figure in text. June 22, 1965. 

13. A new species of turtle. Genus Kinostemon, from Central America, by John 
M. Legler. Pp. 615-625, pis. 26-28, 2 figures in text. July 20, 1965. 

14. A biogeographic account of the herpetofauna of Michoacan, Mexico. By 
WiUiam E. Duellman. Pp. 627-709, pis. 29-36, 5 figures in text. December 
30, 1965. 

15. Amphibians and reptiles of Mesa Verde National Park, Colorado. By Charles 
L. Douglas. Pp. 711-744, pis. 37, 38, 6 figures in text. March 7, 1966. 
Index in course of preparation. 

Vol. 16. 1. Distribution and taxonomy of Mammals of Nebraska. By J. Knox Jones, Jr. 
Pp. 1-356, pis. 1-4, 82 figures in text. October 1, 1964. $3.50. 

2. Synopsis of the lagomorphs and rodents of Korea. By J. Knox Jones, Jr., 
and David H. Johnson. Pp. 357-407. February 12, 1965. 

3. Mammals from Isla Coztunel, Mexico, with description of a new species of 
harvest mouse. By J. Knox Jones, Jr., and Timothy E. Lawlor. Pp. 409-419, 
1 figure in text. April 13. 1965. 

4. The Yucatan deer mouse, Peromvscus yucatanicus. By Timothy E. Lawlor. 
Pp. 421-438, 2 figures in text. July 20, 1965. 

More nvunbers wiU appear in volmne 16. 
Vol. 17. 1. Localities of fossU vertebrates obtained from the Niobrara Formation (Creta- 
ceous ) of Kansas. By David Bardack. Pp. 1-14. January 22, 1965. 

2. Chorda tympani branch of the facial nerve in the middle ear of tetrapods. 
By Richard C. Fox. Pp. 15-21, May 22, 1965. 

3. Fishes of the Kansas River System in relation to zoogeography of the Great 
Plains. By Artie L. Metcalf. Pp. 23-189, 4 figinres in text, 51 maps. 
March 24, 1966. 

More numbers wUl appear in volume 17. 



S-WP" 



University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 4, pp. 191-256, 13 figs. 

^ ,^„„ "iUS. COM'r'. ZQOI 
June 6, 1966 ^ LIBR^Y 

JUN^a 19156 

Factors Affecting Growth and Production 
of Channel Catfish, Ictalurus punctata s 



BY 



BILL A. SIMCO AND FRANK B, CROSS 



University of Kansas 

Lawrence 

1966 



UNIVERSITY OF KANSAS PUBLICATIONS 
MUSEUM OF NATURAL HISTORY 

Institutional libraries interested in publications exchange may obtain this 
series by addressing the Exchange Librarian, University of Kansas Library, 
Lawrence, Kansas, 66044. Copies for individuals, persons vi'orking in a par- 
ticular field of study, may be obtained by addressing instead the Museum of 
Natural History, University of Kansas, Lawrence, Kansas, 66044. When in- 
dividuals request copies from the Musetun, 25 cents should be included, for 
each 100 pages or part thereof, for the purpose of defraying the costs of wrap- 
ping and mailing. For certain longer papers an additional amount, indicated 
below, toward some of the costs of production, is to be included. 

* An asterisk designates those nximbers of which the Museum's supply is exhausted. 
Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. 

•Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 
figures in text. April 9, 1948. 
Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and distribution. By Rol- 
lin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951. 
"2. A quantitative study of the nocturnal migration of birds. By George H. 
Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951. 
3. Phvlogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473- 
530. 49 figiu-es in text, 13 tables. October 10, 1951. 
"4. Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and 
Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, 
1951. 
Index. Pp. 651-681. 
"Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 
figures in text. December 27, 1951. 
1-37 and index. Pp. 1-676, 1951-1953. 

nplete) Mammals of Utah, taxonomy and distribution. By Stephen D. 
Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952. 

1-15 and index. Pp. 1-651, 1952-1955. 

1-10 and index. Pp. 1-675, 1954-1956. 

1-23 and index. Pp. 1-690, 1955-1960. 

Vol. 10. Nos. 1-10 and index. Pp. 1-626, 1956-1960. 

Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. 

Vol. 12. "1. Functional morphology of three bats: Eumops, Myotis, Macrotus. By Terry 
A. Vaughan. Pp. 1-153, pis. 1-4, 24 figures in text. July 8, 1959. 
*2. The ancestry of modem 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 Sidney 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. 
.5. Natural history of the Bell Vireo, Vireo bellii Audubon. By Jon C. Barlow. 
Pp. 241-296, 6 figures in text. March 7, 1962. 

6. Two new pelycosaiu'S from the lower Permian of Oklahoma. By Richard C. 
Fox. Pp. 297-307, 6 figtu-es in text. May 21, 1962. 

7. Vertebrates from the baiTier island of Tamaulipas, Mexico. By Robert K. 
Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309- 
345, pis. 5-8. June 18, 1962. 

8. Teeth of edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 figures 
in text. October 1, 1962. 

9. Variation in the muscles and nerves of the leg in two genera of grouse (Tyni- 
panuchus and Pedioecetes ) . By E. Bruce Holmes. Pp. 363-474, 20 figures) 
in text. October 25, 1963. $1.00. 

10. A new genus of Pennsylvanian fish ( Crossopterygii, Coelacanthiformes ) from 
Kansas. By Joan Echols. Pp. 475-501, 7 figures in text. October 25, 1963. 

11. Observations on the Mississippi kite in southwestern Kansas. By Henry S. 
Fitch. Pp. 503-519. October 25, 1963. 

12. Jaw musculature of the Mourning and White-winged doves. By Robert L. 
Merz. Pp. 521-551, 22 figures in text. October 25, 1963. 

13. Thoracic and coracoid arteries in two families of birds, Columbidae and 
Hirundinidae. By Marion Anne Jenkinson. Pp. 553-573 7 figures in text. 
March 2, 1964. 

14. The breeding birds of Kansas. By Richaid F. Johnston. Pp. 575-655, 10 
figures in text. May 18, 1964. 75 cents. 

15. The adductor muscles of the jaw in some primitive reptiles. By Richard C. 
Fox. Pp. 657-680, 11 figures in text. May 18, 1964. 

Index. Pp. 681-694. 

(Continued on inside of back cover) 



Vol. 


5. 


Nos. 


•Vol. 


6. 


(Coi 


Vol. 


7. 


Nos. 


Vol. 


8. 


Nos. 


Vol. 


9. 


Nos. 



University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 4, pp. 191-256, 13 figs. 
June 6, 1966 



Factors Affecting Growth and Production 
of Channel Catfish, Ictalurus punctatus 



BY 



BILL A. SIMCO AND FRANK B. CROSS 



University of Kansas 

Lawrence 

1966 



UNrVERSITY OF KaNSAS PUBLICATIONS, MuSEUM OF NATURAL HiSTORY 

Editors: E. Raymond Hall, Chairman, Henry S. Fitch, 
Frank B. Cross 



Volume 17, No. 4, pp. 191-256, 13 figs. 
Published June 6, 1966 



MUS. COMP. ZOOL. 
LIBRARY 

JUN^^1966 

UNIVERSITY 

University of Kansas 

Lawrence, Kansas 



PRINTED BY 

ROBERT R. (BOB) SANDERS. STATE PRINTER 

TOPEKA. KANSAS 

1966 



31-3428 



Factors Affecting Growth and Production 
of Channel Catfish, Ictalurus punctatus 

BY 
BILL A. SIMCO AND FRANK B. CROSS 

CONTENTS 

PAGE 

Introduction 194 

Methods 195 

Terminology and abbreviations 196 

Mortality and Reproduction of Channel Catfish in Ponds. 197 

Production of Ch-\nnel Catfish on "Natural" Foods 197 

Nutritional Aspects of Feeding Channel Catfish 199 

Growth on Feeds Containing Different Amounts of Pro- 
tein 205 

Production on same ration in difiFerent years 210 

Growth by subadults fed experimental rations in a second 

year 210 

Feeding different amounts of protein in same pond 212 

Summary of effects of various levels of protein on growth . . 213 

Effect of Supplemental Feeding on Condition 215 

Effect of Daily Feeding-rate on Production 219 

Effect of Stocking-rate on Production 224 

Effect of Crowding on Growth 226 

Carrying CAPAcnr of Ponds for Channel Catfish 230 

Physical-chemical Conditions in Experimental Ponds 235 

Methods 235 

Temperature 236 

Dissolved oxygen 238 

pH values 241 

Alkalinity 241 

Total dissolved solids 244 

Correlation of Physical-chemical Condfiions With Pro- 
duction 244 

Correlations with average weight 247 

Correlations with condition-index 250 

Correlations with changes in weight 251 

Summary 251 

Acknowledgments 253 

Literature Cited 254 

(193) 



194 University of Kansas Publs., Mus. Nat. Hist. 

INTRODUCTION 

The production of channel catfish, Ictalurus punctatus (Rafi- 
nesque), in small ponds has been under study by the State Bio- 
logical Survey of Kansas since 1954. With the steadily increasing 
number of farm ponds in Kansas, there has been a growing interest 
in raising channel catfish for sport, food and profit. Six advantages 
of channel catfish as a pond-fish are: 

1. Channel catfish are native, well-known to Kansas anglers, and 
are highly popular both as a sport- and table-fish. 

2. They are tolerant of varied environmental conditions, includ- 
ing turbid water. Buck (1956:33) has shown that turbid water is 
not conducive to good growth and reproduction of bluegill and 
bass but that catfish do well in both clear and muddy water. 
Hastings and Cross (1962:20) reported that most ponds in Douglas 
County, Kansas, are too small and muddy to provide satisfactory 
angling for bass and bluegill, and suggested substitution of channel 
catfish in such ponds. 

3. Channel catfish attain a large size; the record for the species 
exceeds 60 pounds. Occasionally, channel catfish weighing 25 
pounds or more are caught from lakes in Kansas, and farm ponds 
have been known to produce channel catfish as heavy as 17 pounds. 
Although such large fish are unusual, they indicate the great ca- 
pacity for growth that is an important attribute of any sport-fish. 
At normal rates of stocking, channel catfish attain a desirable size 
early in Hfe; therefore, they are available for capture by anglers 
during a large part of their life-span. 

4. Channel catfish are omnivorous and opportunistic in their 
feeding habits (Davis, 1959:6). Consequently they readily use 
supplemental feeds introduced in the ponds, making it possible to 
increase pond-production greatly. 

5. Natural reproduction of channel catfish in ponds is not usually 
excessive (Marzolf, 1957:28; Davis, 1959:20-21; and others). Thus, 
knowledge of the approximate number of catfish in a pond is pos- 
sible, and there is little likelihood of stunted growth due to over- 
population. Supplemental feed is utilized eflBciently in rearing the 
fish to usable size. Techniques of propagation of catfish are well- 
developed (Morris, 1939; Brown, 1942; Canfield, 1947; Lenz, 1947; 
Clemens and Sneed, 1957; and Marzolf, 1957). Marzolf (1957) 
discusses methods by which natural reproduction in ponds can be 
assured if desired. 



Production of Cel\nnel Catfish 



195 



6. Fingerling channel catfish for stocking-purposes are available 
from State (Kansas Forestry, Fish and Game Commission, Pratt, 
Kansas), Federal (U. S. Fish and Wildlife Service, Farlington, 
Kansas), and several private hatcheries. 

We investigated growth of channel catfish on "natural" food and 
on supplemental feeds, the ejffects of feeds differing in protein-level 
and source of carbohydrate, and the effects of stocking-rates and 
feeding-rates on production. Also, the physical-chemical situations 
existing in ponds will be considered and correlated with the growth 
of the catfish. 

METHODS 

The experiments to be discussed were made in ponds of the State 
Biological Survey of Kansas in Lawrence. These earth-fill, hatch- 
ery-style ponds are adjacent to one another and are ahke in size 
and shape (%o acre, approximately 114 x 40 feet). Nine ponds 
(eight constructed in 1954 and one built in 1962) slope from a 
depth of two feet at one end to five feet at a drain-box in one 
comer of the opposite end. Two other ponds constructed in 1962 
are subdivided transversely by means of hardware cloth (M inch 
mesh, 6 feet high) recessed at least 12 inches in the pond-bottoms 
and dikes to prevent passage of fish; each of these two ponds con- 
tains four sections, 20 x 40 feet, has open areas at each end of the 
pond (Figure 1) and a bottom that slopes only one foot from the 
shallow end to the drain. The 11 ponds are filled and drained 
independently; the source of water is a reservoir of 20 acre-feet 
that receives runoff from grassland. 



13' 



K- 



-2 0- 



*i 



y////////////yy/////////////^^^^ 



^ 



^ 



B 



D 



g 40- 






H 114' H 

Fig. 1. Diagram of One of the Experimental Ponds (Ponds 10 and 11) that 
Were Subdivided by Hardware Cloth (Partitions Indicated by Vertical Lines). 

Area of Each Pond is %o acre. 



196 University of Kansas Publs., Mus. Nat. Hist. 

The ponds remained dry overwinter, except for ponds used to 
"hold" fish in that season. Ponds were filled and stocked with a 
known number of fish in spring, and the rate of growth was esti- 
mated by means of samples of fish seined at two-week intervals in 
summer. Replicated samples indicated that our sampling proce- 
dure was relatively consistent; t-tests detected no significant differ- 
ences, at the 95 per cent level of probability, between samples 
taken from the same pond on the same day. 

In autumn the ponds were drained and all surviving fish were 
recovered; their number and total weight were recorded, and indi- 
vidual lengths and weights were taken of 25 or more fish from each 
pond. Waterlevels and other factors subject to our control were 
the same in all ponds, except for variables (such as kind and rate 
of feeding) being tested. 

In experiments on feeding of catfish, food was introduced at the 
same places in the ponds once daily at sundown. Channel catfish 
feed most actively from sundown to midnight (Bailey and Harri- 
son, 1948:135-136). In experiments involving fertilization, liquid 
ammonium phosphate (8-25-0 formulation) was used at a rate of 
50 pounds per acre per application. 

Aquatic vegetation rarely grew excessively and was removed 
manually. Liquid fertilizer broadcast on the surface of ponds 
usually resulted in the formation of dense plankton-blooms within 
four days after application. High turbidity caused by the blooms 
usually restricted development of rooted and filamentous vegeta- 
tion. The large poundages of fish present, and frequent seining of 
the ponds, also deterred the development of excessive vegetation. 

Terminology and Abbreviations 

Most of the terminology used in this report is the same as that 
in other works on fishery management; the more important terms, 
as they relate to our studies, can be defined as follows: 

"Carrying capacity" is the maximum poundage of fish (one or 
more species) that a pond will support during a stated interval of 
time. 

"Standing crop" is the poundage of fish (one or more species) 
that is present in a pond at a given time. 

"Production" (gain) is the increase in the weight of fish during 
a designated period (normally one growing season). 

"Yield" is the total poundage of fish obtained when a pond is 
drained. The above terms are expressed as pounds of fish per 
surface-acre of water (Ibs./A). 



Production of Channel Catfish 197 

"Fingerling" refers to a fish of the previous year's spawn; "sub- 
adult" refers to a two-year-old fish (one year older than fingerling). 

In statistical analyses, the symbols "n.s.," "*," and "**" signify, 
respectively, non-significance, significance at the 95 per cent level 
of probability, and significance at the 99 per cent level of prob- 
ability. 

MORTALITY AND REPRODUCTION OF CHANNEL 
CATFISH IN PONDS 

Mortalities in 41 experiments varied from to 22 per cent (aver- 
age of 5 per cent); mortalities in 1961-64 varied from to 7 per 
cent. 

Although spawning by channel catfish in small impoundments 
has been reported (Marzolf, 1957:22; Tiemeier, 1957:295; Davis, 
1959:9), and has occurred in several private ponds under study by 
the State Biological Survey in eastern Kansas, no spawns of channel 
catfish have occurred in our experimental ponds. No suitable nest- 
sites are available in the experimental ponds, and most of the cat- 
fish used were less than three years old, the youngest age at which 
channel catfish normally spawn (Brown, 1942:311). 

PRODUCTION OF CHANNEL CATFISH ON "NATURAL" 

FOODS 

To ascertain production of channel catfish that subsisted solely 
on foods naturally available in our ponds, nine experiments were 
conducted without the addition of supplemental feed or fertiHzer 
(Table 1). 

Stocking-rates varied from 500 to 2250 fingerling catfish per acre. 
Except as noted (two cases, Table 1), channel catfish were the 
only fish in these ponds. In one growing season standing crops of 
98 to 162 pounds of catfish per acre were obtained. The average 
standing crop was 131 pounds per acre. In four experiments con- 
ducted for two growing seasons, stocking-rates of 500, 510, 1000, 
and 1000 catfish per acre yielded 183, 182, 170 and 135 pounds per 
acre, respectively (average 168 pounds). 

Several workers (Ball, 1952; Brown, 1951; Hansen et al., 1960; 
Swingle, 1947) have reported increases in the standing crops of 
fishes, following the addition of various inorganic and organic fer- 
tiHzers to ponds. Some kinds of fish receive greater benefits, or 
more direct benefits, from fertihzation than do other species. We 
attempted to determine the eflFect of fertihzation on the production 
of channel catfish (Table 1). Two ponds were stocked at rates of 



198 



University of Kansas Publs., Mus. Nat. Hist. 



Table 1. — Production of Channel Catfish in Ponds 


[Each Mo Acre). 




Without Extensive Supplemental Feeding. 
















•-^ 










<u 




<o 


v 










(h 




u 


(> 










O 




c 


^-s « 






bC 




03 




03 


a « 






c 




;-i 


■1.3 


bl 


■3 >- 






• (.^ 




<U 


.J= 


a> 


S. <u 




^ 


o 4) 


■4-> 


a 


.£P^ 


P. 


bC ci, 




-Q 


O >-< 


c 


CD 


OJ (n 


aj 


rl <» 




a 


06 C3 




73 

5n = 


^'2 




.s-H 




a 






.2 o 




3 
O 


s§ 


^ 


Tl 


a, »• 


-S a 


:3 a 


S c 


13 CL 


-S o< 






c 


1 


> 

< 








1963 
1955 
1963 
1959 
1958 
1959 
1963 
1958 
1958 
1959 
1959 



9 


2250 


2 


500 


6 


750 


6 


1000 


5 


700 


7 


1000 


1 


2250 


3 


700 


2 


700 


5 


750 


4 


700 



At end of one growing season 



7 
(18) 

3 
(22) 

4 
(20) 

2 

1 

1 

4 

1 



t 

t 

200 

500 



0.05 


108.7 


.24 


98.0 


.15 


115.3 


.14 


109.01 


.16 


107.5 


.14 


112.02 


.07 


162.5 


.21 


147.0 


.23 


155.0 


.23 


162.5 


.24 


162.5 



At end of two or more growing seasons 



78. 
91. 
92. 



96.0 
98.4 
99.0 
132.3 
137.9 
145.9 
152.8 
153.4 



1959-60. . . . 


7 


1000 


20 




.17 


135.02 


122.0 


1959-60.... 


6 


1000 


22 




.22 


170.31 


157.3 


1955-56.... 


1 


510 


12 




.40 


182.0 


175.4 


1955-57. . . . 


2 


500 


18 




.47 


183.0 


176.5 



f Small quantities of feed placed on trays in the ponds were not eaten but may have had 
a fertilizing eflEect. 

1. Contained bass, as well as channel catfish, in both first and second growing seasons, 
and bluegill in second growing season. 

2. Contained bass and bluegiU, in addition to channel catfish, in both first and second 
growing seasons. 



700 to 750 fingerling catfish/A and were fertilized periodically in 
summer. On draining after one growing season, each pond yielded 
162.5 pounds of catfish per acre. In two other ponds stocked at a 
rate of 700 catfish per acre, small amounts of feed (totaling ap- 
proximately 30 pounds, or 300 pounds per acre, for the entire year) 
were placed on trays in the ponds. Observations of the trays indi- 
cated that this feed was not eaten, but it may have had a fertilizing 
effect in the ponds. These ponds yielded 147 and 155 pounds of 
catfish per acre when drained after one growing season. 

In 1963 natural production of fish in relatively infertile ponds 
differed from that in fertile ponds. Pond No. 1 (built in 1954) 
had been fertilized in some years, and in feeding-experiments had 
yielded the highest productions obtained in any of our ponds; 



Production of Channel Catfish 199 

therefore, it is considered a fertile pond. Pond No. 9 (constructed 
in 1962) was filled with water for the first time on the date of 
stocking, April 2, 1963. No fertihzer nor feed was added to either 
pond in 1963. The yields of catfish in these two ponds, when 
drained on October 16-17, 1963, were 162 pounds per acre in Pond 
No. 1, and 109 pounds per acre in Pond No. 9 (Table 20, p. 247). 
In Pond 1 catfish did not grow after August 12, when 193 pounds 
per acre were estimated to be present. The 162 pounds of fish per 
acre obtained in Pond 1 equals the poundage obtained earlier in 
ponds that were fertilized several times in the growing season of 
the fish being raised in those ponds. Seemingly, the production of 
catfish varied with the basic fertility of the ponds, and the time of 
application of fertilizer was unimportant in that the effects of fer- 
tilization carried over from previous years. The duration of such 
effects probably is limited; Tanner (1960) reports that ponds 
undergo reduction in their standing crop three or more years after 
fertilization is stopped. 

The different stocking rates that were used did not affect the 
yields significantly. The highest poundages obtained on "natural" 
foods were in ponds stocked with 700 and 2250 catfish per acre. 
(Table 1). High stocking rates yielded large numbers of small 
fish having approximately the same total weight as fish stocked at 
lower rates. 

The greatest poundage of catfish that our ponds will support 
without supplemental feeding (their carrying capacity) is less than 
200 pounds per acre. 

NUTRITIONAL ASPECTS OF FEEDING CHANNEL 

CATFISH 

The nutritional requirements of fish only recently have been 
considered in detail, and such studies have dealt primarily with 
sahnonids (Phillips et al., 1963 and previous reports of the Cortland 
Hatchery; Halver and Coates, 1957). A few publications exist on 
the nutritional requirements of channel catfish (Nail, 1965; Shell, 
1963). 

Effects of synthetic diets on the growth of channel catfish were 
investigated in a concrete feeding-trough, 36 feet long. The trough 
is divided by screens into 24 compartments, each of which is 15 
inches wide, 5 feet long and approximately 30 inches deep. Fil- 
tered tapwater flows continuously through the trough, entering at 
several points along the bottom and overflowing through several 
outlets near the top of the trough. 



200 



University of ICansas Publs., Mus. Nat. Hist. 



In preliminary investigations conducted in 1960, channel catfish 
were fed rations containing 10, 20, 30 and 40 per cent protein. The 
rations consisted of vitamin-free casein and dextrin in amounts that 
gave the desired protein-levels, plus a vitamin-mixture and Salts IV. 
The vitamin-mixture used was that recommended by Halver and 
Coates (1957) for rainbow trout; subsequently, we learned that 
the vitamin-requirements of channel catfish have been investigated 
by Dupree (unpublished doctoral dissertation, Auburn University, 
1960). The amounts of casein and dextrin were adjusted to give 
the various protein-levels, with vitamins and salts constant in all 
rations. 

Two methods of force-feeding were used — capsules and syringes. 
Gelatin capsules (size 00) filled with 1.5 grams of ration were 
forced into the stomach of the fish by means of forceps. Syringes 
used were of the 10 cc veterinary type having graduated plungers. 
These were modified by drilling out the needle-adapter so that 
alloy tubing of relatively large bore could be fitted to the syringe. 
The short section of metal tubing was tipped with flexible plastic 
tubing to minimize injury to the fish. The dry rations were diluted 
with distilled water to a semi-solid consistency that could be ex- 
truded through the tube, into the stomach of the fish. 

Three channel catfish having an average weight of 94 grams 
were placed in each of 18 compartments, and fed once daily from 
August 13 to October 11, 1960. Each of the four rations was fed 
to fish in four compartments, two groups of fish by syringes and 
two by means of capsules. Two other groups of fish received a 
commercial food ( Clark's Pond-fish Food, containing approximately 



Table 2. — Average Gains (in Grams) of Channel Catfish Confined in 
Troughs, on Synthetic Diets Used in 1960. 


Per Cent Protein 


Feeding method 


in Ration 


Syringe 


Capsule 


10 


-11.3 

-2.8 

7.1 

7.1 

-11.9 


17 9 


20 


19 


30 


14 2 


40 


26 6 


35 (Clark's) 


21.3 







Production of Channel Catfish 



201 



35 per cent protein), which was administered by both feeding- 
methods. 

Growth by all fish was too slight (Table 2) to evaluate the ef- 
fects of different protein-levels, possibly due to the short duration 
of the experiment (55 days) late in the growing season. Fish that 
were fed capsules consistently gained more weight than fish re- 
ceiving the same food by means of syringe. The reasons for su- 
periority of capsules may have been 1) undetected regurgitation 
of food administered by syringe, or 2) injury to the stomach-lining 
and interference with digestive processes of fish fed by syringe. 
When all fish were sacrificed after completion of tiie experiment, 
injury to the wall of the stomach was noted in only one fish fed by 
syringe. Our results indicate that in projects requiring force-feed- 
ing, capsules are preferable to syringes. 

In 1961, synthetic diets containing 20, 40 and 60 percent protein 
were used. Fish that received the 10 per cent protein-ration in 



Table 3. — Composition of Synthetic Diets Used in Experimental 
Trough, 1961 (Quantities Expressed as Per Cent). 


Constituent 


Protein-level 


20 


40 


60 


Starch, sucrose, dextrin or cellulose 

Casein 


70.44 

20.00 

3.97 

4.96 

0.79 


50.6 
39.6 
3.97 
4.96 
0.79 


30.75 
59.52 


Salts IV 


3.97 


Corn Oil .... 


4.96 


Choline 


0.79 


Vitamins t . . 













\ Vitamins as recommended by Dupree, unpublished doctoral dissertation. Auburn Uni- 
versity, 1960. 





Amount 








Thiamin 


6 


Mg/lOOgm of 


casein 




Riboflavin 


20 






B« 


4 








Niacin 


80 






Mixed with casein 


Calcium pantothenate 


28 








Inositol 


200 








Biotin 


0.6 








A 


1 


Gm/450gm oil 


Mixed with oil 


E 


8 


}> 


>} 





202 University of Kansas Publs., Mus. Nat. Hist. 

1960 had friable, discolored livers indicative of fatty metamorphosis; 
hence, w^e discontinued testing at this level and increased the upper 
limit of protein to 60 per cent. In addition, the carbohydrate con- 
stituent of the feed varied in 1961; cellulose, sucrose, dextrin, and 
starch were used (Table 3). The vitamin-mixture was that of 
Dupree {loc. cit.) rather than Halver and Coates (1957). 

Because of poor growth by fish that were force-fed in the previous 
year, "ad-lib" feeding was attempted. The rations were diluted with 
distilled water, pressed through a pastry decorator (or caulking 
gun) into long strips and then allowed to dry on screens. When 
dry, the strips were broken into small pieces by hand. The rations 
were then weighed and placed loosely in each section of the trough. 

Five fish ( average weight = 17.7 grams, average length — 132 
mm) were placed in each of 24 sections of the experimental trough. 
Each of the 12 rations was fed to fish in two compartments (a total 
of 10 fish) twice daily at the rate of approximately 5 per cent of 
body weight per day, from July 1 until October 16, 1961. 

Gains made on the various diets are shown in Table 4. An analy- 
sis of variance demonstrated a significant difference (at the 95 per 
cent level of probability) in growth associated wdth the different 
protein-levels as well as wdth the type of carbohydrate present. 

Source of variation Degrees of freedom Mean square F 

Among Protein-levels 2 2444.69 7.216 •• 

Among Carbohydrates 3 1830.52 5.403 " 

Interaction 6 478.24 1.411 n.s. 

Error 12 338.79 

Total 23 

Growth by our catfish correlated directly with the protein-level 
in most diets (Table 4). Differences in growth were mainly be- 
tween the 20 and 40 per cent protein-rations (202 grams), while 
only 66 more grams were gained on the 60 per cent ration than on 
the 40 per cent ration. In studies by Nail ( 1965), the highest gains 
were made on a diet containing 34.8 per cent protein (the highest 
protein-level that Nail used), but growth of his fish on this feed 
was not significantly higher (P ?^ .95) than that made on a diet 
containing 25.3 per cent protein. 

Catfish that received feeds containing cellulose best reflected the 
positive effect of increased protein-content in the synthetic rations 
that we used (Table 4, Figure 2). We assume that channel cat- 
fish derive little energy from cellulose; thus, growth made on the 
diets containing cellulose was due to constituents other than car- 
bohydrates. The other carbohydrates used in our rations (starch, 
dextrin, sucrose) modified the effect of increased protein-content, 



Production of Channel Catfish 



203 



Table 4. — ^Weight-gains (Grams) by Channel Catfish, Confined in 
Experimental Trough, on Synthetic Diets Used in 1961. 



Protein-level 


Source of carbohydrate 


Cellu- 
lose 


Sucrose 


Dextrin 


Starch 


Total 


Average 


20 


24.3 
34.9 


67.0 
46.9 


57.7 
77.0 


89.7 
122.1 










40 


59.2 

78.0 
60.7 


113.9 

106.8 
65.1 


134.7 

113.3 
101.5 


211.8 

124.4 
71.8 


519.6 


65.0 






60 


138.7 

91.0 
96.0 


171.9 

86.4 
107.8 


214.8 

102.7 
75 1 


196.2 

113.6 
115.4 


721.6 


90.2 








187.0 


194.2 


177.8 


229.0 


788.0 


98.5 


Total 


384.9 


480.0 


527.3 


637.0 


2029.2 








Average 


64.1 


80.0 


87.9 


106.2 




84.6 







presumably due to their protein-sparing action. Nail {loc. cit.) 
calculated that 0.23 grams of carbohydrate spared 0.05 grams of 
protein per hundred grams of fish from use for energy by his chan- 
nel catfish. The kind of carbohydrate had httle eflFect on the growth 
made by channel catfish on our diets containing 60 per cent protein, 
presumably because that amount of protein was so high that addi- 
tional energy in the form of carbohydrates was not needed. In 
rations containing lower levels of protein, the amount and kind of 
carbohydrate were important. 

Starch, dextrin, and sucrose were utihzed by channel catfish, but 
were not equally beneficial. More growth was made by our fish 
on diets containing starch than on diets containing dextrin, and 
growth on the dextrin-rations exceeded growth on diets containing 
sucrose (Table 4, Figure 2). This relationship is opposite to that 
reported for chinook salmon by Buhler and Halver (1961:316), who 
found that "fish growth decreased considerably with increasing 
carbohydrate molecular weight." 

In explanation of the differences in growth that were associated 
with the kind of carbohydrate in our tests two possibihties occur 
to us. 

First, more rapid dissolution of sucrose than of the less soluble 
dextrin and starch, before ingestion of the rations by the fish, may 



204 



Untvebsity of Kansas Publs., Mus. Nat. Hist. 



120 



100 



< 80 



Z 60 
< 



40 



20 



STARCH . 



DEXTRIN 



SUCROSE 



CELLULOSE 




A, 



20 



40 



60 



PER CENT OF PROTEIN IN FOOD 



Fig. 2. Gains (in Grams) Made by Fingerling Channel Catfish on Synthetic 
Rations Containing Different Forms of Carbohydrate, while Confined in Ex- 
perimental Trough from July 1 to October 16, 1961. Lines are Regressions on 
Absolute Gains Made by Duplicate Groups of Catfish on Each Protein-level 
(20, 40, and 60 Per Cent). Open Circles Represent Rations Containing Starch; 
Open Triangles, Dextrin; Solid Circles, Sucrose; Solid Triangles, Cellulose. 



have reduced the food-value actually received by fish on the sucrose 
diet. Tunison et al. (1939:11) noted that loss of sucrose by disso- 
lution affected their results in feeding tests using brook trout. 

Second, the amount of digestible carbohydrate used in our low- 
protein rations may have been excessive in relation to the capacity 
of channel catfish to utilize carbohydrate efficiently. Some authors 
(Phillips et al, 1940:24-25, 1963:39-44; McLaren et al, 1946:440) 
have concluded that trout have limited tolerance for carbohydrate, 
because their fish grew poorly and/or had liver-abnormalities when 
fed diets high in carbohydrates. In contrast, Buhler and Halver 
(1961:307) and DeLong et al (1958) found that chinook salmon 
use carbohydrates at levels of 48 to 63 per cent of the diet with- 
out ill effects or decrease in growth. Nail (loo. cit.) reported that 
carbohydrate-levels of at least 18.6 per cent are beneficial to channel 
catfish and suggested that higher levels probably can be utilized 
efficiently by that species. The carbohydrate-levels in diets that we 



Production of Channel Catfish 205 

used were approximately 30, 50, and 70 per cent in the rations con- 
taining 60, 40, and 20 per cent protein, respectively. 

Large variations in the digestion- and absorption-rates of diJBFerent 
forms of carbohydrate were reported by Tunison et al. (1943:10); 
on the average, 75 per cent of total sucrose in rations was absorbed 
by brook trout, but only 65 per cent of total dextrin and 40 per 
cent of total starch were utilized. Therefore, if high carbohydrate- 
levels are harmful rather than beneficial to catfish, the deleterious 
effect of diets containing starch might be less than that of diets con- 
taining sucrose, accounting for the better gains by fish receiving high 
levels of starch than by those receiving high levels of sucrose in our 
tests (Figure 2). 

In an unrelated set of experiments, we have noted that the degree 
of enlargement and vacuolar degeneration in livers of fed channel 
catfish varied inversely with the protein-level in their rations (hence 
tissue-damage correlated with carbohydrate-content). These re- 
sults were obtained with pondfish fed pelleted rations composed of 
commercial foodstuffs, and will be reported elsewhere by C. E. Judd 
and Cross; but, the findings seem pertinent to the above discussion 
in suggesting that a high intake of digestible carbohydrate may be 
detrimental to channel catfish. 

GROWTH ON FEEDS CONTAINING DIFFERENT 
AMOUNTS OF PROTEIN 

Channel catfish are omnivorous, at least in some degree (Davis, 
1959:6). In many natural waters insects are the primary food 
of channel catfish (Bailey and Harrison, 1948:125-130), but the 
variety of items other than insects that have been found in stomachs 
of channel catfish attests that they are opportunistic in their feed- 
ing habits. Perhaps because of this lack of selectivity, channel cat- 
fish respond well to supplemental feeding ( Swingle, 1959; Tiemeier, 
1962), making it possible to increase pond-production by several 
hundred pounds per acre. Although several investigators ( Swingle, 
1957, 1959; Tiemeier, 1957, 1962, 1965; Nail, 1965) reported the re- 
sults of supplemental feeding, relatively little has been pubHshed 
concerning the effects of supplemental feeds containing various 
levels of protein. 

In order to test the response by channel catfish stocked in earthen 
ponds, to feeds containing different levels of protein, studies were 
made in 1961 and 1962 using supplemental feeds having 20, 30, 40 
and 50 per cent protein ( see Table 5 for composition of feeds ) . 



206 



University of Kansas Publs., Mus. Nat. Hist. 



Table 5. — Composition of Pelleted Feeds Used in Ponds in 1961 and 1962 

(Expressed in Per Cent). 



Source 


Per cent protein 


20 


30 1 


40 1 


50 


Blood meal 


1.5 
7.4 
7.4 
32.5 
44.8 
4.9 
1.5 
0.1 


3.5 

17.25 

17.25 

23.7 

31.8 

4.9 

1.5 

0.1 


5.5 

27.2 

27.2 

13.85 

19.8 

4.9 

1.5 

0.1 


7.9 


Fish meal 


39.45 


Soybean meal 


39.45 


Corn, ground yellow 


2.8 


Wheat branft 


3.9 


Brewer's dried yeast 


4.9 


Salt 


1.5 


Vitamin A concentrate^ 


0.1 







f In 1961, 30 and 40 per cent protein-feeds were formed by mixing the 20 and 50 per 
cent protein-feeds in the desired proportions. 

f t Wheat bran used in 1961, wheat brown shorts used in 1962. 
I Vitamin A Concentrate added only in 1962. 



The growth-rates of catfish receiving these feeds are represented 
by the lower set of 4 lines in Figure 3, for biweekly intervals in the 
summer of 1961. The weight of fish in each pond was the same 
when the ponds were filled and stocked on May 13-15, 1961. 
Variation in growth associated with the difiFerent feeds was evident 
when the fish were checked in June, The estimated weights of 
catfish fed 20, 30, 40 and 50 per cent protein-feeds were, respec- 
tively, 73, 90, 121, and 79 Ibs./A on June 12, and 93, 107, 141, and 
127 Ibs./A on June 26. On these two dates, there seemed to be a 
correlation between growth-rate of the fish and the protein-level 
of the feed. Thereafter, the standing crops fluctuated but growth 
by fish receiving 30, 40, or 50 per cent protein differed only slightly 
through August 8. Growth on the 20 per cent protein-feed was 
consistently lower than that on the other rations during this period. 
The standing crops on August 8, 1961, of fish receiving 20, 30, 40 
and 50 per cent protein feeds were 315, 467, 455 and 433 Ibs./A re- 
spectively. Subsequently, greater divergence in growth-rates oc- 
curred. Yields when the ponds were drained in November were 
550, 703, 783, and 624 Ibs./A, in order of increasing protein-level in 
the feed. 



Production of Channel Catfish 



207 



1600 
1400 



a: 

^ 1000 



20% 

. 30% 

.--.40% 
.50% 




1962 



1961 



September October 



Fig. 3. Standing Crops ( in Pounds Per Acre ) of Channel Catfish that Received 

Feeds Containing Different Amounts of Protein, in Open Ponds in 1961 ( Lower 

Set of Four Lines ) and 1962 ( Upper Set of Four Lines ) . 

An analysis of variance, based on individual weights of 25 fish 
from each pond when drained in November, gave the following 
results : 



Source of variation Degrees of freedom Mean square 

Among Feeds 3 0.0353 

Within Feeds 96 0.0105 



F 

3.53* 



Variation among feeds was significant at the 95 per cent probability- 
level. In order to determine where the differences lay, a Duncan's 
Multiple Range test was applied (Steel and Torrie, 1960:107-109), 
using the mean weight obtained on each feed. The results are as 
follows: 



Feed (% Protein) 20 

Mean Weight (Pounds) 0.24 



50 

0.27 



30 
0.30 



40 

0.33 



Weights underlined are not significantly different at the 95 per cent 
level of probabiHty. Note that 50 per cent protein-ration is listed 
out of sequence; the groups are arranged in increasing order of 
weights of fish. Factors other than protein-level are thought to 
have influenced the results of this experiment, and are discussed on 
page 240. 



2—3428 



208 University of Kansas Publs., Mus. Nat. Hist. 

The 30 per cent protein-ration was used in a second pond in 1961. 
(This pond will henceforth be designated Pond SOB and the pond 
discussed above, where the same 30 per cent protein-ration was 
used at the same daily rate, will be designated Pond 30A. ) Pond 
SOB was a holding-pond that contained 20,000 fingerling catfish 
ovenvinter, whereas the other ponds were dry in that season. Pond 
30B was stocked two weeks later with smaller fish than those stocked 
in the other ponds. Thus the experimental conditions in Pond SOB 
did not duplicate those in Pond 30A. The growth-rates of fish in 
the two ponds receiving 30 per cent protein feeds are diagrammed 
in Figure 4. Throughout late spring and summer, the initial dif- 
ference in size of the two groups of fish persisted, but the rates of 
growth by fish in both ponds seemed similar. Growth by catfish 
in Pond 30B continued later into fall than did growth of fish in Pond 



1000 

LlI 

cc 

o 

< 8001- 

oc 

LU 

a. 
m 

Q 

-z. 
r) 
o 

Q. 



600 
400 
200 



POND 30A- 




POND 30B 



August September October 



Fig. 4. Standing Crops (in Pounds Per Acre) of Channel Catfish in Two 
Ponds Where the Same Feed, Containing 30 Per Cent Protein, Was Used 

in 1961. 

SOA, where no gains in weight were recorded after September 21. 
Thus the fish in Pond SOB almost overcame their initial deficit and 
attained sizes almost as large as those of fish in Pond SOA. Values 
of "t" contrasting these two ponds on three dates, based on in- 
dividual lengths of fish were: June 26—8.75 **, October 4—2.807 *, 
and November 4 — 2.025 n.s. An analysis of variance based on in- 
dividual weights at the time of draining (November 4) was also 
non-significant ( F = 2.02, d.f . r= 1/48 ) . Although our analysis de- 
tected no significant difference in the average size of the fish in the 
two ponds (as compared to the variability among fish wdthin each 
pond), the actual yield was 13 per cent (81 pounds per acre) higher 
in Pond SOA than in SOB. Values of "t" obtained in a comparison 
of the yield of Pond SOB with yields of other ponds used in 1961 are 
given below, based on weights of individual fish in each pond: 



Production of Channel Catfish 209 

Feed Used (% Protein) . 20 30A 40 50 

Yield (Pounds Per Acre) . 550 703 783 624 

30B, 622 pounds 1.59 n.s. -1.69 n.s. -3.27 "* -0.13 n.s. 

The poundage of fish obtained from Pond SOB was significantly 
lower than that obtained in the pond where fish received a 40 per 
cent protein ration; differences in yields between Pond SOB and the 
other ponds were not significant. In contrast, the yield in Pond 
SOA was significantly higher than that in the pond where a 20 per 
cent protein-ration was used, and was not significantly lower than 
that in the pond where a 40 per cent protein ration was fed. In 
both ponds where 30 per cent protein-feed was used, absolute yields 
were higher (by 72 and 15S Ibs./A) than those obtained from the 
20 per cent protein-feed. 

Experimental feeding of rations containing different levels of 
protein were repeated in 1962 (Table 5). The patterns of growth 
in the different ponds are shown by the upper set of four lines in 
Figure 3. Fish receiving the 20 per cent ration grew slowly in late 
June and early July, but on August 13, our samples indicated that 
these fish had attained weights approximating those in the other 
ponds. The estimated standing crops of fish on three dates in 
summer are given below: 

Feed used 
(% protein) June 18 July 16 August 13 

20 390 520 890 

30 388 705 964 

40 394 734 922 

50 457 770 882 

Through August IS, differences in weight-gain on SO, 40, and 50 
per cent protein were small. Subsequently, differences in growth- 
rates increased. The fish fed the SO per cent protein-ration had 
the highest standing crop on August 13, and maintained the fastest 
growth-rate thereafter. Drainage of the ponds in October yielded 
poundages of fish on the 20, SO, 40, and 50 per cent protein-rations 
respectively, as follows: llOS, 1515, 1314, and 1245 pounds per 
acre. An analysis of variance based on mean weights of fish in 14 
subsamples from each pond gave the following results: 

Source of variation Degrees of freedom Mean square F 

Among Feeds 3 0.113 20.46 ** 

Within Feeds 52 0.00552 

Variation among feeds was highly significant. A Duncan's Multiple 
Range test, based on the average weight of the fish receiving each 
feed, indicated highly significant differences (P = .99) in the 
weights obtained on all feeds, with one exception — the results on 



210 University of Kansas Publs., Mus. Nat. Hist. 

the 40 and 50 per cent protein rations were not significantly dif- 
ferent (P?^.95). 

Production on Same Ration in Different Years 

Productions obtained in 1962 were much higher than those ob- 
tained on the same feeds in 1961 ( Figure 3 ) . The highest standing 
crop present at any time in 1961 was lower than the lowest standing 
crop on the same date in 1962. An analysis of variance indicated 
highly significant differences between gains made by fingerHng cat- 
fish on the same experimental rations, in 1961 and 1962: 

Source of variation Degrees of freedom Mean square F 

Between years 1 791,911 97.1 ** 

Among Feeds 3 31,310 3.84 n.s. 

Discrepance 3 8,156 

Thus, factors other than the type of feed affected the standing 
crops obtained in these two years. The same stocking-rates were 
used in both years, but the fingerling catfish stocked in 1962 were 
larger than those stocked in 1961; not until early July did the 1961- 
fish attain a size equal to that of the 1962-fish when the latter were 
stocked in May, 1962. The fish in 1962 maintained their size- 
advantage throughout the growing season. Factors other than 
protein-level and initial size of fish (for example, variation in 
weather and differences in water-chemistry) may have contributed 
to the differences obtained in the two years; some of these factors 
will be discussed later. 

Growth by Subadults Fed Experimental Rations in a Second Year 
In order to test the effect of different protein-rations in two grow- 
ing seasons, each of two ponds was stocked 1962 with an equal 
number of subadult catfish that had received 40 per cent protein 
and 20 per cent protein in 1961. The adipose fin was clipped on 
the fish that had received the 40 per cent protein-feed, for identi- 
fication. In 1962, the 20 per cent ration was used in one pond, and 
the 40 per cent ration in the other. Thus, half of the fish in each 
pond received the same feed, and half received a different protein- 
level than that fed to them in the previous year. Some fish (an 
equal number from each group ) were removed for observation dur- 
ing the summer. The poundages of fish at the time of stocking 
(April 17) and draining (October 2-4, 1962) are presented in 
Table 6. 

An analysis of variance was performed based on random sub- 
samples of all fish taken at the time of draining: 



Production of Channel Catfish 



211 



Sotirce of variation Degrees of freedom Mean square F 

Between Ponds 1 0.019 0.359 n.s. 

Between Groups 

Within Each Pond 1 0.2173 7.148 » 

Interaction 1 0.0469 1.543 n.s. 

Error 28 0.0304 

Table 6. — Gains (in Pounds) Made by Catfish Receiving Experimental 

Rations Over a Two-year Period. 



Feed used 
(% protein) 


Pond 

number 

(1962) 


Poundage 
stocked 
per acre 
in 1962 


Yield 
(lbs. 'A) 
in 1962 


Gain 
(lbs. /A) 
in 1962 


Proportional 
gain 


1961 


1962 


(in per cent) 


20 
40 


20 
20 

40 
40 


4 
4 

3 
3 


187 
260 


658 

682 


471 
422 


252 

162 


20 
40 


447 
166 
262 


1,340 
617 

784 


893 
451 
522 


271 
200 




428 


1,401 


973 





The yields of the two ponds in 1962 did not differ significantly; 
therefore, the advantage of a 40 per cent protein-ration over a 20-per 
cent protein-ration seemed far less among subadult catfish (0.20- 
1.40 pounds) than among fingerling catfish. The fish that were fed 
40 per cent protein as fingerlings were larger when stocked in the 
two ponds than were the fish previously fed 20 per cent protein; 
the significant difference persisted to the time when the two ponds 
were drained in 1962. The two groups of fish that gained the 
most weight in 1962 were those that received the same ration in 
both years — whether their ration contained 40 per cent protein or 
20 per cent protein. This result is consistent with the findings by 
Shell (1963), who reported that growth-rates of channel catfish de- 
creased temporarily after a change in the protein-level of their feed 
— whether protein was increased or decreased. However, the 
period of feeding in our experiments was much longer than the term 
of the residual effect reported by Shell. 

Further interpretation of our results is complicated by differences 
in size among the four groups of fish when they were stocked in 
1962, relative to known variation in the capacity for growth by 
fish at different stages in their life-histories. As a general rule, a 
large fish can (potentially) gain more weight in a brief interval 



212 University of Kansas Publs., Mus. Nat. Hist. 

than can a small fish; but, the gain made by the small fish generally 
is higher proportional to its initial weight than that made by the 
large fish. From the standpoint of this rule, the order of absolute 
gains by the four groups of fish (from largest to smallest) should 
have been as follovi^s if the protein-level in their feed in 1962 were 
not a factor: group 40-40, group 40-20, group 20-20, group 20-40. 
The expected order of proportional gains, again from greatest to 
least, should have been as follows for fish on the four rations: 20-40, 
20-20, 40-20, 40-40. The absolute and the proportional gains 
actually made by our fish fall in the expected order, with the ex- 
ception of the group fed 40 per cent protein in 1961 and 20 per 
cent protein in 1962. That group gained less than expected, both 
proportionally and absolutely, in 1962 ( Table 6 ) . 

Seemingly, the advantage of 40 per cent protein over 20 per cent 
protein in feeding fingerling fish, which increased their weight 20 
or more times during one growing season, is much less pronounced 
in feeding larger (subadult) catfish, which increased their weight 
less than threefold in one season. 

Feeding Different Amounts of Protein in Same Pond 

On August 16, 1962, each of five sections in the two ponds sub- 
divided by screens ( Figure 1 ) was stocked with 40 channel catfish. 
The fish stocked had been held together in another pond for several 
months previously. A 20, 30, 40, or 50 per cent protein-ration was 
fed to fish in one section of each pond; the "end" section was used 
as a "control" where fish received no supplemental feed. Selection 
of the feed used in each section was by bHnd drawings. Eight 
ounces of feed were fed daily to each group of fish; therefore the 
feeding-rate, relative to fish-weight, declined during the course of 
the experiment (5 per cent of body- weight per day on August 16; 
1.8 to 2.3 per cent of body- weight on October 18-19). The experi- 
ment continued 64-65 days, until the ponds were drained on Oc- 
tober 18-19, 1962. The results are shown in Table 7 and Figure 5. 

Experimental feeding obviously increased production, because the 
fish that were fed gained 12 to 17 times as much as the unfed "con- 
trol" fish. An analysis of variance indicated that gains in Pond 10 
did not differ significantly from gains in Pond 11. Absolute gains 
made on each ration ( Table 7 ) correlated directly with the amount 
of protein in the rations, and differed significantly ( P == .95 ) : 

Source of variation Degrees of freedom Mean square F 

Among feeds 3 9.3992 14.352 * 

Between ponds 1 .6786 1.036 n.s. 

Remainder 3 .6549 



Production of Channel Catfish 



213 



Table 7. — Gains Made by Yearling Channel Catfish on Experimental 
Rations, in Subdivided Ponds in 1962. 



Feed used 
(% protein) 


Pond 
number 


Section 
number 


Pounds 
stocked 


Pounds 
harvested 


Pounds 
gained 


No feed 


10 
11 

10 
11 

10 
11 

10 
11 

10 

11 


end 
end 

D 
B 

B 
A 

A 

C 

C 
D 


9.49 
9.54 

9.65 
9.72 

9.81 
9.79 

9.77 
9.57 

9.68 
9.75 


9.69 
11.44 

21.65 
22.44 

25.13 
24.38 

25.38 
27.19 

26.69 
27.00 


0.20 




1.90 


20 


2.10 
11.91 




12.72 


30 


24.63 
15.32 




14.59 


40 


29.91 
15.61 




17.62 


50 


33.23 
17 01 




17.25 




34.26 



18 

16 

(/) 
QI4 - 

z 

3l2h 
O 

z 

~ 8 

2 

< 6 
o 

4 - 
2- 



NO 
FEED 



2 % 



PER CENT PROTEIN IN FEED 

3 % 4 % 5 % 



■AAA^U^U^ 



Fig. 5. Gains in Pounds by Channel Catfish in Subdivided Ponds, on Feeds 
Containing Different Amounts of Protein, August 16-October 18, 1962. 

Summary of EfiFects of Various Levels of Protein on Growth 

The cumulative results of experiments involving variable protein- 
levels cannot be summarized in terms of absolute gains made by 



214 



University of Kansas Publs., Mus. Nat. Hist. 



fish on each ration, because of other variables such as 1 ) differences 
in size of the open ponds and the sections of the subdivided ponds, 

2) differences in the number and size of fish stocked therein, and 

3) differences between years. But results can be summarized if 
gains by the catfish are expressed on a relative basis, using the fol- 
lowing procedure. The results are treated as independent sets, de- 
pending on the year of study and whether the experimental units 
were open ponds or sections of subdivided ponds. Within each 
set of comparable experiments, the weight gained on each of the 
four feeds is converted to a percentage of the total gain in weight 
by all fish on all rations in that set. For example, the gain of 522 
pounds per acre by fish that were fed a 20 per cent protein-feed in 
an open pond in 1961 represented 20.5 per cent of the total gain 
(2660 pounds) made by fish on all four feeds in open ponds in 
1961. Relative gains on each protein-level, calculated in this man- 
ner, are shown in Figure 6. 



40 




Fig. 6. Relative Gains Made by Channel Catfish on Feeds Containing DiflFerent 
Amounts of Protein (20 to 50 Per Cent). Height of Bar Indicates the Per- 
centage of Total Gain Made by Fish on All Rations, under Similar Experimental 
Conditions. Within Each Set of Four Bars, Bar 1 ( left ) = Open Ponds in 
1961; Bar 2 = Open Ponds in 1962; Bar 3 = Subdivisions of Pond 10 in 1962; 
Bar 4 = Subdivisions of Pond 11 in 1962. 



Gains made on the 20 per cent protein-feed were consistently 
less than those on feeds containing higher levels of protein, but 
were much greater than gains where no feed was used. Thus feeds 
containing 20 per cent protein will produce satisfactory yields of 
channel catfish in ponds. Gains on the 30 per cent ration were 
usually as good as or better than gains on 40 per cent and 50 per 
cent protein. 

Feeds containing high protein-levels cost more than those con- 
taining less protein. Our 30 per cent feed cost approximately 4.5 
cents per pound, whereas the 20 per cent feed cost about 3.5 cents 
per pound, for a saving of 22 per cent. In four comparable sets of 



Productiox of Channel Catfish 



215 



experiments, our fish-production on the 20 per cent protein-feed 
was 17 per cent to 29 per cent lower (average 23 per cent lower) 
than on the 30 per cent ration. Thus, the increased cost of the 
feed containing 30 per cent protein was nearly balanced by in- 
creased yields. Taking account of other factors such as labor-costs 
and the length of time required to raise the fish to the desired size 
(with attendant risk of mortality), we conclude that the 30 per 
cent protein-feed was the best of those that we used. 

EFFECT OF SUPPLEMENTAL FEEDING ON CONDITION 

Does supplemental feeding of channel catfish merely increase 
their rate of growth, or is the condition (or relative plumpness) of 
the catfish altered also? 

Length-weight relationships were calculated by the procedure 
of Swingle (1964) for 141 groups of catfish (864 individuals) from 
various reservoirs in Kansas (Davis, 1959:16-17), and for 338 groups 
of channel catfish (7,791 individuals) that had received supple- 
mental feeds in our experimental ponds. The length-weight curve 
was calculated using the following formula: 

where W is the weight in pounds, a is the Y-intercept, b is the slope 
and L is the length in inches. The correlation coeflBcient for each 
curve exceeded 0.99. For fish that received supplemental feed, the 
average weight at 0.2-inch intervals of length is given in Table 8. 



CO 
Q 

=) 
o 

Q. 



X 
LlI 



0.45 




0.40 


/ ' 


.35 


/ /■ 


.30 


/ / 


.25 


// 


.20 


FED FISt-TX^^X 


.15 


^//Mrt/ILD FISH . 


.10 


^^.^^^^^^^^^ 


.05 


^.^^---^^'^^^^^ 



5 6 7 8 

TOTAL LENGTH IN INCHES 



10 



II 



Fig. 7A. Length-weight Relationships of Channel Catfish That Have Been 
Fed Intensively in Ponds, and of Wild Channel Catfish from Impoundments 

in Kansas. 



216 



University of Kansas Publs., Mus. Nat. Hist. 



The length- weight curve of channel catfish from Oklahoma (Fin- 
nell and Jenkins, 1954) closely approximates that of catfish from 
reservoirs in Kansas (Figure 7). At any given length, catfish that 
were fed in our experimental ponds were slightly heavier than "wild" 
fish from either state. The slope of the length- weight curve (the 
exponent b, above) for catfish that received supplemental feeds, for 
catfish from reservoirs in Kansas, and for catfish from Oklahoma, 
was 3.099, 3.304, and 3.407, respectively. Thus the weight of chan- 
nel catfish increases by a power slightly greater than the cube of 
its length. Catfish that were fed most closely approached the 
theoretical cubic relationship that is often used in obtaining indices 
of plumpness of fish ( see below ) . 

In order to express the relative plumpness of a fish as a numerical 
value, the ratio of volume (weight) to surface (length) is calcu- 
lated. This value is known as the "condition index"; a fat fish will 



3.5 



3.0 



2.5 



(/) 2.0 

Q 

o 

1.5 



X 

o 

[D 1.0 



0.5 




'-WILD nsH 



9 II 13 15 17 

TOTAL LENGTH IN INCHES 



21 



Fig. 7B. Length-weight Relationships of Channel Catfish That Have Been 

Fed Intensively in Ponds and of Wild Channel Catfish from Impoundments in 

Kansas. Dots Indicate Values for Wild Channel Catfish from Oklahoma. 



Production of Channel Catfish 



217 



Table 8. — Length-weight Relationship of Channel Catfish That 
Received Supplemental Feeds. 



Length 


Weight 


Number per 


Length 


Weight 


(inches) 


(pounds) 


pound 


(inches) 


(pounds) 


3.0 


0.008 


125 


14.2 


1.006 


3.2 


0.010 


100 


14.4 


1.050 


3.4 


0.012 


83.3 


14.6 


1.096 


3.6 


0.014 


71.4 


14.8 


1.143 


3.8 


0.017 


58.8 


15.0 


1.192 


4.0 


0.020 


50 


15.2 


1.242 


4.2 


0.023 


43.3 


15.4 


1.293 


4.4 


0.027 


37.3 


15.6 


1.346 


4.6 


0.031 


32.2 


15 8 


1.400 


4.8 


0.035 


28.5 


16 


1.456 


5.0 


0.040 


25 


16.2 


1.513 


5.2 


0.045 


22.2 


16 4 


1.571 


5.4 


0.050 


20 


16.6 


1.631 


5.6 


0.056 


17.7 


16.8 


1.693 


5.8 


0.063 


15.8 


17.0 


1 . 756 


6.0 


0.070 


14.2 


17.2 


1.821 


6.2 


0.077 


12.9 


17.4 


1.887 


6.4 


0.085 


11.7 


17.6 


1.956 


6.6 


0.094 


10.6 


17.8 


2.025 


6.8 


0.103 


9.7 


18.0 


2.097 


7.0 


0.112 


8.9 


18.2 


2.170 


7.2 


0.123 


8.1 


18.4 


2.244 


7.4 


0.133 


7.5 


18.6 


2.321 


7.6 


0.145 


6.2 


18.8 


2 400 


7.8 


0.157 


6.04 


19.0 


2 479 


8.0 


0.170 


5.8 


19.2 


2.561 


8.2 


0.183 


5.4 


19.4 


2.644 


8.4 


0.198 


5.05 


19 6 


2.730 


8.6 


213 


4.7 


19.8 


2.817 


8.8 


0.228 


4.3 


20.0 


2 906 


9.0 


0.245 


4.08 


20.2 


2.997 


9.2 


0.262 


3 8 


20.4 


3.090 


9.4 


0.280 


3.5 


20.6 


3.185 


9.6 


0.299 


3.3 


20.8 


3.281 


9.8 


0.319 


3.1 


21.0 


3.380 


10.0 


0.339 


2.9 


21 2 


3.481 


10.2 


0.360 


2.7 


21.4 


3.584 


10.4 


0.383 


2.6 


21.6 


3.689 


10 6 


0.406 


2.4 


21.8 


3.795 


10.8 


0.^31 


2.3 


22.0 


3.904 


11.0 


0.456 


2.2 


22.2 


4.015 


11.2 


0.482 


2.07 


22 4 


4.129 


11.4 


0.509 


1.9 


22 6 


4 244 


11.6 


0.537 


1.8 


22.8 


4.361 


11.8 


0.567 


1.7 


23.0 


4.481 


12 


0.597 


1.6 


23 2 


4.603 


12.2 


0.628 


1.59 


23.4 


4.727 


12.4 


0.661 


1.5 


23.6 


4.853 


12.6 


694 


1.41 


23.8 


4. 982 


12.8 


0.729 


1.37 


24.0 


5.112 


13.0 


0.765 


1.3 


24.2 


5.246 


13.2 


0.802 


1.2 


24.4 


5 381 


13.4 


0.840 


1.18 


24.6 


5.519 


13.6 


0.880 


1.1 


24.8 


5.660 


13 8 


0.920 


1 08 


25.0 


5.802 


14.0 


962 


1 04 







218 University of Kansas Publs., Mus. Nat. Hist. 

have a higher condition index than a thin fish of the same species 
and length (Bennett, 1962:76). The weight of a fish varies with 
the cube of its length provided the shape and specific gravity re- 
main the same; any change in the shape or relative plumpness of a 
fish will cause a change in the value of C in the formula: 

W 10^ 
C = 

u 

(Carlander, 1950:8). In our study the value of "C" was calculated 
from total lengths (L) in inches and weights (W) in pounds. For 
convenience, the weight is raised to a power suflBcient to provide for 
two digits to the left of the decimal. 

The condition indices of fish that received experimental feeds in 
our ponds in 1961 and 1962 are shown in Figure 8; the rates of 
growth by the same fish were presented in Figure 3. Also shown 
in Figure 8 are average condition indices of wild channel catfish, 
from reservoirs in Kansas, at sizes identical to those of the fed fish 
in biweekly samples from our ponds. 

Throughout late spring and summer, the condition index of the 
fed fish was higher than that of wild fish of comparable size (the 
"fed" fish had been fed for some time before their condition index 
was calculated for the first time in May ) . But, the condition index 
of wild fish increases as their lengths increase, whereas no such 



May June July August September October 

Fig. 8. Values of Condition Index Among Fish Being Fed in Ponds (Solid 
Lines) Compared with Average Condition Index of Wild Channel Catfish of 
Similar Size (Dash-lines). Vertical Bars Indicate Range of Values for Indi- 
vidual Fish, on Each Date when Samples Were Obtained. Upper Part of 
Graph Represents Data Obtained from All Ponds in 1961, Lower Part, Data 

Obtained in 1962. 



Production of Channel Catfish 219 

trend existed among the fish being fed in our ponds ( Figures 3 and 
8). Therefore, the condition advantage of the fed fish diminished 
as they grew larger, and as their standing crops increased in the 
ponds. Additionally, the condition index of the fed fish actually 
decreased in late summer. When the ponds were drained in Oc- 
tober of 1961 and 1962, the condition indices of the fed fish approxi- 
mated those of wild fish of similar size (Figure 8). Samples of 
fish that were weighed biweekly in 1961 and 1962 indicated that 
the catfish lost weight in late summer and early autumn in most 
ponds (Figure 3). The concomitant decrease in condition index 
adds evidence that a decline in standing crops actually occurred 
(was not an artifact attributable solely to sampling-error). Pre- 
sumably, the fish ceased to grow and utilized fat for energy, caus- 
ing loss in their weight and reduction in their condition index. 

To obtain maximum production from ponds, the fish should be 
harvested prior to any loss of condition. The period in which 
weight-loss occurs may vary with the prevalent temperature and 
the standing crop of fish being held in a pond. In our ponds, such 
losses usually have occurred in late September or October. 

EFFECT OF DAILY FEEDING-RATE ON PRODUCTION 

In order to obtain the most eflScient utiHzation of supplemental 
feeds by channel catfish, a satisfactory feeding-rate, as well as a 
satisfactory kind of feed, must be determined. In ponds used for 
commercial production in Alabama, Swingle (1959:66) reported 
feeding at rates principally between two and five per cent of the 
weight of fish in the ponds; the fish were fed six days each week. 
He indicated that the maximum amount that could be fed without 
mortality due to oxygen-deficiency was 30 pounds per acre per day. 
Snow (1962:114) fed channel catfish fry at an initial rate of 5 per 
cent of body-weight per day, but the rate he used decreased to 
about one percent by the time his ponds were drained after 109- 
136 days of feeding. Snow judged a rate of 20 pounds per acre per 
day to be about as high a feeding-rate as could be used safely for 
a sustained period. 

In our experiments, 35 fingerling catfish were stocked on July 
5, 1963, in each section of the subdivided ponds, and were fed at 
various rates from July 8 to October 10 (Table 9). 

The feeding-rates cited are maximal rates, restored after bi- 
weekly samples of fish from each section permitted recalculation of 
the rates, based on gains made by the fish in each two-week interval. 
Average feeding rates would be less than those shown in Table 9. 



220 



University of Kansas Publs., Mus. Nat. Hist. 



Table 9. — Feeding-rates (Per Cent of Body Weight per Day) and 
potrndage of fish stocked, in sections of subdivided ponds, 1963. 



Feeding-rate 


Section 


Total poundage of fish stocked 


(% of body weight) 


Pond 10 


Pond 11 


None 


E 
E 

A 
D 

B 
C 

C 
B 

D 

A 


3.49 
3.48 
3.43 
3.37 
3.50 




2 


3.57 


3.5 


3.47 


5 


3.39 


7 


3.34 




3.44 





DAILY FEEDING RATE 


16 


NO 2% 3 1/2% 
FEED 


5 % 7 % 


14 


. 


. 


POUNDS 


■ 


fc 




isl^^ 












- 


H 8 


■ 
















- 


5 6 
< 

^ 4 


■ 


■ 








■;'rv:-'-x'!;!v'x:' 






■ 


2 








• -x-: 








'!•!■!•;•"•;•:•: -i-x ■:-:•" 






- 















Fig. 9. Gains in Pounds Made by Channel Catfish that Were Fed at Different 
Daily Rates in Subdivided Ponds, July 8 to October 10, 1963. Feed Used 

Contained 30 Per Cent Protein. 

The experiment was terminated after 95 days (ponds drained 
October 10-11, 1963). The gains made on each feeding-rate are 
presented in Figure 9. An analysis of variance based on large 
samples from each of the sections (including the "control" group) 
indicated no significant difference between ponds, but a highly 
significant difference among gains by the five groups of fish within 
each pond: 

Source of variation Degrees of freedom Mean square 

Between Ponds 1 0.00037 n.s. 

Among Feeding Rates 4 0.596 *" 

Interaction 4 0.033 n.s. 

Remainder 140 0.0226 



Production of Channel Catfish 221 

An analysis of variance performed on individual weights, from sec- 
tions where fish were fed (excluding "controls"), detected no sig- 
nificant differences among gains made on the four feeding-rates. 

Source of variation Degrees of freedom Mean square 

Among Rates 7 0.0264 n.s. 

Within Ponds 112 0.0276 

Thus, where supplemental feeds were used, the weights of fish were 
significantly higher than weights of fish that received no supple- 
mental feed. Although higher feeding-rates generally resulted in 
higher yields (Figure 9) these differences were non-significant 
(P5^.95). 

The eflBciency of different feeding-rates can be measured by cal- 
culating the conversion of feed to flesh. The conversion-factors 
(feed used divided by weight gained by fish) of the fish that were 
fed at different daily rates are given in Table 10. For each period 
in summer, less eflBcient conversions usually were associated with 
higher feeding rates. Conversion-values for the entire summer for 
each feeding-rate were as follows: 2 per cent — 1.25; 3.5 per cent — 
2.20; 5 per cent— 3.20; 7 per cent— 4.28. 

Different feeding-rates have been used in various experiments in 
open ponds; the conversion-factors and the productions obtained 
are presented in Table 11. 

As one might expect, the efficiency of conversion of supplemental 
feed to flesh decreases as the feeding-rate increases. Thus the lowest 
feeding rate ( 2 per cent ) resulted in the best conversion, while the 
highest rate (7 per cent) provided the least eflBcient conversion. 

The productions obtained are not directly correlated with the 
feeding-rates. Our results agree with findings by Swingle (1959: 
66), who reported ineflBcient conversions when feeding-rates higher 
than five per cent were used. Swingle reported also a decrease 
in grov/th-rate associated with a decrease in feeding-rates from five 
to one per cent. Similar results were obtained in our subdivided 
ponds (see Figure 9). However, in the open ponds discussed 
above, productions were highest where the feeding-rate was 3 
per cent. Productions obtained with feeding-rates of 7 per cent 
were generally lower than those obtained on rates of 2.5, 3, and 3-5 
per cent. The low production obtained with the seven per cent 
feeciing-rate (as well as the poor conversions) may be due to 
excessive feed that fouled the water. But, in four of the five ponds 
where fish were fed at the rate of seven per cent, diurnal oxygen- 
levels were consistently higher than 4 ppm throughout the summer, 
and poor growth did not seem to be caused by oxygen-depletion. 



222 



University of Kansas Publs., Mus. Nat. Hist. 



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Production of Channel Catfish 



223 



Table 11. — Production (Pou>fDS per Acre) and Conversion (Feed Used 
Divided by Weight Gained by Fish) Using Various Feeding-rates in Open 
Ponds. Expeu\iments Were Conducted in Different Years, and Different 
Age Groups are Involved. Feeding Rates Listed are Initial Rates That 
Were Adjusted Periodically, and Are Thus the Maximal Feeding Rate 
During the Period Rather Than the Average Rate. 



2.5% 


3% 


3-5%t 


7% 


Produc- 
tion 


Conver- 
sion 


Produc- 
tion 


Conver- 
sion 


Produc- 
tion 


Conver- 
sion 


Produc- 
tion 


Conver- 
sion 


691 


1.94 


1835 


2.12 


1054 


3.35 


522 


6.50 


755 


2.00 


1936 


2.07 


1217 


3.61 


596 


6.79 


721 


2.10 


1224 


3. 74 J 


1303 


3.52 


610 


5.28 


1359 


1.54 


947 


4.6U 


1504 


3.17 


675 
751 


6.67 
6.67 

















} 5%, May 21-August 27; 4%, August 28-September 10; 3%, September 11-October 
8-11. 

t Subadult catfish; all other groups were fingerlings. 

Whatever the reason, the poor conversion-values and poor produc- 
tion obtained with the seven per cent rate indicate that feeding at 
tliis level is uneconomical. The productions that we obtained on 
the 3 per cent feeding-rate demonstrate that this rate is sufficient 
to produce crops that approach the carrying capacity of our ponds. 

The subdivided ponds were each stocked with 1620 fish per acre 
in 1964. Both ponds were fed at the rate of 3 per cent of fish-weight 
per day from May 11 to August 24, 1964. On the latter date the 
estimated poundages of fish in the two ponds were nearly identical 
(1201.0 Ibs./A in Pond 10; 1208.2 Ibs./A in Pond 11). For the re- 
mainder of the season (August 25 to October 4) the feeding-rate 
in Pond 11 was reduced from 3 per cent to 2 per cent. The feeding- 
rate in Pond 10 remained at 3 per cent. The yields obtained on 
October 8-9 in Ponds 10 and 11 were 1316.9 and 1461.2 Ibs./A, 
respectively (see Table 15). In this instance, a late-season reduc- 
tion in the daily feeding rate proved advantageous. 

Limitation of the feeding-rate to an amount less than 30 lbs. /A/ 
day (pounds per acre per day) as suggested by Swingle (1959:66) 
does not seem necessary. In one pond in which a production of 
1936 pounds per acre was obtained (Table 11), our feeding rate 
of three per cent sometimes amounted to more than 45 lbs. /A/day, 
and did not create unfavorable conditions for growth by fish in the 

3—3428 



224 University of Kansas Publs., Mus. Nat. Hist. 

pond. Use of large amounts of feed requires a full knowledge of 
the number and weight of fish in the pond, and of prevalent water- 
conditions, but may result in increased production. 

Experiments in subdivided and open ponds indicate that feeding- 
rates of two to three per cent of fish-weight per day probably are 
the most favorable rates for economical production of channel 
catfish. 

EFFECT OF STOCKING-RATE ON PRODUCTION 

Various studies (Swingle, 1957, 1959; Hickling, 1962:237-245; 
Tiemeier, 1957; and our own results) indicate that the number of 
fish stocked influences production. We stocked fingerling catfish 
(average length 3.1 inches, average weight 0.009 lbs.) in 1960 at 
rates of 1000 per acre (Pond 3) and 5000 per acre (Ponds 2 and 5). 
Their growth in early and late summer is presented in Table 12. 
The stocking-rate of 5000 per acre in Pond 2 was more efficient 
than the rate of 1000 per acre until midsummer, when fish stocked at 
the rate of 5000 per acre were almost as large as those stocked at 
the lower rate; thereafter, growth by fish in Pond 3 was more rapid, 
resulting in fish much larger than those in the other ponds. Never- 
theless, the highest yields were obtained with the higher rate of 
stocking (Ponds 2 and 5). 

Two ponds that were stocked in 1963 with 6500 fingerlings per 
acre, the highest stocking-rate that we have used, yielded the 
highest poundages that we have obtained in one growing season 
( 1922 Ibs./A, av. wt. = 0.33 lbs.; and 2023 Ibs./A, av. wt. = 0.33 

lbs.). 

These results parallel findings reported by Swingle (1959:68-70) 
in that an increase of poundage and a decrease in average size 
usually are associated with increased stocking-rates. But, yields 
varied considerably in adjacent ponds that were stocked at the same 
rate in the same year (Table 12). An examination of results re- 
ported by Swingle (1959), shows that one pond stocked with 3000 
catfish per acre yielded more pounds than were obtained with a 
stocking-rate of 5000 per acre, and another pond stocked with 3000 
catfish per acre yielded fewer pounds than a pond where 1032 cat- 
fish per acre were stocked. 

The variabiHty in yields on the same stocking-rate is evident in re- 
sults that we obtained with stocking rates of 2000 to 2250 per acre 
in different years (Table 13). Our formulated feeds (Table 5) 
were used in 1961 and 1962. Purina Pondfish Feed was used in 



Production of Channel Catfish 



225 



Table 12. — Growth by Catfish Stocked at Different Rates in Open 

Ponds in 1960. 





Stocking 
rate/acre 


July 29 


November 7-9 


Pond 


Average 

length 

(inches) 


Average 

weight 

(pounds) 


Average 

length 

(inches) 


Average 

weight 

(pounds) 


3 
2 
5 


1000 
5000 
5000 


7.8 
7.3 
5.5 


0.17 
0.14 
0.07 


11.8 
9.7 
7.9 


0.54 
0.24 
0.15 



1964. A feeding rate of 7 per cent was used in 1961, 3-5 per cent 
in 1962, and 2J2 per cent in 1964. 

Approximately equal yields of channel catfish were obtained in 
the three ponds stocked with 2000 channel catfish per acre in 1964 
(Table 13). Two of these ponds also contained bass ( largemouth ) 
and bluegill. When drained, these two ponds yielded the follow- 
ing poundage of each species: channel catfish, 726 and 755 lbs. /A; 
bass, 40.6 and 43.8; bluegill, 66.9 and 63.1; the total poundages of 
all species were 833.1 and 861.9 Ibs./A. Additionally, several hun- 
dred young-of-year bluegill, 0.8-3.0 inches long, occurred in both 

Table 13. — Yields of Channel Catfish in One GRO^vING Season, at 
Similar Stocking-rates ( Fingerlings ) . 



Year 


Number 
stocked 
per acre 


Feed 

used 

(% protein) 


Average 

weight 

(pounds) 


Yields 
(lbs. A) 


1961 


2250 
2250 
22.50 
2250 
2250 


20 
30 
50 
30 
40 


0.25 
0.26 
0.28 
0.31 
0.33 


550 




622 
624 
703 

783 


1962 


2250 
2250 
2250 
2250 


20 
50 
40 
30 


0.58 
66 
70 
81 


1103 




1245 
1314 
1515 


1964 


2000 1 
2000 1 
2000 


32 
32 
32 


38 
0.39 
0.40 


726 




755 
789 



f 350 Bass and 690 bluegill also present per acre. 



226 University of Kansas Publs., Mus. Nat. Hist. 

ponds but were not enumerated nor weighed. The feed-conversion 
values by catfish in these two ponds were 1.94 and 2.11 as com- 
pared to 2.00 in the pond containing 2000 catfish alone. Another 
pond that was stocked with 3370 fingerling catfish yielded 1399 
Ibs./A in 1964; the feed-conversion value was 1.60, As stated above, 
large differences in production were associated with the number of 
catfish in the ponds (Tables 12 and 13); but the presence or 
absence of bass and bluegill scarcely affected the production of cat- 
fish. 

Although the stocking-rate importantly affects the yield that can 
be obtained from stocks of fingerling catfish at the end of one grow- 
ing season, other factors influence yields to an even greater extent 
than does the stocking-rate (see subsequent discussion of carrying 
capacity and physical-chemical factors in our ponds). Because of 
wide variation in the yields obtained with similar stocking-rates, 
our data are inadequate to demonstrate the most advantageous 
rate. Some generaHzations seem possible, however: if the main 
objective is to obtain a maximum poundage of small fish, a stock- 
ing-rate of 5000 or more fingerlings per acre may be desirable. 
On the other hand, our results indicate that a stocking-rate of 2250 
per acre often gives yields almost as high as do higher stocking- 
rates; the 2250-rate permitted yields that approached the carrying 
capacity of our ponds. The results obtained in 1960 (Table 12), 
as well as data in more recent years, suggest that the most eflBcient 
method would be to stock at a high rate (5000-6500 per acre), and 
reduce the population by partial cropping when the fish are 6 to 
8 inches long (approx. 0.15 lbs.), and thereafter as they grow larger. 

EFFECT OF CROWDING ON GROWTH 

The tendency toward decrease in size of fish associated with in- 
creased stocking-rates may be due to the effect of physical crowd- 
ing, or to some minimal space requirement of channel catfish that 
must be met in order for normal growth to occur. Studies in which 
fish are grown in aquaria indicate that limited space is a deterrent 
to growth; but decrease in growth may be dependent on the amount 
and quality of water present, rather than the dimensions of the 
tank. Thus catfish might grow well in confined quarters if large 
amounts of water circulated through the tank. 

In 1962, 5, 20, or 50 fish were stocked in three sections (each 15 
inches wide, 5 feet long, 30 inches deep) of an experimental trough 
having a continuous exchange of water, and 5 and 20 fish were 
stocked in tanks (6 feet long, 4/2 feet wide, 27 inches deep) that 



Production of Channel Catfish 



227 



were continuously aerated but lacked exchange of water. All fish 
were fed a 30 per cent protein-ration at a daily rate of about 5 per 
cent of fish-weight. The number stocked in each unit and the 
average weight of the fish on various dates are shown in Table 14. 

Growth was better in the units having exchange of water rather 
than aeration. But, even with continuous exchange of water, the 
highest stocking-rate (50 per section) resulted in decreased growth 
by the fish. Thus a continuous influx of fresh water seemingly will 
not allow maximal growth by unlimited numbers of channel cat- 
fish ( 350,000 per acre in this instance ) . Growth by fish stocked at 
the rate of 20 per unit ( 140,000 per acre ) was as good as that made 
by fish stocked at the rate of five per unit ( 35,(XX) per acre ) . How- 
ever, this growth was much less than that made by fish of com- 
parable size that were stocked in open ponds. 

Table 14. — Average Weight (in Grams) of Chantstel Catfish Confined 

IN Experimental Trough and Tanks, 1962. Numerals in Parentheses 

Indicate Number of Fish in Each of the Five Units. 





Trough section 


Tank number 


Date 


1 
(50) 


2 

(20) 


3 

(5) 


4 
(20) 


5 

(5) 


June 11 


23.8 
29.7 
30.7 
37.1 
44.2 
46.3 
52.4 


28.1 
31.4 
37.3 
41.7 
51.4 
55.0 
75.6 


22.9 
26.3 
38.2 
42.5 
49.5 
61.3 
74.1 


24.0 
26.3 
26.7 
34.5 

t 


22.1 


June 25 


24.5 


July 9 


29.3 


July 23 


30.7 


August 6 

Auarust 20 


33.3 

i 


September 3 











f Fish died August 3-4. 
t Fish died August 7. 

Various numbers of fish were stocked in sections of our sub- 
divided ponds (Ponds 10 and 11; Figure 1) in 1964, to ascertain 
the effects of different degrees of physical crowding of fish in 
earthen ponds. Each pond was stocked at a relatively low rate 
of 1620 catfish per acre. However, different sections of the ponds 
were stocked with 10, 20, 42, or 90 fish so that the degree of crowd- 
ing varied within each part of the pond. Water mixes freely 
throughout these ponds so that the fish in each section occupied 



228 



University of Kansas Publs., Mus. Nat. Hist. 



water of similar quality. Presumably, metabolites were exchanged 
tliroughout the pond and did not accumulate in sections containing 
more fish. Therefore the effect of crowding itself was tested, rather 
than the more complex effects that may arise when different stock- 
ing-rates are used in separate, open ponds. 

The growth-histories (expressed as average weights) of fish in 
each section in the summer of 1964 are presented in Figures 10 and 
11. The weights of fish when stocked and harvested are given in 
Table 15. When the two ponds were drained, the sizes of the catfish 
in each section were highly variable. An analysis of variance de- 
tected no significant difference ( P 5=^ .95 ) in growth associated with 
the various degrees of crowding, although catfish stocked at the 
two lower rates (10 and 20 per section) grew slightly larger than 
those stocked at the higher rates (42 and 90 per section). On the 
other hand, the total yield and total gain in weight were eight times 
as much in sections stocked with 90 fish as in sections stocked with 
10 fish. An analysis of variance indicated that growth in Pond 11 
was significantly higher ( P =: .99 ) than that in Pond 10. Gains 
were consistently higher and conversion of feed to flesh was con- 
sistently better for each stocking-rate in Pond 11 than in Pond 10 
(Table 15), but the trends in growth under different degrees of 
crowding were the same in both ponds (Figures 10 and 11). 



Sotirce of variation Degrees of freedom Mean square 

Between Ponds 1 3.629 

Among Different 

Numbers Stocked 3 0.530 

Interaction 3 0.0256 

Error 55 0.2108 



F 
17.2** 

2.51 n.s. 
0.12 n.s. 




August September 



Fig. 10. Patterns of Growth by Channel Catfish in Subdivided Pond 10 in 
1964, through 21 September. Within the Pond, Enclosures of Equal Size 
Contained 10, 20, 42, or 90 Fish. Fish Were Fed (30 Per Cent Protein) at 
Equivalent Daily Rates (Approximately 3 Per Cent of Weight of Fish in 

Each Enclosure ) . 



Production of Channel Catfish 



229 



§0.90 
3 080 
°'- 0.70 




August 



September 



Fig. 11. Patterns of Growth by Channel Catfish in Subdivided Pond 11 in 
1964, through 21 September. Within the Pond, Enclosures of Equal Size 
Contained 10, 20, 42, or 90 Fish. Fish Were Fed (30 Per Cent Protein) at 
Equivalent Daily Rates (Approximately 3 Per Cent of Weight of Fish in 

Each Enclosure). 

Table 15. — Growth by Sxjbadult Channel Catfish in Subdivided Ponds 
IN 1964. Term of Experiment 147 Days; Ponds Dr.ajned 7 October. 





Pond 
number 


Number of fish stocked 
in each section 


Totals 

or 
average 




10 


20 


42 


90 


162 


Number of fish 
recovered 


10 
11 


10 
10 


20 
20 


42 
42 


90 

89 


162 
161 


Total weight of 
fish stocked 


10 
11 


0.72 
0.77 


1.-48 
1.59 


3.11 
3.21 


6.94 

7.15 


12.25 
12.72 


Average weight of 
fish stocked 


10 
11 


0.072 
0.077 


0.074 
0.079 


074 
076 


0.077 
0.079 


0.076 
0.079 


Total weight of 
fish recovered 


10 
11 


8.75 
10.00 


17.69 
19.31 


33.56 
36.81 


71.69 
80.00 


131.69 
146.12 


Average weight of 
fish recovered 


10 
11 


0.88 
1 00 


0.88 
0.97 


0.80 
0.88 


0.80 
0.90 


0.81 
0.91 


Gain in weight 
(production) 


10 
11 


8.03 
9.23 


16.21 
17.72 


30.45 
33.60 


64.75 

72.85 


119.44 
133.40 


Pounds of feed 
used 


10 
11 


18.97 
17.97 


39.58 
34.26 


76.86 
65.85 


169.66 
141.52 


305 07 
259.60 


Conversion 

feed /gain (lbs.) 


10 
11 


2.36 
1.95 


2.44 
1.93 


2.52 
1.96 


2.62 
1.94 


2.55 
1.95 



Observations of channel catfish in our ponds indicate that they 
are gregarious. Also, circular depressions visible in the bottom of 
our ponds when drained, and samples taken with seines, indicate 



230 University of Kansas Publs., Mus, Nat. Hist. 

that catfish aggregate in some parts of the ponds. We found es- 
sentially no evidence of agonistic behavior, even under crowded 
conditions in holding tanks. 

The relatively minor effect of physical crowding on the growth 
of catfish might be expected, considermg this gregarious behavior, 
which may provide new management techniques. Fish might be 
held in a confined area within a large body of water, simphfying 
their harvest and, perhaps, enabling more eflBcient consumption of 
feed with the advantage of a low stocking-rate in relation to the 
area of the entire body of water. 

CARRYING CAPACITY OF PONDS FOR CHANNEL 

CATFISH 

Little is known about the carrying capacity of waters for different 
species of fish. European fish-farmers have recognized that there 
are production-limits of ponds for carp (Bennett, 1962:61). Com- 
mercial fish-farmers commonly stock ponds with only enough fish 
to produce a marketable product at the end of one or two growing 
seasons. Boccius recognized in 1841 that water as well as land is 
limited in the quantity it can produce (Bennett, 1962:61). 

Swingle and Smith reported in 1938 that ponds stocked with 6500 
bluegill-fry yielded approximately 300 Ibs./A after one year (Ben- 
nett, 1962:61-62). A second year provided no additional growth. 
In another series of ponds 1300, 3200 and 6500 bluegill-fry yielded 
fish averaging 4 ounces, 2 ounces and 1 ounce, respectively. The 
total weight produced was approximately 300 Ibs./A in each pond. 

One pond stocked with bluegill-fry at the rate of 26,000 per acre 
(2 lbs. 5 ounces), and another stocked with year-old finger Hngs at 
the rate of 13,000 per acre ( 180 lbs. ) yielded 105 and 92 pounds 
per acre respectively after one growing season ( Swingle and Smith, 
1939). Thus the fish in one pond gained approximately 100 pounds 
per acre, while those in the other lost about 90 Ibs./A due to over- 
stocking. 

Results like those reported by Swingle and Smith were obtained 
in our ponds containing channel catfish. The maximum poundage 
of catfish that our ponds will support naturally is less than 200 
Ibs./A (Table 1). The variation in total number and weight of 
fish stocked had little effect upon yields. Total weights of popula- 
tions apparently fluctuate about a rather uniform level that repre- 
sents the carrying capacity of that particular environment. 

The carrying capacity of a body of water is dependent mainly 
on its surface area rather than its depth or volume, and is probably 



Production of Channel Catfish 231 

related to the amount of food produced by the body of water 
(Bennett, 1962:61). The area and depth of the zone of Hght 
penetration, where the bulk of the food-supply is produced for 
fishes, is most important. Carlander (1955:550) found no signifi- 
cant correlation between standing crop (per acre) and area of 
several lakes and reservoirs. In contrast, the productivity of a body 
of water apparently is inversely correlated with its depth. Car- 
lander reported a significant (P =: .95) negative regression of stand- 
ing crops of fish on the maximum depths of several warm-water 
lakes. 

Theoretically, a pond containing fishes of many kinds and sizes 
affords the most eflBcient utilization of available food. Carlander 
(1955:553-558) reported an increase in standing crop with increase 
in the number of species present. The total poundage of a given 
species may be affected by the presence or absence of other species 
of fish. But in our experiments in which channel catfish were 
stocked along with bass and bluegill, and did not receive supple- 
mental feeds, the carrying capacity of the ponds for channel cat- 
fish was not affected. Growth in these ponds was comparable to 
that made by catfish stocked alone in other ponds; yields were 110 
to 170 pounds of catfish per acre without supplemental feeding 
(Table 1). Similarly, the presence of bass and bluegill in ponds 
where catfish were fed did not alter the yields of catfish appreciably 
(Table 13). 

Many factors other than the number of fish present affect the 
carrying capacity of a pond: age of the pond and changes asso- 
ciated with its aging; changes in conditions of the watershed; sea- 
sonal changes; and artificial effects of man — all of which alter the 
fertility of a pond. 

The addition of supplemental feed to a pond containing channel 
catfish artificially increases the poundage of fish that a pond can 
support by introducing into the system nutrients that are produced 
from an outside source. Supplemental feeding allows for standing 
crops several times greater than those produced without feeding 
(Tables 12 and 13). Obviously when high poundages are present 
in a pond the fish must be dependent primarily on the feed added 
to the pond. 

In our early experiments, approximately 1200 Ibs./A was the 
maximum yield that we could obtain vdth supplemental feeding. 
Swingle (1957:161) reported yields up to 1242 lbs. of channel cat- 
fish per acre in some Alabama ponds, and in later experiments 
(Swingle, 1959:69) he reported productions as high as 2347 pounds 



232 University of Kansas Publs., Mus. Nat. Hist, 

per acre. Persons who raise channel catfish commercially in Kan- 
sas have told us that annual productions in their ponds sometimes 
exceed 3000 pounds per acre. 

The carrying capacity apparently was reached in two of our 
ponds containing subadult catfish in 1961. All of the channel cat- 
fish in both ponds had been held overwinter in Pond 1. On June 
13, 1961, half of the fish were transferred to Pond 8 resulting in a 
stocking rate of 2160 per acre in each pond. Both ponds received 
a 40 per cent protein feed at a rate of 4-6 per cent of body weight. 
(The rate was adjusted to 6 per cent at two-week intervals, and 
was never less than 4 per cent at the end of any two- week period. ) 

On July 24, 1961, the average lengths and weights of catfish in 
Ponds 1 and 8 were 11.5 inches and 0.50 pounds, and 11.8 inches 
and 0.57 pounds, respectively; therefore growth from June 13 to 
July 24 was almost aHke in both ponds. The same amounts of feed 
were used in both ponds until July 24. Growth proceeded rapidly 
in early summer, from standing crops of about 500 pounds per acre 
to 1100 or 1200 pounds per acre. However, from July 10 to August 
8, growth apparently ceased despite continued feeding and rela- 
tively high oxygen-content (mean oxygen- values in afternoon were 
above 7.2 ppm in Pond 1, and 6.5 ppm in Pond 8 in this period). 
The average temperature between July 10 and August 8 was 84° F. 

In an attempt to induce further growth by the fish two metliods 
were employed: (1) reduction of the number of fish in Pond 8, 
and (2) installation of a mechanical aerator in Pond 1 without re- 
moving any fish. 

On August 8, half the fish were removed from Pond 8, decreasing 
the stocking-rate to 1080 per acre. The remaining fish resumed 
growth, and by October 4 almost as many pounds per acre (1100) 
existed in the pond as were present before thinning. This pond 
was drained on November 1 and yielded 906 pounds per acre. 
Thus cropping of the fish in July resulted in a large increase in the 
total harvest obtained from the pond. 

In Pond 1 a mechanical aerator (of a type manufactured by 
Hinde Engineering Co., Highland Park, Illinois) was installed on 
August 1. From August 8 to November 1 the aerator was operated 
approximately 12 hours each night. The growth-rate of the fish 
increased sharply until October 4, when an estimated 2300 pounds 
per acre existed in the pond — almost twice the previous "carrying 
capacity" of 1200 pounds per acre. Pond 1 was drained November 
1, 1961, and yielded 2215 pounds per acre. 

The high production enabled by aeration of Pond 1 seemingly 



Production of Channel Catfish 233 

rules out any "space factor" as a cause for the limited production 
of 1200 pounds per acre obtained previously in these ponds. Re- 
moval of half the fish in Pond 8 and aeration of Pond 1 may have 
had a similar efiFect in reducing the concentration of volatile wastes 
that (earlier) limited production to about 1200 pounds per acre. 

Subsequent experiments utilizing aerators fail to substantiate this 
hypothesis. Two ponds (2 and 4) were stocked with 840 two-year- 
old channel catfish per acre, and two ponds ( 3 and 5 ) were stocked 
with 6500 fingerlings per acre in 1963 (Table 16). Ponds 2 and 3 
were aerated continuously from June 4 until the date of draining 
in fall by means of aerators similar to that used in 1961. Some fish 
were removed in summer for internal examination, mainly for fat- 
tiness of Uvers; thus, the stocking-rates declined gradually (Table 
16 ) . The fish in all ponds were fed a ration containing 30 per cent 
protein, at a daily rate of 2.5 per cent of fish-weight. 

Aeration may have been beneficial in ponds containing two-year- 
old catfish. When drained on September 27, the aerated pond 
(Pond 2) yielded 267 lbs. /A more than the pond that was not 
aerated ( Pond 4 ) . The fish in Pond 2 averaged 0.47 pound heavier, 
and usually had a slightly higher condition index, than fish in 
Pond 4. 

Aeration seemingly had little efiFect on growth by fingerling cat- 
fish in 1963. Fish in the aerated and non-aerated ponds had the 
same average weight (0.33 pound) when those ponds were drained 
on October 3 and 4; however. Pond 5 (not aerated) yielded 97 
Ibs./A more tiian Pond 3 because of lesser mortality. 

In the ponds that were stocked with fingerlings, the standing 
crops increased steadily throughout summer, and were higher at 
the time of draining than were standing crops in ponds containing 
two-year-old catfish. No increase in standing crops of the latter 
ponds was evident after mid-July; thereafter, gains made by indi- 
vidual catfish approximately equaled the weight of fish that were 
removed periodically. Because of the large size of the two-year-old 
fish, the summer-harvest of them greatly exceeded, in weight, the 
harvest from ponds stocked with fingerlings. The total yield 
(poundage of fish removed in summer plus that recovered when 
ponds were drained) was of the same general magnitude in all four 
ponds. All four had standing crops, on one or more dates, that 
exceeded the 1200-pound-level which seemed to be the approxi- 
mate "carrying capacity" of our ponds in previous experiments. 

Artificial aeration did not account in any major way for the high 
productivity of our ponds in 1963. Chemical data obtained from 



234 



University of Kansas Publs., Mus. Nat. Hist. 



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Production of Channel Catfish 235 

these ponds indicate that aeration was not needed, and that the 
aerators used had scant effect on water-quaHty. ( For example, the 
amount of dissolved oxygen in afternoon remained high in all ponds 
tliroughout the summer of 1963, and diurnal variations in dissolved- 
oxygen w^ere great in aerated as well as in non-aerated ponds; see 
Figure 13.) Aeration did prevent stratification in Ponds 2 and 3; 
stratification occurred daily in Ponds 4 and 5 but was interrupted 
by nocturnal mixing. 

Aeration is beneficial only when some factors, presumably chemi- 
cal, limit the poundage of fish that a pond will support. When 
dissolved oxygen remains low, or growth of fish is retarded by 
substances that may be oxidized, aeration can increase the carrying 
capacity of a pond. Nocturnal aeration, such as employed in 1961, 
seems most important. 

Once the carrying capacity of a pond has been reached, con- 
tinued feeding (without cropping or aeration) is ineflBcient, waste- 
ful, and potentially harmful by pollution of the water. This prin- 
ciple applies generally, although the carrying capacity varies from 
time to time as well as from pond to pond. 

Reduction of fish-growth as the carrying capacity is reached is 
not due to hmited "growth-ability" of channel catfish during one 
growing season. Conditions existing within a pond collectively 
limit the poundage of catfish that the pond will support. Manipu- 
lation of these conditions (as by cropping or aeration) may result 
in raising, or maintaining at a high level, the carrying capacity of 
a pond. 

PHYSICAL-CHEMICAL CONDITIONS IN 
EXPERIMENTAL PONDS 

The variability of productions of channel catfish in different ponds 
indicates that the nature of the ponds themselves influences pro- 
duction. Although our ponds are structurally alike and have a 
common water-source, the physical-chemical properties of the ponds 
vary temporally as well as from pond to pond. From 1961 through 
1964, records were obtained of temperature, dissolved oxygen, alka- 
linity, pH and dissolved solids by the methods described below. 

Methods 

Thermistors (Whitney, or Jarrel-Ash Model 26-601) were used 
to measure temperatures at different depths in the ponds. Mea- 
surements usually were taken three times each week near the sur- 
face, mid-depth (0.7 m.), and bottom of the ponds (approximately 



236 University of Kansas Publs., Mus. Nat. Hist. 

1.4 m.). Early-morning and mid-afternoon readings were taken 
in 1962 and 1963, to approximate the maximum and minimum 
temperatures of the ponds. In 1963, readings were taken once each 
week at three-hour intervals for a 24-hour period. A recording 
thermometer operated continuously in one of the ponds. 

The dissolved oxygen-content of the ponds was determined by 
titration using the Winkler iMethod (Faber, 1955:250-255) in 1961. 
Subsequently the amount of dissolved oxygen was measured directly 
in the ponds by means of an oxygen analyzer (Jarrel-Ash Model 
26-601). Readings were taken in per cent of saturation, and con- 
verted to ppm (Stand. Meth., Faber, 1955:254). Corrections were 
made for altitude. Values obtained by concurrent use of both 
methods (analyzer and titration) corresponded closely. All read- 
ings above 120 per cent of saturation were evaluated by titration. 
Oxygen-determinations were made at approximately the same inter- 
vals and depths as were the temperature-measurements discussed 
above, except in 1961 when oxygen-readings were taken only near 
the surface. 

A Beckman Model N meter was used for pH-determinations; 
readings were taken of the surface-water at intervals discussed 
under temperature. 

Phenolphthalein alkalinity and methyl-orange alkahnity (Stand. 
Meth., Faber, 1955:35-38) were ascertained by titration of water- 
samples taken approximately one foot beneath the surface of the 
ponds. Titrations were performed in the early morning in 1962, 
1963, and 1964; in 1963, titrations were made also in mid-afternoon, 
as well as at three-hour intervals over a 24-hour period, one day a 
week. 

Measurements of the total dissolved solids in the ponds were 
taken weekly by means of a portable conductivity bridge (Indus- 
trial Instruments, Model RC 7P). Specific conductance indicates 
the total concentration of the ionized constituents of waters. It is 
clearly related to the sum of the anions, or cations, determined 
chemically and usually correlates closely with the dissolved residue 
(Faber, 1955:89-92). The electrical conductivity was corrected to 
25° C. and multipHed by a constant (0.65) to obtain the dissolved 
sohds in parts per million (Rainwater and Thatcher, 1960:270). 

Temperature 

The average afternoon temperatures at the surface of our ponds 
in 1961, 1962, and 1963 are presented in Figure 12A. Temperatures 
remained above 25° C. (77° F.) during the greater part of our 



Production of Channel Catfish 237 

experiments ( June to mid-September ) . Subsequently, temperatures 
dropped considerably (to approximately 15° C. by late October 
1961). The least variation in temperature among the three years 
was in August; temperatures in early and late summer varied con- 
siderably in the different years. 







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JUNE JULY AUGUST SEPTEMBER 

Fig. 12. (A) Surface-temperatures in Experimental Ponds in 1961 (Dash- 
line), 1962 (Dotted Line), and 1963 (Solid Line). Lines Join Average Values 
Within Biweekly Intervals in which Data on Growth by Catfish Were Obtained. 
(B) Surface-temperatures (Dash-line) and Bottom-temperatures (Dotted Line) 
in Midafternoon, and Temperatures at both Surface and Bottom (SoUd Line) 
in Early Morning, in Experimental Ponds in 1962. Ponds Stratified Daily, but 
Mixing Overnight Restored Thermal Uniformity. Lines Join Average Tempera- 
tures Within Biweekly Periods in which Data on Growth by Catfish Were 

Obtained. 

The surface-temperatures of our ponds in afternoon remained 
relatively stable during most of our experiments (between 25° C. 
and 30° C. in 1962); however, temperatures varied several degrees 
diumally. The average surface- and bottom-temperatures of all 
ponds in early morning and mid-afternoon in 1962 are presented in 
Figure 12B. Although our ponds are relatively small and shallow, 
their water usually stratified each afternoon and mixed each night. 
Morning-temperatures were essentially the same from the surface 
to the bottom of the ponds. Water near the bottom retained approxi- 
mately the same temperature throughout the day, but the tempera- 



238 University of Kansas Publs., Mus. Nat. Hist. 

ture of the water near the surface increased, resulting in stratification 
and a definite thermocline in the shallow zone. The degree of 
stratification decreased in late summer and essentially no stratifica- 
tion occurred at the end of September in 1962. Although fish in 
our ponds could have remained in water having nearly the same 
temperature throughout any 24-hour cycle, we doubt that they did 
so, because of daily oxygen-depletion in the deepest, coolest zone. 

When fish are acclimated at various temperatures, metabolic 
rates do not increase as much as twofold with each 10° C. increase 
in temperature, and increases in metabolism diminish as the upper 
lethal temperature is reached (Moss and Scott, 1961:390). Moss 
and Scott reported that resting channel catfish showed no appre- 
ciable increase in their metabolic rate as water-temperatures in- 
creased from 25° C. to 35° C. The metabolic mechanisms of chan- 
nel catfish appear to be highly adaptive. If given sufficient oppor- 
tunity for temperature-acchmation they tend to operate at the same 
rate regardless of temperature, at least within the range of 25-35° C. 
(Moss and Scott, 1961:390). Assuming that these experimental re- 
sults are applicable to our ponds, the fluctuations in temperature 
during the summer may have had little effect on periodic growth 
by channel catfish in these ponds. 

In our ponds, growth by catfish in 1961 and 1962 ( Figure 3 ) de- 
creased in late-September of each year, when surface-temperatures 
were below 25° C. Swingle (1959:66) reported that channel cat- 
fish responded best to feeding when surface-waters were warmer 
than 70° F. (21° C), and recommended that supplemental feeding 
be stopped while water temperatures were below 60° F. ( 15.5° C. ). 
Feeding when water temperatures were below 20° C. in our ponds 
was inefficient; feeding at such temperatures should only be at a rate 
sufficient to maintain the fish in satisfactory condition. 

Dissolved Oxygen 

Dissolved oxygen-levels at the surface of our ponds remained 
relatively high throughout summer, the average for two-week- 
intervals being generally higher than six ppm (see Figure 13 for 
oxygen-levels in four ponds in 1963). Tiemeier and Moorman 
(1957:169-170) and Tiemeier and Elder (1957:381-383) reported 
similar oxygen-levels in several farm ponds in Kansas. Weather- 
conditions ( whether day was clear, rainy, windy, calm ) and density 
of plankton-blooms in the ponds influenced the daily variation in 
the oxygen-levels of our ponds. 



Production of Channel Catfish 



239 



The dissolved oxygen of the surface-waters varied a great deal 
diumally. The lowest levels, in early morning (approximately 6:30 
a.m.), usually were 4-6 ppm in June and July of 1963; subsequently, 
morning oxygen-levels were usually less than 4 ppm ( Figure 13 ) . 




JUNE 



JU LY 



AUGUST 



SEPT. 



Fig. 13. Dissolved Oxygen in Early Morning (a. m.) and Midaftemoon (p. m.) 
in Four Experimental Ponds in 1963. Lines Join Average Values Within Bi- 
w^eekly Periods in which Data on Growth by Catfish Were Obtained. Solid 
Line Joining Dots Represents Pond 2 (Aerated, Large Fish); Sohd Line Join- 
ing x-marks, Pond 4 (Not Aerated, Large Fish); Dash-hne, Pond 3 (Aerated, 
Small Fish); Dotted Line, Pond 5 ( Not Aerated, Small Fish). 

Stratification and mixing occurred daily. Oxygen-concentrations 
frequently were less than 2 ppm in the deepest part of the ponds 
during the afternoon. Respiration in the ponds, as well as mixing 
at night of the oxygen-depleted waters from the bottom with oxy- 
genated surface-waters, contributed to the low oxygen-levels pres- 
ent in the ponds in early morning. 

Resting fish of most species consume oxygen at rather constant 
rates over a wide range of ambient oxygen-tensions (Moss and 
Scott, 1961:392). Moss and Scott (1961:385-393) found no change 
in the oxygen-consumption of resting channel catfish that were 
exposed to oxygen-levels varying from approximately 8 ppm to less 
than 1 ppm over a 15-day period. Critical dissolved oxygen-levels 

4—3428 



240 University of Kansas Publs., Mus. Nat. Hist. 

were usually slightly less than 1 ppm but increased slightly with 
rising temperature. 

At low levels of dissolved oxygen, oxygen-consumption by fish 
becomes dependent upon the concentration of oxygen in the en- 
vironment. The oxygen-level at which this shift to oxygen-depen- 
dent metabolism occurs (the incipient lethal level) depends upon 
the degree of activity of the fish, which influences their oxygen- 
requirement. Fish entering this phase of metabolism v^ll even- 
tually die unless their activity is reduced or the oxygen-level is 
increased. 

The amount of oxygen necessary to sustain the normal activity 
that is associated with rapid growth is not known, but seems low 
in channel catfish. Perhaps the extent of daily fluctuation in dis- 
solved oxygen, as well as the minimal daily oxygen-level, impor- 
tantly influences the rate of growth. In two ponds in 1962, morning 
oxygen-levels were less than 2 ppm in the first half of August, but 
fish in those ponds grew satisfactorily (Figure 3). In 1963, morn- 
ing oxygen-levels in Pond 4 averaged less than 2 ppm in mid-Au- 
gust, while the average was 4 ppm in Pond 3 ( Figure 14 ) . Growth 
was rapid in both ponds at this time, and was better than the 
growth-rate in Ponds 2 and 5, which had oxygen-levels interme- 
diate to those in Ponds 3 and 4 during this period (Table 15). 
But, morning oxygen-levels in Pond 4 remained low for an extended 
interval, and seemingly aflFected growth adversely. Morning-read- 
ings in this pond were less than 3 ppm through most of the growing 
season. When Pond 4 was drained, its fish weighed approximately 
0.5 pound less than those in a comparable pond (Pond 2) in which 
oxygen-levels remained above 3 ppm. 

We think that low levels of dissolved oxygen inhibited production 
in an experimental pond where feed containing 50 per cent protein 
was used in 1961. The maximal dissolved-oxygen-level remained 
below 4 ppm in the pond through most of August and September. 
Although the minimal daily levels of dissolved oxygen in this pond 
were not known, subsequent morning and afternoon determinations 
suggest that the oxygen-levels at night were critically low. The 
fish probably entered an oxygen-dependent phase of metabolism, 
and their activity decreased correspondingly. During this period 
the catfish apparently stopped eating, because feed accumulated 
in the pond; and, decomposition of the unused feed undoubtedly 
contributed to the oxygen-deficit. Fish in this pond gave no evi- 
dence of stress by surfacing, but their growth essentially stopped. 



Production of Channel Catfish 241 

Observations of similar eflFects of low oxygen-levels on other species 
have been reported. For example, Shepard (1955:428) reported 
that speckled trout held at oxygen-levels slightly above the incipient 
lethal level were sluggish, fed poorly and were susceptible to dis- 
ease. 

The effects of mechanical aeration of ponds, on the production 
of channel catfish, were discussed under the heading "Carrying 
Capacity of Ponds." 

pH-Values 

Average pH-values of seven experimental ponds in 1963 are 
presented in Table 17. In early morning, pH-readings were usually 
slightly below 8, and pH-levels in afternoon were usually less than 9. 
Swingle (1961:72) reported that waters ranging in pH from 6.5 
to 9.0 before daybreak are generally the most desirable for pondfish 
production; waters in this range are capable of giving highest pro- 
ductions if the level of fertility is adequate. No seasonal trend in 
pH-values was evident in our ponds. The most obvious changes in 
the pH of our ponds occurred diurnally. Afternoon-values were 
consistently higher than morning-readings (Table 17), due to utili- 
zation of free CO, and HCO3 in photosynthesis in daytime. Thus 
the measure of the diurnal fluctuation of pH gives some indication 
of the basic productivity of a pond, although buffering action within 
ponds influences fluctuations in pH. Ponds that received supple- 
mental feeds (and consequently had high standing crops of fish) 
tended to show larger diurnal fluctuations in pH than ponds in 
which fish were not fed. The feed may have had both direct and 
indirect fertilizing effects that resulted in an increase in natural 
productivity in the ponds, reflected by increased diurnal fluctuations 
in pH. In ponds having comparable stocks of fish, aeration reduced 
the diurnal fluctuation of pH. 

Alkalinity 

Total alkalinity expresses concentrations of calcium and carbon 
dioxide, two substances directly necessary to plant-life; on the other 
hand, the alkalinity is itself affected by the entire biological and 
chemical systems of waters. Therefore total alkaUnity has been 
used as a rough index of the productivity of waters (Moyle, 1956: 
310). 

Water having a low total alkalinity usually has low concentrations 
of other salts including phosphorus compounds. Such water usually 
is infertile and lacks dense plankton-populations, according to Moyle. 



242 



University of Kansas Publs., Mus. Nat. Hist. 



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Production of Channel Catfish 



243 



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244 University of Kansas Fuels., Mus. Nat. Hist. 

He reported (1949:324) that ponds where total alkaHnity was less 
than 40 ppm did not yield as many pounds of fish as ponds with 
harder waters. He considered a total alkalinity of 40 ppm to be a 
useful natural separation-point between soft and hard waters. 
Carlander (1955:551-552) found a significant increase in standing 
crops per acre with increased alkalinity (up to 200 ppm) in trout- 
lakes, in warm-water lakes and in reservoirs. 

Phenolphthalein alkalinity was usually absent in our ponds, and 
when present occurred only in mid-afternoon, associated with high 
pH-values. Thus, bicarbonates essentially comprised the total alka- 
linity in our ponds. The average alkahnities for two-week periods 
in early morning, and the change by mid-afternoon, are presented 
in Table 18. Total alkalinity usually remained within the range of 
65-150 ppm, and seemingly was at levels conducive to good growth 
by fish. In most ponds the total alkalinity decreased in the course 
of the summer. This decrease was especially prominent in ponds 
that did not receive supplemental feeds. 

Diurnal variation in alkalinity usually was less than 10 ppm; 
morning-readings usually were slightly higher than those in mid- 
afternoon. 

Total Dissolved Solids 

The total dissolved solids remained within the range of 200-500 
ppm in all of our ponds (see Table 19 for levels that occurred in 
1963). In most instances, all ponds had similar levels of dissolved 
solids at a given time. The range of dissolved solids increased from 
200-300 ppm during June and July to levels of 300-450 ppm in 
August and September. The highest levels occurred in early Sep- 
tember, the last period in which measurements were taken. It is 
noteworthy that the seasonal trend in total dissolved solids was 
opposite to that in alkalinity — the former increasing as the latter 
decreased. 

CORRELATION OF PHYSICAL-CHEMICAL CONDITIONS 

WITH PRODUCTION 

At the start of each set of experiments in our ponds, many factors 
that influence production are subject to our control. The size and 
location of the ponds are similar, and the following factors can be 
constant: source of water and date of filling of ponds, kind, num- 
ber, size, and date of stocking of fish; kind of feed and daily feed- 
ing-rate. In practice one (or more) of the factors listed has been 



Production of Channel Catfish 



245 



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246 University of Kansas Publs., Mus. Nat, Hist. 

varied from pond to pond, for purposes of testing the efiFect of that 
factor on fish-production. 

As experiments progress in earthen ponds hke ours, additional 
variables arise. Some of these new variables may be correlated 
with the principal variable being tested. For example, if feeds 
having different protein levels are being tested, and one of the feeds 
is superior to others, differences in weight-gain by the fish, their 
average weight, and the standing crop follow automatically. These 
differences may induce others, especially in chemical quaHties of 
the water, and the latter differences may in turn affect subsequent 
growth by the catfish. 

An understanding of the interrelationships of these changes 
would have value in pond-management. If the physical-chemical 
conditions that are most favorable for growth of fish can be identi- 
fied, attempts to establish and maintain those conditions are pos- 
sible. Even if such attempts fail, knowledge of the relationship of 
fish-growth to water-quality would enable appropriate adjustments 
( of feeding-rates, for example ) in response to uncontrolled changes 
in water-quality. 

Correlation-coeflBcients (r) were calculated for variable factors 
in seven of our ponds in 1963. In that year, we obtained data on 
growth of fingerling catfish that were not fed (were reliant on food 
that occurred naturally in the ponds), and on fingerling and sub- 
adult catfish that were fed equivalent daily amounts of pelleted 
rations (Tables 16 and 20). 

In Table 21, correlation-coeflBcients are presented for several 
variables in ponds in which fish were not fed (Ponds 1, 6, and 9); 
and, separately, the corresponding correlation-coeflBcients are given 
for data from all ponds in which supplemental feed was used 
(Ponds 2, 3, 4, and 5). The data pertaining to fed fish are subdi- 
vided in Table 22, to permit separate analysis of the results in ponds 
that contained large numbers of small fish (Ponds 3 and 5), and in 
ponds stocked with small numbers of larger fish (Ponds 2 and 4). 

Each variable factor compared is listed in the left-hand column 
of each of the two tables (21 and 22), together with a numeral 
that identifies that same factor at the top of each table. "Average 
weight" is equivalent in effect to the current standing crop where 
equal numbers of fish were present, as in Ponds 2 and 4, and Ponds 
3 and 5 (Table 22). "Condition index" reflects variations in the 
"plumpness" of fish on the different dates when ponds were sam- 
pled, "a weight" represents the gain (or loss) from one sampling- 
date to the next (a 2- week interval). "Former standing crop" 



I 



Production of Channel Catfish 



247 



Table 20. — Growth by Fingerling Channel Catfish That Received No 

Supplemental Feed in 1963. (Ponds 1 and 9 Contained 2250 Catfish Per 

Acre; Pond 6 Contained 600 to 750 Catfish per Acre. ) 



Date 



April 2 

June 3 

June 17 

July 1 

July 15 

July 29 

August 12. . . . 

August 26 

September 9 . . 
October 16-17 



Pond 1 



Average 
weight 



0.013 
.050 
.064 
.073 
.066 
.068 
.086 
.075 
.074 
.074 



Total 
weight 



3.02 
11.34 
14.40 
16.38 
14.94 
15.39 
19.35 
16.92 
16.56 
16.25 



Pond 9 



Average 
weight 



0.013 
.038 
.046 
.050 
.057 
.054 
.058 



.059 
.052 



Total 
weight 



3.02 
8.64 
10.44 
10.89 
11,88 
12.15 
13.05 



13.20 
10.87 



Pond 6 



Average 
weight 



013 
.064 
.100 
.103 
.109 
.078 
.097 
.129 
.133 
.154 



Total 
weight 



1.01 
4.80 
7.50 
7.21 
7.35 
5.07 
6.44 
8.40 
7.98 
8.93 



represents the total poundage of fish on the last previous date of 
sampling. The "a. m." and "p. m." values of temperature and other 
factors approximate the minimal and maximal daily values, and 
are averages of values recorded on several days within the intervals 
between sampHng-dates for catfish. "A temperature" ( or A oxygen, 
A pH, A alkalinity) represents the average difference between morn- 
ing- and afternoon-values for that factor. 

The correlation-coeflBcients were calculated from absolute values 
of the data. Correlation-coeflScients were calculated using logarith- 
mic transformations also (log y = x, y = log x, and log y = log x), 
but such transformations yielded no information additional to that 
obtainable without transformation. 

Correlations with Average Weight 

In ponds where fish were not fed, diurnal change in the amount 
of dissolved oxygen was the only variable tested that was signifi- 
cantly correlated with the average weight of the fish (r = ,44); 
these same two factors were correlated ( r = .62 to .68 ) in ponds 
where fish were fed. In the latter ponds, average weights were 
negatively correlated (-.48 to -.66) with morning-oxygen, but posi- 



248 



University of Kansas Publs., Mus. Nat. Hist. 






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250 University of Kansas Publs., Mus. Nat. Hist. 

tively correlated (.24 to .51) with afternoon-oxygen. These rela- 
tionships probably indicate effects of (1) plankton-blooms that 
commonly develop where standing crops of fish are high, causing 
high afternoon-oxygen through photosynthesis, and (2) high 
respiration-demands by catfish and associated organisms, causing 
oxygen-depletion nocturnally. 

Temperatures and daily change in temperature correlated nega- 
tively (-.36 to -.76; -.54 to -.67) with size of fish, simply because 
the catfish were largest in late-season when temperatures were 
declining and when daily fluctuations in temperature were least. 

Alkalinities in morning and afternoon were significantly correlated 
( negatively, r = -.71 and -.79 ) with the average weights of catfish 
in ponds 3 and 5 but not in other ponds. Correlations of size with 
diurnal change in alkalinity were not significant in any ponds, and 
were especially low where fish were fed. 

Diurnal change in pH and morning-pH were correlated with the 
size of catfish receiving supplemental feeds (r= -f-.57 and +.63; 
-.42 and -.70 ) . The afternoon-pH was less significant, perhaps be- 
cause upward trends in pH in daytime were inhibited by buffering 
agents in the ponds. Correlations of pH with average weights were 
low for fish reliant on natural foods. The pH in early morning was 
lowest in ponds where fish were fed, due to indirect effects of respi- 
ration by the far greater poundages of fish in those ponds than in 
ponds 1, 6, and 9. Afternoon-pH was approximately the same in 
all ponds; thus, the diurnal change in pH was less in the latter group 
of ponds. 

Total dissolved solids were highly correlated with the average 
weights of catfish that were fed (r= .67 and .91), but not with 
weights of those subsisting on natural food (r= .19). The high 
poundages of fish in ponds 2, 3, 4, and 5, as well as the increasing 
amounts of supplemental feed that were used in those ponds, prob- 
ably contributed to the total dissolved soHds, explaining their 
increase as the average weight (and standing crop) of the fish 
increased. 

Correlations with Condition-index 

The condition index of catfish may indicate whether conditions 
are favorable or unfavorable for growth; catfish living in situations 
favorable for growth should be more plump (in better condition) 
than those living under conditions unfavorable for growth. 

Afternoon-temperatures and diurnal changes in temperature were 
significantly correlated ( r =: .66 and .60 ) with condition indices of 
channel catfish that were not fed. Oxygen-levels in afternoon and 



Production of Channel Catfish 251 

diurnal changes in oxygen content were negatively correlated (r = 
-.56) with the condition index of fish that were fed (Ponds 2, 3, 
4, 5). This eflFect was most pronounced in ponds that contained 
fingerling catfish. Alkalinity and diurnal changes in alkalinity were 
highly correlated (r = .75 and .78; r = .43) with the condition index 
of catfish that were not fed. Fish that were fed reflected positive 
but nonsignificant correlations of condition index with alkalinity in 
ponds having fish of comparable size ( 2 and 4; 3 and 5 ) . 

Diurnal changes in pH were significantly correlated (negatively, 
-.49) with condition indices among fed fish, but only when Ponds 
2, 3, 4, and 5 were considered together. As in the case of afternoon 
oxygen-levels, changes in pH were most important in those ponds 
containing fingerling catfish (3 and 5). 

Total dissolved solids were significantly correlated (negatively, 
r = -.63 ) with condition indices only in ponds where supplemental 
feeds were not used. 

Correlations with Changes in Weight 

Periodic changes in the weight of the fish were highly correlated 
(r=.62) with the diurnal change in alkalinity in ponds where 
fish were not fed. Correlations were low for these variables in the 
ponds where supplemental feed was used. Absolute values of 
morning and afternoon alkalinity had low but positive correlation 
(r=.30 and .23) with changes in weight among unfed fish. In 
ponds where fish were fed, these correlations were negative (r = 
-.32 to -.37). As mentioned before, alkalinity is often used as an 
indicator of the productivity of waters. In Ponds 1, 6, and 9, where 
only natural food was available to the catfish, their growth-rate 
declined as alkalinity declined. The opposite relationship existed in 
ponds where catfish were fed; large gains in weight occurred despite 
declining levels of alkahnity during summer. 

Diurnal changes in temperature were correlated (r — .51) with 
changes in weight of the subadult catfish. Among the other groups 
of fish, correlations with change in temperature were not significant; 
and, absolute temperatures were not significantly correlated vdth 
change in weight of fish in any ponds, within the term ( 3 months ) 
of our records. 

SUMMARY 

Production of channel catfish under various experimental condi- 
tions was carried on in Ho-acre earthen ponds and other facilities 
of the State Biological Survey at Lav^rence, mostly from 1960 
through 1964. 



252 University of Kansas Publs., Mus. Nat. Hist. 

1. The weight of channel catfish that our ponds supported with- 
out supplemental feeding was 98 to 183 pounds per acre. 

2. By daily feeding of pelleted rations, yields of 550 to 2215 
pounds of channel catfish per acre were obtained. 

3. In tests of synthetic diets, force-feeding of gelatin capsules to 
channel catfish was more successful than force-feeding by means of 
syringes; but, neither method enabled rapid gains by fish confined 
in concrete troughs. 

4. Ad lib feeding of synthetic diets to catfish in troughs indi- 
cated that the carbohydrate constituent as well as the percentage 
of protein in the diet influences weight-gain by these fish. In diets 
having low protein-content, growth was better on rations contain- 
ing starch than on rations containing dextrin, sucrose, or cellulose. 
DiflFerential effects of the carbohydrate constituent diminished as 
the protein content increased. 

5. In open ponds and subdivided ponds, supplemental feeds 
containing 30 per cent protein produced as many pounds of catfish 
as did feeds containing larger amounts of protein. Gains made on 
feeds containing 20 per cent protein were consistently less than 
those obtained on feeds containing higher levels of protein. 

6. Catfish that received supplemental feeds had a higher condi- 
tion index than did wild catfish. The difference in condition index 
of fed vs. wild catfish diminished as the fed fish grew larger and 
their standing crop increased in ponds. 

7. Experiments in subdivided ponds and in open ponds indicated 
that feeding-rates of two to three per cent of fish-weight per day 
probably are the most favorable rates for economic production of 
channel catfish. 

8. High stocking-rates (5,000 or more fingerlings per acre) usu- 
ally resulted in higher yields of fish than did lower stocking-rates 
(2,000-2,500 fingerlings per acre), at some sacrifice in the average 
size of individual fish. 

9. The carrying capacity of our ponds varied from year to year. 
Catfish ceased to grow, and lost weight, in late summer and early 
autumn in most ponds — when their standing crops were highest 
and the water-temperature began to decline. Weight-loss occurred 
in mid-summer in some ponds having high standing crops of fish. 

10. Stresses caused by critically low oxygen-levels (less than 3 
ppm) seemingly retarded growth by channel catfish, although they 
signaled their distress by surfacing only when oxygen-levels were 



Production of Channel Catfish 253 

less than 1 ppm. Fish affected by these conditions grew rapidly 
after partial cropping, or nocturnal aeration, of the ponds. 

11. The pH of our ponds was consistently between 7 and 9. 
Diurnal changes in pH often exceeded seasonal changes, and ex- 
ceeded differences in pH between ponds where fish were fed or 
not fed. 

12. The total alkalinity of our ponds varied from approximately 
50 ppm to approximately 150 ppm. Diurnal change in alkalinity 
was slight. The alkalinity of most ponds declined in late summer. 

13. Total dissolved soHds in our ponds varied from approxi- 
mately 220 ppm to approximately 450 ppm, and were highest in 
late summer. 

14. Where fish were not fed, their growth-rate and condition 
index were positively correlated with total alkalinity of the ponds, 
and with the extent of diurnal change in alkalinity. As the aver- 
age weight of the fish increased the diurnal fluctuation of dissolved 
oxygen increased. Generally, the growth-rate and condition index 
of the catfish declined (1) as the standing crop increased, (2) as 
the alkalinity decreased, and (3) as the oxygen-level became more 
variable within each 24-hour cycle. Levels of pH and dissolved 
oxygen were highly correlated in these ponds. 

15. WTiere fish were fed, their growth-rate was not found to 
correlate highly with alkalinity nor with other chemical factors that 
were analyzed. But, the condition index of the fish was lowest when 
daily values of dissolved oxygen and of pH were most variable. 
When the average weight ( and standing crop ) of fish was greatest, 
daily fluctuations in pH and oxygen-content were extreme, total 
dissolved soHds were highest, and alkalinity was lowest. 

16. Where fish were fed, production was poor (feed was not 
utilized well) when water-temperatures fell below 70° F. in early 
autumn. 

ACKNOWLEDGMENTS 

These investigations were supported financially by the State Biological Sur- 
vey of Kansas and the Kansas Forestry, Fish and Game Commission (in part 
under Dingell-Johnson Project F-12-R). Professor E. Raymond Hall and As- 
sociate Professor Kenneth B. Axmitage read the manuscript and offered helpful 
suggestions. The following members of the State Biological Survey aided in 
various phases of the investigation: Messrs. J. C. Barlow, W. Berg, F. Busey, 
D. A. Distler, C. E. Judd, E. E. Klaas, G. Philhps, J. H. Vandermeer, J. Well- 
man, and M. L. Wiley. Miss Betty Sue Giesler prepared the figures. Com- 
puter-time was made available by the Computation Center, The University of 
Kansas. Mr. Vandermeer WTrote or modified the computer-programs. 



254 University of Kansas Publs., Mus. Nat. Hist. 

LITERATURE CITED 

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Canfield, H. L. 

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Faber, H. a. (Editor) 

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Production of Channel Catfish 255 

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Shepard, M. p. 

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256 University of Kansas Publs., Mus. Nat. Hist. 

1957. Preliminary results on the commercial production of channel cat- 
fish in ponds. Proc. 10th Annual Coru. Southeastern Game and 
Fish Comm., 1956:160-162. 

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Fish Comm., 1958:63-72. 

1961. Relationship of pH of pond waters to their suitabihty for fish cul- 
ture. Proc. Pacific Sci. Congress 9 (1957), Vol. 10, Fisheries, 
72-75. 

Swingle, H. S., and Smith, E. V. 

1939. Increasing fish production in ponds. Trans. North American Wildl. 
Conf., 4:332-338. 

Swingle, W. E. 

1964. Length-Weight Relationships I and II, IBM 1620, Fortran/Format. 
Trans. Amer. Fish. Soc, 93(3):318-319. 

Tanner, H. A. 

1960. Some consequences of adding fertilizer to five Michigan trout lakes. 
Trans. Amer. Fish. Soc, 89(2):198-205. 

TiEMEIER, O. W. 

1957. Notes on stunting and recovery of the charmel catfish. Trans. Kan- 
sas Acad. Sci., 60(3):294-296. 

1962. Increasing size of fingerling channel catfish by supplemental feed- 
ing. Trans. Kansas Acad. Sci., 65(2): 144-153. 

TiEMEIER, O. W., Deyoe, C. W., and Weardon, S. 

1965. Effects on growth of fingerfing channel catfish of diets containing 
two energy and two protein levels. Trans. Kansas Acad. Sci., 
68(4):379-392. 

TiEMEIER, O. W., and Elder, J. B. 

1957. Limnology of Fhnt Hills farm ponds for 1956 and preliminary re- 
port on growth studies of fishes. Trans. Kansas Acad. Sci., 60(4): 
379-392. 

TiEMEIER, O. W., and Moorman, R. B. 

1957. Limnological observations on some FUnt Hills farm ponds. Trans. 
Kansas Acad. Sci., 60 (2): 167-173. 

TuNisoN, A. v., et al. 

1939. Carbohydrate utifization by trout. Cortland Hatchery Report, 

8:9-12. 
1943. Carbohydrate digestion. Cortland Hatchery Report, 12:9-10. 

Transmitted March 14, 1966. 



n 

31-3428 



(Continued from inside of front cover) 

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, 
Mexico. By William E. Duelhnan. Pp. 19-72, pis. 1-8, 3 figures in text. 
August 16, 1960. 30 cents. 
3. A new subspecies of the slider turtle (Pseudemys scripta) from Coahuila, 
Mexico. Bv John M. Legler. Pp. 73-84, pis. 9-12, 3 figures in text. August 
16, 1960. 
«4. Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pis. 13-20, 
26 figures Ln text. November 30, 1960. 

5. Occurrence of the garter snake, Thamnophis sirtalis, in tlie Great Plains and 
Rocky Mountains. By Henry S. Fitch and T. Paul Maslin. Pp. 289-308. 
4 figures in te.vt. Februaiy 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 cvprinid fish, Hybopsis gracilis. 
By Leonard J. Olund and Frank B. Cross. Pp.'323-348, pis. 21-24, 2 figures 
in text. February 10, 1961. 

8. Descriptions of two species of fi"Ogs, genus Ptychohyla; studies of American 
hylid frogs, V. By William E. Duellman. Pp. 349-357, pi. 25, 2 figures 
in tex-t. 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, pis. 26-30, 3 fig- 
ures in text. August 11, 1961. 75 cents. 

10. North American recent soft-shelled turtles (Family Trionychidae ) . By 
Robert G. Webb. Pp. 429-611, pis. 31-54, 24 figures in text. February 
16, 1962. $2.00. 

Index. Pp. 613-624. 

Vol. 14. 1. Neotropical bats from western Mexico. By Sydney Anderson. Pp. 1-8. 
October 24, 1960. 

2. Geographical 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 figwe in text. July 24, 1961. 

3. Mammals of Mesa Verde National Park, Colorado. Bv Sydnev Anderson. 
Pp. 29-67, pis. 1 and 2, 3 figures in tex-t. Jvily 24, 1961. 

4. A new subspecies of the black myotis (bat) from eastern Mexico. By E. 
Raymond Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December 
29, 1961. 

5. North American yellow bats, "Dasypterus," and a list of the named kinds 
of the genus Lasiurus Gray. By E. Raymond Hall and J. Knox Jones, Jr. 
Pp. 73-98, 4 figures in text. December 29, 1961. 

6. Natural history of the brush mouse ( Peromyscus boylii ) in Kansas with 
description of a new subspecies. Bv Charles A. Long, Pp. 99-110, 1 figvue 
in text. December 29, 1961. 

7. Taxonomic status of some mice of the Peromyscus boylii group in eastern 
Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 111- 
120, 1 figure in text. December 29, 1961. 

8. A new subspecies of ground squirrel ( Spermophilus spilosoma ) from Ta- 
maulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962. 

9. Ta.\onomic status of the free-tailed bat, Tadaiida yucatanica Miller. By J. 
Knox Jones, Jr., and Ticul Alvarez. Pp. 12S-133, -1 figure in text. March 7, 
1962. 

10. A new doglike carnivore, genus Cynarctus, from the Clarendonian, Pliocene, 
of Texas. Bv E. Ravmoiid Hall and Walter W. Dalquest. Pp. 135-138, 
2 figures in te.xt. April 30, 1962. 

11. A new subspecies of wood rat (Neotoma) from northeastern Mexico. By 
Ticul Alvarez. Pp. 139-143. April 30, 1962. 

12. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul 
Alvarez, and M. Ravmond Lee. Pp. 145-159, 1 figure in text. May 18, 
1962. 

13. A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 
1 figTU-e in text. May 21, 1962. 

"14. The mammals of Veracruz. By E. Raymond Hall and Walter W. Dalquest. 
Pp. 165-362, 2 figures in text. May 20, 1963. $2.00. 

15. The Recent mammals of Tamaulipns, Mexico. By Ticul Alvarez. Pp. 363- 
473, 5 figures in text. May 20, 1963. $1.00. 

16. A new subspecies of the fruit-eating bat, Stumira ludovici, from western 
Mexico. Bv J. Knox Jones, Jr., and Gary L. Phillips. Pp. 475-481, 1 figure 
in text. March 2, 1964. 

17. Records of the fossil mammal Sinclair^lla, Family Apatemyidae, from the 
Chadronian and Orellan. By William A. Clemens, Jr. Pp. 483-491, 2 figures 
in text. March 2, 1964. 

18. The mammals of Wvoming. By Charles A. Long. Pp. 493-758, 82 figures 
in text. July 6, 1965. $3.00. 

Index. Pp. 759-784. 

(Continued on outside of back cover) 



(Continued from inside of back cover) 

Vol 15. 1. The amphibians and reptiles of Michoacdn, Mexico. By William E. Duell- 
man. Pp. 1-148, pis. 1-6, 11 figures in text. /December 20, 1961. $1.50. 

2. Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox 
Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962. 

3 A new species of frog (genus Toniodactylus ) from western Mexico. By 
Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962. 

4. Type specimens of amphibians and reptiles in the Museum of Natural His- 
tory, The University of Kansas. By William E. Duellman and Barbara Berg. 
Pp. 183-204. October 26, 1962. 

5. Amphibians and reptiles of the rainforests of southern El Peten, Guatemala. 
By William E. Duellman. Pp. 205-249, pis. 7-10, 6 figures in text. October 
4, 1963. 

6. A revision of snakes of the genus Conophis (Family Colubridae, from Middle 
America). By John Wellman. Pp. 251-295, 9 figures in text. October 4, 
1963. 

7. A review of the Middle American tree frogs of the genus Ptychohyla. By 
William E. Duellman. Pp. 297-349, pis. 11-18, 7 figures in text. October 
18, 1963. 50 cents. 

8. Natural history of the racer Coluber constrictor. By Henry S. Fitch. Pp. 
351-468, pis. 19-22, 20 figures in text. December 30, 1963. $1.00. 

9. A review of the frogs of the Hyla bistincta group. By William E. Duellman. 
Pp. 469-491, 4 figures in text. March 2, 1964. 

10. An ecological study of the garter snake, Thamnophis sirtalis. By Henry S. 
Fitch. Pp. 493-564, pis. 23-25, 14 figures in text. May 17, 1965. 

11. Breeding cycle in the ground skink, Lygosoma laterale. By Henry S. Fitch 
and Harry W. Greene. Pp. 565-575, 3 figures in text. May 17, 1965. 

12. Amphibians and reptiles from the Yucatan Peninsula, Mexico. By William 
E. Duellman. Pp. 577-614, 1 figure in text. June 22, 1965. 

13. A new species of turtle, genus Kinosternon, from Central America. By John 
M. Legler. Pp. 615-625, pis. 26-28, 2 figures in text. June 20, 1965. 

14. A biogeoaraphic account of the herpetofauna of Michoacan, Mexico. By 
WilUam E. Duellman. Pp. 627-709, pis. 29-36, 5 figures in text. December 
30, 1965. 

15. Amphibians and reptiles of Mesa Verde National Park, Colorado. By Charles 
L. Douglas. Pp. 711-744, pis. 37 and 38, 6 figures in text. March 7, 1966. 
Index in preparation. 

Vol. 16. 1. Distribution and taxonomy of mammals of Nebraska. By J. Knox Jones, Jr. 
Pp. 1-356, plates 1-4, 82 figures in text. October 1, 1964. $3.50. 

2. Synopsis of the lagomorphs and rodents of Korea. By J. Knox Jones, Jr., 
and David H. Johnson. Pp. 357-407. February 12, 1965. 

3. Mammals from Isla Cozumel, Mexico, with description of a new species of 
harvest mouse. By J. Knox Jones, Jr., and Timothy E. Lawlor. Pp. 409-419, 
1 figure in text. April 13, 1965. 

4. The Yucatan deer mouse, Peromyscus yucatanicus. By Timothy E. Lawlor. 
Pp. 421-438, 2 figures in text. July 20, 1965. 

5. Bats ftom Guatemala. By J. Knox Jones, Jr. Pp. 439-472. April 18, 1966. 
More numbers will appear in volume 16. 

Vol. 17. 1. Localities of fossil vertebrates obtained from the Niobrara Formation (Cre- 
taceous) of Kansas. By David Bardack. Pp. 1-14. January 22, 1965. 

2. Chorda tympani branch of the facial nerve in the middle ear of tetrapods. 
By Richard C. Fox. Pp. 15-21. June 22, 1965. 

3. Fishes of the Kansas River System in relation to zoogeography of the Great 
Plains. By Artie L. Metcalf. Pp. 23-189, 4 figmes in text, 51 maps. March 
24, 1966. 

4. Factors affecting growth and production of channel catfish, Ictalurus punctatus. 
By Bill A. Simco rjid Frank B. Cross. Pp. 191-256, 13 figures in text. 
June 6, 1966. 

More numbers will appear in volume 17. 



JUL 121966 

UNIVER ilTY 

University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 5, pp. 257-262, 1 Fig. 
June 17, 1966 



A New Species of Fringe-limbed Tree Frog, 
Genus Hyla, from Darien, Panama 



BY 



WILLIAM E. DUELLMAN 



University of Kansas 

Lawrence 

1966 



University of Kansas Publications, Museum of Natural History 

Editors: E. Raymond Hall, Chairman, Henry S. Fitch, 
Frank B. Cross 



Volume 17, No. 5, pp. 257-262, 1 Fig. 
Published June 17, 1966 



University of Kansas 
Lawrence, Kansas 



PRINTED BY 

ROBERT R. (BOB) SANDERS. STATE PRINTER 

TOPEKA. KANSAS 

1 966 



31-3431 



A New Species ot Fringe-limbed Treei Frog^^, ^^'^ 
Genus Hyla, from Darien, Paiifania, ^ 

WILLIAM E. DUELLMAN UNi\/[~.S^ '^ 

Exploration in the mountainous areas of Darien Province in 
eastern Panama has revealed the presence of several previously 
unknown amphibians. One of the most unusual species found to 
date is a large fringe-limbed tree frog, which, like other fringe- 
limbed species, has large hands and fully webbed feet. In allusion 
to the fringelike dermal folds on the limbs, I propose that the new 
species be named 

Hyla thysanota new species 

Holotype. — United States National Museum No. 151080 from Cerro Mali, 
Darien Province, Panama (elevation 1265 meters), obtained by Dr. Charles 
O. Handley, Jr., on February 11, 1964. 

Diagnosis. — Size large (95.7 mm. snout- vent length); dermal fringe on 
outer edge of forearm and outer edge of tarsus; skin of head and dorsum 
granular and uniformly green. 

Description of holotype. — Adult female containing ovarian eggs and having 
snout-vent length of 95.7 mm.; tibia length 51.0 mm., 52 per cent of snout-vent 
length; foot length (measured from proximal edge of inner metatarsal tubercle 
to tip of longest toe) 44.6 mm., 46 per cent of snout-vent length; head length 
31.1 mm., 32 per cent of snout-vent length; greatest width of head just anterior 
to angle of jaws 36.5 mm., 36 per cent of snout-vent length; snout, in lateral 
profile, rounded, in dorsal profile, broadly rounded; canthus heavy, rounded; 
loreal region deeply concave; lips broad, flaring. Nostrils protuberant laterally; 
intemarial distance 7.0 mm., 19 percent of head width; head flat above; 
interorbital distance 12.0 mm., 33 per cent of head width; width of eyelid 
8.1 mm., 22 per cent of head width; diameter of eye 8.3 mm.; pupil horizontally 
ovoid, palpebrum unmarked; diameter of tympanum 4.8 mm., 58 per cent of 
eye, situated directly posterior to eye, separated from eye by 4.9 mm.; upper 
edge of tympanum covered by dermal fold extending posteriorly from eye to 
point above insertion of arm. 

Upper arm short, slender; forearm only slightly robust; two transverse folds 
on wrist; thin, scalloped dermal fringe along outer edge of forearm from elbow 
to disc of fourth toe; hand large, 32.0 mm. from proximal edge of prepoUex 
to tip of longest toe; length of fingers from shortest to longest, 1-2-4-3, fourth 
only shghdy longer than second; fingers webbed to bases of discs, except web 
between first and second fingers extending only to bases of penultimate pha- 
langes; discs large, all except that on first finger larger than tympanum; distal 
subarticular tubercle flat, others conical; supernumerary tubercles small, conical, 
present only on proximal segments, especially numerous on base of fourth 

(259) 



260 University of Kansas Publs., Mus. Nat. Hist. 

finger and on thumb, larger and fewer on second finger; prepollex elongate, 
flat, broadly visible from above, supported by flat bony spine; numerous small 
conical tubercles on ventral surface of prepollex and palm (Fig. la); heels 
overlap by about one-third length of tibia when hind limbs adpressed; tibio- 
tarsal articulation extending to eye; distinct dermal folds on knees and heels; 
thin tarsal fold curving from heel to inner metatarsal tubercle; thin, scalloped 
dermal fringe along outer edge of foot from heel to disc of fifth toe; inner 
metatarsal tubercle flat, broadly elliptical, visible from above; outer metatarsal 
tubercle absent; length of toes from shortest to longest, 1-2-5-3-4; toes fully 
webbed; discs smaller than on fingers; subarticular tubercles conical; super- 
numerary tubercles small, conical, in one row on each proximal segment 
(Fig. lb). 

Anal opening directed posteriorly at level of middle of thighs; anal sheath 
short, granular; distinct granular dermal fold on each side of anus. Skin on 
top of head and body finely granular (perhaps containing osteoderms); skin 
on belly and median posteroventral surfaces of thighs granular; other surfaces 
smooth. Tongue broadly cordiform, shallowly notched posteriorly, barely free 
behind. Vomerine ridges transverse, narrowly separated medially, situated 
between round choanae; vomerine teeth, 10-11; premaxillary teeth, 15-16; 
maxillary teeth, 73-71; all teeth strongly spatulate and bifid. 

Color (in alcohol): dark purplish brown on dorsal surfaces of head, body, 
limbs, outer three fingers, and outer two toes; lower flanks, anterior and posterior 
siu-faces of thighs, and ventral surfaces of hind limbs pale brown; webs dark 
brown; belly creamy yellow with brown spots anteriorly; chin brown; upper 
lip grayish white. Color ( in life, according to collector's field notes ) : dorsum 
green: ventral surfaces pinkish white; eyes brown. 

Skull (skin peeled back from head permitting superficial examination of 
skull) broader than long; having only moderate amount of ossification — no 
well-developed maxillary flanges (pars facialis) nor supraorbital processes; 
alary processes of premaxillary high; intemasal septum cartilaginous; nasals 
slender, separated, long axes parallel to maxillaries, anterior ends not extending 
beyond sphenethmoid, not forming complete bony anterior margin to orbit; 
sphenethmoid large, bony in entire interorbital distance, anterolateral margins 
attached to nasals, posterior margin overlain by frontoparietals, extending 
posteriorly three-quarters length of orbit; frontoparietals slender, converging 
medially posterior to frontoparietal fontanelle, having cartilaginous flange over 
posterior part of orbit; frontoparietal fontanelle ovoid, about thrice as long 
as wide; quadratojugal robust; squamosal having extremely short anterior arm, 
moderately long posterior arm, well-developed ventral arm; prevomers large, 
forming anterior, but not posterior, margins of choanae; palatines well devel- 
oped, toothless, extending medially to sphenethmoid, lacking prepalatine 
processes. 

Discussion. — Htjla thysanota is another species in that group of 
bizarre hylids usually referred to as the fringe-limbed tree frogs. 
In size it approaches Htjia immensa Taylor from Costa Rica and 
H. phantasmagoria Dunn from Colombia, but both of these species 
have tuberculate skin on the dorsum, shorter and more blunt snouts, 
relatively larger tympani, and mottled brown dorsal coloration. 



New Species of Hyla From Panama 



261 




Fic. 1. Ventral views of right hand (a) and right foot (h) of holotype of 
Hyla thysanota (USNM 151080). X 2. 



262 University of Kansas Publs., Mus. Nat, Hist, 

Hyla miliaria Cope from Nicaragua, H. richardtaylori Taylor and 
H. fimbrimembra Taylor from Costa Rica are smaller species hav- 
ing less webbing, more scalloped dermal fringes, and brown dorsal 
coloration, Hyla valancifer Firschein and Smith and H. echinata 
Duellman from Mexico have smooth skin on the dorsum, narrower 
fringes on the limbs, and brown dorsal coloration. The uniformly 
green dorsum of H. thijsanota is unique among the fringe-limbed 
hylids. 

Present knowledge of the fringe-limbed hylids, of which there 
are nearly as many names as known specimens, is insufficient to 
account for relationships. Perhaps the several species are members 
of two or more phyletic lines. It is noteworthy that the sizes and 
relationship of the cranial roofing bones of Hyla thysanota closely 
resemble those in Hyla maxima. 

The type was shot from a tree-top at night, after its eye-shine 
was observed. The specimen was found in humid montane forest 
near the headwaters of the Rio Pucro ( Pacific drainage via the Rio 
Tuira) on the east slope of Cerro Mali, which lies south of Cerro 
Tacarcuna in the Serrania del Darien. 

Acknowledgments. — In January of 1964 I was in Panama and planned to ac- 
company Dr. Handley to Cerro Mali. Political events delayed our departure 
and I had to return to the United States. Dr. Handley generously took time 
from his collecting of mammals to secure amphibians and reptiles. I am grate- 
ful to him for his eflForts and to James A. Peters for the loan of the specimens 
from the United States National Museum. I am indebted to Linda Trueb for 
the delineation of the hands and feet reproduced here as Figure 1. This paper 
is one result of the author's studies on Middle American hylid frogs supported 
by a grant from the National Science Foundation ( NSF GB-1441 ) and a part 
of a survey of the herpetofauna of Panama supported by a grant from the 
National Institutes of Health (NIH GB-12020), 

Transmitted March 14, 1966. 



n 



31-3431 



I • I «_> ~' v— ^-^ ■ 



J l H r44U 966 

University of Kansas Publications'"'^^^^'^'^ 

UNIVERSITY 
Museum of Natural History 



Volume 17, No. 6, pp. 263-279 
June 17, 1966 



Taxonomic Notes on Some Mexican and 
Central American Hylid Frogs 



BY 



WILLIAM E. DUELLMAN 



University of Kansas 

Lawrence 

1966 



University of Kansas Publications, Museum of Natural History 

Editors: E. Raymond Hall, Chairman, Henry S. Fitch, 
Frank B. Cross 



Volume 17, No. 6, pp. 263-279 
Published June 17, 1966 



University of Kansas 
Lawrence, Kansas 



PRINTED BY 

ROBERT R. (BOB) SANDERS. STATE PRINTER 

TOPEKA. KANSAS 

1966 



31-3429 



JUL 121966 

Taxonomic Notes on Some Mexiq^'j',^|^j^'|lj:^Y 
Central American Hylid Frogs 

BY 

WILLIAM E. DUELLMAN 

The acquisition of series of many species of hylid frogs from 
Mexico and Central America and the comparison of these speci- 
mens with the types of many named taxa have provided the basis 
for certain taxonomic conclusions. The purpose of this paper is to 
comment on certain species and to make necessary nomenclatural 
changes. In this report, eleven taxa are placed in the synonymy 
of nine others, one previously recognized species is regarded as a 
subspecies, and one species is placed in a different genus. The 
genera and species, as now recognized, are treated in alphabetical 
order. 

Tlie abbreviations used for the various collections are: AMNH 
(American Museum of Natural History), CNHM (Chicago Nat- 
ural History Museum), FAS (Frederick A. Shannon collection), 
KU (University of Kansas Museum of Natural History), MCZ 
(Museum of Comparative Zoology), RCT (Richard C. Taylor col- 
lection), UMMZ (University of Michigan Museum of Zoology), 
and USNM (United States National Museum). 

For the loan of specimens or for permitting me to work in their 
respective institutions, I am grateful to Charles M. Bogert, Doris 
M. Cochran, Robert F. Inger, Edmund Malnate, Jay M. Savage, the 
late Frederick A. Shannon, Hobart M. Smith, Charles F. Walker, 
Ernest E. Williams, and Richard G. Zweifel. This paper is a re- 
sult of research on Middle American hylid frogs supported by the 
National Science Foundation (Grant No. GB-1441). 

Gastrotheca ceratophrys (Stejneger) New combination 

Hyla ceratophrys Stejneger, Proc. U. S. Natl. Mus., 41:286, August 14, 1911 
[Holotype. — USNM 47705, Upper Rio Pequeni, Panama Province, Pan- 
ama; A. H. Jennings collector]. 

This large and distinctive species heretofore has been known 
only from the holotype. The acquisition of three additional speci- 
mens, one a female carrying nine eggs, confirms a suspicion that 
the species belongs in the genus Gastrotheca. I have examined 
USNM 141795 from Tacarcuna, Darien, KU 77016 from Laguna, 

(265) 



266 University of Kansas Publs., Mus. Nat. Hist. 

Darien, KU 95794 from 5 kilometers west of Almirante, Bocas del 
Toro, Panama, and the holotype. Three males have snout- vent 
lengths of 68, 72, and 74mm., and the female (KU 77016) has a 
snout-vent length of 71 mm. The specimens are alike in having 
only a vestige of a web between the fingers and in having the toes 
webbed to the bases of the discs, except the fourth toe, which is 
webbed to the base of the antepenultimate phalanx. The anal 
opening is directed posteriorly at the level of the upper surfaces of 
the thighs. Contrary to the statement by Stejneger (1911:286), the 
skin on the head is not co-ossified with the underlying cranial bones. 
There are seven to nine transverse rows of minute tubercles on the 
dorsum of the head and body. The outer edge of the upper eye- 
lid projects in the form of a triangular "horn," which is pointed in 
three specimens, but terminally notched in KU 95794. 

Males are tan on the dorsum with darker brown transverse bands 
of varying widths (diflFuse in USNM 141795) and have narrow 
brownish-black transverse stripes on the limbs. The belly is creamy 
white. The female is much darker than the males and has a dark 
brown dorsum, darkest laterally; the limbs are brown with dark 
brown transverse markings; the ventral surfaces are grayish tan. 
All specimens have a small white spot on each side ventrolateral to 
the anal opening and a pale bar extending from the orbit to the 
lip; in the female this bar is an enamellike white, whereas in the 
others the bar is creamy tan. 

In Hfe the female was pale tan when found at night; by day she 
changed to dull brown above and creamy-tan below with pale 
orange-tan thighs. The subocular bar, labial flecks, and subanal 
spots were white, and the iris was a dull bronze-color. The male 
from five kilometers west of Almirante had an olive-tan dorsum 
with brown transverse markings; the Hmbs were yellowish tan 
above with grayish brown bands, and the webs and flanks were 
yellowish tan. The upper lip was barred with olive-tan and brown; 
the venter was white, and the iris was a coppery bronze-color 
medially and a golden bronze-color peripherally. 

Gastrotheca cornutmn (Boulenger, 1898:124), which occurs on 
the Pacific slopes of Eucador and Colombia, has superciliary tri- 
angular appendages like those in G. ceratophrys. Perhaps the two 
are conspecific, but on the basis of the limited number of specimens 
and the lack of an opportunity to examine the holotype of G. 
cornutum, I tentatively retain G. ceratophrys as a species. 



Taxonomic Notes on Hylid Frogs 267 

Hyla colymba Dunn 

Hyla colymba Dunn, Occas. Pap. Boston Soc. Nat. Hist., 5:400, August 18, 
1931 [Holotype.— MCZ 10234, La Loma, Bocas del Tore Province, 
Panama; Chester Duryea and E. R. Dunn collectors], 

Hyla alvaradoi Taylor, Univ. Kansas Sci. Bull., 35:882, July 1, 1952 [Holo- 
type. — KU 30886, Moravia, Cartago Province, Costa Rica; Edward H. 
Taylor collector]. 

Dunn (1924:3) originally reported this species from La Loma as 
Hyla albomarginata Spix; liis specimens later formed the type series 
of Hyla colymba (Dunn, 1931a:400) and were compared with Hyla 
albomarginata. The hototype and an adult paratype each possesses 
a circular mental gland encompassing most of the anterior edge 
of the chin; this gland was not mentioned by Dunn. Taylor 
(1952:882) based the description of Hyla alvaradoi on one adult 
male and subsequently (1954:625) reported two additional speci- 
mens from the t>'pe locality. Taylor (1952:882) gave a detailed 
description of the t}^pe of Hyla alvaradoi, and he noted the circular 
mental gland, but he did not compare Hyla alvaradoi with H. 
colymba. 

Comparison of the holotypes of H. alvaradoi and H. colymba 
shows that the two specimens are nearly identical in structure; the 
former has fewer supernumerary tubercles on the hand than the 
latter, but a paratype of H. colymba (MCZ 10235) has as many 
supernumerary tubercles as the holotype of H. alvaradoi. The 
coloration of the types of the two alleged species and other speci- 
mens examined agrees and is distinctive. Although in life, the 
frogs are greenish yellow above, in preservative they fade to a 
creamy white with chromatophores forming flecks, or clusters, 
on the dorsum, especially anteriorly. A white or yellowish-white 
line from the canthus along the outer edge of the eyelid to a point 
above the insertion of the arm is a diagnostic feature of this species. 

Hyla colymba ranges from low to moderate elevations on the 
Caribbean lowlands of Costa Rica to the Serrania del Darien in 
extreme eastern Panama. I have examined specimens from La Lola 
and Moravia in Costa Rica and from the following localities in 
Panama: La Loma, Chiriqui Province; EI Valle, Code Province; 
Altos de Pacora, Panama Province, and Laguna and Cerro Mali, 
Darien Province. 

Hyla ebraccata Cope 

Hyla ebraccata Cope, Proc. Acad. Nat. Sci. Philadelaphia, 26:69, 1874 
[Holotype. — unknown, Nicaragua; John F. Bransford collector]. 

Hyla weyerae Taylor, Univ. Kansas Sci. Bull., 36:633, June 1, 1954 [Holo- 
cype. — KU 34850, Esquinas Forest Preserve, Las Esquinas, betweer» 



268 University of Kansas Publs., Mus. Nat. Hist. 

Palmar and Golfito, Puntarenas Province, Costa Rica; Mrs. Albert E. 
Weyer collector]. 

In his description of HyJa tveyerae Taylor (1954:635) did not 
compare the frog with Hyla ebraccata, but instead stated: "This 
small species may belong in a group of small frogs that includes 
Hyla alleei, Hyla rufiocula (sic), and Hyla uranochroa. . . ." 
There are no morphological characters to associate Hyla tveyerae 
with H. uranochroa and its allies, but there exist no morphological 
characters by which to separate the holotype of H. tveyerae from 
H. ebraccata. All individuals of the latter are like the type of 
H. tveyerae in possessing an extensive axillary membrane, white 
labial stripe expanded below the eye, and unpigmented thighs. 
Most specimens ( 462 of 549 examined from Middle America ) have 
a dorsal dark-mark roughly in the form of an hour-glass. The 
holotype of H. tveyerae differs by lacking any dorsal markings. 

Examination of series of specimens from throughout the range 
of the species reveals the presence of plain ( unmarked ) individuals 
from several localities, but they always are in the minority as com- 
pare with individuals having the hour-glass mark on the back. 
Some individuals lack the hour-glass mark but have small dark 
brown spots on the dorsum; some specimens from the northern part 
of the range have the hour-glass mark fragmented into an anterior 
triangular mark and one or two spots in the sacral region (Table 
1 ) . The highest incidence of plain individuals is in the Palmar- 
Golfito area of Costa Rica (including the type locality of H. 
tveyerae), where six of 13 specimens have no dorsal markings. 

I have collected plain and "normal" individuals from the same 
pond on the same night and can discern no differences in the breed- 
ing call. These observations and the occurrence of plain individuals 
at several localities on both Caribbean and Pacific lowlands of 
Central America indicate that the name Hyla tveyerae is based on 
a color variant of Hyla ebraccata. 

Dunn (1931b: 407, 1933:63) and Breder (1946:417) used the 
name Hyla leucophyllata (Beireis) for Panamanian frogs currently 
assigned to this species. Likewise, Taylor ( 1942:80) used that name 
for specimens from Piedras Negras, Guatemala. I am now con- 
vinced that all known specimens from Mexico and Central America 
are H. ebraccata, which ranges into northern Colombia and differs 
from H. leucophyllata in having a white labial mark expanded 
below the eye. The lips of H. leucophyllata are uniform brown. 



Taxonomic Notes on Hylid Frogs 



269 



Table 1. — Color Pattern Variants in Hyla ebraccata 



Locality 


Hour-glass 


Triangle 


Spotted 


Plain 


Total 


Valle Nacional, Oaxaca, 

Mexico 

Teapa, Tabasco, 

^lexico 


19 
13 
47 
87 
48 
21 
45 
50 
7 
46 
16 
63 


2 
6 


6 
3 

16 
3 


1 

14 

15 

6 

15 


19 
16 


Toocog, El Peten, 

Guatemala 

Turrialba, Cartago, 

Costa Rica 


53 

87 


Pacuare, Cartago, 

Costa Rica 


48 


Moravia, Cartago, 

Costa Rica 


21 


Suretka, Limon, 

Costa Rica 


65 


Tilardn, Guanacaste, 

Costa Rica 


68 


Palmar — Golfito, Puntarenas 
Costa Rica 


13 


Anchiote, Col6n, 

Panamd 


77 


Cerro la Campana, 

Panamd, Panamd 

Tacarcuna-Rio Mono, 

Darien, Panamd 


19 
63 






Totals 


462 


8 


28 


51 


549 







Several species in the Hyla leucophyllata group in the upper 
Amazon Basin (H. bifurca, frontalis, laynei, memhranacea, reti- 
culata, rossalleni, and sarayacuensis) diflFer only in minor features 
of the color pattern. Hyla memhranacea is an unmarked species 
colored like the holotype of H. weyerae, and Hyla laynei is spotted 
much hke many individuals of H. ebraccata. The evidence pre- 
sented here concerning variation in color-pattern in Hyla ebraccata 
suggests that detailed studies of hving frogs and series of specimens 
of the Hyla leucophyllata group in the upper Amazon Basin might 
reveal an unwarranted multiplicity of named taxa. 



270 University of Kansas Publs., Mus. Nat. Hist. 

Hyla elaeochroa Cope 

Hyla elaeochroa Cope, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 8:105, 
1876 [Lectotype. — USNM 30689, east foot of mountains near Sipurio, 
Limon Province, Costa Rica; William M. Gabb collector]. 

Hyla quinquevittata Cope, Proc. Amer. Philos. Soc, 23:273, April 1886 
[Holotype. — USNM 14187, Nicaragua; J. F. Bransford collector]. 

Hyla dvlcensis Taylor, Univ. Kansas Sci. Bull., 39:37, November 18, 1958 
[Holotype. — KU 32168, Golfito, Puntarenas Province, Costa Rica; Ed- 
ward H. Taylor collector]. 

Dunn and Emlen (1932:25) placed H. elaeochroa and H. quin- 
quevittata as synonyms of H. rubra. Taylor (1952:861) concurred 
that H. elaeochroa and H. quinquevittata were conspecific but 
thought that Hijla rubra was another species. Taylor (1958:37) 
described Hyla dulcensis from Golfito on the Pacific lowlands of 
Costa Rica; he diagnosed the new species (based on two males and 
one female) as differing from H. elaeochroa on the Caribbean low- 
lands in having "somewhat larger size, smaller finger and toe discs, 
the obsolete canthus rostralis, the loreal region concave, and the 
choanae larger." 

Examination of several series of specimens from Costa Rica and 
Panama reveals that individuals from the Golfo Dulce area on the 
Pacific lowlands of Costa Rica attain a slightly larger size than do 
the frogs elsewhere in the range of the species, but that the differ- 
ence is minor. For example, fifteen males chosen at random from 
each of three localities in Costa Rica vary in snout-vent length as 
follows: Golfo Dulce area, Puntarenas Province 33.8 (29.6-38.8 
mm.); Puerto Viejo, Heredia Province 29.6 (27.9-32.3 mm.); Tur- 
rialba, Cartago Province 30.5 (27.9-32.6 mm.). Sixteen males (in- 
cluding the holotype) from the Golfo Dulce region display varia- 
tion in the canthus, loreal region, size of discs on digits and size 
of choanae that extends from the condition described in H. dulcensis 
to that ordinarily found in H. elaeochroa on the Caribbean low- 
lands. The nearly identical breeding calls, close resemblance of 
tadpoles, and lack of any definitive morphological criteria indicate 
that Hyla dulcensis Taylor is a synonym of H. elaeochroa Cope. 

Until Taylor named Hyla dulcensis, Hyla elaeochroa was not 
known from the Pacific coast of Central America. Subsequent col- 
lecting has extended the known range of H. elaeochroa westward 
through the Arenal depression in Guanacaste and southward along 
the Pacific slopes to the Golfo Dulce area. 

Dunn and Emlen (1932:25) incorrectly placed H. elaeochroa 
in the synonymy of H. rubra. The former has no pattern on the 
posterior surfaces of the thighs, whereas the bold black and creamy 



Taxonomic Notes on Hylid Frogs 271 

yellow pattern is present in H. rubra. A gap of about 225 kilo- 
meters separates the known ranges of the two species. Hyla 
elaeochroa ranges from east-central Nicaragua southward to the 
Golfo Dulce area of Costa Rica and on the Caribbean lowlands to 
Laguna de Chiriqui in western Panama, whereas Hyla rubra is 
widespread in South America, but in Panama ranges no farther west 
than the Canal Zone. 

Hyla lancasteri Barbour 

Hyla lancasteri Barbour, Proc. New England Zool. Club., 10:31, Pi. 4, fig. 2, 

March 2, 1928 [Holotype.— MCZ 13062, Peralta, Cartago Province, 

Costa Rica; C. R. Lancaster collector]. 
Hijla moraviaensis Taylor, Univ. Kansas Sci. Bull., 35:865, fig. 57, July 1, 

1952 [Holotype. — KU 30284, Moravia, Cartago Province, Costa Rica; 

Edward H. Taylor collector]. 

Barbour (1928:31) named Hyla lancasteri on the basis of one 
juvenile having a snout-vent length of 19.1 mm. Dunn and Emlen 
(1932:25) placed H. lancasteri in the synonymy of H. boulengeri. 
Taylor (1952:858) questioned that allocation of H. lancasteri. A 
comparison of the holotypes of H. lancasteri and H. moraviaensis 
with well-preserved specimens of H. boulengeri (including juve- 
niles) reveals that the latter is specifically distinct from both H. 
lancasteri and H. moraviaensis, but that the holotype of H. lancasteri 
is conspecific with the frogs that Taylor (1952:865) named Hyla 
moraviaensis. Although the type of H. lancasteri is in poor condi- 
tion and the only markings present are the dark bars on the hind 
limbs, the species can be associated with H. moraviaensis by the 
broad head, short truncate snout, short fingers, and amount of 
webbing. In all of these characters H. lancasteri differs notably 
from H. boulengeri, which has a relatively narrow head, long 
pointed snout, and relatively long fingers. Dunn and Emlen 
(1932:25) apparently synonymized H. lancasteri with H. boulengeri 
solely on the basis of the strongly barred thighs, a coloration known 
at that time, in Central American hylids, only in H. boulengeri. 

Although many specimens of Hyla lancasteri are in collections, 
all are from the Caribbean slopes of Cartago Province, Costa Rica, 

Hyla loquax Gaige and Stuart 

Hyla loquax Gaige and Stuart, Occas. Pap. Mus. Zool. Univ. Michigan, 
281:1, June 9, 1934 [Holotype.— UMMZ 75446, Ixpuc Aguada, north 
of La Libertad, Departamento El Peten, Guatemala; L. C. Stuart collec- 
tor]. 

Hyla stadelmani Schmidt, Proc. Biol. Soc. Washington, 49:45, May 1, 1936 
[Holotype. — MCZ 21310, Subirana Valley, Departamento Yoro, Hon- 
duras; Raymond E. Stadelman collector.] 



272 University of Kansas Publs., Mus. Nat. Hist. 

Hyla axillamembrana Shannon and Werler, Trans. Kansas Acad. Sci., 58:383, 
fig. 6, September 24, 1955 [Holotype.— FAS 4083, 5 mi. south of Lake 
Catemaco on San Andres Tuxtla-Minatitlan road, Veracruz, Mexico; John 
Werler and Jack Reid collectors]. 

Hyla loqttax is one of the most distinctive Middle American tree 
frogs; in living and freshly preserved specimens the thighs and 
webs are red, but in old preserved specimens the red fades to 
creamy white. Gaige and Stuart's (1934:2) description included 
an account of the coloration of living frogs ( collected and observed 
by Stuart), but Schmidt (1936:45) had no acquaintance with the 
frogs in life that he named Hyla stadelmani. 

I have compared the type and a paratype of H. stadelmani ( MCZ 
21310-11) with two paratypes of H. loqimx (MCZ 19754, 21456) 
and several other specimens of H. loqtmx from Central American 
and southern Mexico. These specimens obviously represent one 
taxon. The older preserved specimens of H. stadelmani and H. 
loquax, from which the colors have faded, are indistinguishable. 

Duellman (1960:62) placed Hyla axillamembrana in the syn- 
onymy of Hyla loquax, which is widely distributed in the forested 
Gulf and Caribbean lowlands from southern Veracruz to Costa Rica. 

Hyla melanomma bivocata Duellman and Hoyt, new combination 

Hyla bivocata Duellman and Hoyt, Copeia, no. 4:414, figs. 1-3, December 
19, 1961 [Holotype. — KU 58446, stream above (6.2 kilometers by road 
south of) Rayon Mescalapa, Chiapas, Mexico; William E. Duellman and 
Dale L. Hoyt collectors]. 

Examination of specimens of Hyla melanommn. (Taylor, 1940) 
and the acquisition of specimens from the Mexican state of Oaxaca 
provide evidence of close relationship of H. bivocata to H. mela- 
nomma. Both are small (snout-vent lengths of breeding males 
26.1-29.9 mm.), yellowish-tan frogs having axillary membranes, 
acuminate snouts in dorsal view, and breeding calls consisting of a 
primary note followed by shorter secondary notes. These resem- 
blances notwithstanding, certain difiPerences exist between the frogs 
in the Chiapan highlands and those in the Mexican highlands west 
of the Isthmus of Tehuantepec. The most noticeable difference be- 
tween H. bivocata from Chiapas and H. melanomma from Guerrero 
and Oaxaca is the much smaller tympanum of the former. Seven 
males of H. bivocata from above Rayon Mescalapa, Chiapas, have 
tympanum/eye ratios of 0.276-0.323 (mean 0.298), whereas the ratio 
in three samples of H. melanomma is higher — eight males from 
Agua del Obispo, Guerrero, 0.500-0.531 (0.514), seven from 12 
kilometers north-northwest of San Gabriel Mixtepec on the Pacific 



Taxonomic Notes on Hylid Frogs 273 

slopes of Oaxaca, 0.478-0.550 (0.511), and two from Campamento 
Vista Hermosa on the Atlantic slopes of Oaxaca, 0.368-0.419 (0.394). 
Aside from this measurable difference, the Chiapan frogs have 
slightly less webbing on the hands and fewer dark flecks on the 
dorsal surfaces, especially on the shanks, as compared with those 
specimens from Oaxaca and Guerrero. 

Subspecific recognition of the populations seems to be warranted. 
Thus, Hyla bivocata Duellman and Hoyt becomes Hijla melanomma 
bivocata. It is known only from the northern slopes of the high- 
lands of Chiapas, whereas H. in. melanomma occurs on the coastal 
slopes of the Sierra Madre del Sur in Guerrero and Oaxaca. Four 
specimens (KU 87112-5) are from Campamento Vista Hermosa on 
the northern slopes of the Sierra de Juarez in northern Oaxaca. 
The two adult males are intermediate between H. m. bivocata and 
H. m. melanomma in respect to the size of the tympanum relative 
to the eye, but in other characters are more nearly like H. m. 
melanomma. 

Hyla pictipes Cope 

Hylu punctariola pictipes Cope, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 
8:106, 1876 [Syntypes.— USNM 30631, 30652, Pico Blanco, Limon Prov- 
ince, Costa Rica; W. M. Gabb collector]. 

Hula punctariola moesta Cope, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 
^8:106, 1876 [Holotype.— USNM 30660, Pico Blanco, Limon Rov Prov- 
ince, Costa Rica; W. M. Gabb collector]. 

Hyla punctariola monticola Cope, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 
8:106, 1876 [Holotype.— USNM 30661, Pico Blanco, Limon Province, 
Costa Rica; W. M. Gabb collector]. 

Cope (1876:106) erroneously associated his specimens of hylids 
with Peter's Hijla punctariola, which actually is an Eleutherodac- 
tylus (Dunn, 1940:109). Taylor (1952:855, 878) is the only worker 
who subsequently treated these names, and he placed each as a 
distinct species of Hyla, although he had no new material and had 
not examined the types of H. moesta or monticola. 

In April, 1964, Jay M. Savage and I examined the types of the 
three subspecies described by Cope in an attempt to determine 
which names, if any, were applicable to various hylids that we had 
collected in the highlands of Costa Rica. All four types are soft, 
badly faded, and partially deteriorated, thereby making accurate 
comparisons and determinations impossible. We were unable to 
distinguish these types from one another by any seemingly mean- 
ingful character and agreed that the diflPerences observed in the 
types were encompassed in the range of variation displayed by 
series of a species that we had collected in Costa Rica. The names 



274 University of Kansas Publs., Mus. Nat. Hist. 

and descriptions of all three taxa appear on the same page (Cope, 
1876:106). The name that appeared first on the page is here 
chosen. Hyla punctariola moesta and H. p. monticola are relegated 
to the synonymy of Hyla pictipes. 

Since this species have never been adequately described, the following 
description (based on KU 64643-87 from Rio Poasito, Alajuela Province, 
Coast Rica) is provided: snout-vent length of 10 males 34.6 (32.0-36.7 
mm.), of six females 42.2 (40.6-43.2 mm.); tibia sHghtly longer than half 
snout-vent length; snout truncate viewed laterally and acuminate viewed 
dorsally; top of head flat; diameter of tympanimi about half diameter of eye; 
fingers long, stout, about one-fourth webbed; toes short, about two-thirds 
webbed; inner metatarsal tubercle elevated, flat, elliptical; outer metatarsal 
tubercle smaller, conical; tarsal fold lacking; palms, soles, ventral surfaces of 
proximal segments of digits bearing numerous conical supernumerary tubercles; 
anal sheath short; anal opening at level of middle of thighs, directed ventrally; 
skin or dorsum and limbs smooth; skin or belly granular. 

The coloration of preserved specimens is dull brown or tan on dorsal sur- 
faces of females and either uniform dark brown or dull brown with many 
darker spots dorsally in males. In females the flanks and anterior and posterior 
surfaces of the thighs are tan with cream-colored spots, whereas in males these 
surfaces are dark brown or black with smaller cream-colored spots. The ventral 
surfaces are creamy gray with moderate (usually in females) or heavy (usually 
in males) sufi^usion of dark brown. 

In life, males have an olive-green dorsum with dark olive-green 
or brown spots; in females the dorsum is uniform pale green. In- 
dividuals of both sexes have brown flanks and thighs with yellow 
spots and a golden tan iris. 

This stream-breeding Hyla occurs at elevations above 1500 meters 
in the Cordillera Central and in the Cordillera de Talamanca in 
Costa Rica. 

Hyla staufferi Cope 

Hyla staufferi Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:195, October 

1865 [Holotype.— USNM 15317, Orizaba, Veracruz, Mexico; Francis 

Sumichrast collector]. 
Hyla culex Dunn and Emlen, Proc. Acad. Nat. Sci. Philadelphia, 84-24, 

March 22, 1932 [Holotype.— MCZ 16098, Tela, Departamento Atlantidad, 

Honduras; Raymond E. Stadelman collector]. 
Hyla altae Dunn, Occas. Papers Boston Soc. Nat. Hist., 8:61, lune 7, 1933. 

[Holotype. — MCZ 17972, Summit, Canal Zone, Panama]. 

After comparing the types of the three proposed names with 
series of well-preserved specimens from throughout the range of 
the species, I concur with Taylor (1952:865) that only one species 
is involved. Hyla culex was named and described on the basis of 
an adult male having a snout-vent length of 25.1 mm. In his key 
to the species of the Hyla rubra group in Central America, Dunn 
(1933:62) separated Hyla culex from other members of the group 
by the former having a tympanum that was one-third the size of 



Taxonomic Notes on Hylid Frogs 275 

the eye; in the other recognized taxa the tympanum is larger. 
Examination of series of Htjla staiifferi from throughout the range 
of the species reveals that the size of the tympanum is variable, and 
that the frogs in northern Honduras do not differ significantly from 
individuals from other parts of the range. In the description of 
the t\pe, Dunn and Emlen (1932:24) mentioned the presence of 
a dark interorbital triangle, dorsolateral marks, and transverse dark 
marks on the limbs, but Dunn (1933:62) stated that H. cidex lacked 
stripes. The holotype of H. culex is soft and faded, but dark 
dorsolateral stripes, a canthal stripe, and transverse marks on the 
limbs are faintly visible. 

Dunn (1933:61) named Hyla altae on the basis of five males ob- 
tained at Summit, Canal Zone, Panama, and compared these speci- 
mens with Hyla rubra and boulengeri. The holotype of H. altae 
is indistinguishable from individuals of Hyla staufferi, which lack 
dark transverse marks on the limbs. The pattern on the dorsal sur- 
faces of the hind limbs varies from unicolor grayish tan to a pattern 
involving numerous dark flecks, a dark longitudinal stripe, or one 
or two dark transverse marks. Unicolor hind limbs are more prev- 
alent in Panama and Costa Rica than in other parts of the range. 

The taxonomy of Hyla staufferi has been confused for many years. 
Cope (1887:14) placed H. staufferi as a subspecies of H. eximia. 
Giinther (1901:262) considered H. staufferi to be a synonym of 
H. eximia. Gaige (1936:293) suggested that H. culex and H. altae 
might be conspecific, but regarded H. staufferi as a different species, 
although she correctly intimated that H. staufferi belonged in the 
rubra group and not with H. eximia. The only subsequent de- 
parture from this arrangement was that of Blair ( 1960 ) , who placed 
H. staufferi in the eximia group on the basis of the similarity in 
breeding calls. 

Skeletal features, especially the characteristics of the broad nasals 
and slender tegmen tympani, the body form of the tadpoles, and 
the absence of webbing between the first and second toes are some 
of the more significant characters that indicate relationships of Hyla 
staufferi with the Hyla rubra group. Hyla staufferi is the northern- 
most species in the group, which is predominately South American; 
H. staufferi ranges from Guerrero and southern Tamaulipas, Mexico 
southeastward to the Bayano Valley of east-central Panama at low 
and moderate elevations throughout Central America, exclusive of 
the wet tropical forests of the Caribbean lowlands. 



276 University of Kansas Publs,, Mus. Nat. Hist. 

Hyla uranochroa Cope 

Hyla uranochroa Cope, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 8: 103, pi. 
27, fig. 4, 1876 [Holotype.— USNM 30651, near Sipurio, Limon Province, 
Costa Rica; W. M. Gabb collector]. 

Hyla alleei Taylor, Univ. Kansas Sci. Bull., 35:831, fig. 48, July 1, 1952 
[Holotype. — RCT 775, Isla Bonita, Heredia Province, Costa Rica; Richard 
C. Taylor collector]. 

Taylor (1952:837) examined the holotype of H. uranochroa and 
MCZ 10249 from La Loma, Panama, but referred two specimens 
from Isla Bonita to a new species, Hyla alleei, which he distin- 
guished from H. uranochroa on the basis of (1) lacking a white 
stripe above the anus, (2) having the diameter of the tympanum 
about one-half that of the eye (larger in H. uranochroa), (3) hav- 
ing only a vestige of web on hand ( outer-fingers one-fourth webbed 
in H. uranochroa), and (4) lacking a distinct inner tarsal fold 
(moderately distinct in H. uranochroa). I have examined 139 
specimens from Costa Rica and western Panama, including the 
holotype of H. uranochroa and the paratype of H. alleei (CNHM 
101375, formerly RCT 774 ) . Of 20 specimens from the type locaHty, 
only four lack a white stripe above the anus; six of 15 from 
Moravia, Cartago Province, Costa Rica, apparently lack the white 
stripe. The nature of the tarsal fold is variable; in some individuals 
the fold is lacking and in others it is weak. In those specimens 
having a better developed tarsal fold, the fold is weak and extends 
only about half the length of the tarsus. The amount of webbing 
in the hand is slight; in no specimen are the fingers more than one- 
fourth webbed and some specimens have only a vestigial web be- 
tween the outer fingers. The diameter of the tympanum of most 
specimens is 60 to 65 per cent of the diameter of the eye; I have 
seen no specimens having a tympanum/eye ratio of less than 53 
per cent. 

The absence of distinguishing characters of morphology and 
coloration together with the presence of only one kind of breeding 
call and one kind of tadpole is indicative of the presence of only 
one species. Hijla uranochroa is a distinctive stream-breeding frog 
Hving at elevations usually above 1100 meters, although in Costa 
Rica, individuals have been found at 650 meters at Ciudad Quesada, 
Alajuela Province and at 680 meters at El Tigre, Limon Province. 
The species occurs on the Caribbean and Pacific slopes of Costa 
Rica and is known from La Loma on the Caribbean slopes of west- 
ern Panama. 



i 



Taxonomic Notes on Hylid Frogs 277 

Hyla uranochroa is the only member of the genus in Central 
America having a bright green dorsum, creamy-yellow belly, white 
lateral stripe, and bright red eye. Two other apparently related, 
red-eyed species ( H. legleri and H. rtt-fioculis ) differ in having dark 
olive-green or brown dorsal colors and noticeably different breeding 
calls. 

Phrynohyas spilomma (Cope) 

Hyla spilomma Cope, Proc. Amer. Philos. Soc, 17:86, August 1877 [Holo- 
type. — unknown, Cosamaloapam, Veracruz, Mexico; Francis Sumichrast 
collector]. 

Acrodytes modesta Taylor and Smith, Proc. U. S. Natl. Mus., 95:594, pi. 
27, fig. 2, pi. 28, figs. 2-3, June 30, 1945 [Holotype.— USNM 115013, 
Cruz de Piedra, near Acacoyagua, Chiapas, Mexico; Dr. and Mrs. Hobart 
M. Smith collectors]. 

In his revision of Phrynohyas, Duellman (1956) regarded P. 
spilomma and P. modesta as distinct species. The former was char- 
acterized by an irregular dorsal dark mark and distinct dark trans- 
verse bands on the limbs, whereas P. modesta was characterized by 
the absence of any dorsal markings, except scattered black spots. 
In 1956 P. modesta was known from eight localities in southern 
Veracruz and the Pacific lowlands of Chiapas, Guatemala, and 
El Salvador. Fugler (1960:10) reported P. modesta from British 
Honduras, and Neill (1965:88) noted that the color pattern in 
Phrynohyas from British Honduras varied from the nearly unicolor 
pattern of P. modesta to the pattern described for P. spilomma. 

Examination of specimens of Phrynohyas collected since 1956 
reveals two additional specimens from Veracruz, 19 specimens from 
Tabasco, and one from Honduras, in addition to those reported 
from British Honduras. Seventy-two specimens (AMNH 74377-90) 
from Cuyuta, 20 kilometers north of San Jose, Departamento Es- 
cuintla, Guatemala, show a range in variation encompassing that 
previously known in both P. spilomma and P. modesta. Some 
specimens in the series have a unicolored tan dorsum; in others the 
tips of the warts are dark brown. Some specimens have dark brown 
dorsolateral bands, and in others the mid-dorsal region also is dark 
brown. Therefore, it seems that the uniform tan dorsum is only an 
extreme, but common and widespread, color variant of Phrynohyas 
spilomma. 

This intraspecific variation in coloration is like that noted in 
Amazonian P. venulosa by Duellman (1956:39). 



278 University of Kansas Publs., Mus. Nat. Hist. 

LITERATURE CITED 

Barbour, T. 

1928. New Central American frogs. Proc. New England Zool. Club, 
10:25-31, pis. 1-4, March 2. 

Blair, W. F. 

1960. Mating call as evidence of relations in the Hyla eximia group. 
Southwest. Nat., 5:129-135, November 1. 

BotTLENGER, G. A. 

1898. An account of the reptiles and batrachians collected by Mr. W. F. H. 
Rosenberg in western Ecuador. Proc. Zool. Soc. London, 1898: 
107-126, pis. 10-18, June 1. 

Breder, C. M., Jr. 

1946. Amphibians and reptiles of the Rio Chucunaque Drainage, Darien, 
Panama, with notes on their life histories and habits. Bull. Amer. 
Mus. Nat. Hist., 86:375-436, pis. 42-60, August 26. 

Cope, E. D. 

1876. On the Batrachia and Reptilia of Costa Rica. Jour. Acad. Nat. 

Sci. Philadelphia, ser. 2, 8:93-154, pis. 23-28. 
1887. Catalogue of batrachians and reptiles of Central America and 
Mexico. Bull. U. S. Natl. Mus., 32:1-98. 

DUELLMAN, W. E. 

1956. The frogs of the hylid genus Phrynohyas Fitzinger, 1843. Misc. 

Publ. Mus. Zool. Univ. Michigan, 96:1-47, pis. 1-6, February 21* 
1960. A distributional study of the amphibians of the Isthmus of Tehuan- 

tepec, Mexico. Univ. Kansas Publ. Mus. Nat. Hist., 13:19-72, 

pis. 1-8, August 16. 
Dunn, E. R. 

1924. Some Panamanian frogs. Occ. Papers Mus. Zool. Univ. Michigan, 

151:1-12, pis. 1-2, July 1. 

1931a. New frogs from Panama and Costa Rica. Occ. Papers Boston Soc. 
Nat. Hist., 5:385-401, August 18. 

1931b. The amphibians of Barro Colorado Island. Occ. Papers Boston Soc. 
Nat. Hist, 5:403-421, October 10. 

1933. A new Hyla from the Panama Canal Zone. Occ. Papers Boston Soc. 
Nat. Hist, 8:61-64, June 7. 

1940. New and noteworthy herpetological material from Panama. Proc. 
Acad. Nat Sci. Philadelphia, 92:105-122, pi. 2, November 18. 

Dunn, E. R., and Emlen, J. T., Jr. 

1932. Reptiles and amphibians from Honduras. Proc. Acad. Nat. Sci. 
Philadelphia, 84:21-32, March 22. 

FUGLER, C. M. 

1960. New herpetological records for British Honduras. Texas Jour. Sci.„ 
12:8-13. 

Gaige, H. T. 

1936. Some reptiles and amphibians from Yucatan and Campeche, Mexico. 
Carnegie Inst. Washington Publ. 457:289-304, February 5. 

Gaige, H. T. and Stuart, L. C. 

1934. A new Hyla from Guatemala. Occ. Papers Mus. Zool. Univ. 
Michigan, 281:1-3, June 9. 

GiJNTHER, A. C. L. G. 

1885-1902. Biologia Centrah-Americana. Reptilia and Batrachia. Taylor 
and Francis, London, xx + 326 pp., 76 pis. 



Taxonomic Notes on Hylid Frogs 279 

Neill, W. T. 

1965. New and noteworthy amphibians and reptiles from British Honduras. 
Bull. Florida State Mus., 9:77-130, April 9. 

Schmidt, K. P. 

1936. New amphibians and reptiles from Honduras in the Museum of 
Comparative Zoology. Proc. Biol. Soc. Washington, 49:43-50, 
May 1. 

Stejneger, L. 

1911. Descriptions of three new batrachians from Costa Rica and Panama. 
Proc. U. S. Natl. Mus., 41:285-288, August 14. 

Taylor, E. H. 

1940. Herpetological Miscellany, Univ. Kansas Sci. Bull., 26:489-571, 

November 27. 
1942. New tailless Amphibia from Mexico. Univ. Kansas Sci. Bull., 

28:67-89, May 15. 
1952. A review of the frogs and toads of Costa Rica. Univ. Kansas Sci. 

Bull., 35:577-942, July 1. 
1954. Additions to the known herpetological faima of Costa Rica with 

comments on other species. No. I. Univ. Kansas Sci. Bull., 36:597- 

639, June 1. 
1958. Additions to the known herpetological fauna of Costa Rica with 

comments on other species. No, HI. Univ. Kansas Sci. Bull., 

39:3-40, November 18. 

Transmitted March 14, 1966. 



I 



n 

■■31-3429 



University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 7, pp. 281-375, pis. 1-12, 17 figs. 
July 14, 1966 



Neotropical Hylid Frogs, Genus Smilisca 



BY 



WILLIAM E. DUELLMAN AND LINDA TRUEB 



urNr'ivE'RSfT 



University of Kansas 

Lawrence 

1966 



UNIVERSITY OF KANSAS PUBLICATIONS 
MUSEUM OF NATURAL HISTORY 

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345, pis. 5-8. June 18, 1962. 

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in text. October 1, 1962. 

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Hii-undinidae. By Marion Aime Jenkinson. Pp. 553-573 7 figures in text. 
March 2, 1964. 

14. The breeding birds of Kansas. By Richard F. Johnston. Pp. 575-655, 10 
figures in text. May 18, 1964. 75 cents. 

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Fox. Pp. 657-680, 11 figures in text. May 18, 1964. 

Index. Pp. 681-694. 

(Continued on inside of back cover) 



University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 7, pp. 281-375, pis. 1-12, 17 figs. 
July 14, 1966 



Neotropical Hylid Frogs, Genus Smilisca 



BY 



WILLIAM E. DUELLMAN AND LINDA TRUEB 



University of Kansas 

Lawrence 

1966 



University of Kansas Publications, Museum of Natural History 

Editors: E. Raymond Hall, Chainnan, Henry S. Fitch, 
Frank B. Cross 



Volume 17, No. 7, pp. 281-375, pis. 1-12, 17 figs. 
Published July 14, 1966 



University of Kansas 
Lawrence, Kansas 



LIBRARY 

MAR 2 1967 

UNIVERSITY. 



PRINTED BY 

ROBERT R. (BOB) SANDERS. STATE PRINTER 

TOPEKA. KANSAS 

1966 



31-3430 



Neotropical Hylid Frogs, Genus Smilisca 

BY 
WILLIAM E. DUELLMAN AND LINDA TRUEB 

CONTENTS 

PAGE 

Introduction 285 

Acknowledgments 286 

Materials and Methods 287 

Genus Smilisca Cope, 1865 287 

Key to Adults 288 

Key to Tadpoles 289 

Accounts of Species 289 

Smilisca baudini ( Dumeril and Bibron ) 289 

Smilisca cyanosticta (Smith) 303 

Smilisca phaeota ( Cope ) 308 

Smilisca piimu ( Cope) 314 

Smilisca sila New species 318 

Smilisca sordida ( Peters ) 323 

Analysis of Morphological Characters 330 

Osteology 330 

Descriptive Osteology of Smilisca baudini 331 

Developmental Cranial Osteology of Smilisca baudini . . 333 

Comparative Osteology 336 

Musculature 341 

Skin 342 

Structure 342 

Comparative Biochemistry of Troteins 343 

External Morphological Characters 343 

Size and Proportions 343 

Shape of Snout 344 

Hands and Feet 344 

Ontogenetic Changes 344 

Coloration 344 

Metachrosis 345 

Chromosomes 345 

(283) 



284 University of Kansas Publs., Mus. Nat. Hist. 



PAGE 



Natural History 345 

Breeding 345 

Time of Breeding 345 

Breeding Sites 346 

Breeding Behavior 346 

Breeding Call 351 

Eggs 356 

Tadpoles 357 

General Structure 357 

Comparison of Species 357 

Growth and Development 361 

Belmvior 365 

Phylogenetic Relationships 366 

Interspecific Relationships 366 

Evolutionary History 369 

Summary and Conclusions 371 

Literature Cited 372 



INTRODUCTION 

The family Hylidae, as currently recognized, is composed of 
about 34 genera and more than 400 species. Most genera (30) and 
about 350 species live in the American tropics. Hyla and 10 other 
genera inhabit Central America; four of those 10 genera {Gastro- 
theca, Hemiphr actus, Phrynohyos, and Phyllomedusa) are widely 
distributed in South America. Tlie other six genera are either re- 
stricted to Central America or have their greatest differentiation 
there. Plectrohyla and Ptychohyla inhabit streams in the highlands 
of southern Mexico and northern Central America; Diaglena and 
Triprion are casque-headed inhabitants of arid regions in Mexico 
and northern Central America. Anotheca is a tree-hole breeder in 
cloud forests in Middle America. The genus Smilisca is the most 
widespread geographically and diverse ecologically of the Central 
American genera. 

The definition of genera in the family Hylidae is diflBcult owing 
to the vast array of species, most of which are poorly known as 
regards their osteology, colors in life, and modes of life history. 
The genera Diaglena, Triprion, Tetraprion, Osteocephalus, Trachy- 
cephalus, Aparasphenodon, Corythomantis, Hemiphractus, Pterno- 
hyla, and Anotheca have been recognized as distinct from one 
another and from the genus Hyla on the basis of various modifica- 
tions of dermal bones of the cranium. Phyllomedusa is recognized 
on the basis of a vertical pupil and opposable thumb; Plectrohyla 
is characterized by the presence of a bony prepollex and the ab- 
sence of a quadratojugal. Gastrotheca is distinguished from other 
hylids by the presence of a pouch in the back of females. A pair 
of lateral vocal sacs behind the angles of the jaws and the well- 
developed dermal glands were used by Duellman (1956) to dis- 
tinguish Phrynohyos from Hyla. He (1963a) cited the ventrolateral 
glands in breeding males as diagnostic of Ptychohijla. Some species 
groups within the vaguely defined genus Hyla have equally dis- 
tinctive characters. The Hyla septentrional is group is characterized 
by a casque-head, not much different from that in the genus Osteo- 
cephalus (Trueb, MS). Males in the Hijla maxima group have a 
protruding bony prepollex like that characteristically found in 
Plectrohyla. 

Ontogenetic development, osteology, breeding call, behavior, and 
ecology are important in the recognition of species. By utilizing 

(285) 



286 University of Kansas Publs., Mus. Nat. Hist. 

the combination of many morphological and biological factors, the 
genus Smilisca can be defined reasonably well as a natural, phyletic 
assemblage of species. Because the wealth of data pertaining to 
the morphology and biology of Smilisca is lacking for most other 
tree frogs in Middle America it is not possible at present to com- 
pare Smilisca with related groups in more than a general way. 

Smilisca is an excellent example of an Autochthonous Middle 
American genus. As defined by Stuart (1950) the Autochthonous 
Middle American fauna originated from "hanging relicts" left in 
Central America by the ancestral fauna that moved into South 
America and difiPerentiated there at a time when South America 
was isolated from North and Middle America. The genus Smilisca, 
as we define it, consists of six species, all of which occur in Central 
America. One species ranges northward to southern Texas, and 
one extends southward on the Pacific lowlands of South America 
to Ecuador. We consider the genus Smilisca to be composed of 
rather generalized hylids. Consequently, an understanding of the 
systematics and zoogeography of the genus can be expected to be 
of aid in studying more specialized members of the family. 

Acknowledgments 

Examination of many of the specimens used in our study was possible only 
because of the cooperation of the curators of many systematic collections. For 
lending specimens or providing working space in their respective institutions 
we are grateful to Doris M. Cochran, Alice G. C. Grandison, Jean Guibe, Rob- 
ert F. Inger, Giinther Peters, Gerald Raun, William J. Riemer, Jay M. Savage, 
Hobart M. Smith, Wilmer W. Tanner, Charles F. Walker, Ernest E. Williams, 
and Richard G. Zweifel. 

We are indebted to Charles J. Cole and Charles W. Myers for able assist- 
ance in the field. The cooperation of Martin H. Moynihan at Barro Colorado 
Island, Charles M. Keenan of Corozal, Canal Zone, and Robert Hunter of San 
Jose, Costa Rica, is gratefully acknowledged. Jay M. Savage turned over to 
us many Costa Rican specimens and aided greatly in our work in Costa Rica. 
James A. Peters helped us locate sites of collections in Ecuador and Coleman 
J. Goin provided a list of localities for the genus in Colombia. 

We especially thank Charles J. Cole for contributing the information on the 
chromosomes, and Robert R. Patterson for preparing osteological specimens. 
We thank M. J. Fouquette, Jr., who read the section on breeding calls and 
ofiFered constructive criticism. 

Permits for collecting were generously provided by Ing. Rodolfo Hernandez 
Corzo in Mexico, Sr. Jorge A. Ibarra in Guatemala, and Ing. Milton Lopez in 
Costa Rica. This report was made possible by support from the National 
Science Foundation (Grants G-9827 and GB-1441) and the cooperation of the 
Museiun of Natural History at the University of Kansas. Some of the field 
studies were carried out in Panama under the auspices of a grant from the 
National Institutes of Health (NIH GM- 12020) in cooperation with the Gorgas 
Memorial Laboratory in Panama. 



Neotropical Hylid Frogs, GE^fus Smilisca 287 

Materials and Methods 

In our study we examined 4151 preserved frogs, 93 skeletal preparations, 
88 lots of tadpoles and young, and six lots of eggs. We have collected speci- 
mens in the field of all of the species. Observations on behavior and life his- 
tory were begun by the senior author in Mexico in 1956 and completed by us 
in Central America in 1964 and 1965. 

Osteological data were obtained from dried skeletons and cleared and 
stained specimens of all species, plus serial sections of the skull of Smilisca 
baudini. Developmental stages to which tadpoles are assigned are in accord- 
ance with the table of development published by Gosner (1960). Breeding 
calls were recorded in the field on tape using Magnemite and Uher portable 
tape recorders. Audiospectrographs were made by means of a Vibralyzer ( Kay 
Electric Company). External morphological features were measured in the 
manner described by DueUman (1956). In the accounts of the species we 
have attempted to give a complete synonymy. At the end of each species 
account the localities from which specimens were examined are listed alpha- 
betically within each state, province, or department, which in trnm are listed 
alphabetically within each country. The countries are arranged from north 
to south. Abbreviations for museum specimens are listed below: 

AMNH — American Musemn of Natural History 

BMNH — British Museum ( Natural History ) 

BYU — Brigham Young University 

CNHM — Chicago Natural Histor>' Museum 

KU — University of Kansas Museum of Natural History 

MCZ — Museum of Comparative Zoology 

MNHN — Museu National d'Histoire Naturelle, Paris 

UF — University of Florida Collections 

UIMNH — University of Illinois Museum of Natural History 

UMMZ — University of Michigan Musemn of Zoology 

use — University of Southern Cahfomia 

USNM — United States National Museum 

TNHC — Texas Natural History Collection, University of Texas 

ZMB — Zoologisches Museum Berfin 

Genus Smilisca Cope, 1865 

Smilisca Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:194, Oct., 1865 [Type 
species Smilisca daulinia Cope, 1865 = Hyla baudini Dmneril and Bib- 
ron, 1841]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:75, June 17, 
1948. Starrett, Copeia, 4:300, December 30, 1960, Coin, Ann. Car- 
negie Museum, 36:15, July 14, 1961. 

Definition. — Medium to large tree frogs having: (1) broad, well ossified 
skull (consisting of a minimum amount of cartilage and/or secondarily ossified 
cartilage), (2) no dennal co-ossification, (3) quadratojugal and intemasal 
septum present, (4) large ethmoid, (5) M. depressor mandibulae consisting 
of two parts, one arising from dorsal fascia and other from posterior arm of 
squamosal, (6) divided M. adductor mandibulae, (7) paired subgular vocal 
sacs in males, (8) no dermal appendages, (9) pupil horizontally eUiptical 
(10) small amounts of amines and other active substances in skin, (11) 
chromosome number of N = 12 and 2N = 24, ( 12 ) breeding call consisting of 
poorly modulated, explosive notes, and ( 13 ) /3 tooth-rows in tadpoles. 

Composition of genus. — As defined here the genus Smilisca contains six 
recognizable species. An alphabetical fist of the specific and subspecific names 



288 University of Kansas Publs., Mus. Nat. Hist. 

that we consider to be applicable to species of Smilisca recognized herein is 
given below. 

Names proposed Valid names 

Htjla haudini Dumeril and Bibron, 1841 = S. haudini 

Hyla haudini dolomedes Barbour, 1923 = S. phaeota 

Hyla heltrani Taylor, 1942 —s. haudini 

Hyh. gahbi Cope, 1876 = s! sardida 

Hyla lahialis Peters, 1863 rz S. pJiaeota 

Hyla manisorum Taylor, 1954 = s! haudini 

Hyla muricolor Cope, 1862 = S. haudini 

Hyh nigripes Cope, 1876 = S. sordida 

Hyla pansosana Brocchi, 1877 = S. haudini 

Hyla phaeota Cope, 1862 = S. phaeota 

Hyla phaeota cyanosticta Smith, 1953 —S. cyanosticta 

Hyla puma Cope, 1885 = S. puma 

Hyla salvini Boulenger, 1882 = S. sordida 

Hyla sordida Peters, 1863 =5. sordida 

Hyla vanvlietii Baird, 1854 = S. haudini 

Hyla vociferans Baird, 1859 =S. haudini 

Hyla wellmanorum Taylor, 1952 —S. puma 

Distrihution of genus. — Most of lowlands of Mexico and Central America, 
in some places to elevations of nearly 2000 meters, southward from southern 
Sonora and Rio Grande Embayment of Texas, including such continental is- 
lands as Isla Cozumel, Mexico, and Isla Popa and Isla Cebaco, Panama, to 
northern South America, where known from Caribbean coastal regions and 
valleys of Rio Cauca and Rio Magdalena in Colombia, and Pacific slopes of 
Colombia and northern Ecuador. 



Key to Adults 

1. Larger frogs (cT to 76 mm., 9 to 90 mm.) having broad flat heads and a 

dark brown or black postorbital mark encompassing tympanum 2 

Smaller frogs ( cf to 45 mm., 9 to 64 mm.) having narrower heads and lack- 
ing a dark brown or black postorbital mark encompassing tympanum ... .4 

2. Lips barred; flanks cream-colored with bold brown or black mottling in 
groin; posterior surfaces of thighs brown with cream-colored flecks, 

S. haudini, p. 289 
Lips not barred; narrow white labial stripe present; flanks not cream- 
colored with bold brown or black mottling in groin; posterior surfaces of 
thighs variable 3 

3. Flanks and anterior and posterior surfaces of thighs dark brown with 
large pale blue spots on flanks and small blue spots on thighs, 

S. cyanosticta, p. 303 
Flanks cream-colored with fine black venation; posterior surfaces of 
thighs pale brown with or without darker flecks or small cream-colored 
spots S. phaeota, p. 308 

4. Fingers having only vestige of web; diameter of tympanum two-thirds 
that of eye; dorsum pale yellowish tan with pair of broad dark brown 
stripes S. puma, p. 314 

Fingers about one-half webbed; diameter of tympanum about one-half 
that of eye; dorsum variously marked with spots or blotches 5 

5. Snout short, truncate; vocal sacs in breeding males dark gray or brown; 

blue spots on flanks and posterior surfaces of thighs S. sila, p. 318 

Snout long, sloping, rounded; vocal sacs in breeding males white; cream- 
colored or pale blue flecks on flanks and posterior surfaces of thighs, 

S. sordida, p. 323 



Neotropical Hyldd Frogs, Genus Smilisca 289 

Key to Tadpoles 

1. Pond tadpoles; tail about half again as long as body; mouth anteroven- 

tral 2 

Stream tadpoles; tail about twice as long as body; mouth ventral 5 

2. Labial papillae in two rows 3 

Labial papillae in one row 4 

3. First upper tooth row strongly arched medially; third lower tooth row 
much shorter than other rows; dorsal fin deepest at about two-thirds 
length of tail; tail cream-colored with dense gray reticulations, 

S. puma, p. 314 

First upper tooth row not arched medially; third lower tooth row nearly 
as long as others; dorsal fin deepest at about one-third length of tail; tail 
tan with brown flecks and blotches S. baudini, p. 289 

4. Dorsal fin extending onto body S. phacota, p. 308 

Dorsal fin not extending onto body S. cyanostida, p. 303 

5. Mouth completely bordered by two rows of papillae; inner margin of up- 
per beak not forming continuous arch with lateral processes; red or red- 
dish brown markings on tail S. sordida, p. 323 

Median part of upper lip bare; rest of mouth bordered by one row of 
papillae; inner margin of upper beak forming continuous arch with lat- 
eral processes; dark brown markings on tail S. sila, p. 318 

ACCOUNTS OF SPECIES 
Smilisca baudini (Dumeril and Bibron) 

Hyla baudini Dumeril and Bibron, Erpetologie general, 8:564, 1841 [Holo- 
type. — MNHN 4798 from "Mexico;" Baudin collector]. Giinther, Cata- 
logue Batrachia SaUentia in British Museum, p. 105, 1858. Brocchi, 
Mission scientifique au Mexique . . ., pt. 3, sec. 2, fitudes sur les 
batrachiens, p. 29, 1881. Boulenger, Catalogue Batrachia Salientia in 
British Museum, p. 371, Feb. 1, 1882. Werner Abhand. Zool-Bot. 
Gesell. Wien., 46:8, Sept. 30, 1896. Giinther, Biologia Centrali- Ameri- 
cana: Reptilia and Batrachia, p. 270, Sept. 1901. Werner, Abhand. 
Konigl. Akad. Wiss. Munchen, 22:351, 1903. Cole and Barbour, Bull. 
Mus. Comp. Zool., 50(5) :154, Nov. 1906. Gadow, Through southern 
Mexico, p. 76, 1908. Ruthven, Zool. Jahr. 32(4) :310, 1912. Decker, 
Zoologica, 2: 12, Oct., 1915. Stejneger and Barbour, A checklist of North 
American amphibians and reptiles, p. 32, 1917. Noble, Bull. Amer. Mus. 
Nat. Hist., 38(10):341, June 20, 1918. Nieden, Das Tierreich, Am- 
phibia, Anura I, p. 243, June, 1923. Gadow, JoruUo, p. 54, 1930. Dunn 
and Emlen, Proc. Acad. Nat. Sci. Philadelphia, 84:24, March 22, 1932. 
Kellogg, Bull. U. S. Natl. Mus., 160:160, March 31, 1932. Martin, 
Aquarien Berlin, p. 92, 1933. Stuart, Occas. Papers Mus. Zool., Univ. 
Michigan, 292:7, June 29, 1934; Misc. Publ. Mus. Zool. Univ. Michigan, 
29:38, Oct. 1, 1935. Gaige, Carnegie Inst. Washington, 457:293, Feb. 
5, 1936. Gaige, Hartvveg, and Stuart, Occas. Papers Mus. Zool. Univ. 
Michigan, 360:5, Nov. 20, 1937. Smith, Occas. Papers Mus. Zool. Univ. 
Michigan, 388:2, 12, Oct. 31, 1938; Ann. Carnegie Mus., 27:312, March 
14, 1939. Taylor, Copeia, 2:98, July 12, 1939. Hartweg and Oliver, 
Misc. Publ. Mus. Zool. Univ. Michigan, 47:12, July 13, 1940. Schmidt 
and Stuart, Zool. Ser. Field Mus. Nat. Hist., 24(21):238, August 30, 
1941. Schmidt, Zool. Ser. Field Mus. Nat. Hist., 22(8):486, Dec. 30, 

1941. Wright and Wright, Handbook of frogs and toads, Ed. 2, p. 134, 

1942. Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 471:15, May 
17, 1943. Bogert and Oliver, Bull. Amer. Mus. Nat. Hist., 83(6):343, 
March 30, 1945. Taylor and Smith, Proc. U. S. Natl. Mus., 95(3185) :590, 
June 30, 1945. Smith, Ward's Nat. Sci. Bull., 1, p. 3, Sept., 1945. Schmidt 
and Shannon, Fieldiana, Zool. Chicago Nat. Hist. Mus., 31(9) :67, Feb. 



290 University of Kansas Publs., Mus, Nat, Hist. 

20, 1947. Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 69:26, June 
12, 1948. Wright and Wright, Handbook of frogs and toads, Ed. 3, p. 
298, 1949. Stuart, Contr. Lab. Vert. Biol. Univ. Michigan, 45:22, May, 
1950. Mertens, Senckenbergiana, 33:170, June 15, 1952; Abhand. 
Senckenb. Naturf. Gesell., 487:28, Dec. 1, 1952. Schmidt, A checkHst 
of North American amphibians and reptiles, Ed. 6, p. 69, 1953. Stuart 
Contr. Lab. Vert. Biol. Univ. Michigan, 68:46, Nov. 1954. Zweifel and 
Norris, Amer. Midi. Nat, 54(1):232, July 1955. Martin, Amer. Nat, 
89:356, Dec. 1955. Duelhnan, Copeia, 1:49, Feb. 21, 1958. Coin, 
Herpetologica, 14:119, July 23, 1958. Turner, Herpetologica, 14:192, 
Dec. 1, 1958. Conant, A field guide to reptiles and amphibians, p. 284, 
1958. Duellman, Univ. Kansas Publ., Mus. Nat Hist., 13(2):59, Aug. 
16, 1960; Univ. Kansas Publ., Mus. Nat. Hist, 15(1): 46, Dec. 20, 1961. 
Porter, Herpetologica, 18:165, Oct. 17, 1962. 

Hyla vanvlietii Baird, Proc. Acad. Nat. Sci. Philadelphia, 7:61, April 27, 
1854 [Holotype. — USNM 3256 from Brownsville, Cameron County, 
Texas; S. Van Vhet collector]. Baird, United States and Mexican bound- 
ary survey, 2:29, 1859. Smith and Taylor, Univ. Kansas Sci. Bull., 33: 
361, March 20, 1950. Cochran, Bull. U. S. Natl. Mus., 220:60, 1961. 

Hyla vociferans Baird, United States and Mexican bovmdary survey, 2:35 
1859 [nomen nudum]. Diaz de Leon, Indice de los batracios que se 
encuentran en la Repiibhca Mexicana, p. 20, June 1904, 

Hyla muricolor Cope, Proc. Acad. Nat Sci. Philadelphia, 14(9) :359, 1862 
[Holotype. — USNM 25097 from Mirador, Veracruz, Mexico; Charles 
Sartorius collector]. Smith and Taylor, Univ. Kansas Sci. Bull., 33:349, 
March 20, 1950. Cochran, Bull. U. S. Nad. Mus., 220:56, 1961. 

Smilisca daulinia Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:194, Oct. 
1865 [Holotype. — "skeleton in private anatomical museum of Hyrtl, Pro- 
fessor of Anatomy in the University of Vienna"]. Smith and Taylor, Univ. 
Kansas Sci. Bull., 33:347, March 20, 1950. 

Smilisca daudinii [lapsus for haudini]. Cope, Proc. Acad. Nat. Sci. Philadel- 
phia, 23, pt 2:205, 1871. 

Smilisca baudini, Cope, Bull. U. S. Nat. Mus., 1:31, 1875; Jour. Acad. Nat 
Sci. Philadelphia, 8, pt 2:107, 1876; Proc. Amer. Philos. Soc., 18:267, 
August 11, 1879. Yarrow, BuU. U. S, Nat. Mus., 24:176, July 1, 1882. 
Cope, Bull. U. S. Nat Mus., 32:13, 1887; Bull. U. S. Nat Mus., 34:379, 
April 9, 1889. Dickerson, The frog book, p. 151, July, 1906. Smith and 
Taylor, Univ. Kansas Sci. Bull., 33:442, March 20, 1950; Taylor, U. Kan. 
Sc. Bull., 34:802, Feb. 15, 1952; Univ. Kansas Sci. Bull., 35:794, July 1, 
1952. Brattstrom, Herpetologica, 8(3) :59, Nov. 1, 1952. Taylor, U. Kan. 
Sci. Bull., 35:1592, Sept. 10, 1953. Peters, Occas. Papers Mus. Zool, 
Univ. Michigan, 554:7, June 23, 1954. Duellman, Occas. Papers Mus. 
Zool. Univ. Michigan, 560:8, Oct. 22, 1954. Chraphwy and Fugler, 
Herpetologica, 11:122, July 15, 1955. Smith and Van Gelder, Herpeto- 
logica, 11:145, July 15, 1955. Lewis and Johnson, Herpetologica, 11: 
178, Nov. 30, 1955. Martin, Misc. Publ. Mus. Zool. Univ. Michigan, 101: 
53, April 15, 1958. Stuart, Contr. Lab. Vert. Biol. Univ. Michigan, 75:17, 
June, 1958. Minton and Smith, Herpetologica, 17:74, July 11, 1961. Nelson 
and Hoyt, Herpetologica, 17:216, Oct. 9, 1961. Hohnan, Copeia, 2:256, 
July 20, 1962. Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 122:41, 
April 2, 1963. MasUn, Herpetologica, 19:124, July 3, 1963. Holman 
and Birkenholz, Herpetologica, 19:144, July 3, 1963. Duellman, Univ. 
Kansas Publ. Mus. Nat. Hist, 15(5) :228, Oct. 4, 1963. Zweifel, Copeia, 
1:206, March 26, 1964. Duelhnan and Klaas, Copeia, 2:313, June 30, 
1964. Davis and Dixon, Herpetologica, 20:225, January 25, 1965. Neill, 
Bull. Florida State Mus., 9:89, April 9, 1965. 

Hyla pansosana Brocchi, Bull. Soc. Philom., ser. 7, 1:125, 1877 [Holotype. 
— MNHN 6313 from Panzos, Alta Verapaz, Guatemala; M. Bocourt col- 
lector]; Mission scientifique au Mexique . . ., pt. 3, sec. 2, fitudes 
sur les batrachiens, p. 34, 1881. 



Neotropical Hylid Frogs, Genus Smilisca 291 

Hyla baudini baudini, Stejneger and Barbour, A checklist of North American 
amphibians and reptiles, Ed. 3, p. 34, 1933. Wright and Wright, Hand- 
book of frogs and toads, p. 110, 1933. Stejneger and Barbour, A check- 
list of North American amphibians and reptiles, Ed. 4, p. 39, 1939; A 
checklist of North American amphibians and reptiles, Ed. 5, p. 49, 1943. 
Smith and Laufe, Trans. Kansas Acad. Sci., 48(3) :328, Dec. 19, 1945. 
Peters, Nat. Hist. Misc., 143:7, March 28, 1955. 

Hyla beltrani Taylor, Univ. Kansas Sci. Bull. 28(14):306, Nov. 15, 1942 
[Holotype.— UIMNH 25046 (formerly EHT-HMS 29563) from Tapa- 
chula, Chiapas, Mexico; A. Magana collector]. Smith and Taylor, Bull. 
U. S. Natl. Mus. 194:87, June 17, 1948; Univ. Kansas Sci. Bull., 33:326, 
March 20, 1950. Smith, iHinois Biol. Mono., 32:23, May, 1964. 

Smilisca baudini baudini. Smith, Join:. Washington Acad. Sci., 37(11):408, 
Nov. 15, 1947. Smith and Taylor, BuU. U. S. Natl. Mus., 194:75, June 
17, 1948; Univ. Kansas Sci. Bull., 33:347, March 20, 1950. Brown, 
Baylor Univ. Studies, p. 68, 1950. Smith, Smith, and Werler, Texas 
Jour. Sci., 4(2):254, June 30, 1952. Smith and Smith, Anales Inst. Biol., 
22(2):561, Aug. 7, 1952. Smith and Darhng, Herpetologica, 8(3):82, 
Nov. 1, 1952. Davis and Smith, Herpetologica, 8(4): 148, Jan. 30, 1953. 
Neill and Allen, Publ. Res. Div. Ross Allen's Reptile Inst, 2(1):26, Nov. 
10, 1959. Mashn, Univ. Colorado Studies, Biol. Series, 9:4, Feb. 1963. 
Hoknan, Herpetologica, 20:48, April 17, 1964. 

Hijla manisorum Taylor, Univ. Kansas Sci. Bull, 36:630, June 1, 1954 
[Holotype. — KU 34927 from Batan, Limon Province, Costa Rica; Ed- 
ward H. Taylor collector]. Duellman and Berg, Univ. Kansas Publ. Mus. 
Nat. Hist, 15(4): 193, Oct 26, 1962. 

Diagnosis. — Size large ( $ 76 mm., $ 90 mm.); skull noticeably wider than 
long, having small frontoparietal fontanelle (roofed with bone in large indi- 
viduals); postorbital processes long, pointed, curving along posterior border of 
orbit; squamosal large, contacting maxillary; tarsal fold strong, full length of 
tarsus; inner metatarsal tubercle large, high, elliptical; hind limbs relatively 
short, tibia length less than 55 per cent snout-vent length; lips strongly barred 
with brown and creamy tan; flanks pale cream with bold brown or black 
reticulations in groin; posterior surfaces of thighs brown with cream-colored 
flecks; dorsal surfaces of limbs marked with dark brown transverse bands. 
(Foregoing combination of characters distinguishing S. baudini from any other 
species in genus.) 

Description and Variation. — Considerable variation in size, and in certain 
proportions and structural characters was observed; variation in some charac- 
ters seems to show geographic trends, whereas variation in other characters 
apparently is random. Noticeable variation is evident in coloration, but this 
will be discussed later. 

In order to analyze geographic variation in size and proportions, ten adult 
males from each of 14 samples from various locahties throughout the range of 
the species were measured. Snout-vent length, length of the tibia in relation 
to snout-vent length, and relative size of the tympanum to the eye are the only 
measurements and proportions that vary noticeably (Table 1). The largest 
specimens are from southern Sinaloa; individuals from the Atlantic lowlands 
of Alta Verapaz in Guatemala, Honduras, and Costa Rica are somewhat smaller, 
and most specimens from the Pacific lowlands of Central America are slightly 
smaller than those from the Adantic lowlands. The smallest males are from 
the Atlantic lowlands of Mexico, including Tamaulipas, Veracruz, the Yucatan 
Peninsula, and British Honduras. 



292 



University of Kansas Publs., Mus. Nat. Hist. 



Table 1. — Geographic Variation in Size and Proportions in Males of 

Smilisca baudini. ( Means in Parentheses Below Observed Ranges; Data 

Based of 10 Specimens From Each Locality.) 



Locality 



Southern Sinaloa. 



Ocotito, Guerrero. 



Pochutla, Oaxaca. 



San Salvador, El Salvador. 



Managua, Nicaragua 


(62.1) 

52.9-63.6 
(57.3) 




Esparta, Costa Rica 


57.6-66.0 
(61.3) 




Ciudad Victoria, Tamaulipas. . 


50.6-56.9 
(53.7) 


C6rdoba, Veracruz 


53.8-63.4 
(57.5) 




Isla del Carmen, Campeche. . . 


47.3-56.6 
(50.9) 


Chich^n-Itzd, Yucatdn 


49.6-57.1 

(53.8) 


British Honduras 


49,0-59.6 
(54.9) 




Chinajjl, Guatemala 


56.8-67.6 
(63.2) 


Atlantidad, Honduras 


52.5-65.1 
(57.6) 


Lim6n, Costa Rica 


57.7-71.3 

(62.4) 





Snout-vent 
length 



62.3-75.9 

(68.6) 

55.6-64.0 

(58.7) 

56.1-65.1 
(60.2) 

57.0-68.0 



Tibia length/ 
snout-vent 



Tympanum/ 
eye 



43.2-46.7 

(44.9) 

46.1-51.2 

(47.8) 

44.7-49.4 
(47.5) 



84.2-94.4 

(87.8) 

66.7-82.8 
(74.6) 

73.0-84.2 

(77.4) 



42.1-46.1 

(44.9) 


74.6-83.3 

(77.6) 


45.6-49.4 
(47.5) 


73.7-89.7 
(79.4) 


44.6-49.3 

(47.3) 


65.5-83.6 

(75.2) 


44.5-48.7 
(46.6) 


67.2-84.3 
(73.9) 


43.9-48.4 
(45.6) 


66.1-75.9 
(70.0) 


44.7-48.9 
(47.6) 


61.5-72.6 
(65.7) 


45.2-53.4 

(49.5) 


62.7-80.7 
(72.6) 


47.5-50.7 
(49.1) 


67.9-76.8 

(72.2) 


47.0-51.0 
(49.5) 


70.0-82.8 
(73.6) 


49.8-53.6 
(51.5) 


56.1-76.5 
(67.0) 


50.4-.52.3 
(51.2) 


63.9-73.0 

(68.5) 



The ratio of the tibia to the snout-vent length varies from 42.1 to 53.6 in 
the 14 samples analyzed. The average ratio in samples from the Pacific low- 
lands varies from 44.9 in Sinaloa and El Salvador to 47.8 in Guerrero; on the 
Gulf lowlands of Mexico the average ratio varies from 45.6 in Veracruz to 47.6 
on Isla del Carmen, Campeche. Specimens from the Yucatan Peninsula and 
the Caribbean lowlands have relatively longer legs; the variation in average 
ratios ranges from 49.1 in British Honduras to 51.2 in Costa Rica and 51.5 in 
Honduras. 

Specimens from southern Sinaloa are outstanding in the large size of the 



Neotropical Hylid Frogs, Genus Smilisca 293 

tympanum; the tympanum/eye ratio varies from 84.2 to 94.4 (average 87.8). 
In most other samples the variation in average ratios ranges from 72.2 to 79.3, 
but specimens from Veracruz have an average ratio of 70.0; Campeche, 65.7; 
Honduras, 67.0; and Limon, Costa Rica, 68.5. 

No noticeable geograpliic trends in size and proportions are evident. Speci- 
mens from southern Sinaloa are extreme in their large size, relatively short 
tibia, and large tympani, but in size and relative length of the tibia the Sina- 
loan frogs are approached by specimens from such far-removed localities as 
San Salvador, El Salvador, and Chinaja, Guatemala. Frogs from the Carib- 
bean lowlands of Honduras and Costa Rica are relatively large and have rela- 
tively long tibiae and small tympani. 

The inner metatarsal tubercle is large and high and its shape varies. The 
tubercle is most pronounced in specimens from northwestern Mexico, Tamauli- 
pas, and the Pacific lowlands of Central America. Possibly the large tubercle 
is associated with drier habitats, where perhaps the frogs use the tubercles for 
digging. 

The ground color of Smilisca baudini is pale green to brown dorsally and 
white to creamy yellow ventrally. The dorsum is variously marked with dark 
brown or dark olive-green spots or blotches (Pi. 6A). In most specimens a 
dark interorbital bar extends across the head to the lateral edges of the eye- 
lid; usually this bar is connected medially to a large dorsal blotch. There is no 
tendency for the markings on the dorsum to form transverse bands or longi- 
tudinal bars. In specimens from the southern part of the range the dorsal 
dark markings are often fragmented into small spots, especially posteriorly. 
The limbs are marked by dark transverse bands, usually three on the forearm, 
three on the thigh, and three or four on the shank. Transverse bands also 
are present on the tarsi and proximal segments of the fingers and toes. The 
webbing on the hands and feet is pale grayish brown. The loreal region and 
upper lip are pale green or tan; the lip usually is boldly marked with broad 
vertical dark brown bars, especially evident is the bar beneath the eye. A 
dark brown or black mark extends from the tympanum to a point above the 
insertion of the forearm; in some specimens this black mark is narrow or in- 
distinct, but in most individuals it is quite evident. The flanks are pale gray 
to creamy white with brown or black mottling, which sometimes forms reticu- 
lations enclosing white spots. The anterior surfaces of the thighs usually are 
creamy white with brown mottling, whereas the posterior surfaces of the thighs 
usually are brown with small cream colored flecks. A disrinct creamy-white 
anal stripe usually is present. Usually, there are no white stripes on the outer 
edges of the tarsi and foreamis. In breeding males the throat is gray. 

Most variation in coloration does not seem to be correlated with geography. 
The lips are strongly barred in specimens from throughout the range of the 
species, except that in some specimens from southern Nicaragua and Costa 
Rica the lips are pale and in some specimens the vertical bars are indistinct. 
Six specimens from 7.3 kilometers southwest of Matatan, Sinaloa, are distinc- 
tively marked. The dorsum is uniformly grayish green with the only dorsal 
marks being on the tarsi; canthal and post-tympanic dark marks absent. A 
broad white labial stripe is present and interrupted by a single vertical dark 
mark below the eye. A white stripe is present on the outer edge of the foot. 
The flanks and posterior surfaces of the thighs are creamy white, boldly marked 
with black. Two specimens from Alta Verapaz, Guatemala (CNHM 21006 



294 University of Kansas Publs., Mus. Nat. Hist. 

from Coban and UMMZ 90908 from Finca Canihor), are distinctive in having 
many narrow transverse bands on the limbs and fine reticulations on the flanks. 
Two specimens from Limon Province, Costa Rica (KU 34927 from Batan and 
36789 from Suretka), lack a dorsal pattern; instead these specimens are nearly 
uniform brown above and have only a few small dark brown spots on the back 
and lack transverse bands on the limbs. The post-tympanic dark marks and 
dark mottling on the flanks are absent. Specimens lacking the usual dorsal 
markings are known from scattered localities on the Caribbean lowlands from 
Guatemala to Costa Rica. 

The coloration in life is highly variable; much of the apparent variation is 
due to metachrosis, for individuals of Smilisca haudini are capable of under- 
going drastic and rapid change in coloration. When active at night the frogs 
usually are pale bright green with olive-green markings, olive-green with brown 
markings, or pale brown with dark brown markings. The dark markings on 
the back and dorsal surfaces of the limbs are narrowly outlined by black. The 
pale area below the eye and just posterior to the broad suborbital dark bar is 
creamy white, pale green, or ashy gray in life. The presence of this mark is 
an excellent character by which to identify juveniles of the species. The flanks 
are creamy yellow, or yellow with brown or black mottling. In most individ- 
uals the belly is wliite, but in specimens from southern El Peten and northern 
Alta Verapaz, Guatemala, the belly is yellow, especially posteriorly. The iris 
varies from golden bronze to dull bronze with black reticulations, somewhat 
darker ventrally. 

Natural History. — Throughout most of its range Smilisca baudini occurs in 
subhumid habitats; consequently the activity is controlled by the seasonal na- 
ture of the rainfall and usually extends from May or June through September. 
Throughout Mexico and Central America the species is known to call and 
breed in June, July, and August. Several records indicate that the breeding 
season in Central America is more lengthy. Gaige, Hartweg, and Stuart 
(1937:4) noted gravid females collected at El Recreo, Nicaragua, in August 
and September. Schmidt (1941:486) reported calling males in February in 
British Honduras. Stuart (1958:17) stated that tadpoles were found in mid- 
February, juveniles in February and March and half-grown individuals from 
niid-March to mid-May at Tikal, El Peten, Guatemala. Stuart (1961:74) 
reported juveniles from Tikal in July, and that individuals were active at night 
when there had been light rain in the dry season in February and March in 
El Peten, Guatemala. Smilisca baudini seeks day-time retreats in bromeliads, 
elephant-ear plants (Xanthosoma), and beneath bark or in holes in trees. By 
far the most utilized retreat in the dry season in parts of the range is beneath 
the outer sheaths of banana plants. Large numbers of these frogs were found 
in banana plants at Cuautlapan, Veracruz, in March, 1956, in March and De- 
cember, 1959. 

Large breeding congregations of this frog are often foimd at the time of 
the first heavy rains in the wet season. Gadow (1908:76) estimated 45,000 
frogs at one breeding site in Veracruz. In the vicinity of Tehuantepec, Oaxaca, 
large numbers of individuals were found around rain pools and roadside ditches 
in July, 1956, and July, 1958; large concentrations were found near Chinaja, 
Guatemala, in June, 1960, and near Esparta, Costa Rica in July, 1961. Usu- 
ally males call from the ground at the edge of the water or not infrequently 
sit in shallow water, but sometimes males call from bushes and low trees 



Neotropical Hylid Frogs, Genus Smelisca 295 

around the water. Stuart (1935:38) recorded individuals calling and breed- 
ing throughout the day at La Libertad, Guatemala. Smilisca haudini usually 
is absent from breeding congregations of hyhds; frequently S. baudini breeds 
alone in small temporary pools separated from large ponds where numerous 
other species are breeding. In Guerrero and Oaxaca, Mexico, S. haudini breeds 
in the same ponds with Rhinophrtjmis dorsalis, Bufo marmoreus, Engystomops 
pustulosits, and Diaglena reticulata, and in the vicinity of Esparta, Costa Rica, 
S. baudini breeds in ponds with Bufo coccifer, Hyla staufferi, and Phrynohyas 
venulosa. In nearly all instances the breeding sites of S. baudini are shallow, 
temporary pools. 

The breeding call of Smilisca baudini consists of a series of short explosive 
notes. Each note has a duration of 0.09 to 0.13 seconds; two to 15 notes make 
up a call group. Individual call groups are spaced from about 15 seconds to 
several minutes apart. The notes are moderately high-pitched and resemble 
"wonk-wonk-wonk." Little vibration is discernible in the notes, which have 
140 to 195 pulses per second and a dominant frequency of 2400 to 2725 cycles 
per second (Pi. lOA). 

The eggs are laid as a surface film on the water in temporary pools. The 
only membrane enclosing the individual eggs is the vitelline membrane. In 
ten eggs (KU 62154 from San Salvador, El Salvador) the average diameter 
of the embryos in first cleavage is 1.3 mm. and of the vitelline membranes, 
1.5 mm. Hatchling tadpoles have body lengths of 2.6 to 2.7 mm. and total 
lengths of 5.1 to 5.4 mm. The body and caudal musculature is brovim; the 
fins are densely flecked with brown. The gills are long and filamentous. 
Growth and development of tadpoles are summarized in Table 9. 

A typical tadpole in stage 30 of development (KU 60018 from Chinaja, 
Alta Verapaz, Guatemala) has a body length of 8.7mm., a tail length of 
13.6 mm., and a total length of 22.3 mm.; body slightly wider than deep; 
snout rounded dorsally and laterally; eyes widely separated, directed dorso- 
laterally; nostril about midway between eye and tip of snout; mouth antero- 
ventral; spiracle sinistral, located about midway on length of body and slightly 
below midline; anal tube dextral; caudal musculature slender, shghtly curved 
upward distally; dorsal fin extending onto body, deepest at about one-third 
length of tail; depth of dorsal fin slightly more than that of ventral fin at mid- 
length of tail; dorsal part of body dark brown; pale crescent-shaped mark on 
posterior part of body; ventral surfaces transparent vdth scattered brown pig- 
ment ventrolaterally, especially below eye; caudal musculature pale tan with 
a dark brown longitudinal streak on middle of anterior one-third of tail; dor- 
sum of anterior one-third of tail dark brown; brown flecks and blotches on 
rest of caudal musculature, on all of dorsal fin, and on posterior two-thirds of 
ventral fin; iris bronze in life (Fig. 11). Mouth small; median third of upper 
lip bare; rest of mouth bordered by two rows of conical papillae; lateral fold 
present; tooth rows %; two upper rows about equal in length; second row 
broadly interrupted medially, three lower rows complete, first and second equal 
in length, shghtly shorter than upper rows; third lower row shortest; first upper 
row sharply cturved anteriorly in midUne; upper beak moderately deep, form- 
ing a board arch vdth slender lateral processes; lower beak more slender, 
broadly V-shaped; both beaks bearing blunt serrations (Fig. 15A). 

In tadpoles having fully developed mouthparts the tooth-row formula of 
% is invariable, but the coloration is highly variable. The color and pattern 



296 University of Kansas Publs., Mus. Nat. Hist, 

described above is about average. Some tadpoles are much darker, such as 
those from 11 kilometers north of Vista Hermosa, Oaxaca, (KU 87639-44), 
3.5 kilometers east of Yokdzonot, Yucatan (KU 71720), and 4 kilometers west- 
southwest Puerto Juarez, Quintana Roo, Mexico (KU 71721), whereas others, 
notably from 17 kilometers northeast of Juchatengo, Oaxaca, Mexico (KU 
87645), are much paler and lack the dark markings on the caudal musculature. 
The variation in intensity of pigmentation possibly can be correlated with en- 
vironmental conditions, especially the amount of light. In general, tadpoles 
that were found in open, sunlit pools are palhd by comparison with those from 
shaded forest pools. These subjective comparisons were made with preserved 
specimens; detailed comparative data on hving tadpoles are not available. 

The relative length and depth of the tail are variable; in some individuals 
the greatest depth of the tail is about at mid-length of the tail, whereas in 
most specimens the tail is deepest at about one-third its length. The length 
of the tail relative to the total length is usually 58 to 64 per cent in tadpoles in 
stages 29 and 30 of development. In some individuals the tail is about 70 
per cent of the total length. On the basis of the material examined, these 
variations in proportions do not show geographical trends. Probably the pro- 
portions are a reflection of crowding of the tadpoles in the pools where they 
are developing or possibly due to water currents or other environmental factors. 

Stuart (1948:26) described and illustrated the tadpole of Smilisca baudini 
from Finca Chejel, Alta Verapaz, Guatemala. The description and figures 
agree with ovurs, except that the first lower tooth row does not have a sharp 
angle medially in Stuart's figure. He (1948:27) stated that color in tadpoles 
from different locahties probably varies with soil color and turbidity of water. 
Mashn (1963:125) described and illustrated tadpoles of S. baudini from Piste, 
Yucatan, Mexico. These specimens are heavily pigmented like specimens that 
we have examined from the Yucatan Peninsula and from other places in the 
range of the species. Maslin stated that the anal tube is median in the speci- 
mens that he examined; we have not studied Maslin's specimens, but all tad- 
poles of Smilisca that we have examined have a dextral anal tube. 

Newly metamorphosed young have snout-vent lengths of 12.0 to 15.5 mm. 
(average 13.4 in 23 specimens). The largest young are from La Libertad, 
El Peten, Guatemala; these have snout-vent lengths of 14.0 to 15.5 mm. ( aver- 
age 14.5 in five specimens). Young from 11 kilometers north of Vista Hermosa, 
Oaxaca, Mexico, are the smallest and have snout-vent lengths of 12.0 to 12.5 
mm. (average 12.3 in three specimens). Recently metamorphosed young 
usually are dull olive green above and white below; brown transverse bands 
are visible on the hind limbs. The labial markings characteristic of the adults 
are represented only by a creamy white suborbital spot, which is a good diag- 
nostic mark for young of this species. In life the iris is pale gold. 

Remarks: The considerable variation in color and the extensive geographic 
distribution of Sm-ilisca baudini have resulted in the proposal of eight specific 
names for the frogs that we consider to represent one species. Dumeril and 
Bibron (1841:564) proposed the name Hyla baudini for a specimen (MNHN 
4798) from Me.xico. Smith and Taylor (1950:347) restricted the type locahty 
to Cordoba, Veracruz, Mexico, an area where the species occurs in abundance. 
Baird (1854:61) named Htjla vanvlieti from Brownsville, Texas, and (1859:35) 
labelled the figures of Hyla vanvlieti [— Hyla baudini] on plate 38 as Hyla 
vociferans, a nomen nudum. Cope (1862:359) named Hyla muricolor from 
Mirador, Veracruz, Mexico, and (1865:194) used the name Smilisca daulinia 



PLATE 1 



r 





\ 




■<€.- 



^y\ 



.HkJS- 



^if^^f^ 




%,. -. 




.^\ / 



gca^ 



# 



.-e''"' 
•/f' 




^^. 



Dorsal views of skulls of young Smilisca haudini: (A) recently metamorphosed 
young (KU 60026), snout-vent length 12.6 mm. x23; (B) young (KU 85438), 

snout-vent length 32.1 mm. x9. 



PLATE 2 





Skull of adult female Smilisca baudini (KU 68184): (A) Dorsal: 

(B) Ventral. x4.5. 



PLATE 3 




B 





Skull of adult female Smilisca buuciini (KU 68184): (A) Lateral; 
(B) Dorsal view of left mandible; (C) Posterior. x4.5. 



PLATE 4 







r^ 



/lA ,^, 



m 











^i-y 



% **i Et If 











.f :^, 






p ■'^mi 



Palmar views of right hands of Smilisca: (A) S. baudini (KU 

87177); (B) S. phueota (KU 64276); (C) S. cyanosticta (KU 

87199); (D) S. 5orc/i(fa (KU 91761; (E) S. puma (KU 91716), 

and (F) S. sila (KU 77408). X3. 



PLATE 5 



^'- 




HV^^ 






^ 



B 















If': 



■ 'r ff" 






»/ 






/ ^^ 



-W '■:*^. 









w 






•'0 .$ 









T'ss^ %5^ 



#: 




«!v 



->;i--.^it.,_,-i 



X-*: 



Ventral aspect of right feet of Smilisca: (A) S. haudini (KU 87177); (B) 
S. phaeota (KU 64276); (C) S. ajanosticta (KU 87199); (D) S. sorJ/f/a 
(KU 91761); (E) S. puma (KU 91716), and (F) S. s/Za (KU 77408). x3. 



PLATE 6 







Living Smilisca: (A) S. haudmi (UiMMZ 115179) from 1.7 km. W 
Xicotencatal, Tamaulipas, Mexico; (B) S. cyanosticta (UMMZ 118163) 
trom V'olcan San Martin, Veracruz, Mexico; (C) S. phaeota (KU 64282) 
from Barranca del Rio Sarapiqui, Heredia Prov., Costa Rica. All approx. 

nat. size. 



PLATE 7 




B 





Living Srnilisca: (A) S. puma (KU 65307) from 5.9 km. 
W. Puerto Viejo, Heredia Prov., Costa Rica; (B) S. 
sila (KU 77407) from Finca Palosanto, 6 km. WNW El 
Volcan, Chiriqui, Panama; (C) S. sordida (KU 64257) 
from 20 km. WSW San Isidro el General, San Jose 
Prov., Costa Rica. All approx. nat. size. 



PLATE 8 




Fig. 1. Breeding site of Smilisca baudini, 4 km. WNW of Esparta, Puntarenas 

Prov., Costa Rica. 




Fig. 2. Breeding site of Smilisca phaeota, Puerto Viejo, Heredia Prov., Costa 

Rica. 



PLATE 9 




Fig. 1. Breeding site of Smilisca puma, 7.5 km. W of Puerto Viejo, Heredia 

Prov., Costa Rica. 




Fig. 2. Breeding site of Smilisca sordida, Rio La Vieja, 30 km. E of Palmar 

Norte, Puntarenas Prov., Costa Rica. 



PLATE 10 




0.2 03 0.4 

TIME IN SECONDS 



SECTION 



Audiospectrographs and sections of breeding calls of Smilisca: (A) S. haudinl 
(KU Tape No. 74); (B) S. cijanosticta ( KU Tape No. 373); (C) S. phaeota 

(KU Tape No. 79). 



PLATE 11 



8- 



Q 8- 



O 
O 
UJ 

cr 

Q. 

CO 

Ld 

_l 

o 
> 
o 
o 



6- 



4- 



2- 



8- 



6- 



4- 



2- 






» * * 




B 




.iki. 




f*«*** 



#?*^!« 



;?*^- 




I i lW l ii ^mi i i ii tll i jaiiu awfl ll fa il l l Wl l i i 



¥>^^^ 




0.1 



0.2 



0.3 



^ 



0.5 





-as! 




TIME IN SECONDS 



SECTION 



Audiospectrographs and sections of breeding calls of Smilisca: (A) S. puma 
(KU Tape No. 382); (B) S. *i7fl (KU Tape No. 385); (C) S. sordida 

(KU Tape No. 398). 



PLATE 12 




B 




^ 




f/4 



^.-v---^^l 



V:^--;-' ' 



^..^-tJ^-.: 






"'■'M^ 




Lateral views of the heads of Smilisca: (A) S. baudini (KU 87177); (B) 
S. sordida (KU 91765); (C) S. phaeota (KU 64276); (D) S. puma (KU 
91716); (E) S. ctjanosticta (KU 87199); (F) S. «7fl (KU 77408). X 3.2. 



Neotropical Hylid Frogs, Genus Smilisca 



297 



for a skeleton that he employed as the basis for the cranial characters diagnostic 
of the genus Smilisca, as defined by him. Although we cannot be certain, Cope 
apparently inadvertently used daulinia for baudini, just as he used datidinii for 
baudini (1871:205). Brocchi (1877:125) named Hijla pansosana from Panzos, 
Alta Verapaz, Guatemala. 

Aside from the skeleton referred to as Smilisca daulinia by Cope ( 1865: 194), 
we have examined each of the types of the species synonymized with S. 
baudini. All unquestionably are representatives of S. baudini. 

Taylor (1942:306) named Hyla beltrani from Tapachula, Chiapas. This 
specimen (UIMNH 25046) is a small female (snout-vent length, 44 mm.) of 
S. baudini. Taylor (1954:630) named Hyla manisorum from Batan, Limon, 
Costa Rica. The type (KU 34927) is a large female (snout-vent length, 
75.3 mm.) S. baudini. In this specimen and a male from Sixretka, Costa Rica, 
the usual dorsal color pattern is absent, but the distinctive curved supraorbital 
processes, together with other structural features, show that the two specimens 
are S. baudini. 

Hyla baudini dolomedes Barbour (1923:11), as shown by Dunn (1931a: 
413), was based on a specimen of Smilisca phaeota from Rio Esnape, Darien, 
Panama. 

Distribution. — Smilisca baudini inhabits lowlands and foothills usually cov- 
ered by xerophytic vegetation or savannas, but in the southern part of its 
range baudini inhabits tropical evergreen forest. The species ranges through- 
out the Pacific and Atlantic lowlands of Mexico from southern Sonora and the 
Rio Grande embayment of Texas southward to Costa Rica, where on the 
Pacific lowlands the range terminates at the southern limits of the arid tropical 
forest in the vicinity of Esparta; on the Caribbean lowlands the distribution 




Fig. 1. Map showing locality records for Smilisca baudini. 
2—3430 



298 University of Kansas Publs., Mus, Nat. Hist. 

seems to be discontinuous southward to Suretka (Fig. 1). Most localities 
where the species has been collected are at elevations of less than 1000 meters. 
Three localities are notably higher; calling males were found at small tem- 
porary ponds in pine-oak forest at Linda Vista, 2 kilometers northwest of 
Pueblo Nuevo Solistahuacan, Chiapas, elevation 1675 meters, and 10 kilometers 
northwest of Comitan, Chiapas, at an elevation of 1925 meters. Tadpoles and 
metamorphosing young were obtained from a pond in arid scrub forest, 17 
kilometers northeast of Juchatengo, Oaxaca, elevation 1600 meters. Stuart 
(1954:46) recorded the species at elevations up to 1400 meters in tlie south- 
eastern highlands of Guatemala. 

Specimens examined. — 3006, as follows: United States: Texas: Cam- 
eron County, Brownsville, CNHM 5412-3, 6869, UMMZ 54036, USNM 3256. 

Mexico: Campeche: Balchacaj, CNHM 102285, 102288, 102291, 102311, 
UIMNH 30709-22, 30726; Champoton, UMMZ 73172 (2), 73176, 73180; 16 
km. E Champoton, UMMZ 73181; 5 km. S Champoton, KU 71369-75; 9 km. 
S Champoton, KU 71367-8; 10.5 km. S Champoton, KU 71365-6, 71722 (tad- 
poles), 71723 (yg.); 24 km. S Champoton, UMMZ 73177 (2); Chuina, KU 75101- 
3; Ciudad del Carmen, UIMNH 30703-8; Dzibalchen, KU 75413-31; Encama- 
cion, CNHM 102282, 102289, 102294-5, 102300, 102306-8, 102312, 102314, 
102316-7, 102319, 102322, UIMNH 30727-40, 30836-7; 1 km. W Escarcega, 
KU 71391-6; 6 km. W Escarcega, KU 71397-403; 7.5 km. W Escarcega, KU 
71376-89; 14 km. W Escarcega, KU 71390; 13 km. W, 1 km. N Escarcega, 
KU 71404; 3 km. N Hopelchen, KU 75410-11; 2 km. NE Hopelchen, KU 
75412; Matamoras, CNHM 36573; Pital, UIMNH 30741; 1 km. SW Puerto 
Real, Isla del Carmen, KU 71345-64; San Jose Carpizo, UMMZ 99879; Tres 
Brazos, CNHM 102284, UIMNH 30723-5; Tuxoefia Camp, UMMZ 73239. 

Chiapas: Acacoyagua, USNM 114487-92; 2 km. W Acacoyagua, USNM 
114493-4; 5 km. E Arroyo Minas, UIMNH 9533-7; Berriozabal, UMMZ 
119186(7); Chiapa de Corzo, UMMZ 119185 (2); Cintalapa, UIMNH 50077; 
Colonia Soconusco, USNM 114495-9; 5 km. W Colonia Soconusco, UMMZ 
87885 (7); Comitan, UMMZ 94438; 10 km. NW Comitan, KU 57185; El 
Suspiro, UMMZ 118819 (11); Escuintla, UMMZ 88271 (7), 88278, 88327, 
109233; 6 km. NE Escuintla, UMMZ 87856 (26); 3 km. E Finca Juarez, 
UIMNH 9538; Finca Prussia, UMMZ 95167; Honduras, UMMZ 94434-7; 
LaGrada, UMMZ 87862; 21 km. S La Trinitaria, UIMNH 9540-1; 14.4 km. 
SW Las Cruces, KU 64239-44; Palenque, UIMNH 49286, USNM 114473-84; 
2 km. NW Pueblo Nuevo SoUstahuacan, KU 57182-4, UMMZ 119948 (8), 
121514; 1.3 km. N Puerto Madero, KU 57186-9; 4 km. N Puerto Madero, 
KU 57190-1; 8 km. N Puerto Madero, UMMZ 118379 (2); 12 km. N Puerto 
Madero, KU 57192; 17.6 km. N Puerto Madero, UMMZ 118378; Rancho 
Monserrata, UIMNH 9531-2, UMMZ 102266-7; Region Soconusco, UIMNH 
33542-56; San Bartola, UIMNH 9519-30; San Geronimo, UIMNH 30804; 
San Tuanito, USNM 114485-6; San Ricardo, CNHM 102406; Solosuchiapa, 
KU 75432-3; Tapachula, CNHM 102208, 102219, 102239, 102405, UIMNH 
25046, 30802-3; Tonola, AMNH 531, CNHM 102232, 102416, UIMNH 
30805-9, USNM 46760; Tuina, KU 41593 (skeleton); Tuxtla Gutierrez, CNHM 
102231, 102248; 6 km. E Tuxtla Gutierrez, UIMNH 9539; 10 km. E Tuxtla 
Gutierrez, UMMZ 119949. 

Chihuahua: 2.4 km. SW Toquina, KU 47226-7; Riito, KU 47228. 

Coahuila: mountain near Saltillo, UIMNH 30833-4. 

Colima: No specific locality, CNMH 1632; Colima, AMNH 510-11; Haci- 
enda Albarradito, UMMZ 80029 (2); Hacienda del Colomo, AMNH 6208; Los 
Mezcales, UMMZ 80028; Manzanillo, AMNH 6207, 6209; Paso del Rio, 
CNHM 102207, 102229-30, UIMNH 30819-21, UMMZ 110875 (3); Periquillo, 
UMMZ 80025 (3), 80026 (14); 1.6 km. SW Pueblo Juarez, UMMZ 115564; 
Queseria, CNHM 102204, 102216-7, 102224, UIMNH 30816-8, UMMZ 80023 
(7), 80024 (7); Santiago, UMMZ 80027; 7.2 km. SW Tecolapa, UMMZ 115184. 



Neotropical Hylid Frogs, Genus Smilisca 299 

Guerrero: Acahuizotla, UF 1338 (2), 1339-40, UMMZ 119182 (2), 119184; 
3 km. S Acahuitzotla, KU 87183-7; Acapulco, AMNH 55276, UMMZ 121879 
(4), USNM 47909; 3 km. N Acapulco, UMMZ 110127; 8 km. NW Acapulco, 
UF 11203 (7); 27 km. NE Acapulco, UIMNH 26597-610; Agua del Obispo, 
CNHM 102214, 102290, 102293, 102310, 102413, KU 60413, 87180-2, UIMNH 
30764-6; Atovca, KU 87175-8; Buena Vista, CNHM 102279, 102304, 102313, 
102315, UIMNH 30774; Caculutla, KU 87179; 20 km. S Chilpancingo, CNHM 
102242, 102401, 102410-1, 102415; Colonia Buenas Aires, UMMZ 119189; 
El Limoncito, CNHM 102292, 102303, 102321, 102414; El Treinte, CNHM 
102212, 102221, 102237, 102240-1, UIMNH 30783-5, USNM 114508-10; 
Laguna Coyuca, UMMZ 80960 (2); 3 km. N Mazatlan, UIMNH 30777-9; 9 km. 
S Mazatlan, CNHM 102209, 102215, 102234, 102246, UIMNH 30781-2; 
Mexcala, CNHM 102399, 102403, 102409, 106539-40, UIMNH 30775-6; 
Ocotito, KU 60414-23; 5.4 km. N Ocotito, UMMZ 119181 (4); 1.6 km. N 
Organos, UIMNH 30752-63; Palo Blanco, CNHM 102283, 102286, 102305, 
102320, 102404, UIMNH 30767-70; Pie de la Cuesta, AMNH 55275, 59202-5; 
Puerto Marquez, AMNH 59200-1 (13); 5.6 km. S San Andreas de la Cruz, 
KU 87173-4; San Vincente, KU 87172; Zaculapan, UMMZ 119183. 

Hidalgo: Below Tianguistengo, CNHM 102318. 

Jalisco: Atenqueque, KU 91435-6; 5 km. NE Autlan, UIMNH 30810; 5 
km. E Barro de Navidad, UMMZ 110900; Charco Hondo, UMMZ 95247; 
Puerto Vallarta, UIMNH 41346; between La Huerta and Tecomates, KU 
91437; 3 km. SE La Resolana, KU 27619, 27620 (skeleton); 11 km. S, 1.6 km. 
E Yahualica, KU 29039; Zapotilitic, CNHM 102238. 

Michoacan: Aguililla, UMMZ 119179 (5); Apatzingan, CNHM 38766-90, 
KU 69101 (skeleton); 7 km. E Apatzingan, UMMZ 112843; 11 km. E Apat- 
zingan, UMMZ 112841 (3); 27 km. S Apatzingan, KU 37621-3; 1.6 km. N 
Arteaga, UMMZ 119180; Charapendo, UMMZ 112840; Coahuayana, UMMZ 
104458; El Sabino, CNHM 102205-6, 102210-1, 102220, 102228, 102233, 
UIMNH 30822-3; La Placita, UMMZ 104456; La Playa, UMMZ 105163; 30 
km. E Nueva Italia, UMMZ 120255 (2); 4 km. S Nueva Italia, UMMZ 112842; 
Ostula, UMMZ 104457 (4); Salitre de Estopilas, UMMZ 104459; San Jose de la 
Montana, UMMZ 104461 (2); 11 km. S Tumbiscatio, KU 37626; 12 km. S 
Tzitzio, UMMZ 119178. 

Morelos: 3.5 km. W Cuautlixco, KU 87188-90; 1 km. NE Puente de Ixtla, 
KU 60393-4; 20 km. S Puente de Ixtla, CNHM 102400, UIMNH 30832; 
Tequesquitengo, AMNH 52036-9, 

Nayarit: 3 km. S Acaponeta, UMMZ 123030 (4); 56 km, S Esquinapa 
(Sinaloa), KU 73909; Jesiis Maria, AMNH 58239; San Bias, KU 28087, 37624, 
62360-2, USNM 51408; 8.6 km. E San Bias, UMMZ 115185; Tepic, UIMNH 
30812-5; 4 kiTi. E Tuxpan, KU 67786; 11 km. SE Tuxpan, UIMNH 7329-31, 
7335-59. 

Nuevo Leon: Galeana, CNHM 34389; Salto de Cola de Caballo, CNHM 
30628-31, 30632 (40), 30633-7, 34454-67, 

Oaxaca: 11 km. S Candelaria, UIMNH 9515-8; Cerro San Pedro, 24 km. 
SW Tehuantepec, UMMZ 82156; Chachalapa, KU 38199; 8 km. S Chiltepec, 
KU 87191; 12 km. S Chivela, UMMZ 115182; Coyul, USNM 114512; Garza 
Mora, UIMNH 40967-8; Juchatengo, KU 87193; 17 km. NE Juchatengo, KU 
87645 (tadpoles), 87646 (young); Juchitan, USNM 70400; Lagartero, UIMNH 
9514; Matias Romero, AMNH 52143-5; 25 km, N Matias Romero, KU 33822-8; 
7 km. S Matias Romero, UIMNH 42703; Mirador, AMNH 6277, 13832-9, 
13842-55; Mira Leon, 1.6 km. N Huatulco, UIMNH 9503-4; Mixtequillo, 
AMNH 13924; Pochutla, KU 57167-81, UIMNH 9505-13; Quiengola, AMNH 
51817, 52146; Rio del Corte, UIMNH 48677; Rio Mono Blanco, UIMNH 
36831; Rio Sarabia, 5 km. N Sarabia, UMMZ 115180 (4); 2.5 km. N Salina 
Cruz, KU 57165-6; San Antonio, UIMNH 37286; 5 km. NNW San Gabriel 
Mixtepec, KU 87192; San Pedro del Istmo, UIMNH 37197; Santo Domingo, 
USNM 47120-2; 3.7 km. N Sarabia, UMMZ 115181 (3); Tapanatepec, KU 
37793 (skeleton), 37794, UIMNH 9542, UMMZ 115183; between Tapanatepec 



300 University of Kansas Publs., Mus. Nat. Hist. 

and Zanatepec, UIMNH 42704-25; Tecuane, UMMZ 82163 (3); Tehuantepec, 
AMNH 52625, 52639, 53470, UMMZ 82157-8, 82159 (9), 82160 (4), 82161 
(8), 82162 (12), 112844-5, 118703, USNM 10016, 30171-4, 30188; 4.5 km. W 
Tehuantepec, KU 59801-12 (skeletons), 69102-3 (skeletons); 10 km. S Tehuante- 
pec, KU 57163-4; Temazcal, USC 8243 (3); 3 km. S Tolocita, KU 39666-9; 
Tolosa, AMNH 53605; Tuxtepec, UMMZ 122098 (2); 2 km. S Valle Nacional, 
KU 87194-5; 11 km. N Vista Hermosa, KU 87196, 87639-41 (tadpoles), 87642-3 
(young), 87644 (tadpoles); Yetla, KU 87197. 

Puebla: 16 km. SW Mecatepec (Veracruz), UIMNH 3657-8; San Diego, 
AMNH 57714, USNM 114511; Vegas de Suchil, AMNH 57712; Villa Juarez, 
UF 11205. 

Quintana Roo: Coba, CNHM 26937; Esmeralda, UMMZ 113551; 4 km. 
NNE Felipe Carrillo Puerto, KU 71417-8; Pueblo Nuevo X-Can, KU 71405; 
10 km. ENE Pueblo Nuevo X-Can, KU 71406; 4 km. WSW Puerto Juarez, 
KU 71407-11, 71721 (tadpoles); 12 km. W Puerto Juarez, KU 71412-6; San 
Miguel, Isla de Cozumel, UMMZ 78542 (6), 78543 (10), 78544 (2); 3.5 km. N 
San Miguel, Isla de Cozumel, KU 71419-22; 10 km. E San Miguel, Isla de 
Cozumel, UMMZ 78541; Telantunich, CNHM 26950. 

San Luis Potosi: Ciudad Valles, AMNH 57776-81 (12), CNHM 37193, 
102297, KU 23705; 21 km. N Ciudad Valles, UMMZ 118377; 6 km. E Ciudad 
Valles, UF 3524; 24 km. E Ciudad Valles, UF 7340 (2); 5 km. S Ciudad Valles, 
UIMNH 30751; 16 km. S Ciudad Valles, AMNH 52953; 30 km. S Ciudad 
Valles, CNHM 102394, 102402, 102412, UIMNH 30749-50; 63 km. S Ciudad 
Valles, UIMNH 19247-58; Pujal, UMMZ 99872 (2); Rio Axtla, near Axtla, 
AMNH 53211-5, 59516, KU 23706; Tamazunchale, AMNH 52675, CNHM 
39621-2, 102226, 102281, UF 7615 (2), UIMNH 26596, UMMZ 99506 (9), 
118701 (2), USNM 114468; 17 km. N Tamazunchale, UIMNH 3659; 2.4 km. S 
Tamazunchale, AMNH 57743; 17 km. E Tamuin, UF 11202 (2); Xilitla, 
UIMNH 19259-60. 

Sinaloa: 8 km. N. Carrizalejo, KU 78133; 4 km. NE Concordia, KU 73914; 
5 km. SW Concordia, KU 75438-9; 6 km. E Cosala, KU 73910; Costa Rica, 
16 km. S. Cuhacan, UIMNH 34887-9; 51 km. SSE Culiacan, KU 37792; El 
Dorado, KU 60392; 1.6 km. NE El Fuerte, CNHM 71468; Isla Palmito del 
Verde, middle, KU 73916-7; 21 km. NNE Los Mochis, UIMNH 40536-7; 
Matatan, KU 73913; 7.3 km. SW Matatan, KU 78464, 78466-70; Mazatlan, 
AMNH 12562, UMMZ 115197 (3); 57 km. N Mazatlan, UIMNH 38364; 
Plomosas, USNM 47439-40; Presidio, UIMNH 30811, USNM 14082; Rosario, 
KU 73911-2; 5 km. E Rosario, UIMNH 7360-76; 8 km. SSE Rosario, KU 37625; 
5 km. SW San Ignacio, KU 78465; 1.6 km. ENE San Lorenzo, KU 47917-24; 
Teacapan, Isla Palmito del Verde, KU 73915; 9.6 km. NNW Teacapan, KU 
91410; Villa Union, KU 78471; 9 km. NE Villa Union, KU 75434-7; 1 km. 
W Villa Uni6n, AMNH 59284. 

Sonera: Guiracoba, AMNH 51225-38 (25). 

Tabasco: 4 km. NE Comalcalco, AMNH 60313; Teapa, UMMZ 119943; 
5 km. N Teapa, UMMZ 119940, 119944, 122997 (2); 10 km. N Teapa, UMMZ 
119187, 119188 (2); 13 km. N Teapa, UMMZ 119941 (2), 119945 (3), 120254 
(2); 21 km. N Teapa, UMMZ 119942, 119947; 29 km. N Teapa, UMMZ 119946 
(11); Tenosique, USNM 114505-7. 

Tamaulipas: Acuna, UMMZ 99864; 5 km. S Acuna, UMMZ 101180; 13 
km. N Antiguo Morelos, UIMNH 40532-5; 3 km. S Antiguo Morelos, UF 
11204; 3 km. NE Chamal, UMMZ 102867; 1.6 km. E Chamal, UMMZ 110734; 
Ciudad Mante, UMMZ 80957, 80958 (3), 106400 (3); 16 km. N Ciudad Victoria, 
CNHM 102408; 34 km. NE Ciudad Victoria, KU 60395-411; 8.8 km. S Ciudad 
Victoria, UIMNH 19261-3; 11 km. W Ciudad Victoria, UIMNH 30924; 16 
km. W Ciudad Victoria, UIMNH 30825; 3 km. W El Carizo, UMMZ 111279; 
Gomez Farias, UMMZ 110837-8; 8 km. NE Gomez Farias, UMMZ 102265, 
102916 (4), 102917, 104110 (5), 105493, 110836 (2), 111274-7; 8 km. NW 
Gomez Farias, UMMZ 101178 (7), 101179 (3), 101362-3, 101364 (2), 108799 
(2), 110129, 111278, 111280; 8 km. W Gomez Farias, UMMZ 102859 (2); 16 
km. W Gonzales, KU 37795-6; Jimenez, KU 60412; La Clementina, 6 km. 



Neotropical Hylid Frogs, Genus Smilisca 301 

W Forlan, USNM 106244; Limon, UIMNH 30831; Llera, USNM 140137-40; 
3 km. E Llera, UIMNH 16858; 21 km. S Llera, UIMNH 30828-9; 23 km. 
S Llera, UIMNH 30830; 11 km. SW Ocampo, UMMZ 118956; 22 km. W, 5 km. 
S Piedra, KU 37568-71; Rio Sabinas, UMMZ 97976; 5 km. W San Gerardo, 
UMMZ 110733 (2); Santa Barbara, UMMZ 111272-3; Villagran, CNHM 102280, 
102287, 102299, 102309, UIMNH 30826-7; 1.7 km. W Xicotencatl, UMMZ 
115179. 

Veracruz: 1.6 km. NW Acayucan, UMMZ 115189; 28.5 km. SE Alvarado, 
UMMZ 119933; 2.4 km. SSW Amatitlan, UMMZ 115195; Barranca Metlac, 
UIMNH 38365; Boca del Rio, UIMNH 26619-30, UMMZ 74954 (9); 16 km. 
S Boca del Rio, UIMNH 26631; between Boca del Rio and Anton Lizardo, 
UIMNH 42701; Canada, CNHM 102397; Catemaco, UMMZ 118702 (4); 
Ciudad Aleman, UMMZ 119608 (3); Cordoba, CNHM 38665-7, USNM 30410-3; 
5.2 k-m. ESE Cordoba, KU 71423-35, 89924 (skeleton); 7 km. ESE Cordoba, 
UMMZ 115176 (4); Cosamaloapan, UMMZ 115193 (2); Coyame, UIMNH 
36853-6, 38366, UMMZ 111461 (3), 111462-3; 1 km. SE Coyame, UMMZ 
121202 (3); Cuatotolapam, UMMZ 41625-39; Cuautlapan, CNHM 38664, 
70591-600, 102218, 102398, KU 26300, 26302, 26309, 26312-3, 26315-6, 26321, 
26336, 26339, 26347 (skeleton), 26354, 55614-21 (skeletons), UIMNH 11236-67, 
11269-71, 11273, 26611-8, 30792-5, UMMZ 85466 (6), 115173 (25), 115175 
(7), USNM 114433-57; Dos Rios, CNHM 39623; 5 km. ENE El Jobo, KU 
23843, 23845, 23847; 6.2 km. E Encero, UIMNH 30835; Escamilo, CNHM 
102298, UIMNH 30788; 1 km. N Fortin, UF 11201; 4 km. SW Huatusco, 
UMMZ 115177; 1 km. SW Huatusco, UMMZ 12-3119; 10 km. SE Hueyapan, 
UNLMZ 115190; 20 km. S Jesus Carranza, KU 23844, 23846, 27399; 38 km. 
SE Jesus Carranza, KU 23417; Laguna Catemaco, UMMZ 119932 (62); 1.6 
km. N La Laja, UIMNH 3651; La Oaxaquena, AMNH 43930-1; 17 km. E 
Martinez de la Torre, UIMNH 36630-2; 6.2 km. W Martinez de la Torre, 
UIMNH 3652-4; Minatitlan, AMNH 52141-2; Mirador, USNM 25097-8, 
115178; 6 km. S Monte Blanco, UF 11200 (4); 21 km. E Nanchital, UMMZ 
123004; 2 km. S Naranja, UMMZ 115188 (3); 1.6 km. NE Novillero, UMMZ 
115194 (2); 3 km. NE Novillero, UMMZ 115196; 5.2 km. NE Novillero, 
UMMZ 115192 (4); 6 km. NE Novillero, UMMZ 115191; 5 km. N Nuevo 
Colonia, UMMZ 105066; Orizaba, USNM 16563-6; 4 km. NE Orizaba, UMMZ 
120251 (2); Panuco, UMMZ 118922; Paraje Nuevo, UMMZ 85465 (2), 85467 
(35), 85468 (36); Paso del Macho, UIMNH 49281; Paso de Talaya, Jicaltepec, 
USNM 32365, 84420; Perez, CNHM 1686 (5); 20 km. N Piedras Negras, Rio 
Blanco, KU 23708; Plan del Rio, KU 26310, 26333-5, 26345, 26354, UMMZ 
102069, 102070 (5); Potrero, UIMNH 49282-5, UMMZ 88799, 88805, 88806 
(2), USNM 32391-5: Potrero Viejo, CNHM 102296, KU 26301, 26304-5, 
26307-8, 26311, 26317-20, 26323-25, 26326-8 (skeletons), 26329-31, 26332 
(skeleton), 26337-8, 26340-4, 26346, 26348, 26351, 26353, 27400-12, UIMNH 
30800, UMMZ 88800 (2), 88802 (15), 88803 (9), 88804, USNM 114458-67; 5 
km. S Potrero Viejo, KU 26303, 26314, 26322; Puente Nacional, UIMNH 
21783-8; 3 km. N Rinconada, UMMZ 122099 (5); Rio de las Playas, USNM 
118635-6; Rio Seco, UMMZ 88801 (9); Rodriguez Clara, CNHM 102225; San 
Andres Tuxtla, CNHM 102213, 102222, 102227, 102247, UIMNH 30789-91; 
10 km. NW San Andres Tuxtla, UMMZ 119935; 13.4 km. NW San Andres 
Tuxtla, UMMZ 119939 (2); 19.8 km. NW San Andres Tuxtla, UMMZ 119938; 
27.2 km. NW San Andres Tuxtla, UMMZ 119936 (6); 48 km. NW San Andres 
Tuxtla, UMMZ 119937; 4 km. W San Andres Tuxtla, UMMZ 115187; 37.4 
km. S San Andres Tuxtla, UMMZ 119934 (12); 15 km. ESE San Juan de la 
Punta, KU 23707; San Lorenzo, USNM 123508-12; 3 km. SW San Marcias 
KU 23841; 1.5 km. S Santa Rosa, UIMNH 42702; 2 km. S Santiago Tuxtla, 
UMMZ 121201 (4); Sauzel, UMMZ 121239; 14 km. E Suchil, UIMNH 46880; 
15 km. S Tampico (Tamaulipas), UMMZ 103322 (4); 4 km. N Tapalapan, 
UMMZ 115186 (2); Tecolutla, UIMNH 42677-700; 16 km. NW Tehuatlan, 
UIMNH 3660-3; 5 km. S Tehuatlan, KU 23842; Teocelo, KU 26306; Tierra 
Colorado, CNHM 102393, 102395-6, UIMNH 30789-91; Veracruz, AMNH 
6301-4, 59398-402, UIMNH 30801, UMMZ 115174, 122060 (2); 24 km. W 
Veracruz, CNHM 104570-2. 



302 University of Kansas Publs., Mus. Nat. Hist. 

Yucatan: No specific locality, CNHM 548, 49067, USNM 32298; Chlchen- 
Itza, CNHM 20636, 26938-49, 36559-62, UIMNH 30742-6, UMMZ 73173 
(6), 73174 (14), 73175 (14), 73178-9, 76171, 83107 (2), 83108, 83109 (2), 83915 
(30), USNM 72744; 9 km. E Chichen-Itza, KU 71438-9; 12 km. E Chichen- 
Itza, KU 71440; Merida, CNHM 40659-66, UIMNH 30747-8, UMMZ 73182; 
6 km. S Merida, KU 75194; 8.8 km. SE Ticul, UMMZ 114296; Valladolid, 
CNHM 26934-6; Xcalah-op, CNHM 53906-14; 3.5 km. E Yokdzonot, KU 
71441-3, 71720 (tadpoles). 

British Honduras: Belize, CNHM 4153, 4384-5, 4387, UMMZ 75310, 
USNM 26065; Bokowina, CNHM 49064-5; Cocquercote, UMMZ 75331 (2); 
Cohvme Ridge, UMMZ 80738 (15); Double Falls, CNHM 49066; El Cayo, 
UMMZ 75311; 6 km. S El Cayo, MCZ 37856; Gallon Jug, MCZ 37848-55; 
Manatee, CNHM 4264-7; Mountain Pine Ridge, MCZ 37857-8; San Augustin, 
UMMZ 80739; San Pedro, Columbia, MCZ 37860-2; Valentin, UMMZ 80735 
(4), 80736 (2), 80737 (2); 5 km. S Waha Loaf Creek, MCZ 37859. 

Guatemala: Alta Verapaz: 5.1 km. NE Campur, KU 68464 (tadpoles), 
67465 (young); 28.3 km. NE Campur, KU 64203-22, 68183-4 (skeletons); 
Charna, MCZ 15792-3, UMMZ 90895 (7), 90896 (5), 90897 (29), 90898 (12), 
90899; Chinaja, KU 55939-41, 57193-8, 60018-20 (tadpoles), 60021 (eggs), 
60022 (tadpoles); Coban, CNHM 21006; Cubilquitz, UMMZ 90902 (10); Finca 
Canihor, UMMZ 90908; Finca Chicoyou, KU 57246-8, 60026 (young), 64202, 
68466-7 (tadpoles); Finca Los Alpes, KU 64197-201, 68463 (tadpoles); Finca 
Los Pinales, UMMZ 90903 (2); Finca Tinajas, BYU 16031; Finca Volcan, 
UMMZ 90905 (4), 90906-7; Panzos, MNHN 6313, UMMZ 90904; Samac, 
UMMZ 90900; Samanzana, UMMZ 90901 (6). 

Baja Verapaz: Chejel, UMMZ 90909 (7), 90910 (3); San Geronimo, UMMZ 
84076 (16). 

Chiquimula: 1.6 km. SE Chiquimula, UMMZ 98112; Esquipulas, UMMZ 
106793 (28). 

El Peten: 20 km. NNW Chinaja (Alta Verapaz), KU 57199-240; Flores, 
UMMZ 117985; La Libertad, KU 60024 (young), UMMZ 75313-20, 75323 
(2), 75324 (7), 75325 (13), 75326 (2), 75327 (11), 75328 (12), 75329 (2); 3 km. 
SE La Libertad, KU 57243-4; 13 km. S La Libertad, MCZ 21458 (2); Pacomon, 
USNM 71334; Piedras Negras, USNM 114469-71; Poptun, UMMZ 120475; 
Poza de la Jicotea, USNM 114672; Ramate-Yaxha trail, UMMZ 75321; Rio de 
la Pasion between Sayaxche and Subin, KU 57151; Rio San Roman, 16 km. 
NNW Chinaja (Alta Verapaz), KU 55942-6; Sacluc, USNM 25131; Sayaxche, 
KU 57144-5; Tikal, UMMZ 117983 (7), 117984 (5), 117993 (5), 120474 (5); 
Toocog, KU 57241-2, 60023 (young), 60025 (young); Uaxactun, UMMZ 
70401-3; Yaxha, UMMZ 75322; 19 km. E Yaxha, UMMZ 75330 (4). 

El Quiche: Finca Tesoro, UMMZ 89166 (3), 90549 (tadpoles). 

Escuintla: Rio Guacalate, Masagua, USNM 125239; Tiquisate, UMMZ 
98262 (7). 

Guatemala: 16 km. NE Guatemala, KU 43545-53. 

Huehuetenango: Finca San Rafael, 16 km. SE Barillas, CNHM 40912-6; 
45 km. WNW Huehuetenango, KU 64223-4; Jacaltenango, UMMZ 120080 
(6), 120081 (14), 120082 (13). 

Izabal: 2 km. SW Puerto Matias de Galvez, KU 60027-8 (tadpoles); Quiri- 
gua, CNHM 20587, UMMZ 70060. 

Jalapa: Jalapa, UMMZ 98109, 106792 (11). 

Jutiapa: Finca La Trinidad, UMMZ 107728 (10); Jutiapa, UMMZ 106789; 
1.6 km. SE Mongoy, KU 43069; Santa Catarina Mita, UMMZ 106790. 

Progreso: Finca Los Leones, UMMZ 106791. 
Quetzaltenango: Coatepeque, AMNH 62204. 

Retalhueleu: Casa Blanca, UMMZ 107725 (18); Champerico, UMMZ 
107726 (3). 

San Marcos: TaUsman Bridge, USNM 128056-7. 



Neotropical Hylid Frogs, Genus Smilisca 303 

Santa Rosa: Finca La Guardiana, UMMZ 107727 (6); Finca La Gloria, 
U\L\IZ 107724 (6); 1.6 km. WSW El Molino, KU 43065-8. 

El Salvador: La Libertad: 16 km. NW Santa Tecla, KU 43542-4. 
Morazan: Divisadero, USNM 73284. San Salvador: San Salvador, CNHM 
65087-99, KU 61955-88, 62138-9 (skeletons), 62154 (eggs), 62155-60 (tadpoles), 
68462 (tadpoles), UMMZ 117586 (3), 118380 (3), USNM 140278. 

Honduras: State unknown: Guaimas, UMMZ 58391. Atlantidad: Isla de 
Roatan, CNHM 34551-4; La Ceiba, USNM 64985, 117589-91; Lancetilla, 
MCZ 16207-11; Tela, MCZ 15774-5, 28080, UMMZ 58418, USNM 82173-4. 
Choluteca: 1.5 km. NW Choluteca, KU 64228-32; 10 km. NW Choluteca, KU 
64233; 10 km. E Choluteca, KU 64226-7; 12 km. E Choluteca, KU 64225; 5 
km. S Choluteca, USC 2700 (2). Colon: Bambii, UF 320; Belfate, AMNH 
45692-5; Patuca, USNM 20261. Comayagua: La Mision, 3.5 leagues N 
Siguatepeque, MCZ 26424-5. Copan: Copan, UMMZ 83026 (2). Cortes: 
Cofradia, AMNH 45345-6; Hacienda Santa Ana, CNHM 4724-31; Lago de 
Yojoa, MCZ 26410-1; Rio Lindo, AMNH 54972. El Paraiso: El Volcan, MCZ 
26436. Francisco Morazan: Tegucigalapa, BYU 18301-4, 18837-41, MCZ 
26395-7, USNM 60499. Gracias A Dios: Rio Segovia, MCZ 24543. Santa 
Barbara: Santa Barbara, USNM 128062-5. 

Nicaragua: Chinandega: 4 km. N, 2 km. W Chichigalpa, KU 85385; 
Chinandega, MCZ 2632; Rio Tama, USNM 40022; San Antonio, KU 84944-9 
(skeletons), 85386-403. Chontales: 1 fan. NE Acoyapa, KU 64234. Esteli: 
Finca Daraili, 5 km. N, 15 km. E Condega, KU 85404-8; Finca Venecia, 7 
km. N, 16 km. E Condega, KU 85409. Leon: 1.6 km. ENE Poneloya, KU 
43084-5. Managua: Managua, USNM 79989-90; 8 km. NW Managua, KU 
43094-110; 20 km. NE Managua, KU 85412; 21 km. NE Managua, KU 
85413-4; 5 km. SW Managua, KU 43086-93; 2 km. N Sabana Grande, KU 
85411; 3 km. N Sabana Grande, KU 43070-8; 20 km. S, 0.5 km. W Tipitapa, 
KU 85410. Matagalpa: Guasquahe, UMMZ 116493; Matagalpa, UMMZ 
116492; 19 km. N Matagalpa, UMMZ 116494. Rio San Juan: Greytown, 
USNM 19585-6, 19767-8. Rivas: Javillo, UMMZ 123001; Moyogalpa, Isla 
Ometepe, KU 85428-37, 87706 (tadpoles); Pefias Blancas, KU 85417; Rio 
Javillo, 3 km. N, 4 km. W Sapoa, KU 85418-20, 85438 (skeleton); 13.1 km. 
SE Rivas, KU 85415; 14.8 km. SE Rivas, KU 85421-3; 11 km. S, 3 km. E 
Rivas, KU 85416; 16 km. S Rivas, MCZ 29009-10; 7.7 km. NE San Juan del 
Sur, KU 85426-7; 16.5 km. NE San Juan del Sur, KU 85424-5, 87705 (young); 
5 km. SE San Pablo, KU 43079-83. Zelaya: Cooley, AMNM 7063-8, 8019-20, 
8022, 8034-5; Cukra, AMNH 8016-7; Musahuas, Rio Huaspuc, AMNH 58428- 
31; 11 km. NW Rama, Rio Siquia, UMMZ 79708, 79709 (5), 79710 (2); Rio 
Escondido, USNM 19766, 20701; Rio Siquia at Rio Mico, UMMZ 79707 (10); 
Sioux Plantation, AMNH 7058-61, 8023-33. 

Costa Rica: Alajuela: Los Chiles, AMNH 54639; Orotina, MCZ 7960-1; 
San Carlos, USNM 29991. Guanacaste: La Cruz, USC 8232 (3); 4.3 km. NE 
La Cruz, UMMZ 123002; 18.4 km. S La Cruz, USC 8136; 23.5 km. S La Cruz, 
USC 8094 (4); 3 km. W La Cruz, USC 8233 (4); 2 km. NE Las Caiias, KU 
64235-7; Las Huecas, UMMZ 71212-3; Liberia, KU 36787, USC 8161; 11.5 
km. N Liberia, USC 8149; 13 km. N Liberia, USC 8139; 22.4 km. N Liberia, 
USC 8126; 8 km. NNW Liberia, KU 64238; 8.6 km. ESE Playa del Coco, 
USC 8137; 21.8 km. ESE Playa del Coco, USC 8138; Rio Piedra, 1.6 km. W 
Bagaces, USC 7027; Rio Bebedero, 5 km. S Bebedero, KU 64158; 5 km. NE 
Tilaran, KU 36782-6. Heredia: 13 km. SW Puerto Viejo, KU 64142-6. 
Limon: Batan, KU 34927; Guacimo, USC 621; Pandora, USC 505 (3); Suretka, 
KU 36788-9; Tortugero, UF 7697, 10540-2. Puntarenas: Barranca, CNHM 
35254-6; 15 km. WNW Barranca, KU 6415.5-7, UMMZ 118381; 18 km. WNW 
Barranca, UMMZ 118382 (4); 4 km. WNW Esparta, KU 64159-96, 68178-82 
(skeletons); 19 km. NW Esparta, KU 64147-54. 

Smilisca cyanosticta (Smith), new combination 

Hijla phaeota, Taylor, Univ. Kansas Sci. Bull., 28(5) :80, May 15, 1942. 
Taylor and Smith, Proc. U. S. Natl. Mus., 95(3185):589, June 30, 1945. 



304 University of Kansas Publs., Mus. Nat. Hist. 

Hyla phaeota cyanostica Smith, Herpetologica, 8:150, Jan, 30, 1953 [Holo- 
type. — USNM 111147 from Piedras Negras, El Peten, Guatemala; Hobart 
M. Smith collector]. Shannon and Werler, Trans. Kansas Acad. Sci., 
58:386, Sept. 24, 1955. Poglayen and Smith, Herpetologica, 14:11, April 
25, 1958. Cochran, Bull. U. S. Natl. Mus., 220:57, 1961. Smith, Illinois 
Biol. Mono., 32:25, May, 1964. 

Smilisca phaeota cyanosticta, Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 
122:42, April 2, 1963. Duellman, Univ. Kansas Publ. Mus. Nat. Hist, 
15(5) :229, Oct. 4, 1963. 

Diagnosis. — Size moderately large ( $ 56.0 mm., $ 70.0 mm.); skull as 
long as wide; frontoparietal fontanelle large; narrow supraorbital flanges hav- 
ing irregular margins anteriorly; large squamosal not in contact vidth maxillary; 
tarsal fold moderately wide, full length of tarsus; inner metatarsal tubercle 
moderately large, low, flat, elliptical; hind limbs relatively long; tibia usually 
more than 52 per cent of snout-vent length; labial stripe silvery- white; lips 
lacking vertical bars; loreal region pale green; pale bronze-colored stripe from 
nostril along edge of eyelid to point above tympanum narrow, bordered below 
by narrow dark brown stripe from nostrU to eye, and broad dark brown 
postorbital mark encompassing tympanum and terminating above insertion of 
arm; flanks, dark brown with large pale blue spots; anterior and posterior 
surfaces of thighs dark brown with small pale blue spots on thighs. ( Foregoing 
combination of characters distinguishing S. cyanosticta from any other species 
in genus.) 

Description and Variation. — The largest males are from Piedras Negras, El 
Peten, Guatemala, and they average 52.5 mm. in snout-vent length whereas 
males from Los Tuxtlas, Veracruz, average 50.6 mm. and those from northern 
Oaxaca 50.3 mm. The smallest breeding male has a snout-vent length of 44.6 
mm. The average ratio of tibia length to snout-vent length is 54.8 per cent 
in males from Piedras Negras, and 56.4 and 56.3 per cent in males from Los 
Tuxtlas and Oaxaca, respectively. The only other character showing noticeable 
geographic variation is the size of the tympanum. The average ratio of the 
diameter of the tympanum to the diameter of the eye is 76.3 per cent in males 
from Piedras Negras, 71.8 from Oaxaca, and 66.9 from Los Tuxtlas. 

The dorsal ground color of Smilisca cyanosticta is pale green to tan and 
the venter is creamy-white. The dorsum is variously marked with dark olive- 
green or dark brown spots or blotches (Pi. 6B). An interorbital dark bar 
usually is present. The most extensive dark area is a V-shaped mark in the 
occipital region with the anterior branches not reaching the eyelids; this mark 
is continuous, by means of a narrow mid-dorsal mark, with an inverted V- 
shaped mark in the sacral region. In many specimens this dorsal marking is 
interrupted, resulting in irregular spots. In some specimens the dorsum is 
nearly uniform pale green or tan with a few small, dark spots. The hind limbs 
are marked by dark transverse bands, usually three or four each on the thigh 
and shank, and two or three on the tarsus. The webbing on the feet is brown. 
The loreal region is pale green, bordered above by a narrow, dark brown 
canthal stripe extending from the nostril to the orbit, which is bordered above 
by a narrow, bronze-colored stripe extending from the nostrfl along the edge 
of the eyelid to a point above the tympanum. The upper lip is white. A 
broad dark brown mark extends posteriorly from the orbit and encompasses the 
tympanum to a point above the insertion of the forelimb. The flanks are dark 
brown with many pale blue, rounded spots, giving the impression of a pale 



Neotropical Hylid Frogs, Genus Smilisca 305 

blue ground color with dark browTi mottling enclosing spots. The anterior 
and posterior surfaces of the thighs are dark bro\vn with many small pale 
blue spots. The inner surfaces of the shank and tarsus are colored like the 
posterior surfaces of the thighs. Pale blue spots are usually present on the 
proximal segments of the second and third toes. A distinct white stripe is 
present on the outer edge of the tarsus and fifth toe; on the tarsus the white 
stripe is bordered below by dark brown. A white stripe also is present on the 
outer edge of the forearm and fourth finger. The anal region is dark brown, 
bordered above by a narrow transverse white stripe. The throat in breeding 
males is dark, grayish brown with white flecks. 

No geographic variation in the dorsal coloration is evident. Specimens from 
the eastern part of the range (Piedras Negras and Chinaja, Guatemala) have 
bold, dark reticulations on the flanks enclosing large pale blue or pale green 
spots, which fade to tan in preservative. Specimens from Oaxaca and Veracruz 
characteristically have finer dark reticulations on the flanks enclosing smaller 
blue spots; in many of these specimens the ventrolateral spots are smallest and 
are white. 

All living adults are easily recognized by the presence of pale, usually blue, 
spots on the flanks and tliighs. Individuals under cover by day have a tan 
dorsum with dark brown markings. A hiding individual at Chinaja, Alta 
Verapaz, Quatemala (KU 55936), had a pale tan dorsum when found; later 
the dorsal color changed to chocolate brown. A pale green patch was present 
below the eye; the spots on the posterior surfaces of the thighs were pale blue, 
and those on the flanks were yellowish green. A calling male obtained 10 
kilometers north-northwest of Chinaja (KU 55934) was reddish brown when 
found at night; later the dorsal color changed to pale tan. A green patch be- 
low the eye was persistent. Usually these frogs are green at night. The colora- 
tion of an adult male (KU 87201) from 11 kilometers north of Vista Hermosa, 
Oaxaca, Mexico, was typical: "When calling dorsum pale green; later changed 
to dull olive-green. Flanks dark brown with pale blue spots in axilla and 
groin and bluish white flecks on mid-flank. Anterior and posterior surfaces of 
thighs, inner smfaces of shanks, and median dorsal surfaces of tarsi dark brown 
with blue spots. Canthal and postorbital stripes dark chocolate brown; labial 
stripe creamy white. Forearm, tarsal, and anal stripes pale cream-color. 
Throat dark brown with yellow flecks; belly and ventral surfaces of limbs 
creamy bufi^; webs pinkish tan; iris deep bronze, brown below pupil." (Duell- 
man, field notes, June 24, 1964. ) 

Some individuals have both green and brown coloration in life. An indi- 
vidual obtained at night on the south slope of Volcan San Martin, Veracruz, 
Mexico, had a pale tan dorsum changing peripherally to pale green. The 
dorsal markings were dark brown and dark olive-green. 

In contrast to the color-changes noted above, the labial region below tlie 
eye is always pale green, and pale spots are always present on the flanks and 
thighs in adults. The iris is invariably golden or bronze above and darker, 
usually brown, below. Minute black flecks are present on the iris, and in some 
individuals these flecks are so numerous that the eye appears gray. 

Recently metamorphosed young have pale tan flanks, and the posterior sur- 
faces of the thighs are orange-yellow; pale spots are absent. A juvenile (KU 
55935) from Chinaja, Alta Verapaz, Guatemala, having a snout-vent length 
of 35.0 mm. was pale yellowish tan above with oHve-green markings; the 
flanks were dark brown with pale blue spots, and the anterior and posterior 



306 University of Kansas Publs., Mus. Nat. Hist. 

surfaces of the thighs were uniform bright tomato red. A juvenile (UMMZ 
121298), 18.6 mm. in snout-vent length, from the southeast slope of Volcan 
San Martin, Veracruz, Mexico, had pale tan flanks lacking blue spots, but had 
red thighs. Apparently the ontogenetic changes in coloration proceed as fol- 
lows: ( 1 ) flanks pale tan and thighs orange-yellow, both lacking spots, ( 2 ) 
flanks pale tan and thighs red, both lacking spots, (3) flanks dark brown with 
blue spots and thighs red, lacking spots, and (4) flanks and tliighs dark brown, 
both having pale blue spots. 

Natural History. — Smilisca cijanosticta inhabits humid tropical forest and 
cloud forest from the lowlands to elevations of about 1200 meters in Los Tuxtlas 
and to about 900 meters in northern Oaxaca. In these moist environments 
the frogs apparently are active tliroughout the year. Active individuals have 
been obtained in January, July, and August in Los Tuxtlas, Veracruz, in June 
and July in northern Oaxaca, in February and March at Cliinaja, Guatemala, 
and Taylor and Smith reported (1945:589) activity in May at Piedras Negras, 
Guatemala. Calling males were observed as follows; in a rain barrel 11 kilome- 
ters north of Vista Hennosa, Oaxaca, Mexico, on June 23-28, 1964; in a quiet 
pool in a stream 8 kilometers south of Yetla, Oaxaca, Mexico, in July, 1963 
(Dale L. Hoyt, personal communication); in and near springs flowing into a 
stream at Dos Amates, Veracruz, Mexico, on August 4, 1959 (Douglas Robin- 
son, personal communication); and in a water-filled depression in a log 10 
kilometers west-northwest of Chinaja, Guatemala, on March 13, 1960. Taylor 
and Smith (1945:589) reported that individuals were found at night on the 
ground at the edge of temporary pools at Piedras Negras, Guatemala, on May 
28-29, 1939. A clasping pair was found on a rock at the edge of a small 
stream on the south slope of Volcan San Martin, Veracruz, Mexico, on July 11, 
1959 (Douglas Robinson, personal communication). 

Only one individual has been observed in a tree at night. In the daytime, 
individuals were found in elephant ear plants (Xantliosoma) at Chinaja, 
Guatemala. 

The breeding call consists of one or two moderately short notes that are 
lower pitched than those of S. batidini, but higher pitched than those of S. 
phaeota. Each note has a duration of 0.25 to 0.45 seconds and is repeated at 
intervals of one-half minute to several minutes. Each note is a vibrant 
"waunk," having 110 to 180 pulses per second and dominant frequency of 
1600 to 2100 cycles per second (PL lOB). 

Apparently the eggs are deposited as loose clumps in the water. In eggs 
in the yolk plug stage of development, the diameter of the embryo is about 
2.3 mm.; that of the outer envelope is 4.0 mm. Hatchling tadpoles have total 
lengths of 5.8 to 6.5 mm. and body lengths of 2.8 to 3.1 mm. The external 
gills are moderately long, slender, and filamentous; the yolk sac is stfll moder- 
ately large. The body and anterior part of the caudal musculature are dark 
brown; posteriorly the caudal musculature is pale brown. The caudal fins are 
creamy tan. The oral discs are large and ovoid. The growth of the tadpole 
is summarized in Table 10. 

A typical tadpole in stage 30 of development (KU 87652 from 11 km. 
N Vista Hermosa, Oaxaca, Mexico) can be described as follows: 

Body length 9.5 mm.; tail length 15.5 mm.; total length 25.0 mm.; body 
slightly wider than deep; snout rounded laterally, broadly ovoid dorsally; eyes 
widely separated, directed dorsolaterally; nostril about midway between eye 



Neotropical Hylid Frogs, Genus Smilisca 307 

and tip of snout; mouth anteroventral; spiracle sinistral, slightly posterior to 
midpoint of body and slightly below midline; anal tube dextral; caudal 
musculature slender, barely curved upward distally; dorsal fin not extending 
onto body, depth of dorsal fin shghtly more than that of ventral fin on mid- 
length of tail; dorsal part of body dark brown; ventral surfaces transparent, 
lacking pigment; posterior edge of body pale cream-color; caudal musculature 
creamy white with interconnected brown spots; caudal fins transparent with 
small brown blotches on dorsal fin and posterior half of ventral fin; iris coppery 
bronze in life (Fig. 12). Mouth small, median part of upper hp bare; rest 
of mouth bordered by single row of bluntly rounded papillae; lateral fold 
present; tooth rows %; all tooth-rows approximately equal in length; second 
upper row broadly interrupted medially; other rows complete; upper beak 
moderately deep, forming broad arch with slender lateral processes; lower beak 
slender, broadly V-shaped; both beaks finely serrate ( Fig. 15C ) . 

All tadpoles having fully developed mouth parts have % tooth rows. Little 
variation is noticeable in coloration. In many specimens the posterior edge 
of the body is dark brown instead of pale cream color. Mottling is rather 
dense on the caudal fins in all specimens; in some individuals pigment is con- 
centrated along the anterior one-third of the lateral groove. In life the body 
is dark brown with greenish gold flecks ventraUy; the caudal musculature is 
gray. 

In each of two recently metamorphosed young the snout-vent length is 14.0 
mm. Coloration of young in hfe (KU 87653 from II km. N Vista Hermosa, 
Oaxaca, Mexico): "Dorsum pale tan with dark broNvn markings. Thighs 
orange-yellow; labial stripe white; iris bronze" (Duellman, field notes, July 10, 
1964.) 

Remarks. — Smith (1953:150) named cyanosticta as a subspecies of Hyla 
phaeota. The difi^erences in cranial characters and certain external characters 
between phaeota and cyanosticta indicate that they are distinct species. Fur- 
thermore, a gap of about 350 kilometers exists between the known geographic 
ranges of the two kinds. 

Distribution. — Smilisca cyanostica inhabits wet forests on the Atlantic slope 
of southern Mexico and northern Central America from northern Oaxaca and 
southern Veracruz through northern Chiapas in Mexico and into El Peten and 
northern Alta Verapaz in Guatemala (Fig. 2), Apparently the range is dis- 
continuous, for in southern Mexico the species is found in cloud forest at 
elevations of 830 to 900 meters on the northern slopes of the Sierra de Juarez. 
In the Sierra de Los Tux-tlas in southern Veracruz the species is found in wet 
forest at elevations of 300 to 1200 meters, but is absent in the intervening 
lowlands characterized by drier forest. In the west forests of northern Alta 
Verapaz and El Peten, Guatemala, the species is found at low elevations. 

Specimens examined. — 78, as follows: Mexico: Chiapas: Monte Libano, 
MCZ 28271-9; 8 km. N San Fernando, 24 km. NE Tuxtla Gutierrez, UIMNH 
41588. Oaxaca: 11 km. N Vista Hermosa, KU 84918-20 (skeletons), 87198-212, 
87647 (eggs), 87648-52 (tadpoles), 87653 (young), UIMNH 57199-201; 8 km. 
S Yetla, KU 87213, UMMZ 124838 (8). Veracruz: Coyame, UMMZ 111459- 
60; between Coyame and Tebanco, UMMZ 121198; Dos Amates, UMMZ 
121297; between Laguna de Catemaco and Volcan San Martin, UMMZ 
121200; Volcan San Martin, UIMNH 35403-4, 35408-12, UMMZ 118163; SE 
slope Volcan San Martin, UMMZ 121199, 121295 (2), 121296, 121298. 

Guatemala: Alta Verapaz: Chinaja, KU 55935-7, 55938 (skeleton). El 



308 



University of Kansas Publs., Mus. Nat. Hist. 



Peten: 10 km. NNW Chinaja (Alta Verapaz), KU 55934; Piedras Negras, 
CNHM 99006-7, 99227, UIMNH 28853, USNM 111139-41, 111143-7; 8 km. 
S Piedras Negras, CNHM 99008; Semicoch, USNM 35907. 



= zv 




Fig, 2. Map showing locality records for Siiiilisca cyanosticta (triangles) 

and Smilisca phaeota (circles). 



Smilisca phaeota (Cope) 

Hyla phaeota Cope, Proc. Acad. Nat. Sci. Philadelphia, 14 (9):358, 1862 
[Holotype. — USNM 4347 from Turbo, Colombia; J. Cassin collector]. 
Boulenger, Catalogue Batrachia Salientia in British Museum, p. 402, 
Feb. 1, 1882. Werner, Sitzungsb. Akad. Wiss. Miinchen, 27:215, 1897. 
Giinther, Biologia Centrali- Americana : Reptilia and Batrachia, p. 269, 
Sept. 1901. Nieden, Das Tierreich, Amphibia, Anura I, p. 261, Tune 
1923. Dimn, Occas. Papers Boston Soc. Nat. Hist., 5:413, Oct. 10, 1931. 
Gaige, Hartweg, and Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 
357:4, Oct. 26, 1937. Cooper, Copeia, 2:122, June 30, 1944. Breder, 
Bull. Amer. Mus. Nat. Hist., 86(6):416, Aug. 26, 1946. Smith and 
Taylor, Bull. U.S. Natl. Mus., 194:88, June 17, 1948; Univ. Kansas 
Sci. Bull., 33:364, March 20, 1950. Taylor, Univ. Kansas Sci. Bull., 
35(1):837, July 1, 1952. Brattstrom and Howell, Herpetologica, 10:117, 



Neotropical Hylid Frogs, Genus Smilisca 



309 



Aug. 1, 1954. Goin, Herpetologica, 14:120, July 23, 1958. Cochran, 
BuU. U. S. Natl. Mus., 220:57, 1961. 

Hyla labialis Peters, Monats. Konigl. Akad. Wissen. Berlin, p. 463, 1863 
[Holotype. — ZMB 4913 from "region of Bogota," Colombia]; Monats. 
Konigl. Akad. Wissen. Berlin, p. 618, Oct. 16, 1873. Boulenger, Catalogue 
Batrachia and Salientia in British Museum, p. 397, Feb. 1, 1882. 

Hyla haudini dolomedes Barbour, Occas. Papers Mus. Zool. Univ. Michigan, 
129:11, Jan. 25, 1923 [Holotype.— MCZ 8539 from Rio Esnape, Sambu 
Vallev, Darien, Panama; Barbour and Brooks collectors]. Barbour and 
Loveridge, Bull. Mus. Comp. Zool. Harvard, 69:278, June, 1929. 

Hyla phaeota phaeota. Smith, Herpetologica, 8:152, Jan. 30, 1953. Minton 
and Smith, Herpetologica, 16:103, June 17, 1960. 

Smilisca phaeota, Starrett, Copeia, 4:303, Dec. 30, 1960. 

Diagnosis. — Size large ( $ 65 mm., 9 78 mm.); skull as long as wide, lack- 
ing frontoparietal fontanelle; large supraorbital flanges having straight edges 
and extending posterolaterally; large squamosal not in contact with maxillary; 
tarsal fold moderately wide, full length of tarsus; inner metatarsal tubercle 
moderately large, low, flat, elliptical; hind limbs relatively long, tibia averaging 
more than 53 per cent of snout-vent length; labial stripe silvery white; lips 
lacking vertical bars; loreal region pale green; dark brown or black tympanic 
mark dispersing into brown vena ted pattern on flanks; posterior surfaces of 
thighs pale brown, with or without darker flecks or small cream-colored flecks. 
(Foregoing combination of characters distinguishing S. phaeota from any other 
species in genus.) 

Description and Variation. — For the purposes of analyzing geographic varia- 
tion in size and proportions, measurements were taken on ten adult males from 
each of five samples throughout the range of the species. Aside from the data 
summarized in Table 2, the average ratio of tibia length to snout-vent length 
is noticeably less in Colombian specimens (53.4 per cent, as compared with 
54.8 to 57.8 per cent in the other samples) and the ratio of head length to 

Table 2. — Geographic Variation in Size and Proportions in Males of 

Smilisca phaeota. (Means in Parentheses Below Observed Ranges; 

Data Based of Ten Specimens From Each Locality.) 



Locality 


Snout-vent 
length 


Head width/ 

snout-vent 

length 


Interorbital 

distance/ 

head width 


Bonanza, Nicaraeua 


40.8-47.7 

(43.7) 

46.3-59.0 
(51.7) 

53.6-64.9 
(61.4) 

52.4-65.5 
(56.5) 

52.6-61.0 
(56.0) 


34.1-38.0 
(36.3) 

32.5-36.0 
(35.0) 

32.6-36.1 

(34.5) 

33.5-37.6 
(35.6) 

33.1-37.1 
(35.0) 


31.0-36.1 


Heredia Pro v., Costa Rica 

Puntarenas Prov., Costa Rica 
Canal Zone, Panamd 


(35.4) 

30.5-39.6 

(34.7) 

31.0-38.0 
(34.4) 

31.3-37.2 


Rio Quesada, Colombia 


(34.7) 

30.1-33.9 
(32.1) 



310 University of Kansas Publs., Mus. Nat. Hist. 

snout-vent length is noticeably less in Costa Rican specimens (33.5 per cent 
as compared with 34.9 to 35.1 per cent in the other samples). Also, specimens 
from Heredia Province, Costa Rica, have a relatively smaller tympanum (62.7 
to 80.4 [mean 68.4] per cent of the diameter of the eye, as compared with 
means of 74.0 to 77.9 per cent in the other samples). 

Two populations are distinctive as regards the size of adult males. Speci- 
mens from the northern Caribbean lowlands of Nicaragua (Bonanza, the 
northernmost locality for the species) are remarkably small. Males having 
snout-vent lengths of between 40 and 43 mm. were breeding; the largest male 
found had a snout-vent length of 47.7 mm. The other extreme in size is at- 
tained in specimens from the Pacific lowlands of eastern Costa Rica and west- 
em Panama, where most breeding males have snout-vent lengths of more than 
55 mm.; the largest male had a snout-vent length of 64.9 mm. 

The rather striking differences in size among certain samples and the minor 
differences in proportions among other samples show no geographic trends. 
Instead, the variations apparently are random among the samples. The data 
presented here possibly are the results of inadequate sampHng, but more likely 
reflect actual differences in the populations. 

The dorsal ground color of Smilisca phoeota is pale green to tan; the venter 
is creamy white. The dorsum is variously marked with dark olive-green or 
dark brown spots or blotches (Pi. 6C). A dark interorbital bar is usually 
present. Usually a large dark dorsal mark extends from the occiput to the 
sacral region, but in many individuals this blotch is replaced by two or three 
dark marks. The dorsal markings are irregular in shape and do not tend to 
form transverse bands or longitudinal bars. The hind limbs are marked by 
dark transverse bands, usually four or five on the thigh, five or six on the shank, 
and four on the tarsus. Two or three narrow bands are usually present on the 
proximal part of the fourth toe. The webbing on the feet is brown. The 
loreal region is pale green, bordered above by a narrow dark brown canthal 
stripe extending from the nostril to the orbit. The upper lip is silvery white. 
A broad dark brown or black mark extends posteriorly from the orbit, encom- 
passing the tympanum, to a point above the insertion of the forelimb. The 
flanks are pale green or pale tan and marked with a fine dark brown or black 
venation. The anterior surfaces of the tliighs usually are pale brown or grayish 
tan, sometimes having small, indistinct darker flecks. The posterior surfaces 
of the tliighs are similarly colored, but in most specimens small but distinct 
dark flecks are present; in some specimens small cream-colored spots are also 
present on the posterior surfaces of the thighs. A distinct, narrow creamy- 
white anal stripe usually is present. A distinct white stripe is present on the 
outer edge of the tarsus and fifth toe; on the tarsus the white stripe is bordered 
below by dark brown. A white stripe also is present on the outer edge of the 
forearm and fourth finger. In breeding males the throat is dark gray. 

Little geographic variation in color or pattern is evident. Few, if any, 
specimens from the Pacific lowlands of South America are green in life. (We 
have seen no hving individuals from South America.) Some living individuals 
from Costa Rica and all those seen alive from Nicaragua have a tint of pale 
blue on the flanks. In some specimens the dorsal pattern is so faint as to be 
barely discernible, whereas in most specimens the pattern is bold. 

The coloration in the living frogs is highly variable due to extreme meta- 
chrosis. Individuals of this species are capable of changing the dorsal colora- 



Neotropical Hylid Frogs, Genus Smilisca 311 

tion from green to brown in a short period of time. Both green and brown 
individuals have been found active at night. Usually those individuals found 
hiding by day are brown. One individual from Finca La Sumbadora, Panama 
(now KU 91914), was kept aUve in the laboratory for nearly one month. This 
individual usually was pale green with tan dorsal markings at night and tan 
with pale green markings by day. On occasion the pale green dorsal markings 
were boldly outUned by bright dark green. 

In hving individuals from throughout the range of the species the iris 
is a bronze color, darkest medially with fine black reticulations. 

Natural History. — Smilisca phaeota inhabits humid lowland tropical forest 
and seldom ascends the foothills to more than 1,000 meters. The rather 
equable climatic conditions, especially more or less evenly distributed rainfall 
throughout the year, permit this frog to be active most of the year. Dunn 
(1931:413) reported males calling on Barro Colorado Island, Panama, in Feb- 
ruary and in July, and Breder (1946:416) noted calling individuals in the 
Chucanaque drainage of Darien, Panama in January, March, July, August and 
October and in Costa Rica in April through August inclusively. Calling males 
were found at Bonanza, Nicaragua in March and in July. 

At all times of year the usual daytime retreats for these frogs are near water; 
the frogs have been found in elephant ear plants (Xanthosoma) and in bro- 
meliads; occasional individuals have been found sitting on shaded branches of 
bushes and trees. None has been observed on the ground or beneath ground- 
cover by day. 

The length of the breeding season cannot be determined definitely. The 
earhest date on which eggs have been found is May 23; Gaige, Hartweg, and 
Stuart (1937:5) reported a gravid female taken at El Recreo, Nicaragua, in 
September, and we have a gravid female taken at Almirante, Panama, in 
March. 

Males usually call from secluded spots at the edge of water. All calhng 
males that we observed were on the ground within a few centimeters of the 
water. The males usually are hidden beneath an over-hanging leaf or some 
other cover; they definitely do not sit in the open like Smilisca baudini. Most 
calhng males and clasping pairs have been found at the edges of small pools 
or shallow ditches, although occasional individuals are found at the edges of 
large ponds or streams. 

The breeding call consists of one or two moderately short, low-pitched notes 
(duration 0.33 to 0.42 seconds), repeated at intervals of about 20 seconds to 
several minutes. Each note is a low, vibrant "wauk," having 100 to 130 pulses 
per second and a dominant frequency of 330 to 420 cycles per second ( Pi. IOC ) . 

The eggs are deposited in loose clumps amidst vegetation in the water. 
Hatchling tadpoles have total lengths of 8.7 to 10.6 mm., and body lengths 
of 4.1 to 4.5 mm. The external gills are long and filamentous, and the yolk 
sac is large. The head and caudal musculature are dark bro\\Tiish black, and 
the caudal fins are gray. The oral discs are large and roughly circular. The 
growth and development of the tadpoles are summarized in table 11 and 
figure 16. 

A typical tadpole in stage 30 of development (KU 68482 from the Rio 
Chitaria, Cartago Province, Costa Rica) may be described as follows: body 
length 9.7 mm.; tail length 14.6 mm.; total length 24.3 mm.; body as vdde 
as deep; snout rounded dorsally and laterally; eyes widely separated, directed 



312 University of Kansas Publs., Mus, Nat. Hist. 

dorsolaterally; nostril about midway between eye and tip of snout; mouth 
anteroventral; spiracle sinistral, about midway on length of body and slightly 
below midline; anal tube dextral; caudal musculature slender, curved upward 
distally; dorsal fin extending onto body; depth of dorsal fin slightly less than 
that of ventral fin at mid-length of tail; dorsal part of body pale brown; ventral 
surfaces transparent with scattered pigment; pale cream-colored, crescent- 
shaped mark on posterior edge of body; caudal musculature pale creamy tan 
with scattered pale brown spots; caudal fins transparent with scattered small 
brown blotches on dorsal and ventral fins; iris pale bronze in life (Fig. 13); 
mouth small; median part of upper lip bare; rest of mouth bordered by one 
row of pointed papillae; lateral fold present; tooth-rows %, first upper row 
longest; second upper row slightly shorter, broadly interrupted medially; three 
lower rows complete, equal in length, slightly shorter than second upper row; 
upper beak moderately deep, forming broad arch with slender lateral proc- 
esses; lower beak slender, broadly V-shaped; both beaks serrate (Fig. 15E). 
In tadpoles having fully developed mouth-parts the tooth-row formula of 
% is invariable. The pale crescent-shaped mark on the posterior part of the 
body curves anterodorsally on the dorsal surface of the body. These marks 
in dorsal view give the appearance of a pair of short, pale stripes on the pos- 
terior part of the body. Most specimens from Costa Rica have the pale colora- 
tion like that described above, but some individuals (notable KU 87683 from 
Guapiles, Costa Rica, KU 87707 from Finca Tepeyac, Nicaragua, and KU 87708 
from Bonanza, Nicaragua) have much more pigment. In these specimens 
the same color pattern obtains as in the pallid individuals, but the pigmentation 
is dense. This is especially noticeable on the tail. 

Recently metamorphosed young have snout-vent lengths of 12.7 to 16.7 mm. 
(average, 14.3 mm. in eleven specimens). Coloration of young in life (KU 
68484 from Rio Chitaria, Cartago Province, Costa Rica): "Dorsum pale tan; 
side of head and flanks darker brown, separated from tan dorsum by an 
indistinct cream stripe. Limbs pale yellow; thighs flecked with brown; shank 
and tarsus yellowish tan with indistinct brown bars. Soles of feet brown. Belly 
white; throat dusty cream flecked with silvery white. Upper lip silvery white. 
Iris bright gold with black flecks. Heels, tarsal and anal stripes white" ( Duell- 
man, field notes. May 23, 1961). 

Remarks. — Peters (1863:463) named Hyla labialis from the "region of 
Bogota, Colombia", but in 1873 regarded his new species as identical with 
Hyla phaeota Cope, 1862, from Turbo, Colombia. The name Hyla labialis 
has been used for frogs from the northern Andes in Colombia (see Dunn, 
1944:72, and Stebbins and Hendrickson, 1959:522, for discussion of nomen- 
clature). Rivero (1961:131) used the name Hyla vilsoniana Cope, 1899, for 
the frogs from the northern Andes previously referred to Hyla labialis. A 
review of the nomenclature and taxonomy of these frogs, which superficially 
resemble Smilisca but are unrelated, is beyond the scope of the present study. 
Hyla baudini dolomedes Barbour, 1923, is based on a small Smilisca phaeota 
(MCZ 8539) having a snout-vent length of 45.5 mm. Dimn (1931a:413) 
placed dolomedes in the synonymy of Smilisca phaeota. We have examined 
the holotype of dolomedes and agree with Dimn's assignment. 

Smith (1953:150) described Hyla phaeota cyanosticta from Guatemala. 
Our studies on the external morphology, coloration, and especially the cranial 
osteology provide evidence that cyanosticta is a species distinct from phaeota. 



Neotropical Hylid Frogs, Genus Smilisca 313 

Distribution. — Smilisca phaeota inhabits humid tropical forests from north- 
eastern Nicaragua southward on the Caribbean lowlands to elevations of about 
1000 meters and on the Pacific lowlands of Costa Rica, exclusive of the arid 
regions of Guanacaste, throughout the lowlands of Panama, exclusive of the 
savannas of the Pacific lowland and the Azuero Peninsula, and southward on 
the Pacific slopes of South America through Colombia to west-central Ecuador; 
also the valleys of the Rio Cauca and Rio Magdalena in Colombia (Fig. 2). 

Specimens examined. — 528, as follows: Nicaragua: Matagalapa: Finca 
Tepeyac, 10 km. N, 9 km. E Matagalpa, KU 85439, 87707 (tadpoles); Mata- 
galpa, MCZ 3546-7, UMMZ 92367; 19 km. N Matagalpa, UMMZ 116495-6. 
Zelaya: Bonanza, KU 84854-62, 84950-2 (skeletons), 85440-50, 87708-9 (tad- 
poles); Cukra, AMNH 80618; Rio Mico, 16 km. E Recreo, UMMZ 79711 (6), 
79712 (4); junction Rio Mico and Rio Siguia, UMMZ 79713 (10); Rio Siguia, 
11 km. NW Rama, UMMZ 79714 (14), 79715 (11), 79716 (21), 79717, 79718 (3). 

Costa Rica: Alajuela: Cinchona, KU 32255, 64286-8; 5 km. S Ciudad 
Quesada, USC 8077; Laguna Monte Alegre, KU 64289-90; Las Playuelas, 11 
km. S Los Chiles, USC 7216; San Carlos, USNM 29961. 

Cartage: Moravia de Turrialba, KU 32212-47, 37133-5, 41093 (skeleton), 
64280-1, USC 7243 (3); Peraha, KU 32271-2; Rio Chitaria, 3 km. NNE Pa- 
vones, KU 64273-9, 68477 (eggs), 68478-83 (tadpoles), 68484 (young); Rio 
Reventazon, MCZ 29196-203, UMMZ 117677 (9); Turrialba, KU 25720-2, 
32209-11, 32266-8, 32273-4, 37136-67, 41090-2 (skeletons), 64270-2, MCZ 
29221, 29222 (tadpoles), 29269-70, USNM 29934. 

Guanacaste: Tilaran, KU 36805-7; 8 km. NE Tilaran, KU 36803-4. 

Heredia: Barranca del Rio Sarapiqui below Isla Bonita, KU 64282-3; Cari- 
blanco, KU 32256-60, 41094 (skeleton), 64284, MCZ 7967; Isla Bonita, KU 
32250-4; 4.2 km. W Puerto Viejo, KU 64285, 68485; 7.5 km. W Puerto Viejo, 
KU 68486; 1 km. S Puerto Viejo, KU 86518. 

Limon: Bambu, USC 7182 (4); Batan, UMMZ 118582; Coen, MCZ 9825; 
La Lola, KU 32262-4, UF 4029, UMMZ 117678 (3); Los Diamantes, CNHM 
101295-8, KU 25723-4, 32265, 64267-9; Pandora, UMMZ 122650 (2), USC 
7188 (3), 7190; Puerto Limon, KU 32261; Rio Lari at Rio Dipari, 21 km. SW 
Amubre, USC 7177; Rio Toro Amarillo, 7 km. W Guapiles, KU 86519, 87683 
(tadpoles); Suretka, KU 36808-10, 37168. 

Funtarenas: Agua Buena, KU 36790; 1.6 km. E Buenos Aires, UMMZ 
117578; 3 km. NW Buenos Aires, KU 64304; 4 km. N, 15 km. W Dominical, 
KU 68491-2 (tadpoles); Esparta, MCZ 8029-30, 8032; Golfito, KU 32270; 6 
km. E Golfito, KU 84999-500 (skeletons); Gromaco, UMMZ 123677 (4); Pal- 
mar, KU 32269; 4 km. ESE Palmar Sur, KU 64305-6; 5.6 km. SE Palmar Sur, 
KU 68489 (tadpoles); 7.0 km. SE Pahnar Sur, KU 68490 (young); 8.5 km. SE 
Piedras Blancas, KU 64292-303; Quebrada Boruca, 22 km. E Pahnar Norte, 
KU 64291; Rincon, "Camp Seattle," Peninsula de Osa, UMMZ 123676 (3), 
USC 7254; Rio Ferruviosa, 7 km. S Rinc6n, USG 7235; 1.6 km. WNW Villa 
Neily, KU 68493 (young), 68494 (tadpoles). 

San Jose: San Isidro el General, KU 32249, UMMZ 75025; 10 km. N San 
Isidro el General, MCZ 29099-103; 13 km. WSW San Isidro el General, KU 
86517; 15 km. WSW San Isidro el General, KU 68487 (tadpoles), 68488 
(young), 68495 (young); 20 km. WSW San Isidro el General, KU 32248. 

Panama: No province: Cano Saddle, USNM 69588; Punta de Pena, 
USNM 38733; Quipo, AMNH 18925-6. Bocas del Toro: Almirante, KU 
80080, 91835-6; 1.6 km. W Almirante, KU 91837; 3 km. W Almirante KU 
91824 (skeleton), 91838-43, 91906-7; 11 km. NW Almirante, CNHM 67853-61; 
13 km. W Almirante, KU 91825-7 (skeletons), 91844-9; Fish Creek, KU 92329; 
Isla Popa, KU 91850-1. Canal Zone: Barro Colorado Island, CNHM 6007, 
13316, 13325, 13331, 13360-2, 13377-8, MCZ 24191-5, UF 7523, UMMZ 
63547-60, 64457, 69497 (3); 3.7 km. W Cocoli, KU 67916; Fort Sherman, 
MCZ 10139; Gatun, MCZ 35644; Junction roads C25B and C16, TNHC 23839; 

3—3430 



314 University of Kansas Publs., Mus. Nat. Hist. 

Madden Forest Preserve, TNHC 23837-8. Code: El Valle, KU 77521-4, 
77649 (tadpole), TNHC 23369. Comarca del Baru: Progreso, UMMZ 61085-9. 
Colon: Achiote, KU 77516-20, 77648 (young); Rio Candelaria, CNHM 67- 
851-2. Darien: Rio Esnape, Sambii Valley, MCZ 8539; Rio Sucubti, Cha- 
lichiman's Creek, AMNH 40512; Camp Creek, AMNH 40758-9; Camp Creek, 
Camp Townsend, AMNH 40988. Panama: NW slope Cerro Prominente, KU 
80459; Finca La Sumbadora, KU 91914 (skeleton). Chiriqui: 2 km. W Con- 
cepcion, AMNH 68910. 

Colombia: Antioqnia: Puerto Berrio, CNHM 30805 (Coin); Turbo, USNM 
39899. Caldas: Pueblorrica, Santa Cecilia, CNHM 54768-71 (Coin). Choco: 
No specific locality, AMNH 3984-6; Andagoya, BMNH 1915. 10. 21. 69-70, 
CNHM 81857 (Coin); Golfo de Uraba, CNHM 63881 (Coin); Peiia Lisa, 
Condoto, BMNH 1913. 11. 12. 118-125, 1913. 11. 12. 137-146 (Coin); Pizarro, 
CNHM 4451-3, 4455-61 (Coin); Rio San Juan, Playa del Oro, CNHM 54772 
(Coin); Rio Quesada, AMNH 13615-77; 37 km. up Rio Pune, AMNH 13688; 
48 km. up Rio Pune, AMNH 13689. Narino: Tumaco, Rio Rosario, CJG 
2310-13 (Coin). Valle: Buenaventura, BMNH 1895. 11. 16. 82 (Coin); Raposa, 
WAT 166, 346-47, 388 (Coin); Rio Calima above Cordoba, CJG 2249-57 
(Coin). 

Ecuador: No province: Bulun, AMNH 10620. Esmeraldas: Cachabe, 
AMNH 10625-8; Rio Capayas, CNHM 35712; Rio Sapaya, UMMZ 58910 (5); 
Salidero, AMNH 10623-4; San Javier, AMNH 10618. Guayas: Hacienda 
Balao Chico, UMMZ 123904. Imbabura: Pambelar, AMNH 10629, 10631. 
Pichincha: Hacienda Espinosa, 9 km. W Santo Domingo de los Colorados, 
KU 40220. 

Smilisca puma (Cope), new combination 

Hyla puma Cope, Proc. Amer. Philos. Soc, 22:183, 1885 [Holotype. — USNM 
13735 from Nicaragua; J. F. Moser collector], Giinther, Biologia Centrali- 
Americana: Reptilia and Batrachia, p. 270, Sept., 1901. Nieden, Das 
Tierreich, Amphibia, Anura I, p. 251, June, 1923. Cochran, Bull. U. S. 
Nad. Mus., 220:58, 1961. 

Hyla wellmanorum Taylor, Univ. Kansas Sci. Bull. 25(1) :843, July 1, 1952 
[Holotype. — KU 30302 from Batan, Limon, Costa Rica, Edward H. Tay- 
lor collector]; Univ. Kansas Sci. Bull., 36(1):626, June 1, 1954. Duell- 
man and Berg, Univ. Kansas Publ. Mus, Nat. Hist., 15:194, Oct. 26, 1962. 

Smilisca welljnanorum, Starrett, Copeia, 4:303, Dec. 30, 1960. 

Diagnosis. — Size small ( $ 38.0 nmi., $ 46.0 mm.), differing from other 
species in the genus by the following combination of characters: skull about 
as long as broad; frontoparietal fontanelle keyhole-shaped; supraorbital flanges 
absent; squamosal small, not in contact with maxiUary; bony portion of ethmoid 
terminating at anterior edge of orbit; tarsal fold weak, two-thirds length of 
tarsus; inner metatarsal tubercle small, low, flat, elliptical; snout rounded in 
dorsal profile; lips thin and flaring; fingers having only vestige of web; toes one- 
half webbed; diameter of tympanum about two-thirds that of eye; narrow 
labial stripe white; pair of dark brown (sometimes interconnected) stripes on 
tan dorsum; no blue spots on flanks or thighs; vocal sac in breeding males 
pale brown. (Foregoing combination of characters distinguishing S. puma 
from other species in genus.) 

Description and variation. — Ten breeding males from the vicinity of Puerto 
Viejo, Heredia Province, Costa Rica, have snout-vent lengths of 32.5 to 37.9 
mm. ( 34.8 mm. ) . In these specimens, the length of the tibia to the snout-vent 
length is 0.48 to 0.53 (0.51), and the tympanum/eye ratio is 0.52 to 0.72 
( 0.65 ) . Seven females have snout-vent lengths of 40.8 to 45.8 mm. ( 43.9 mm. ). 



Neotropical Hyled Frogs, Genus Smilisca 315 

No individual has more than a vestige of a web between the second and third 
and fourtli fingers. None has a web between the first and second fingers. 
Breeding males lack nuptial excrescences on the tliumbs. The vocal sac is 
moderately large and bilobed. 

In preserved specimens the dorsal ground color varies from yellowish tan 
to grayish browTi. All specimens have dark brown dorsal markings in the form 
of a pair of dorsal stripes, variously modified (Pi. 7A). In some specimens, 
such as KU 91716, the stripes are discrete and extend from the postorbital 
region nearly to the vent. In most specimens the stripes are connected by a 
transverse mark in the scapular region and in many others also by a trans- 
verse mark in the sacral region. In some specimens the stripes are fragmented 
posteriorly; fragmentation is extreme in KU 30300, in which the dorsal pat- 
tern consists of two series of dark longitudinal dashes. The other extreme is 
a nearly complete fusion of the stripes, as in KU 91714. A dark brown inter- 
orbital bar usually extends onto the eyeUds, but in some specimens this is 
reduced to a short V-shaped mark or small spot between the eyes. There is 
no dark post-tympanic mark, but dark brown pigment forms a venated pattern 
from the axilla to the mid-flank; the inguinal region is white, finely mottled with 
dark bro%vn. The dorsal surfaces of the hind limbs are colored like the body 
and have two or three dark brown transverse marks on the thighs, three to five 
marks on the shanks, and one or two marks or irregularly arranged dark flecks 
on the tarsi. The anterior and posterior surfaces of the thighs are pale tan to 
brown. The webbing of the feet is tan to grayish brown. A narrow white 
labial stripe, white anal stripe, and narrow white stripes on the tarsi and outer 
edges of the forelimbs are invariably present. The ventral surfaces are creamy 
white. 

In life the dorsum is tan or pale brown with dark brown markings. Some 
individuals have scattered metallic green flecks on the dorsum. The flanks 
are mottled dark brown and creamy white. The posterior surfaces of the 
thighs are dark brown. The vocal sacs are grayish brown, and the iris is a 
deep bronze color. 

Natural Histortj. — Smilisca puma inhabits humid lowland tropical forests 
having more or less evenly distributed rainfall throughout the year. The 
equable climatic conditions seemingly permit these frogs to be active through- 
out most of the year. Taylor (1952:846) found calling males at Batan, Costa 
Rica, on July 20, 1951. We found the species breeding near Puerto Viejo, 
Costa Rica, on February 19, June 18, July 13, and July 31. Specimens of 
calling males from Costa Rica in the collection at the University of Southern 
California were obtained in Febniary at La Fortuna, on August 22 at Los 
Diamantes, on August 30 at Jabillos, and on September 5 at La Lola. Gravid 
females were collected in June, July and August. 

Males call from shallow water. All breeding congregations of this species 
that we have found were in a grassy marsh, 7.5 kilometers west of Puerto 
Viejo, Costa Rica. Tadpoles were found in water-fiUed depressions in the 
marsh at night. When first observed, tadpoles were near the surface of the 
water; they responded to light by quickly taking refuge in the dense grass. 
No tadpoles were observed by day. 

The breeding call consists of a low squawk, usually followed by a series 
of one or more rattling secondary notes (duration of primary notes, 0.06-0.35 



316 University of Kansas Publs., Mus. Nat. Hist. 

seconds; of secondary notes, 0.10 to 0.47 seconds), repeated at intervals of 
5 to 55 seconds. The primary notes have 187 to 240 pulses per second and 
major frequencies of about 740 to 1870 cycles per second (Pi. IIA). 

Only six tadpoles are available for study. Four of them in stage 34 of 
development have body lengths of 9.0 to 9.5 mm., tail lengths of 14.0 to 15.0 
mm., and total lengths of 23.0 to 24.5 mm. One tadpole in stage 38 and one 
in stage 40 have total lengths of 31.0 mm. A typical tadpole in stage 34 of 
development (KU 91807 from 7.5 km. W Puerto Viejo, Heredia Province, 
Costa Rica) has a body length of 9.5 mm., tail length of 15.0 mm., and total 
length of 24.5 mm.; body about three-fourths as deep as wide; snout rounded 
dorsally and laterally; eyes widely separated, directed dorsolate rally; nostril 
about midway between eye and tip of snout; mouth antero ventral; spiracle 
sinistral, about two-thirds distance from snout to posterior end of body and 
slightly below midline; anal tube dextral; caudal musculature slender, barely 
curved upward distally; dorsal fin extending onto body; at mid-length of tail, 
depth of caudal musculature equal to that of dorsal fin and ventral fin; body 
grayish brown, palest ventrally; caudal musculature pale creamy yellow with 
bold gray reticulations; caudal fins transparent with gray reticulations anteriorly 
and black flecks posteriorly on both fins (Fig. 14A). Median part of upper 
lip bare; rest of mouth bordered by two rows of short blunt papillae; lateral 
fold present; tooth-rows %; upper rows equal in length; second upper row 
broadly interrupted medially; three lower rows complete, first and second 
rows equal in length, shghtly shorter than upper rows; third lower row notice- 
ably shorter; upper beak shallow, forming broad, continuous arch with slender 
lateral processes; lower beak slender, broadly V-shaped, both beaks finely 
serrate (Fig. 15B). 

All six tadpoles are colored alike, except that in the larger specimens scat- 
tered white flecks are present on the ventral surface of the body, and the dark 
reticulations continue farther posteriorly on the caudal fins than in the smaller 
tadpoles. In two specimens the third lower tooth-row is only about one-half 
the length of the other lower rows, and in one specimen the second lower 
tooth-row is shorter than the first. Coloration of tadpoles in life: "Body olive- 
brown with silvery green flecks laterally. Caudal musculature olive-brown with 
greenish tan flecks. Fins browTi with greenish gold flecks. Iris deep bronze." 
(Duellman, field notes, February 19, 1965). 

One recently metamorphosed young (KU 91808) has a snout-vent length 
of 12.4 mm. In life this frog had a pale tan dorsum with dark bro\vn mark- 
ings, yellowish tan posterior surfaces of thighs, grayish brown throat, and 
bronze iris. 

Remarks. — The identity of Cope's Hyla puma has not been known. The 
name has appeared in various compilations, but no workers have referred any 
of their specimens to that species. Examination of the holotype (USNM 
13735), an adult female, revealed the presence of the following combination 
of characters: snout-vent length 45.8 mm., snout blunt above and rounded 
laterally, nostrils close to tip of snout, lips thin and flaring, a vestige of a 
web on the hands, feet about one-half webbed, tarsal fold weak and extending 
about two-thirds length of tarsus, dorsal markings consisting of a faded dark 
interorbital bar and a pair of faded longitudinal brown marks connected by 
a transverse band in the scapular region. The type agrees well with speci- 
mens of Smilisca ivellmanorum (Taylor, 1952); the vestigial webbing on the 



Neotropical Hylid Frogs, Genus Smilisca 



317 



hands and the dorsal coloration are especially significant. Consequently, we 
consider Hyla wellmanorum Taylor, 1952, to be a synonym of Hyla puma Cope, 
1885. Cope gave only "Nicaragua" as the locality for Hyla puma. The speci- 
men was part of a collection received at the United States National Museum 
from Lt. J. F. Moser. Among the species in the collection are Dentrobates 
pumilio, Phyllomedusa helenae, Conjthophanes cristatus, Pliocercus dimidatus, 
Tretanorhinus nigroluteus, and others characteristically found on the Carib- 
bean lowlands of Central America. Thus, it seems reasonable to assume that 
the type specimen of Hyla puma came from the Caribbean lowlands. Though 
no other Nicaraguan specimens have been found by us, numerous specimens 
are known from the Caribbean lowlands of Costa Rica. 

Cochran (1961:58), in her catalogue of type specimens in the United 
States National Museum, listed Hyla puma Cope, 1885, as a synonym of Hyla 
molitor Schmidt, 1857. She made no qualifying statements. Schmidt (1858: 
246), in his descriptions of the species in the year following his publication 
of the names and Latin diagnoses, stated: "Dorsum uniformly gray, more 
intensive on back, fading away laterally and on extremities; in every-day-life 
this blue would be called Mueller's Blaii. A delicately dotted black line nms 
on the canthus rostralis from the opening of the nose to the comer of the eye. 
In the armpits, on the flanks and the thighs two of our three specimens have 
black marblings." [Free translation] Certainly on the basis of coloration 
Hyla puma is distinctly different from Hyla molitor. 

Distribution. — This species lives in the wet, forested regions of the Carib- 
bean lowlands of Costa Rica and presumably southern Nicaragua (Fig. 3). 
All specimens are from low elevations; the highest known elevation for the 
occurrence of this frog is 285 meters at Lagima Bonilla. 




Fig. 3. Map showing locaUty records for Smilisca puma (triangles) and 

Smilisca sila (circles). 



Specimens examined. — 62, as follows: Nicaragua: No specific locality, 
USNM 13735. 

Costa Rica: Alajuela: Jabillos, 5 km. N Santa Clara, USC 8058 (6); 5 
km. W La Fortuna, USC 8078 (2); Rio La Fortuna at La Fortuna, USC 7151 
(3). Cartage: Laguna Bonilla, tunnel camp near Peralta, KU 32171. Heredia: 
Puerto Viejo, KU 86521; 5.9 km. W Puerto Viejo, KU 64307; 7.5 km. W Puerto 



318 University of Kansas Publs., Mus. Nat. Hist. 

Viejo, KU 64308-10, 64311 (skeleton), 64312-15, 68635-6 (skeletons), 85001-2 
(skeletons), 86520, 87770-1 (skeletons), 91709-16, 91791-2, 91807 (tadpoles), 
91808 (young). Limon: Batan, KU 30300-2; La Lola, KU 32169, USC 141, 
201, 8067; Los Diamantes, KU 32170, UMMZ 118470 (6), USC 212; 2.4 km. 
E Los Diamantes, USC 8049 (5). 

Smilisca sila new species 

Hyla gabbi. Noble, Proc. Biol. Soc. Washington, 37:66, Feb. 21, 1924. 

Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, Oct. 10, 1931. 

Schmidt, Smithsonian Misc. Coll., 89(1 ):6, March 16, 1933. 
Hylu sordida, Dunn, Copeia, 3:166, Nov. 19, 1937. Cooper, Copeia, 2:121, 

June 30, 1944. Breder, Bull. Amer. Mus. Nat. Hist., 86(8):417, Aug. 

26, 1946. 
Hyla phaeota, Breder, Bull. Amer. Mus. Nat. Hist., 86(8): pi. 55, Aug. 26, 

1946. 

Holotijpe. — Adult male, KU 91852 from a small stream at the north edge 
of the village of EI Volcan, Chiriqui Province, Panama, elevation 1280 meters; 
obtained on Feb. 5, 1965, by William E. Duellman. 

Paratypes. — KU 91853-74, collected with tlie holotype. 

Diagnosis. — Size moderate ( S 45.0 mm., $ 62.2 mm.); skull wider than 
long, having large, ovoid frontoparietal fontanelle; supraorbital flanges absent; 
squamosal small, not contacting maxillary; bony section of ethmoid extending 
anteriorly between nasals; tarsal fold weak, full length of tarsus; inner meta- 
tarsal tubercle low, flat, elliptical; lips thick, rounded, not flaring; fingers one- 
third webbed; toes three-fourths webbed; diameter of tympanum about one- 
half that of eye; margin of upper lip faintly marked by interrupted creamy 
white stripe; dark spots on dorsum; pale Hecks on flanks and posterior surfaces 
of thighs; vocal sacs in breeding males dark brown. (Foregoing combination 
of characters distinguishing S. sila from any other species in genus. ) 

Description of holotype. — Snout- vent length 36.6 mm.; tibia length 19.8 
mm., 54.1 per cent of snout-vent length; foot length 15.5 mm., 42.3 per cent 
of snout-vent length; head length 12.7 mm., 34.7 per cent of snout-vent length; 
head width 13.3 mm., 36.8 per cent of snout-vent length; snout short, in lat- 
eral profile truncate, only slightly rounded above, in dorsal profile rovmded; 
canthus rounded; loreal region noticeably concave; hps thick, rounded, not 
flaring; nostrils not protuberant, directed laterally; internarial distance 3.0 mm.; 
intemarial area flat; top of head flat; interorbital distance 3.5 mm., 26.3 per 
cent of head width; diameter of eye 4.2 mm., thrice distance (1.4 mm.) from 
tympanum to eye, and half again distance (2.8 mm.) from orbit to nostril; 
pupil horizontally ovoid; width of eyefid 2.8 mm., 21.1 per cent of head width; 
dermal fold from posterior corner of orbit covering upper edge of tympanum 
to point above insertion of forelimb; diameter of tympanum 2.3 mm., 54.7 per 
cent of diameter of eye; no axillary membrane; arms moderately robust; weak 
fold on wrist; faintly scalloped fold along ventrolateral margin of forearm; 
fingers short, slender; fingers from shortest to longest, 1-2-4-3; vestige of web 
between first and second fingers; others about two-fifths webbed; discs mod- 
erate, diameter of that on third finger about one-third diameter of eye; tri- 
angular outer palmar tubercle; elliptical inner pahnar tubercle on base of 
poUex; subarticular tubercles large, conical, none bifid; supemmnerary tubercles 
few, large, conical; brown nuptial excrescence on prepollex; heels overlap by 
about one-fifth length of shank when hind limbs adpressed; tibiotarsal articu- 
lation extending to nostril; tarsal fold weak, extending nearly full length of 
tarsus; inner metatarsal tubercle elliptical, flat; outer metatarsal tubercle ab- 
sent; toes moderately long; toes from shortest to longest, 1-2-3-5-4, third and 
fifth about equal in length; discs about same size as those on fingers; webbing 



Neotropical Hylid Frogs, Gentjs Smilisca 319 

extending to middle of penultimate phalanx on all toes, except only to distal 
end of antepenultimate phalanx of fourth toe; subarticular tubercles round; 
supernimierary tubercles large, round, present only on proximal digits. Anal 
opening directed posteriorly at level of upper edge of thighs; no noticeable 
anal sheath; flat tubercles ventrolateral to anal opening large; skin of chest, 
belly, and posterior surfaces of thighs granular; other surfaces smooth; tongue 
broadly cordiform, shallowly notched posteriorly, and barely free behind; 
vomerine teeth 4-4, situated on ventral surfaces of separated rounded promi- 
nences between posterior margins of small, ovoid inner nares; vocal slits long, 
each situated along inner margin of ramus; color ( in preservative ) pinkish tan 
above with irregular olive-brown markings forming interconnected spots on 
back; four bars on dorsal surface of each thigh; five bars on shank, and three 
on tarsus; inguinal region white with black mottling; posterior surfaces of 
thighs yellowish tan proximally, dark brown distally; margins of lips grayish 
white with brown markings; ventral surfaces of hands and feet grayish brown; 
belly and posterior part of throat creamy white; anterior part of throat brown. 

Description and variation. — Ten breeding males from Finca La Sumbadora, 
Panama, have snout-vent lengths of 40.0 to 44.8 mm. (42.3 mm.). In these 
specimens the tibia/snout-vent length ratio is 0.50 to 0.57 (0.54), and the 
tympanum/eye ratio is 0.48 to 0.58 (0.53). There is a geographic gradient 
in size; specimens from the western part of the range (southern Costa Rica) 
are smaller than those from the eastern part of the range (eastern Panama). 
Five males from the Pacific lowlands of southern Costa Rica have snout-vent 
lengths of 31.6 to 38.2 mm. (34.7 mm.); ten males from El Volcan, Chiriqui, 
Panama, 32.6 to 37.9 mm. (36.4 mm.), and eight males from Barro Colorado 
Island, Canal Zone, 38.2 to 42.0 mm. ( 35.6 mm. ) . These are smaller than the 
males from Finca La Sumbadora, which is east of the Canal Zone. Ten fe- 
males from El Volcan have snout-vent lengths of 44.2 to 55.6 mm. (49.2 mm.), 
as compared 56.1 to 62.2 mm. (58.2 mm.) in three females from Finca 
La Sumbadora. 

Large females have scattered small tubercles on the head and back; tuber- 
cles occur in males from Costa Rica and in some males from western Panama. 
The truncate snout is characteristic of both sexes. 

The coloration of Smilisca sila consists of a gray, tan, or pale reddish brown 
dorsal ground color and a creamy white venter. The dorsum is marked by 
dark brown, olive-brown, or dark reddish brown spots or blotches (Pi. 7B). 
Usually the blotches are discrete, but in some individuals they are intercon- 
nected and form an irregular dark mark on the dorsum. There is no tendency 
for the blotches to form transverse bars as in Smilisca sordida. In one speci- 
men (KU 80467) the blotches are fused and form two wide irregular longi- 
tudinal stripes, as in Smilisca puma. In some females the dorsal markings are 
reduced to a few small spots or are nearly absent (KU 92332), whereas in 
other females the dorsal markings are bold. In one female (KU 91894) the 
dorsal markings are narrowly bordered by pale blue, and nvunerous pale blue 
flecks are present on the pale brown dorsum. In many individuals of both 
sexes small white flecks are present on the dorsal surfaces. 

Usually the flanks and posterior surfaces of the thighs have black mottling 
enclosing pale blue spots and flecks, respectively. The dorsal surfaces of the 
limbs are marked by dark brown transverse bars; usually three or four bars 
are present on each forearm, thigh, and shank. Tlie coloration of the flanks 
and limbs varies geographically. Specimens from southern Costa Rica and 
western Panama have distinct bars on the limbs; the posterior surfaces of the 
thighs have brown reticulations enclosing small blue flecks in specimens from 



320 University of Kansas Publs., Mus. Nat. Hist. 

Costa Rica and bolder, black reticulations enclosing large pale blue spots in 
specimens from western Panama. In specimens from Costa Rica the flanks are 
brown with pale blue flecks, whereas in those from Chiriqui, Panama, the 
flanks are pale blue with dark brown mottling in the inguinal region. Frogs 
from El Valle and Cerro la Campana usually have distinct bars on the limbs; 
the posterior surfaces of the thighs are colored as in frogs from Chiriqui, and 
the inguinal region is pale blue with coarse brown mottling. Specimens from 
Barro Colorado Island are marked like those from El Valle and Cerro la Cam- 
pana, except that on the posterior surfaces of the thighs fine black reticulations 
enclose many dark blue spots. In specimens from Darien and from Panama 
Province east of the Canal Zone (Altos de Pacora, Cerro Jefe, Finca La Sum- 
badora, and Rio Pacora), the markings on the dorsal surfaces of the limbs are 
indistinct or absent in males, but distinct in some females. Intense brown 
and black piginent forms fine reticulations delimiting bold blue spots on the 
flanks; this coloration extends to the axilla in many specimens. Fine black 
reticulations enclose many dark blue spots on the posterior surfaces of the 
thighs. 

In females, the throat is creamy white; in some specimens scattered brown 
flecks are present on the chin and throat. In breeding males the anterior part 
of the throat is dark gray or dark brown. 

The coloration in hfe is as variable as it is in preservative. In life the 
holotype had a tan dorsum with dark olive-green irregular markings and small 
green flecks. The limbs were tan with dark brown transverse bars. The flanks 
were grayish tan anteriorly; the inguinal region and posterior surfaces of thighs 
were blue with black motthng. The belly was creamy white, and the throat 
was brown with creamy yellow flecks. The iris was a dull bronze color. 
Among the paratypes, some individuals had green flecks, others did not. The 
inguinal region and posterior surfaces of the thighs were pale blue, pale yel- 
lowish green, or grayish tan with black mottling. The blue was most notice- 
able in females. 

Colors of a male from Finca La Sumbadora, Panama, were described as 
follows: "Dorsum olive-brown; irregular dark brown blotches, pale green 
flecks, and raised creamy yellow spots on dorsal surfaces; belly creamy white; 
throat grayish brown; undersides of limbs grayish tan; groin, anterior and 
posterior surface of thigh, inner surface of shank, anterior edge of tarsus, and 
proximal parts of third and fourth toes pale blue marbled with dark brown 
and black; webbing brown; iris pale bronze, finely reticulated with black." 
( Duellman, field notes, January 28, 1964. ) 

A female (now KU 91894) from Altos de Pacora, Panama, was described 
as follows: "An irregular dark brown, green-bordered figure on head and 
back; dark brown, green-bordered bands on limbs — all on a lighter brovvTi and 
heavily green-spotted background. These markings are more vivid at night 
than during the day. Lower sides, from midbody onto front of thighs and 
rear of thighs onto venter of shanks to heels and thence dorsally onto basal 
portions of toes heavily blue spotted on a light brown (front of thighs and 
venter of shanks) to blackish brown background. Venter cream. Iris gray- 
brown, finely veined with dark brown." (Charles W. Myers, field notes, De- 
cember 14, 1964.) Note that in the earlier discussion of coloration of pre- 
served specimens, the green spots and borders have changed to pale blue after 
six months in alcohol. 



Neotropical Hylid Frogs, Genus Smilisca 321 

In living individuals from Costa Rica and Panama west of the Canal Zone, 
the blue coloration on the flanks and thighs is much less conspicuous than in 
specimens from eastern Panama. The color of the iris is variable, even in 
frogs from one locality. The coloration of the iris in 13 Uving frogs (now 
KU 92333-45) from Valle Homito, Chiriqui, Panama, was described as fol- 
lows: "Iris variable — from pale to dark brown; in a few the iris has a golden 
cast to the brown; in a few others the lower half of the iris is pale gray with 
the upper half being light brown." (Charles W. Myers, field notes, April 24, 
1965). 

Natural history. — Smilisca sila inhabits the Pacific slopes of lower Central 
America where a pronounced dry season occurs. We have records of males 
calling in December through May and also in August (latter date from El 
Volcan, Chiriqui, Panama). The breeding season seems to be correlated with 
the time of the year when the water is clear and at a low level in the streams 
where these frogs breed. 

Males call from the edges of small, shallow streams, from rocks in the 
streams, or less frequently from vegetation overhanging the streams. Females 
are most frequently found on the banks of streams, and clasping pairs usually 
are in shallow pools in streams. One individual was found in a bromeliad 
about three meters above the ground in the daytime. 

The breeding call consists of a low squawk, usually followed by a series 
of one or more rattling secondary notes (duration of primary notes, 0.06 to 
0.28 seconds; of secondary notes, 0.14 to 0.48 seconds), repeated at intervals 
of 4 to 20 seconds. The primary notes have 97 to 120 pulses per second and 
major frequencies of about 900 to 2220 cycles per second (Pi. IIB). 

Eggs were obtained artificially in the field; the average length of ten 
embryos in the neural groove stage is 2.4 mm., and the average diameter of 
the outer envelope is 4.9 mm. Hatchlings have large, conical oral discs, heavy 
gills, and a large amount of yolk; their average total length is 6.3 mm. 

Tadpoles have been found in pools in clear streams; some tadpoles have 
been observed to cling by their mouths to rocks in the stream; others were 
found on the bottom where they seek refuge among pebbles or under rocks 
and leaves. A complete developmental series of tadpoles is not available. 
Eleven tadpoles in stage 25 of development have body lengths of 8.3 to 10.2 
mm. (9.3 mm.), tail lengths of 17.3 to 21.0 mm. (18.8 mm.), and total lengths 
of 25.9 to 31.0 mm. (28.1 mm.). One tadpole in stage 41 and one in stage 42 
have body lengths of 11.5 and 12.5 mm., tail lengths of 27.2 and 29.5 mm., 
and total lengths of 38.7 and 42.0 mm., respectively. The snout-vent lengths 
of two specimens in stage 43 and one in stage 45 are 12.7, 13.0, and 13.6 mm., 
respectively. 

A typical tadpole in stage 25 of development (KU 80620 from Finca La 
Sumbadora, Panama) has a body length of 9.5 mm., tail length of 19.0 mm., 
and a total length of 28.5 mm.; body only slightly wider than deep, nearly 
flat dorsally; snout broadly rounded in dorsal view, bluntly rounded in lateral 
view; eyes widely separated, directed dorsolaterally; nostril shghtly closer to 
eye than to tip of snout; mouth ventral; spiracle sinistral, located about two- 
thirds distance from snout to posterior edge of body; anal tube dextral; caudal 
musculature moderately heavy, straight; dorsal fin not extending onto body; 
fins deepest at about two-fifths length of tail, where depth of caudal muscu- 
lature about equal to depth of dorsal and depth of ventral fin; musculature 



322 University of Kansas Publs., Mus. Nat. Hist. 

extending nearly to tip of tail; body dark grayish brown above and pale grayish 
tan below with small dark brown spots dorsally and white flecks laterally; 
caudal musculature pale tan with dark brown flecks over entire surface and 
dark brown streaks on posterior one-half of ventral fin and on all of dorsal 
fin (Fig. 14B). Median one-third of upper lip bare; rest of mouth bordered 
by a single row of conical papillae; lateral fold present; tooth rows %; upper 
rows cone-shaped, about equal in length, broadly /\-shaped; second upper 
row narrowly interrupted medially; lower rows complete, about equal in 
length, but slightly shorter than upper rows; upper beak moderately massive, 
its inner surface forming a continuous arch with short lateral processes; lower 
beak broadly \/-shaped; both beaks finely serrate (Fig. 15D). 

Tadpoles from El Volcan, Chiriqui (KU 91833), are more heavily pig- 
mented than those from Finca La Sombadora; the spots on the tail are larger. 
In life these tadpoles had dark brownish black bodies with golden and green 
lichenous flecks; the tail was tan with dark brown markings, and the iris was a 
grayish bronze color. In hfe tadpoles from Finca La Sumbadora were olive- 
tan above and dark gray with pale bluish gray irridescent spots ventrally. The 
caudal musculature was creamy tan with brown flecks and streaks, and the 
iris was pale bronze. 

Metamorphosing young have been found on vegetation at the edge of 
streams and have been raised in the laboratory. Seven recently metamor- 
phosed young have snout-vent lengths of 13.6 to 15.6 mm. (14.6 mm.). A 
living juvenile (KU 91913) raised in the laboratory from a tadpole obtained 
at Finca La Sumbadora had a brown dorsum with darker brown markings, 
a white spot below the eye, and a narrow white labial stripe. The belly was 
white; the flanks were brown with white spots, and the posterior surfaces of 
the thighs were yellowish tan. The iris was a golden bronze color with much 
black reticulation. 

Remarks. — This species has been confused with Smilisca sordida; most 
authors have referred both species to Hijla (Smilisca) gabhi. Examination of 
the types of Hyla sordida, gabhi, salvini, and nigripes revealed that all of the 
names were referable to a single species (S. sordida), and that the small, blunt- 
snouted species in Panama and southern Costa Rica probably was vvithout a 
name. Possibly Hyla molitor Schmidt (1857) is based on the species that 
we have named S. sila, but several discrepancies in his description, plus the 
unknown provenance of the type, have led us to discount the applicability 
of that name to the species under consideration. 

Distribution, — Smilisca sila ranges along the Pacific slopes and lowlands of 
southern Costa Rica and Panama at elevations from sea level to about 1300 
meters; in northern South America the species occurs in the Carribean low- 
lands and in the valleys of the northward draining rivers of Colombia ( Fig. 3 ) . 

Specimens examined, 234, as follows: Costa Rica: Puntarenas: 6 km. E 
Golfito, KU 91717; Quebrada Boruca, 22 km. E Palmar Norte, KU 64265-6; 
Rio Zapote, 7 km. E Palmar Norte, USC 7100 (2). San Jose: San Isidro el 
General, KU 28200; 14 km. NW San Isidro el General, USC 7098 (2); 15 km. 
WSW San Isidro el General, USC 7097. 

Panama: Canal Zone: Barro Colorado Island, AMNH 62320-3, CNHM 
13324 13326-8, 13330, 13338, 13359, 13423-5, KU 80460-6, 80619 (young), 
80625 (skeleton), UMMZ 63542-6, USC 7051. Chiriqui: Boquete, AMNH 
69815, UMMZ 58441-5; El Volcan, KU 77413, 91828-31 (skeletons), 91852-74, 
91832 (eggs), 91833 (tadpoles); 6 km. S El Volcan, CNHM 60442; 16 Ian. 
NNW El Volcan, KU 91879-90; Finca Palosanto, 6 km. WNW El Volcan, 



Neotropical Hylid Frogs, Genus Smilisca 323 

KU 77406-12, 77692 (skeleton), 91875-7, 92330-1; Rio Colorado, 17 km. 
NNW El Volciin, KU 91878, 92332; Valle Hornito, 19 km. NE Gualaca, KU 
92333-45. Code: El Valle, AMNH 55440-5 (13), 59607-14, CNHM 48140, 
60349-2. 60387-92, 60401-4, 60443, 67842-5, KU 91834 (young), 91902-4, 
TNHC 23751-2, USNM 140653. Colon: Rio Candelaria, AMNH 53708-15, 
CNHM 67826-36. Darien: Camp Creek, Camp Townsend, AMNH 40756-7, 
40936-9, 40992; Rio Chico, AMNH 39784, 40986-7; Rio Pita, CNHM 67823-5; 
Tacarcuna, USNM 141796-802; Three Falls Creek, AMNH 41684, 51788. 
Los Santos: Cerro Hoya, USNM 148213-4; Lajamina, Rio Pm-ia, KU 67915. 
Panama: Altos de Pacora, KU 91894; Cerro Jefe, KU 91895-6; Cerro La 
Campana, CNHM 67846, KU 91897-900, USNM 139689; Finca La Smnba- 
dora, KU 80467-81, 80620 (tadpoles), 91910 (eggs), 91911-2 (tadpoles), 91913 
(young), 91908-9 (skeletons); Rio Calobra, USNM 53722, Rio Pacora, 9 km. 
NNE Pacora, KU 91901. Veraguas: Cerro Carbunco, USNM 129066; Cerro 
Tute, CNHM 67837-41; Isla Cebaco, Rio Platanal, KU 91891-3. 

Colombia: Antioquia: Uraba, Villa Arteaga, CNHM 63893 (Coin). At- 
lantico: Sabanalarga, Rio Causa, AMNH 14506. 

Smilisca sordida (Peters), new combination 

Hylu sordida Peters, Monatsb. Konigl. Akad. Wissen. Berlin., p. 460, 1863 
[Syntypes. — ZMB 3141 (two specimens) from "Veragua," Panama; J. 
von Warszewicz collector]. Brocchi, Mission scientifique au Mexique 
. , ., pt. 3, sec. 2, fitudes sur les batrachiens, p. 42, 1881. Boulenger, 
Catalogue Batrachia Salientia in British Musevmi, p. 393, Feb. 1, 1882. 
Giinther, Biologia Centralia-Americana: Reptilia and Batrachia, p. 273, 
Sept. 1901. Nieden, Das Tierreich, Amphibia, Anura, I, p. 258, June, 
1923. 

Hyla gabbi Cope, Jour. Acad. Nat. Sci. Philadelphia, new ser., 8, pt. 2:103, 
1876 [Syntypes.— USNM 30658-9 from near Sipurio, Limon, Costa Rica; 
William M. Gabb collector]. Brocchi, Mission scientifique au Mexique 
. . ., pt. 3, sec. 2, fitudes sur les batrachiens, p. 37, 1881. Boulenger, 
Catalogue Batrachia Salientia in British Museum, p. 372, Feb. 1, 1882. 
Cope, Bull. U. S. Natl. Mus., 32:32, 1887. Gunther, Biologia Centrali- 
Americana: Reptilia and Batrachia, p. 274, Sept. 1901. Werner, Abhand. 
Konigl. Akad. Wissen. Miinchen., 22:351, 1903. Nieden, Das Tierreich, 
Amphibia, Anura I, p. 252, June, 1923. Taylor, Univ. Kansas Sci. Bull., 
35(1):840, July 1, 1952. Cochran, Bull. U. S. Natl. Mus., 220:54, 1961. 

Hyla nigripes Cope, Jour. Acad. Nat. Sci. Philadelphia, new ser., 8, pt. 2:104, 
1876 [Syntypes.— USNM 30685-6, from Pico Blanco, Costa Rica; William 
M. Gabb collector]. Brocchi, Mission scientifique au Mexique . . ., 
pt. 3, sec. 2, fitudes sur les Batrachiens, p. 38, 1881. Boulenger, Cata- 
logue Batrachia SaHentia in British Museum, p. 394, Feb. 1, 1882. Cope, 
Bull. U. S. Natl. Mus., 32:32, 1887. Gunther, Biologia Centrah- Ameri- 
cana: Reptilia and Batrachia, p. 278, Sept., 1901. Nieden, Das Tier- 
reich, Amphibia, Anura I, p. 253, June, 1923. James, Copeia, 3:147, 
Sept. 30, 1944. Taylor, Univ. Kansas Sci. Bull., 35( 1 ) :853, July 1, 1952. 
Cochran, Bull. U. S. Natl. Mus., 220:56, 1961. 

Hyla salvini Boulenger, Catalogue Batrachia Salientia in British Museum, 
p. 372, Feb. 1, 1882 [Syntypes.— BMNH 1947.2.24.13-14 from Cartago, 
Costa Rica; Osbert Salvin collector]. Giinther, Biologia Centrali- Ameri- 
cana: RepHHa and Batrachia, pi. 71, Fig. B., Sept., 1901. Werner, 
Abhand. Zool.-Bot. Gesell. Wien, 46:8, Sept. 30, 1896. 

Smilisca gabbi, Starrett, Copeia, 4:303, Dec. 30, 1960. 

Diagnosis. — Size moderate ( 3 45 mm., $ 64 mm.); skull slightly wider than 
long, having large and elongate frontoparietal fontanelle; supraorbital flanges 
absent; squamosal small, not contacting maxillary; bony section of ethmoid 
terminating just anterior to anterior edge of orbit; tarsal fold weak, full length 
of tarsus; inner metatarsal tubercle long, low, flat, elliptical; lips thin and flar- 



324 University of Kansas Publs., Mus. Nat. Hist. 

ing; fingers one-half webbed; toes four-fifths webbed; diameter of tympanum 
about one-half that of eye; no white labial stripe; dorsal dark markings ir- 
regular, sometimes forming broad transverse bars; pale flecks on flanks and 
usually on posterior surfaces of thighs; vocal sacs in breeding males white. 
(Foregoing combination of characters distinguishing S. sordida from any other 
species in genus. ) 

Description and variation. — Ten breeding males from 15 to 20 kilometers 
west-southwest of San Isidro el General, San Jose, Costa Rica, have snout-vent 
lengths of 38.1 to 42.6 mm. (40.5 mm.). In these specimens, the tibia snout- 
vent length ratio is 0.50 to 0.54 (0.52), and the tympanum/eye ratio is 0.45 
to 0.57 (0.49). Specimens from the Pacific slopes of Costa Rica are larger 
than those from the Meseta Central and the Caribbean lowlands. Ten males 
from 6 kilometers east of Golfito, Puntarenas, have snout-vent lengths of 38.4 
to 44.6 mm. (41.8 mm.), and five males from Rincon, Peninsula de Osa, have 
snout-vent lengths of 38.8 to 41.6 mm. (40.3 mm.). Snout-vent lengths of 
ten males from La Fortuna, Alajuela, are 31.9 to 36.0 mm. (34.4 mm.), of 
ten males from Pandora, Limon, 33.8 to 37.6 mm. (35.9 mm.), and of ten 
males from Escazu and Rio Jorco on the Meseta Central, 34.3 to 37.6 mm. 
(36.0 mm.). Eight females from the Rio Jorco on the Meseta Central have 
snout-vent lengths of 48.8 to 53.8 mm. (50.4 mm.), and six females from 
various localities on the Pacific slopes of Costa Rica have snout-vent lengths of 
56.5 to 64.0 mm. (59.8 mm.). The only noticeable differences in proportions 
between males and females is in the tjonpanum eye ratio; for example, this 
ratio is 0.47 to 0.53 (0.49) and 0.54 to 0.68 (0.61) in ten males and eight 
females, respectively, from the Meseta Central. 

The shape of the snout and the associated cranial elements of S. sordida 
vary geographically and ontogenetically. Specimens from the Caribbean low- 
lands have blunt snouts in lateral view; those from the Pacific lowlands have 
longer, more slender snouts that are pointed in lateral view, and those from 
the Meseta Central are intermediate in snout shape between the two lowland 
populations (Fig. 4). These differences in shape of the snout are dependent 
on the nature of the underlying cranial bones, principally the maxillaries and 
nasals. In specimens from the Caribbean lowlands the nasals are long, wide, 
and narrowly separated from the ethmoid; the anterior edge is just posterior to 
the nostril. The maxillary flanges are nearly vertical. In specimens from the 
Pacific lowlands the nasals are relatively shorter, narrower, and rather widely 
separated from the ethmoid; the anterior edges of the nasals do not extend so 
far forward as in specimens from the Caribbean lowlands. The maxillary 
flanges slant medially. In these cranial characters, specimens from the Meseta 
Central are intermediate between the two lowland populations. 

Superimposed on this geographic variation are ontogenetic changes, which 
are most noticeable in males. In smaller, and presumably younger, specimens 
the snouts are more pointed than in larger specimens; consequently some small 
males from the Caribbean lowlands resemble larger males from the Pacific 
lowlands, since the nasals and maxillaries of the former are not fully ossified. 
In addition, in small breeding males the ethmoid is only about one-half ossified, 
a large frontoparietal foramen is present, the anterior arm of the squamosal 
extends only about one-fourtli the distance to the maxillary (two-thirds the 
distance in larger specimens), and the tegmen tympani are short, as compared 
with the long, thin elements in larger specimens. 



Neotropical Hylid Frogs, Genus Smilisca 



325 









Fig. 4. Variation in the shape of the snout in Smilisca sordida; left column 
females, right column males; all from Costa Rica: (A) Camp Seattle, Rincon 
de Osa, Puntarenas Prov. (UMMZ 123684); (B) Quebrada Agua Buena, 3 km. 
SW Rincon de Osa, Puntarenas Prov. (USC 7236); (C) Rio Oro, 28.5 km. NW 
Villa Neily, Puntarenas Prov. (KU 91742); (D) Rio Jorco, near Desamparados, 
San Jose Prov. (KU 91765); (E-F) Bambu, Limon Prov. (USC 7183). X3. 

The dorsal ground-color of Smilisca sordida is gray to pale tan or reddish 
brovvn; the venter is white. The dorsum is variously marked with dark gray, 
dark brown, reddish brown, or olive-green spots or blotches (PI. 7C). A dark 
interorbital bar usually is present. The dorsal markings on the body usually 
consist of a blotch, or two or more spots, on the occiput, in the scapular region, 
and in the sacral region. In many specimens, especially females, these mark- 
ings are in the form of broad transverse bars. A female (USC 7164) from 



326 University of Kansas Publs., Mus. Nat. Hist. 

Las Canas, Guanacaste, Costa Rica, has a tan dorsum with many black flecks 
and round brown spots bordered by darker brown. One female (KU 91763) 
from the Rio Jorco, San Jose, Costa Rica, has a unicolor tan dorsum. Some 
individuals have scattered, small white spots on the dorsum; these are most 
evident in a male (USC 7153) from La Fortuna, Alajuela. White labial stripes 
and anal stripes are absent in all specimens. 

The limbs are marked by dark brown transverse bars; these are indistinct 
in some specimens from the Meseta Central and Caribbean lowlands, whereas 
they are distinct in all specimens from the Pacific lowlands. Specimens from 
the Caribbean lowlands have two to six bars on each shank, whereas specimens 
from the Pacific slopes have four to six bars on each shank, and specimens 
from the Meseta Central have as many as eight bars on each shank. A 
narrow, sometimes broken white line is present on the ventrolateral edge of 
the forearm. The webbing on the hand is tan or pale gray, and the ventral 
surfaces of the tarsi and the webbing on the feet are dark gray or brown. 
Breeding males have dark brown nuptial excrescences on the prepoUex. 

The flanks and posterior surfaces of the thighs usually are marked by bluish 
white and creamy tan flecks, respectively, but vary considerably. In specimens 
from the Caribbean lowlands a small amount of flecking is present in the 
inguinal region, and on the posterior surfaces of the thighs flecks are few or 
absent. In specimens from the Meseta Central, numerous large flecks or 
small, round spots (pale bluish white in life) are on the posterior half of the 
flanks; small flecks are on the posterior surfaces of the thighs. Specimens 
from the Pacific slopes and lowlands of southern Costa Rica (Puntarenas and 
San Jose Provinces) have bold mottling of black and bluish wliite on the 
flanks and many bluish white flecks on the posterior surfaces of the thighs. 
The flanks are reticulated from the axilla to the groin in two females (UMMZ 
123684 and USC 7236) from Rincon, Peninsula de Osa, In specimens from 
the Pacific slopes of Guanacaste in northwestern Costa Rica, flecks are present 
in the inguinal region; indistinct flecks are on the posterior surfaces of the 
thighs. 

The throat is immaculate in specimens from the Caribbean lowlands in 
Limon Province; the throats are dusky laterally in most other specimens except 
some from the Meseta Central, in which the throats are heavily flecked with 
black. This variation occiirs in males and females. 

The color and pattern in life are highly variable. A composite description 
of living individuals (now KU 91718-41) from 6 kilometers east of Golfito, 
Puntarenas, Costa Rica, fllustrates the variabihty: "Dorsum pale oUve-green, 
fading to tan posteriorly, or tan all over with dark olive-green or dark brown 
spots on back and bars on Umbs. Flanks dark brown with cream, greenish 
gray, or bluish gray mottling. Posterior surfaces of thighs dark brown with 
pale blue, pale green, or tan flecks. Iris creamy sflver. Throats white with 
some brown flecks peripherally." ( Duellman, Field notes, February 15, 1965. ) 
A male from the Rio Jorco, San Jose, Costa Rica, was dvdl olive-tan above 
with olive-green marks; the flanks were brown with pale tan flecks, and the 
posterior surfaces of the thighs were pale brown with cream-colored flecks. 
Six females from the same locality were reddish brown above with olive-brown 
or dark brown markings; one was uniform orange-tan, and another was dull 
olive-green with darker markings. 

The color of the iris in living frogs varies from creamy silver to grayish 
yellow or bronze with a variable amount of black reticulation. 



Neotropical Hylid Frogs, Genus Smilisca 327 

Natural History. — Smilisca sordida is not associated with any one type of 
vegetation; instead it lives in the vicinity of rocky streams having low gradients. 
Breeding takes place primarily in the dry season, when the water in the 
streams is clear and at a low level. Through most of the range of S. sordida 
showers, or even short heavy rains, occur in the dry season. After such rains 
the breeding activity is maximal. Breeding congregations have been found 
from December through April, but a few calhng males and gravid females 
have been taken in June, July, and August. In the rainy season non-breeding 
individuals are found sitting on bushes near streams at night. Taylor (1952: 
843 ) found specimens in bromeliads by day. 

Males usually call from rocks or gravel bars in, or at the edge of, streams. 
Some individuals perch in low bushes overhanging the streams, and some sit 
in shallows in the streams. Clasping pairs have been found on the banks of 
streams and in shallow water in streams. 

The breeding call consists of one to six moderately short, rather high-pitched 
notes (duration 0.18 to 0.45 seconds) repeated at intervals of 12 seconds to 
several minutes. Each note is a vibrant rattle having 78 to 135 pulses per 
second and major frequences of about 1200 to 2600 cycles per second (PI. IIC). 

The tadpoles live in shallow parts of the streams, where they cling to the 
surfaces of small rocks and hide beneath leaves and rocks. A complete de- 
velopmental series of tadpoles is not available; measurements of those stages 
examined are summarized in Table 12. 

A typical tadpole in stage 36 of development (KU 68475 from 15 km. 
WSW of San Isidro el General, Costa Rica) has a body length of 11.7 mm., 
tail length of 22.8 mm., and a total length of 34.5 mm.; body about three- 
fourths as deep as wide; snout broadly rounded in dorsal view, sloping and 
rounded in lateral view; eyes widely separated, directed dorsolaterally; nostril 
slightly closer to eye than to tip of snout; mouth ventral; spiracle sinistral, 
about two-thirds distance from snout to posterior end of body and slightly 
below mid-line; anal tube dextral; caudal musculature heavy, straight; dorsal 
fin not extending onto body; fins deepest at about mid-length of tail; there 
depth of caudal musculature equal to depth of dorsal fin and half again as 
deep as ventral fin; musculature extending nearly to tip of tail; body reddish 
brown above and pale grayish brown with white flecks below; caudal muscula- 
ture pale tan with brown flecks; a series of reddish brown dashes at base of 
caudal fin separated from others in series and from dashes on other side by 
creamy white; fins transparent with reddish brown flecks on posterior one- 
half of ventral fin and on all of dorsal fin (Fig. 14C). Mouth bordered by 
two rows of short, pointed papillae; lateral fold present; tooth-rows %; upper 
rows equal in length; second upper row narrowly interrupted medially; three 
lower rows complete, nearly as long as upper rows, deeply indented medially; 
upper beak robust, inner surface not forming continuous arch with short lateral 
processes; lower beak deep, V-shaped; both beaks bearing short serrations 
(Fig. 15F). 

Little variation occurs in structure. In some specimens the second upper 
tooth-row is complete; no individuals were fovmd to have the row broadly 
interrupted medially. 

The series of dark dashes on the dorsal edge of the caudal musculature is 
diagnostic of all stages studied. In life, tadpoles from 15 and 20 kilometers 
west-southwest of San Isidro el General, Costa Rica, had a tan body, often 



328 University of Kansas Publs., Mus. Nat. Hist. 

with an olive-tan tinge; the caudal musculature was tan; the flecks and dashes 
were dull red or reddish brown. Tadpoles from 6 kilometers east of Golfito, 
Costa Rica, had bodies with olive-green flecks. The caudal musculature was 
brown with bluish green flecks; the fins were transparent with reddish brown 
flecks. The belly was a silvery golden color. Tadpoles from Bajos de Jorco, 
Costa Rica, had browTi bodies with bluish green flecks; the tail and fins had 
reddish brown flecks and dashes. The iris was a bronze color in specimens 
from all three locahties, as well as in the young mentioned in the following 
paragraph. 

Nine recently metamorphosed young were found on vegetation at the edges 
of streams in April. These specimens have snout- vent lengths of 13.1 to 
15.7 mm. ( 14.9 mm. ) and in life were pale greenish tan or olive-tan above 
and white below. The hands, feet, and thighs were pale yellowish tan. 

Remarks. — The foregoing synonymies indicate that confusion has existed in 
the application of various names to this species, as well as in use of the names 
sordida and gabbi to include the species that we describe and name SmiUsca 
sila. Correct allocation of the names involved was possible only after studying 
and comparing the type specimens, for the descriptions given by the various 
authors are not sufficiently explicit to determine the nature of many essential 
features. 

The presence of a rounded snout and a long white throat in males distin- 
guishes S. sordida from S. sila, which has a high truncate snout and short dark 
throat in males. The two syntypes of Htjla sordida Peters, 1863, (ZMB 3141) 
are males having snout-vent lengths of 36.9 and 37.0 nun. The two syntypes 
of Hyla gabbi Cope, 1876 (USNM 30658-9), are females having snout-vent 
lengths of 52.8 and 53.7 mm., respectively. Also included in the collections 
made by Gabb is eastern Costa Rica are two males (USNM 30685-6), which 
Cope (1876) named and described as Hyla nigripes. These specimens are 
soft and faded, but are recognizable as the same as Hyla sordida Peters; the 
syntypes of Hyla nigripes have snout-vent lengths of 37.6 and 37.7 mm. We 
have examined one of the syntypes of Hyla salvini Boulenger, 1882 (BMNH 
1947.2.24.13), a female having a snout-vent length of 54.6 mm. We are 
convinced that all of these type specimens are representatives of one species, 
the earliest name for which is Hyla sordida Peters, 1863. The type localities 
for three of the named species are in Costa Rica — H. gabbi from Sipurio on 
the Caribbean lowlands, H. nigripes from the Caribbean slopes of Pico Blanco, 
and H. salvini from Cartago on the Meseta Central. The type locality of H. 
sordida was given as "Veraguas" by Peters (1863). At that time Veraguas 
was often considered to be most of western Panama. Though we have not 
seen Panamanian specimens other than the types of S. sordida and one speci- 
men from the Pacific lowlands of western Panama, the species probably occurs 
on the Caribbean slopes of western Panama. The species has been taken on 
the Caribbean lowlands of Costa Rica within a few kilometers of Panama; 
collecting on the Caribbean slopes in the provinces of Bocas del Toro and 
Veraguas should reveal the presence of Smilisca sordida there. 

Distribution. — Smilisca sordida is found along the Pacific slopes and low- 
lands from Guanacaste, Costa Rica, southeastward to extreme western Panama, 
to elevations of about 1200 meters on the Meseta Central in Costa Rica, and 
on the Caribbean slopes and lowlands of Costa Rica and probably adjacent 
Panama (Fig. 5). One specimen purportedly comes from "Rio Grande, 
Nicaragua." 



Neotropical Hylid Frogs, Genus Smilisca 



329 




10 10 20 50 



85' 



Fig. 5. Map showing locality records for Smilisca sordida. 

Specimens examined. — 412, as follows: Nicaragua: "Rio Grande" (? 
Depto. Zelaya), MCZ 2634. 

Costa Rica: Alajuela: Between Atena and Salto de San Mateo, USC 
6185; 8 km. N Ciudad Quesada, USC 7155 (4); La Fortuna, USC 7153 (20); 
3 km. E La Fortuna, USC 7150; San Carlos, USNM 29969; Sarchi, KU 32990-9, 
36792-3. 

Cartage: Cartago, BMNH 1947.2.24.13; headwaters of Rio Pacuare, USC 
119; Instituto Interamericano de Ciencias Agricolas, Turrialba, KU 37012, 
USC 420, 437; Rio Reventazon, Turrialba, MCZ 29268; 10 km. N Rio 
Reventazon bridge, USC 7073; 5 km. SW Rio Reventazon bridge on Paraiso- 
Orosi road, USC 669; Turrialba, UMMZ 118405, USC 455, USNM 29936-9. 

Heredia: Puerto Viejo, KU 36791. 

Guanacaste: Las Canas, USC 7164; Santa Cecilia, MCZ 7924-5; Tilaran, 
USC 7161 (5). 

Limon: Bambu, USC 7171 (2), 7183 (13); La Lola, USC 820 (6), 6083-94, 
8064, 8071; Pandora, USC 7188 (7), 7189, 7190 (3), 7191 (5); Pico Blanco, 
USNM 30685-6; Rio Lari, 14-16 km. SW Amubre, USC 7179, 7180 (10); 
Sipurio, USNM 30658-9; Suretka, KU 36764, 36765 (skeleton), 36766-78. 

Puntarenas: 6 km. N Dominical, KU 91749-50, 91811 (young), 91812 (tad- 
poles); Esparta, MCZ 8028; 6 km. E Golfito, KU 91718-41, 91809 (young), 
91810 (tadpoles), 91816-9 (skeletons), USC 7103 (23); Quebrada Agua Buena, 
3 km. SW Rincon de Osa, USC 7236 (6); Quebrada Boruca, 22 km. E Palmar 
Norte, KU 64264; Rincon de Osa, Camp Seattle, UMMZ 123680-5, S-2792 



4—3430 



330 University of Kansas Publs., Mus. Nat. Hist. 

(skeleton), USC 705 (5), 6023, 7254; Rio Barranca, USC 7119 (2); Rio Ceiba, 
6 km. NW Buenos Aires, KU 91747-8, USC 7112 (7); Rio Ciruelitas, 16 km. 
NW Esparta, USC 7121 (3); Rio Claro, 14.2 km. NW ViUa Neily, USC 7110 
(4); Rio Ferruviosa, 7 km. S Rincon de Osa, USC 7235 (4); Rio Lagarto at 
Pan-American Hwy. (Guanacaste Border), USC 7122 (4); Rio La Vieja, 30 km. 
E Palmar Norte, KU 87684 (tadpoles), 91743-6, USC 7083 (2); Rio Oro, 28.5 
km. NW Villa Neily, KU 91742; Rio Volcan, 10 km. W Buenos Aires, USC 
7113; Rio Zapote, 7 km. E Palmar, USC 7100 (4); 3-5 km. W Palmar, USC 7101 
(18); 7 km. SE Palmar, KU 64261-3; 1.2 km. NW Villa Neily, USC 8032; 
3 km. NW ViUa Neily, USC 7109 (20); 5 km. NW Villa Neily, USC 6176, 
8035. 

San Jose: Bajos de Jorco, KU 91813 (tadpoles); Escazu, KU 34863, 34869- 
75, use 813; between Monrovia and La Hondura, ± 0.5 km. N Santa Rosa, 
USC 302 (2); Paso Ancho, Rio Jorco, UMMZ 122649 (6), USC 530 (3); Rio 
Jorco, near Desamparados, KU 91757-65, 91796-7, 91820-3 (skeletons), USC 
228, 513, 7117 (7); Rio Peje, 10 km. SSE San Isidro el General, USC 7115 
(3); Rio Tirivi, MCZ 7972; San Isidro el General, CNHM 101096, KU 28201, 
32989, UMMZ 72024; 15 km. WSW San Isidro el General, KU 64245-56, 68473 
(tadpoles), 68474 (young), 68475 (tadpoles), 86516, 91754-6, 91793-5, USC 
7097 (6); 17.1 km. WSW San Isidro el General, USC 6047; 18 km. WSW San 
Isidro el General, USC 689; 20 km. WSW San Isidro el General, KU 64257-9, 
64260 (skeleton), 68468 (young), 68469 (tadpoles), 68470 (young), 68471-2 
(tadpoles), 68476 (young), 68633-4 (skeletons), 91751-3; San Jose, AMNH 
7501-4, USC 298; Santa Rosa, Rio Virilla, USC 7145. 

Panama: Chiriqui: Rio Jacu, 5.8 km. ESE Paso Canoas, KU 91905. 
"Veraguas," ZMB 3141 (2). 

ANALYSIS OF MORPHOLOGICAL CHARACTERS 

Osteology 

In attempting to assay the taxonomic significance of skeletal differences 
we are faced with a dearth of data on the skeletons of frogs in general and 
hylids in particular. Recent reviews by Brattstrom (1957) and Hecht (1962, 
1963 ) have been concerned with general saUentian classification and phylogeny, 
principally at the family level. Savage and Carvalho (1953), Griffiths (1959), 
and Baldauf (1959) used osteological characters in determining the taxonomic 
status of the families Pseudidae, Brachycephalidae, and Bufonidae, respec- 
tively. Carvalho (1954) presented osteological evidence for the generic 
separation of New World microhylids. Zweifel ( 1956 ) and Tihen ( 1962 ) 
used osteological characters at the levels of the species-group and species in 
their respective studies on Scaphiopus and Biifo. Little has been recorded 
about the skeletons of the hylids. Coin ( 1961 ) mentioned dentigerous ele- 
ments and cranial co-ossification in his synopsis of the genera of hylids. 
Copland (1957) in his review of the Hyla of Australia, Funkhouser (1957) 
in her revision of Phyllomedusa, and Zweifel (1958) in his review of Nyc- 
timystes did not consider skeletal characters. 

Some osteological studies on hylids have yielded worthwhile information. 
Mittleman and List ( 1953 ) used osteological characters in defining the genus 
Limnaoedtts: Starrett (1960) used cranial characters in combination with jaw 
musculature in defining the genus Smilisca, and Duellman (1964) used cranial 
characters in delimiting the Hylu bistincta group. Brief descriptions of cranial 
structure were given for Phrynohyas (Duellman, 1956) and Ptychohyla 
(Duellman, 1963a); specific and sexual differences in the skulls of Hyla 
chaneque and Hyla taeniopus were pointed out by Duellman (1965). Stokely 



Neotropical Hylid Frogs, Genus Smilisca 331 

and List (1954) described early cranial development in the hylid Pseudacris 
triseriata triseriata. 

Because our knowledge of the skeleton in hylids is so incomplete, we are 
not attempting to place Smilisca in the general scheme of hylid phylogeny on 
the basis of skeletal characters. Instead, our purposes are to describe the 
skeleton and its ontogenetic development in one member of the genus (S. 
baudini), and to make comparisons that show taxonomic differences in osteo- 
logical characters among species of Smilisca. 

The study of 68 dried skeletons and 25 cleared and stained preparations, 
including an ontogenetic series of S. baudini, has resulted in an understanding 
of the progressive development of skeletal elements and a knowledge of inter- 
specific and intraspecific variation in these elements. Furthermore, investiga- 
tions of the osteology have provided correlations between some cranial char- 
acters and certain aspects of external morphology. 

Descriptive Osteology of Smilisca baudini 

The following description is based primarily on an adult female (KU 
68184): 

Skull. — The skull is large, solid, and broader than long; the greatest width 
is between the sutures of quadratojugal and maxillary on either side of the 
skull (Pis. 2-3). The maxiUaries bear well-developed dorsal flanges, curve 
gently, join the moderately convex premaxillaries anteriorly and form a slightly 
truncate snout. The combined premaxillary width is about one-fourth the 
width of the skull. The premaxillaries are separated medially, and laterally 
from the maxillaries by sutures. Each premaxHlary bears a dorsomedial alary 
process, which is anteriorly convex and four times as high as the depth of the 
lateral wing of premaxillary; each premaxillary also has a ventromedial palatine 
process that projects dorsally from the lingual edge of the premaxillary. The 
septomaxillaries are closely associated dorsally with the premaxUlaries imme- 
diately lateral to the prenasal processes. 

The nasals are large, widest anteriorly and narrov^dng posteriorly, parallel to 
maxillaries, and not separated from the ethmoid by cartilage. The nasals bear 
long, dehcate maxillary processes extending nearly to the maxillaries. Anteriorly, 
the nasals are widely separated by the partially ossified internasal septum, 
which is in contact with the premaxillaries between the prenasal processes; the 
anterior points of the nasals lie approximately one-half the distance between 
the anterior ends of the ethmoid and the premaxillaries. The ethmoid is large 
and completely ossified; the margins are smooth. The trunate anterior edge 
lies between the nasals and is in contact with the internasal septum. The 
frontoparietals are large, smooth-margined, and bear large supraorbital flanges 
curving posterolateraUy at the rear of the orbit. A small, oval foramen in- 
volves the posterior part of the ethmoid and anterior portion of frontoparietals; 
continued ossification in older specimens fills in the foramen, thereby resulting 
in a solidly roofed cranium. The auditory regions are relatively massive and 
bear narrow tegmen tympani; the distal ends of the tegmen tympani are 
medial to the lateral edge of the pterygoids in dorsal view. The squamosals 
are large; the long anterior arm is separated from the maxillary by a suture. 
The dehcate, spindle-shaped columellae he ventral to the tegmen tympani and 
squamosals, are spatulate distally, and have a broad basal attachment to the 
auditory region. 

The vomers are moderately large and are in contact anteriorly with the 
premaxillaries and posteriorly with the ethmoid. Each vomer has two wide 
serrated flanges laterally. The tooth-bearing parts of the vomers are widely 
separated and at a slight angle to one another; the vomers terminate medially 
in two pointed processes on the ethmoid. The palatines are edentate, but 
bear strong ridges throughout their lengths. They are broadly in contact with 
the maxillary, are narrow medially, and are attached by pointed processes to 



332 



University of Kansas Fuels., Mus. Nat. Hist. 



the medial part of the ethmoid. The pterygoids are large, attached to the 
maxillaries immediately anterior and medial to the squamosal-maxillary con- 
nection, bear well-developed pedicles, which are broadly attached to the 
prootic, and a wide wing is in contact posteriorly with the distal two-thirds of 
the quadrate. 

The angular makes up most of the lower jaw, bears a broad articular sur- 
face posteriorly, and has a small coronoid process on the Ungual edge; anteriorly 
the angular is separated from the dentary and mentomecklian by Meckel's 
cartilage. The dentary lies external to the angular and extends from the 
mentomeckhan to approximately the mid-length of the angular. The mento- 
mecklians are ossified, but separated by cartilage medially. 

Hyaid. — The hyoid plate is curved, thin, and mostly cartilaginous, but 
calcified posteriorly (Fig. 6). The anterior cornua are slender, cartilaginous, 
and curve anteromedially from the hyoid plate and thence laterally and 
posteriorly, to attach to the posterior surface of the prootics. The lateral 
cornua are broad, flat, cartilaginous lateral extensions from the bases of the 
anterior cornua. The posterior cornua are bony, except distally. 




Gen. 



Fig. 6. Ventral view of hyoid apparatus of an adult male Smilisca 
battdini showing areas of muscle attachment: Gen, L., attachment of 
geniohyoideus laterahs; Gen. M., attachment of geniohyoideus medialis; 
Hyo., attachment of hyoglossus; Omo., attachment of omohyoideus; 
Fet., petrohyoideus; St., attachment of stemohyoideus. KU 64220, X5. 



Neotropical Hylid Frogs, Genus Smilisca 333 

Vertebral Column. — The atlas lacks transverse processes and a neural crest, 
whereas transverse processes are present on the other seven presacral vertebrae, 
and knobhke neural crests are present on the second, third, and fourth verte- 
brae; a faint neural ridge is visible on the fifth vertebra. The transverse 
processes are directed laterally on the second and sixth vertebrae, ventrolaterally 
on the third, posterolaterally on the fourth and fifth, and anterolateral^ on 
the seventh and eighth. The processes are sUghtly expanded on the fourth, 
and more so on the fifth, vertebra. The sacral diapophyses are expanded and 
have a border of calcified cartilage laterally. There are two sacral condyles. 
The slender coccyx has a thin dorsal ridge on the anterior three-foiuths of its 
length. 

Pectoral Girdle. — The omostemum is large, ovoid, and cartilaginous; the 
sternum is a thin cartilaginous sheet deeply notched posteriorly and is not dif- 
ferentiated into epistemal and xiphisternal elements. The coracoids are robust, 
twice as stout as the clavicles. The epicoracoidal cartilages overlap in the 
usual arciferal manner, except that they are fused anteriorly between the slender 
clavicles. The clavicles are strongly arched. The clavicle, coracoid, and 
scapula on each side form a bony articulation at the glenoid fossa. A bifurca- 
tion of the ventral end of the scapula results in a large glenoid foramen. The 
scapula is flat and expanded dorsaUy; the suprascapula is broad, flat, and 
calcified in large adults. In young specimens no distinct ossification of the 
cleithrum or ossification of endochondral centers are evident. 

Arm and Hand. — ^The humerus is equally well-developed in both sexes and 
has a prominent lateral crest. The radius and ulna are completely fused. A 
bony prepollex is present in both sexes. The metacarpals are about equal in 
length. The phalangeal formula is 2-2-3-3; the terminal phalanges are claw- 
shaped. 

Pelvic Girdle. — The ilia are long, slender, and slightly curved. A thin ridge 
projects laterally from the dorsal edge of the posterior one-half of each ilium. 
The ilial prominence is large and knobhke when viewed from above. The 
anterior edge of the ihal prominence is at the level of the anterior edge of the 
acetabular border. The dorsal acetabular expansion is small. The pubis is 
slender, and the ischium is elevated and robust. 

Leg and Foot. — The slightly curved femur has a distinct crest proximally 
on the posterior surface. The nearly straight tibio-fibula is shghtly longer 
than the femur. The tibial and fibial elements are completely fused but have 
a distinct cleft between them. A small foramen exists at the mid-length of 
the tibio-fibula. The fibulare (calcaneum) is much more robust than the 
tibiale (astragalus). The prehallux is large and flat. The metatarsals of the 
third, fourth, and fifth digits are equal in length; the metatarsal of the second 
is somewhat shorter, and that of the first is much shorter. The phalangeal 
formula is 2-2-3-4-3; the terminal phalanges are claw-shaped. 

Developmental Cranial Morphology of Smilisca baudini 

The following description of development of the skull of Smilisca baudini 
is based on the examination of 12 cleared and stained specimens. In table 3 
the cranial bones are listed in the left hand colunm in the approximate order 
of their appearance in the young frogs. Across the top of the table selected 
specimens designated by developmental stage or snout-vent length are listed. 
It should be noted that although each individual, from left to right, has an 
increasing number of ossified bones, the correlation with increasing size is 
imperfect; the precise ages of the individuals are unknown. 

The first bones to appear are the septomaxUlaries, frontoparietals, part of 
the exoccipital, and the parasphenoid in developmental stage 40. The fronto- 
parietals are represented by two slender ossifications dorsomedial to the orbits; 
the septomaxillaries are present as small ossifications anterior to the nasal 
capsules (PI. lA). The parasphenoid is present as a faint median ossification, 
and the exoccipital shows some ossification. 



334 



University of Kansas Publs., Mus. Nat. Hist. 



Table 3. — The Order of Occxjrrence of Cranial Ossifications in the 
Skuul of Smilisca baudini. Where Numbers Are Divided by a Slash 
Mark, the Left and Right Symbols Correspond to the Left and Right 

Sides of the Skull, Respectively. 



Bone 



Frontoparietal 

Parasphenoid 

Septomaxillaries 

Exoccipitals 

Squamosals 

Premaxillaries 

Maxillaries 

Nasals 

Pterygoids 

Vomers 

Palatines 

Quadratojugals 

Ethmoid 

Columellas 

Supraorbital Flanges. 

Prootics 

Vomerine Teeth 

Maxillary Teeth 

Premaxillary Teeth . . 



Stage 
40 



X 
X 
X 
X 



Stage 
44 



X 
X 
X 
X 
X 
X 
X 



12.6 
mm. 



X 
X 
X 
X 
X 
X 
X 
X 
X 



13.9 

mm. 



X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 



32.0 

mm. 



0/7 

2/4 



1/1 
3/5 
3/3 



4/3 
6/5 

5/5 



X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 



27.0 
mm. 



5/5 

30/31 

7/6 



X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

3/3 

30/26 

8/6 



20.1 
mm. 



X 
X 
X 
X 

X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
5/4 
37/36 
8/7 



The dentigerous bones are among the most rapidly developed, although 
not the first to appear. They are present in developmental stage 44 before 
metamorphosis is completed. The maxillaries bear a few teeth anteriorly and 
are ossified posteriorly to a point one-third of the distance from the anterior 
to the posterior edge of the orbit. Ossification lengthens the posterior termini 
of the maxillaries to the posterior edge of the orbit. In front of the anterior 
margin of the orbit, bone is proliferated dorsal to the main axes of the maxil- 
laries and forms moderate dorsal maxillary flanges. The premaxillaries appear 
simultaneously with the maxillaries. Initially they are widely separated 
medially from each other, and laterally from the developing maxillaries; each 
bears two or three teeth, large dorsally blunt alary processes, and small 



Neotropical Hylid Frogs, Genus Smilisca 



335 



palatine processes. The median and lateral edges of the prenasal processes 
lengthen heterochronously, causing the median edges to be longest and to lie 
slightly dorsal to the level of the septomaxillaries. After the maxillaries and 
premaxillaries develop, the vomers appear as small horizontal ossifications 
anterior to the parasphenoid. Ossification begins in the lateral flanges, then 
in the prevomerine processes, and lastly in the posterior dentigerous parts of 
the bones; the prevomerine processes are the last parts of the vomers to ossify 
completely. 

Initially the frontoparietals are present as tliin rods of ossification dorso- 
medial to the orbits; the frontoparietals extend from the anterior to the 
posterior end of the orbit by developmental stage 44. The anterior ends of 
the bones remain thin and pointed; ossification progresses medially from the 
midpoint of the length of the orbit and posteriorly to the level of the exoc- 
cipital; a median center of ossification joins the frontoparietals posteriorly, 
thereby forming the posterior border of the frontoparietal fontanelle. The 
supraorbital flanges of the frontoparietals do not appear until all other cranial 
bones are ossified, or nearly so. The most rapid ossification begins laterally 
at the posterior edge of the orbit and decreases anteriorly over the posterior 
half of the orbit. This differential rate of proliferation of bone results in the 
pattern of development of the supraorbital flanges sho\\Ti in figure 7. The 
nasals appear as tliin slivers of bone half way betvv^een tlie anterior ends of the 
frontoparietals and the end of the snout. As ossification proceeds the nasals 
assume a triangular shape in dorsal view. The anterior ends are pointed; 
the lateral margins are parallel to the maxillaries. The posteromedial points 
do not reach the lateral margins of the ethmoid, and the maxillary processes 
extend about three-fourths the distance from the bodies of the nasals to the 
maxillaries. Following the union of the frontoparietals posteriorly, the nasals 
widen anteriorly and are narrower at the midpoints of tlieir long axes than 
anteriorly or posteriorly. With further ossification the maxillary processes 
extend to the maxillaries and form complete bony anterior margins to the 
orbits; the mid-parts of the nasals widen (PI. IB). 




Fig. 7. Developmental sequence of the frontoparietal fontanelle and associ- 
ated bony elements in Smilisca baudini: (A) KU 60026, x5; (B) KU 85438, 
X4; (C) KU 26328, X3; (D) KU 68184, X2.3. 



The parasphenoid is the first of the palatal bones to appear. At metamor- 
phosis the bone is well developed; the anterior tip is situated just in front 
of the anterior edge of the orbit, and posteriorly the lateral processes extend 
laterally beyond the ossified parts of the auditory region. The pterygoids do 
not appear until metamorphosis, when ossification is evident in only the mid- 
parts of the posterolateral arms. Ossification follows in the mid-parts of the 



336 University of Kansas Publs., Mus. Nat. Hist. 

anterolateral arms and occurs last in the pterygoid pedicles. The palatines do 
not appear until all three arms of the pterygoids are at least partly ossified. 
Ossification proceeds rapidly from the maxillaries medially to the unossified 
ethmoid, which is the last of the cranial bones to appear. Initially it is ex- 
tremely shallow; dorsally it is widely separated from the nasals, and ventrally 
the posterior margin meets the anterior point of the parasphenoid. In dorsal 
view, ossification proceeds anteriorly between the nasals and posteriorly, ventral 
to the frontoparietals; ventrally, ossification proceeds posteriorly dorsal to the 
parasphenoid. 

The ventral arms of the squamosal and the supraoccipital region of the 
exoccipital are the first occipital bones to appear. Ossification follows in the 
regions of the semicircular canals and occipital condyles. The dorsal end of 
the ventral arm of the squamosal and the posterior arm of the squamosal 
ossify as a unit at the same time the quadratojugal appears. Shortly tliere- 
after the anterior arm of the squamosal ossifies, the distal part of the columella 
appears, and the anterior and lateral parts of the auditory region ossify. 

The angiJar and dentary of the lower jaw appear concurrently with the 
dentigerous bones. Initially, the angular is short and broad; the articular sur- 
face is absent, and the anterior end is slightly overlapped by the dentary. 
The mentomeckelians do not ossify until approximately the same time that the 
quadratojugal appears in the upper jaw. 

Comparative Osteology 

The genus Smilisca is characterized by the following combination of cranial 
osteological characters: (1) A large amount of bone is involved in the skull 
and a minimal amount of cartilage and/or secondarily ossified cartilage; co- 
ossification is absent. (2) The skulls are uniformly broad with angular lateral 
margins, and truncate anteriorly. ( 3 ) An intemasal septum and quadratojugals 
are present. (4) A well-developed squamosal minimally extends one-fourth the 
distance from the dorsal end of the quadrate to the maxillary, and maximally 
is separated from the maxillary by a suture. (5) The ethmoid is large; the 
distance between the anterior end of the ethmoid and the anterior edge of 
the premaxillary varies between 15 and 20 per cent of the total length of the 
skull. 

On the basis of cranial osteology two species-groups can be recognized 
within the genus Smilisca. The sordida group, comprising S. sordida and puma, 
is characterized by a broad skull in which the lateral margins of the maxillaries 
are relatively straight anterior to the orbit. The moderate-sized nasals are 
rounded anteriorly, and bear relatively short, sometimes blunt, maxillary 
processes. The long axes of the nasals are not parallel to the maxillaries. 
The ethmoid is proportionaely small in the sordida group. The bony part of 
the ethmoid terminates near the anterior edge of tlie orbits and does not 
extend anteriorly between the nasals; the entire anterior margin of the ethmoid 
is separated from the nasals by cartilage. The squamosals are generally small. 
They are narrow in dorsal view, and minimally extend one-fourth the distance 
from the dorsal end of the quadrate to the maxillary, and maximally, two- 
thirds the distance. The tegmen tympani are relatively small (Fig. 8). 

In contrast to the tendency for reduction of cranial parts in the sordida 
group, the baudini group, constituted by S. cyanosticta, phaeota, and baudini. 



Neotropical Hylid Frogs, Genus Smilisca 



337 



is characterized by more ossification of the cranial elements. The skull is 
broad; the lateral margins are less angular and are gently curved, rather than 
straight as in the sordida group. The nasals tend to be larger with the long 
axes parallel to the maxillar>'. Anteriorly the nasals are pointed, and posteriorly 





Fig. 8. Dorsal views of the skulls of the species of Smilisca: (A) S. haudini 
(KU 68184); (B) S. puma (KU 68636); (C) S. phaeota (KU 41090); (D) 
S. sih (KU 80625); (E) S. cyanosticta (KU 55938), and (F) S. sordida (KU 

36765). Xl.5. 



338 University of Kansas Publs., Mus. Nat. Hist. 

they bear long, delicate palatine processes extending to the maxillary. The 
ethmoid is fully ossified, extends anteriorly between the nasals, and laterally 
is separated by a suture from the nasals if the latter are fully ossified. The 
squamosals are large, and wide in dorsal view. They minimally extend one- 
fourth the distance from the dorsal end of the quadrate to the maxillary, and 
maximally are sutured to the maxillary. The tegmen tympani are massive. 

Smilisca sila is intermediate between the two species-groups described. The 
skull is broad; the lateral margins are gently curved, and have a pronounced 
angularity just anterior to the palatines which results in a broad, truncate 
snout. The nasals are moderate in size; because of the anterior angularity 
of the lateral margins, the long axes of the nasals lie parallel to the maxillary. 
The nasals are only slightly pointed anteriorly, and posteriorly they bear short, 
blunt palatine processes and medial processes in contact with the lateral 
corners of the ethmoid. The ethmoid is fully ossified, but does not extend 
anteriorly between the nasals. The squamosals are moderate in size and ex- 
tend one-fourth the distance from the dorsal end of the quadrate to the 
maxillary. The tegmen tympani are relatively large, but proportionately short. 

The cranial characters utilized in the analysis of species groups (general 
shape, nature of the nasals, ethmoid, squamosals, and tegmen tympani), to- 
gether with other characters, such as the relative height and shape of the 
prenasal processes, the extent of the internasal septum, and the nature of the 
vomers, frontoparietals, maxillaries and pterygoids are useful in distinguishing 
the various species (Table 4, Fig. 8), as well as in establishing relationships 
within the species-groups. 

Within the sordida group, S. sordida and S. puma can be distinguished by 
the following characters: Tlie bony part of the ethmoid terminates posterior to 
the anterior edge of the orbit and is thus widely separated from the nasals by 
cartilage in S. puma. In S. sordida the bony part of the ethmoid always 
terminates at a level equal to, or slightly in front of the anterior edge of the 
orbit; therefore, less cartilage exists between the ethmoid and nasals in S. 
sordida tlian in S. puma. The width of the premaxillary comprises about 30 
per cent of the width of the skull in S. sordida and 20 per cent in S. puma. 
The proportion of the length of the skull anterior to the bony part of the 
ethmoid in S. sordida is approximately 21 per cent, as compared with about 29 
per cent in S. puma. The prenasal processes are convex in S. sordida and 
straight in S. puma. 

The marked ontogenetic variation in S. sordida is considered in more detail 
in the account of that species, but it is pertinent to the present discussion to 
note that with respect to some features of the skull some young breeding speci- 
mens of S. sordida are intermediate in appearance between large females of 
S. sordida and adults of S. puma. In some breeding males (usually the 
smaller individuals) of S. sordida the bony part of the etlimoid terminates at 
the anterior edge of the orbit and is widely separated from the nasals by 
cartilage. In small individuals S. sordida, especially in males, and in adults 
of S. puma the tegmen tympani are relatively short, whereas in adult females 
of S. sordida these elements are long and slender. In the smaller specimens 
of S. sordida and in S. puma the squamosal is small; it extends only about one- 
fourth of the distance to the maxillary in the smaller S. sordida and about one- 
half the distance in S. puma. The more massive squamosal in large adult 
females of S. sordida extends at least two-thirds of the distance to the maxillary. 



Neotropical Hylid Frogs, Genus Smilisca 



339 






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University of Kansas Publs., Mus. Nat. Hist, 



Within the baudini group, the skull of S. cyanosticta is the most generalized 
of the three species; the cranial characters are intermediate between S. phaeota 
and S. baudini. The lateral margins of the skuU in S. cyanosticta are gently 
curved, and have an angularity anterior to the palatine-maxillary suture; the 
anterior margins are less angular in S, phaeota, which has a broader snout. 
Posteriorly in S. baudini the margins are shghtly curved medially, and the 
greatest width of the skuU is between the quadratojugal-maxillary sutures on 
either side of the skull. The frontoparietals of S. cyanosticta bear slightly 
irregular lateral margins and a large fontanelle. There is a tendency for ob- 
literation of the fontanelle with increasing age in both S. baudini and S. 
cyanosticta; the lateral margins of the frontoparietals bear large supraorbital 
flanges in both of these species. In S. phaeota the flanges are most prominent; 
they extend posterolaterally with straight margins along two-thirds of the 
length of the orbit and terminate in rather blunt points. The broad interorbital 
flanges result in a relatively broad external interorbital distance. In S. baudini 
the flanges are curved posterolaterally around the orbit and terminate in sharp, 
thin points. The tegmen tympani of all three species are massive. In S. 
cyanosticta the prootics slope posteriorly, whereas they slope anteriorly in S. 
baudini and S. phaeota. 

The skulls of S. cyanosticta and S. baudini are alike in certain respects. 
The squamosals of both species are large and connected to the maxillary by a 
bony connection; the squamosals of S. phaeota are large, but extend only two- 
thirds of the distance from the dorsal end of the quadrate to the maxillary. 
In S. baudini and S. cyanosticta the nasals are separated throughout their 
lengths from the ethmoid, whereas the nasals of S. phaeota are separated 
from the ethmoid by cartilage. The latter separation is due to an incomplete 
ossification of the nasals in S. phaeota. The bony part of each nasal is con- 
stricted in the middle of the long axis of the bone, and the nasals are widest 
anteriorly; posteriorly each nasal bears a medial process, which is narrowly 
separated from the lateral edge of the ethmoid. 

The teeth of all species of Smilisca are spatulate and bifid. The numbers 
of maxillary, premaxillary, and vomerine teeth are summarized in Table 5. 
Smaller and presumably younger specimens of all species of Smilisca have 
fewer teeth than do larger specimens of the same species. This correlation 

Table 5. — Variation in the Number of Teeth in the Species of Smilisca. 
( All Are Males; N = Number of Jaws, or Twtce the Number of Individ- 
uals; Means Are Given in Parentheses After the Observed Ranges.) 



Species 



S. baudini . . . 
S. cyanosticta 
S. phaeota . . . 
S. puma .... 

S. sila 

S. sordida . . . 



N 



20 
8 

20 
6 
8 

12 



Maxillary 



49-65 (56.0) 
50-64 (57.9) 
50-68 (58.1) 
60-67 (63.6) 
48-60 (52.9) 
39-55 (44.2) 



Premaxillary 



9-16 (13.6) 
10-12 (10.8) 
10-15 (12.1) 
11-13 (12.0) 
10-14 (11.3) 

7-11 (9.3) 



Vomerine 



5-9 (7.2) 

4-11 (7.1) 

5-9 (7.3) 

4-7 (5.3) 

5-7 (5.7) 

4-6 (5.2) 



Neotropical Hylid Frogs, Genus Smilisca 



341 



between size and number of teeth does not exist as an interspecific trend 
within the genus; for example, the smallest species in the genus, S. puma, 
has the highest number of maxillary teeth. In small specimens of a given 
species wide gaps are present between the maxillary teeth posteriorly; in 
large specimens the gaps are filled by teeth, beginning anteriorly and progress- 
ing posteriorly, until the maxillary dentition is continuous. 

Musculature 

No extensive study of the muscular system was undertaken, but certain 
muscles know to be of taxonomic importance were studied. 

Jaw Musculature. — Starrett (1960) pointed out the unique jaw musculature 
in Smilisca. In this genus M. depressor mandibulae consists of two parts, one 
arising from the dorsal fascia and one from the posterior arm of the squamosal. 
Two muscles arise from the anterior arm of the squamosal and insert on the 
lateral face of the mandible. Of these muscles, M. adductor mandibulae 
posterior subexternus lies medial to the mandibular branch of the trigeminal 
nerve; the other, M. adductor mandibulae extemus superficialis, hes lateral to 
the same nerve (Fig. 9). In most other hylids the latter muscle is absent. 
No significant variation in the position of the muscles was noted in the various 
species of Smilisca, though M. adductor mandibulae originate somewhat more 
anteriorly in S. baudini and S. cyanosticta than in the other members of the 
genus, all of which have a shorter anterior arm of the squamosal that does not 
reach the maxillary. The two separate parts of M. depressor mandibulae are 
not so widely separated in members of the sordida group as in the baudini 
group. 




D.Af. 



Fig. 9. Lateral view of the left jaw of Smilisca baudini; A. M. E. S., 
adductor mandibulae extemus superficialis; A. M. P. S., adductor mandib- 
ulae posterior subexternus; Col., columella; D. M. depressor mandib- 
ulae; M. S. T. N., mandibular branch trigeminal nerve; Sq., squamosal. 

KU 64214, X5. 



342 



University of Kansas Publs., Mus. Nat. Hist. 




Fig. 10. Ventral view of throat musculature in an adult male Smi- 
lisca baudini (Superficial musculature on left, deep musculatiure on 
right); A. C. anterior cornua of hyoid; Gen. L., geniohyoideus later- 
alis; Gen. M., geniohyoideus medialis; Hyo., hyoglossus; Omo., omos- 
ternum; Pet., petrohyoideus; S., submentalis; Sm., submaxillar is; St., 
stemohyoideus; V. S., vocal sac. KU 64220, x2.5. 

Throat Musculature. — The frogs that comprise the genus Smilisca are 
characterized by paired subgular vocal sacs, essentially the same as those in 
Triprion (Duellman and Klaas, 1964). The following description is based 
on Smilisca baudini (Fig. 10). 

M. submentalis lies in the anterior angle of the lower jaw, is thick, and 
consists of transverse fibers extending between the dentaries. M. submaxillaris 
is thin and arises from the whole of the inner surface of the lower jaw, except 
for the anterior angle occupied by M. submentalis. Anteriorly M. sub- 
maxiUaris is broadly attached by fascia to M. hyoglossus and M. geniohyoideus, 
which lie dorsal to M. submaxillaris. Medially this attachment continues 
posteriorly for about one-half the length of the hyoglossus. Posteriorly M. 
submaxillaris is folded and attached to M. sternoradiahs of the pectoral girdle. 
The vocal sacs are formed by a pair of posterolateral evaginations of M. 
submaxillaris; a broad connection between the pouches allows free passage 
of air between the pouches. 

The deeper throat musculature is essentially the same as that described 
for Phrtjnohtjos spilomma by Duellman (1956), except for slight differences 
in the place of attachment on the hyoid. 



SKIN 

Structure 

The skin of Smilisca is typical of that of most hyhds in organization and 
structure. Smilisca sila is distinguished from other members of the genus by 
the presence of small wartlike protrusions and peculiar white, pustular spots 
on the dorsum. The wartlike structures are composed of three or four 
epidermal cells, which protrude from the surface of the epidermis; the structures 
are covered by a slightly thickened layer of keratin. The white pustules are 
slightly elevated above the surrounding skin. Internally they consist of ag- 



Neo-eropical Hylid Frogs, Genus Smilisca 



343 



gregations of swollen, granular, pigment-cells (perhaps lipophores) lying be- 
tween the epidermis and the melanophores. 

Biochemical Variations 

Dried skins of all species of Smilisca were sent to Jose M. Cei, Institute 
Nacional de Cuyo, Mendoza, Argentina, for biochemical screening by means 
of the chromatographic techniques described by Erspamer and Cei (1963). 
The species in the baudini group have detectable amounts of penta-hydroxi- 
trypatamine, whereas only a trace is present in the other species. Furthermore, 
species in the baudini group differ from S. sila and the sordida group in lack- 
ing, or having only a trace of, tryptophan-containing polypeptides. These 
superficial biochemical tests support the arrangement of species as ascertained 
by conventional taxonomic characters. 

External Morphological Characters 

The features of external morphology tliat were studied in connection with 
the taxonomy of the genus Smilisca are discussed below. 



Size and Proportions 

The frogs of the genus Smilisca are medium to large tree frogs. The three 
species comprising the baudini group (S. baudini, cyanosticta, and phaeota) 
are notably larger than S. puma, sila, and sordida (Table 6). The largest 
specimen that we examined is a female of S. baudini having a snout- vent 
length of 90 mm. Smilisca puma is the smallest species; the largest male has 
a snout-vent length of 38 mm. and the largest female, 46 mm. 

No outstanding differences in proportions exist between species, although 

Table 6. — Compakison of Sizes and Certain Proportions of the Species 
OF Smilisca. ( Means in Parentheses Below Observed Ranges; Data for 

Males Only.) 



Species 


N 


Snout-vent 
length 


Tibia length/ 
snout-vent 


Tympanum/ 
eye 


S. baudini 


140 


47.3-75.9 
(58.7) 


42.1-53.6 

(47.8) 


56.1-94.4 




(73.5) 


S. cyanosticta 


40 


44.6-56.8 
(50.7) 


51.9-59.7 
(56.0) 


62.7-88.4 




(71.4) 


S. phaeota 


50 


40.8-65.5 
(53.9) 


50.9-60.2 
(55.5) 


62.7-85.5 




(76.6) 


S. puma 


20 


31.9-38.1 
(34.7) 


48.2-53.1 
(51.3) 


52.1-72.2 




(64.9) 


S. sila 


33 


31.6-44.8 
(37.7) 


49.7-58.1 
(54.8) 


47.6-58.3 
(53.2) 


S. sordida 


55 


31.9-44.6 
(37.9) 


50.5-57.1 
(53.4) 


46.5-57.1 
(49.1) 



344 University of Kansas Publs., Mus. Nat. Hist. 

certain proportions are sufficiently different in some species to warrant men- 
tion. Smilisca baudini is a more squat and stocky frog than other members of 
the genus; this is reflected in the somewhat shorter hind legs (Table 6). The 
size of the tympanum relative to that of the eye is highly variable within 
samples of a given species. Even so, noticeable differences in the tympanum/ 
eye ratio are apparent. Members of the baudini group have the largest 
tympani, whereas S. sila and sordida have the smallest, and S. puma is inter- 
mediate (Table 6). 

Shape of Snottt 

Although all members of the genus have rather truncate snouts, subtle dif- 
ferences exist among the species (Pi. 12). Smilisca sila has the shortest snout; 
that of S. baudini is only sHghdy longer. The snouts of S. cyanosticta and 
puma are nearly square in lateral profile, whereas those of S. phaeota and 
sordida are slightly inclined. The shape of the snout is relatively uniform 
within each species and displays no noticeable sexual dimorphism, except in 
S. sordida, in which there are sexual differences and geographic variation 
(seep. 324). 

Hands and Feet 

The characters of the hands and feet are among the most taxonomically 
important external features in Smilisca. Consistent differences exist in relative 
lengths of the digits, size of subarticidar tubercles, size and mmiber of super- 
numerary tubercles, size and shape of the inner metatarsal tubercle, and 
amount of webbing ( Pis. 4 and 5 ) . In the baudini group the series of species 
(baudini-pJiaeota-cyanosticta) show a progressive increase in amount of web- 
bing in the hand and a decrease in number, and corresponding increase in 
size, of supernumerary tubercles. The amount of webbing in the feet of 
S. baudini and phaeota is about the same, but the webbing is slightly more 
extensive in S. cyanosticta. Smilisca puma is unique in the genus in lacking 
webbing in the hand; furthermore, this species is distinctive in having many 
large subarticular tubercles on the hand and a relatively small inner metatarsal 
tubercle. The two stream-inhabitants, S. sila and sordida, have shorter and 
stouter fingers than the other species. The webbing is most extensive in both 
the hands and feet of these species, which also are distinctive in having many 
small supernumerary tubercles on the feet. 

Ontogenetic Changes 

Minor ontogenetic changes in structure involve the shape of the snout, 
relative size of the eye, development of the tympanum, and amount of web- 
bing in the hand. In recently metamorphosed young the snout is more 
rounded than in adults; the canthus and loreal concavity are not evident. 
Usually the tympanum is not differentiated in recently metamorphosed young, 
and the eye is proportionately large. The webbing in the feet is completely 
developed at metamorphosis, but young individuals have noticeably less web- 
bing in the hand than do adults of the same species. 

Coloration 

Some of the most distinctive characters of the species of Smilisca are color 
and pattern of the living frogs. Although many chromatic features are lost or 
subdued in preserved specimens, the patterns usually persist. 



Neotropical Hylid Frogs, Genus Smilisca 345 

Metachrosis 

Change in color, well known in frogs, is common in hylids, especially in 
species having green dorsal surfaces {Phyllomedusa is a notable exception). 
The non-green Smilisca {puma, sila, and sordida) changes color, but this 
mostly is a change in intensity of color. In these species the markings usually 
are most distinct at night; frequently by day the frogs become pallid. The 
most striking examples of metachrosis in Smilisca are found in the baudini 
group, in which the dorsal ground-color changes from green to tan; correlated 
with the change in ground-color may be a corresponding change in the dorsal 
markings, but the dorsal markings may change to the opposite color. 

Chromosomes 

Chromosomes of all six species of Smilisca were studied by means of the 
propriono-orcein squash technique described by Duelhnan and Cole (1965). 
Karyotype analysis was attempted for several species by means of intraperi- 
toneal injections of colchicine, which affected the mitotic cells as desired, but 
the testes examined contained too few mitotic cells to allow accurate determina- 
tion of karyotv'pes. 

Haploid ( n ) chromosome numbers were determined from cells in diakinesis, 
metaphase I, and metaphase II of meiosis. Diploid (2n) chromosome num- 
bers were determined from cells in late prophase and metaphase of mitosis. 
Chromosome counts from as few as 23 meiotic cells of S. phaeota and as many 
as 80 cells of S. sordida reveal a constant haploid (n) number of 12; counts 
of chromosomes in one to five mitotic cells in all species, except S. sila, reveal 
that the diploid (2n) number is 24. 

NATURAL HISTORY 

Breeding 

Like most hylid frogs Smilisca is most readily collected and observed when 
individuals congregate for breeding. 

Time of Breeding 

Smilisca breeds primarily in quiet water and reaches its height of breeding 
activity at times of plentiful rainfall, — usually from May through October. 
Through most of its range Smilisca baudini breeds in those months, but in 
some places where abundant rain falls in other seasons, the species breeds at 
those times. For example, in southern El Peten and northern Alta Verapaz, 
Guatemala, Smilisca baudini has been found breeding in Febmary and March. 
The other pond-breeding species (S. cyanosticta, phaeota, and puma) live in 
regions lacking a prolonged dry season, and possibly they breed throughout 
the year, but breeding activity seems to be greatest in the rainiest months. 

The two stream-breeding species (S. sila and sordida) breed in the dry 
season when the streams are low and clear, principally in December through 
April. At high elevations the species sometimes breed in the rainy season; 
also, individuals sometimes breed in the short dry season (summer canicula) 
in July and August. 

5—3430 



346 University of Kansas Publs., Mus. Nat. Hist. 

At several localities species have been found breeding at different times of 
the year: S. baudini in March and July at Chinaja, Guatemala; S. phaeota in 
April and August at Palmar Sur, Costa Rica; S. puma in February and July 
at Puerto Viejo, Costa Rica; and S. sila in February, April, and August at 
El Volcan, Panama. These observations indicate only that the population 
breeds at more tlian one time in the year, but do not provide any evidence on 
the breeding cycles of the individual frogs. This is one important aspect of 
the natiual history of Smilisca for which we lack data. 

Breeding Sites 

All members of the genus Smilisca presumably deposit their eggs in water. 

Smilisca baudini usually breeds in temporary rain pools; often these are 
nothing more than shallow, muddy puddles. In other instances the sites are 
extensive ditches or large flooded areas (PI. 8, Fig. 1). This species is an 
opportunistic breeder, and males gather at any of a wide variety of suitable 
breeding sites that are formed by torrential rains in the early part of the 
rainy season. Smilisca baudini nearly always breeds in open pools having bare 
earthen edges. Frequently congregations of S. baudini are found at such 
small pools, but are absent from nearby large ponds surrounded by vegetation. 

Little is known of the breeding habits of S. ajanosticta, which inhabits 
humid forests on foothills and lowlands. Apparently its breeding sites are 
not unlike those of S. phaeota, which usually are pools surroimded by vegeta- 
tion (Pi. 8, Fig. 2), although sometimes males of S. cyanosticta call from 
open muddy puddles. In uplands, where standing water is uncommon, this 
species breeds in quiet pools in streams. 

Smilisca puma breeds in grass-choked ponds and marshes, where the males 
call from bases of dense clumps of grass in the water ( Pi. 9, Fig. 1 ) . 

Smilisca sila and S. sordida differ noticably from other species in the genus 
by breeding exclusively in streams, where males usually call from rocks or 
gravel bars in or at the edges of streams (Pi. 9, Fig. 2); sometimes individuals 
perch on bushes overhanging streams. In the streams, or parts of streams, 
utilized by these frogs the water is clear, shallow, and has a slow gradient; 
occasional males have been found calling along cascading mountain streams. 

Breeding choruses composed of ten or more species of frogs are not un- 
common in Middle America, but Smilisca usually breeds alone or with one 
or two other species and at the most five others. This tendency towards soli- 
tary breeding possibly is the result of selection of breeding sites that are 
unsuitable to many other species of frogs. Nevertheless, many other species 
of frogs have been found at the breeding sites with the various species of 
Smilisca; these breeding associates (Table 7) are most numerous for S. 
baudini, which has a broad geographic range, including a variety of habitats. 

Breeding Behavior 

Calling sites. — All species of Smilisca usually call from the ground, including 
rocks and gravel bars; some individuals sit in shallow water near the edge of 
the pool or stream. Sometimes males of S. baudini, sila, and sordida call from 
low bushes or trees near the breeding site. One S. baudini was observed call- 
ing while it was floating on the surface of a pond. Smilisca cyanosticta, 
phaeota, and puma call from secluded places at the edge of the water or in 
the water, whereas S. baudini, sila and sordida call from open situations. 



Neotbopical Hylid Frogs, Genus Smilisca 347 

Table 7. — Breeding Associates of the Vabious Species of Smilisca, 



Associate 





e 




















Vi 










CO 


e 






•f* 




o 


C3 




"a 


e 




S 


9 


e 


^3) 


n^ 


s 




>o 


W 


A 


R. 


CO 


QQ 


CO 


CO 


^ 


CO 



O 



Rhinophrynus dorsalis 

Leptodadylus holivianus ... 

Leptodadylus labialis 

Leptodadylus melanonotus . . 
Leptodadylus occidentalis . . . 
Leptodadylus quadrivitlatus . 
Leptodadylus pentadadylus . 
Engystomops puslulosus . . . . 

Bufo canalifertis 

Bufo cavifrons , 

Bufo coccifer 

Bufo coniferus 

Bufo cristatus , 

Bufo gemmifer 

Bufo haematiticus 

Bufo kellogi 

Bufo luetkeni 

Bufo marinus 

Bufo marmoreus 

Bufo mazatlanensis 

Bufo melanochloris 

Brifo perplexus 

Bufo typhonius 

Atelopus varius 

Diaglena reticulata 

Diaglena spatulata 



X 
X 
X 



X 
X 

X 



X 
X 
X 
X 
X 

X 



X 
X 



X 



X 



X 
X 
X 



X 
X 
X 



X 



X 



X 



X 



X 



X 



X 



X 



X 



X 



X 
X 



X 



X 



348 University of Kansas Publs., Mus. Nat, Hist. 

Table 7. — Continued 



Associate 





e 




















u 








s 




S 






■'^ 


o 


o 


o 




§ 


S 


s 


S 


o 


o 


s> 


■« 


s 




rO 


C 


?5, 


?>. 


« 


=Q 


CQ 


=C 


00 


CQ 



e 
■a 

V. 



Hyla boulengeri 

Hyla colymba 

Hyla ebraccata 

Hyla elaeochroa 

Hyla eximia 

Hyla legleri 

Hyla microcephala 

Hyla phlebodes 

Hyla pida 

Hyla robertmertensi . ... 

Hyla rosenbergi 

Hyla rufioculis 

Hyla smithi 

Hyla staufferi 

Hyla walkeri 

Phrynohyas inflata 

Phrynohyas spilomma . . . 
Phrynohyas venidosa . . . . 
Phyllomedusa callidryas . 
Phyllomedusa dacnicolor . 
Phyllomedusa moreleti. . . 

Pternohyla fodiens 

Smilisca baudini 

Smilisca cyanostida 

Smilisca phaeota 

Smilisca puma 



X 



X 

X 
X 



X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 



X 



X 
X 



X 



X 



X 
X 



X 



X 



X 



X 



X 



X 



X 



Neotropical Hylid Frogs, Genus Smilisca 
Table 7. — Concluded 



349 



Associate 


s 




s 

1 


a. 


S. 






Smilisca silu 


X 

X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 


— 


X 

X 
X 


X 


X 
X 

X 
X 

X 


X 


Smilisca sordida 


X 


Triprion petasatus 




Cochranella Heischmanni 


X 


Cenirolene vrosoblevon 




Gastrophryne elegans 




Gastrophryne olivacea 




Gaslrovhrvne usta 




Hypopachus alboventer 




Hypopachus caprimimus 




Hypopachus inguinalis 

Hypopachus macidatus 

Hypopachus oxyrrhinus 

Hypopachus variolosus 

Rana palmipes 

Rana pipiens 

Rana warschewitschi 


X 



chorus structure. — Limited observations on some of the species of Smilisca 
show a definite organization of the calUng behavior of individuals. Smilisca 
baudini and S. phaeota call in duets. This is especially noticeable in S. baudini, 
in which the members of a duet often call from sites separated by only a few 
centimeters. The call of S. baudini consists of a series of like notes (see 
description of call in following section); the duration of each note is about 
equal to the interval between notes. Normally one individual utters one note, 
pauses, and utters a single note again, or series of two or three notes. If 
there is no response, the first individual often waits several seconds or even 
several minutes and then repeats the call. The second individual usually 
responds after the first or second note of the sequence. The notes of the second 
individual usually are spaced so that they are emitted in the intervals between 
the notes of the first individual. This can be shovvTi diagrammatically by having 



350 University of Kansas Publs., Mus. Nat. Hist, 

the figure "1" represent notes of the first individual and figiu-e "2," the notes 
of the second; an empty interval is represented by "0": 

1-0-1-2-1-2-1-2-1-2-1-2 
Usually a chorus is initiated by one duet and is quickly picked up by other 
individuals also calling in duets. A numerical representation of a chorus of 
eight frogs would approximate the following organization: 
l_0-l-2-l-2-l-2-l-2-l-2-l-2 

3-0^^-^^-3-4-3-4-3-4-3 
5_6_5_6-5-6-5-6-5-6-5-6 
7-8-7-8-7-8-7-8-7-8-7-8 

After the first one or two duets are initiated, the second individuals in 
the following duets usually call immediately after their respective partners 
have given the first notes. The other noteworthy aspect about the organiza- 
tion is that the entire chorus usually stops abruptly. Normally the first duet 
stops calling shortly before the others, but this is not invariable. Often one 
duet or one individual will emit several notes after the rest of the frogs have 
become silent. An interval of several minutes sometimes elapses before the 
chorus begins again. Successive choruses apparently are initiated by the same 
duet. Responses can be initiated artificially by imitating the call, and some- 
times any loud noise will start a chorus. 

Similar duets have been observed in S. piiaeota. In this species the in- 
tervals are often much longer than the notes, and if two males are calling in 
close proximity, their calls can be mistaken for tliose of one individual. 
Smilisca phaeota does not congregate in large numbers; usually only two males 
call from one restricted site. 

Smilisca sila has a call consisting of a primary note followed by one or more 
secondary notes. Males often call in duets, but not necessarily so. In a duet, 
the first male usually utters only primary notes until the second individual re- 
sponds; then each individual produces a rapid series of secondary notes. 

Smilisca puma also produces primary and secondary notes. Although in- 
dividuals sometimes call alone, duets, trios, or quartets were more common. 
The chorus is initiated by one individual uttering primary notes until joined by 
the second, third, and fourth frogs. In one quartet in a marsh 7.5 kilometers 
west of Puerto Viejo, Costa Rica, on February 19, 1965, the same individual 
initiated four consecutive choruses. Each time the second member of the 
chorus was the same; the third and fourth frogs joined the chorus nearly 
simultaneously. 

Individuals of S. sordida are usually irregularly situated along a stream. 
No duets or other combinations of individuals are apparent in the chorus 
structure, but once an individual calls, a frog nearby calls almost unmediately; 
then a frog near the second individual calls, and so on. The resulting series 
of calls gives the impression that the sound is moving along the stream as suc- 
cessive individuals join the chorus and the first callers become quiet. It is 
not known if the same individual initiates successive choruses or if the order 
of calling is the same in subsequent choruses. 

These limited observations on chorus structure in Smilisca show the presence 
of behavioral organization. The methods of establishing the organization and 
the significance of the call-order in breeding have yet to be discovered. 

Calling males of S. baudini are often close together; some individuals have 
been observed almost touching one another, but no indication of territoriality 



Neotropical Hylid Frogs, Genus Smilisca 351 

or aggressive behavior has been witnessed. The more distant spacing of the 
stream-breeding species S. sila and S. sordida may be a function of calHng- 
territories, but no direct evidence is available to substantiate this supposition. 

Sex recognition and amplexus. — Observations on Smilisca baudini indicate 
that the calls of males attract females. At Tehuantepec, Oaxaca, Mexico, a 
female was first observed about two meters away from a male calling at the 
edge of a rain pool; in a series of short hops she progressed directly towards 
the male, although vegetation obscured him until she was less than a meter 
away. When she approached to within about 20 centimeters of the male, 
he took notice of her, moved to her, and clasped her. At Chinaja, Alta 
Verapaz, Guatemala, a female swam directly across a pool about three meters 
wide to a calling male. Her line of movement took her within a few centi- 
meters of a silent male, to whom she paid no attention. She stopped just in 
front of the calling male, which immediately clasped her. At a large muddy 
pond 4 kilometers west-northwest of Esparta, Puntarenas, Costa Rica, a fe- 
male was observed swimming toward a small submerged tree; a male was 
calling from a branch about one meter above the water. The female climbed 
to a branch about 20 centimeters below the male, which upon seeing her 
there immediately jumped down and clasped her. These few observations of 
S. baudini show that in this species females are capable of locating calling 
males by means of phono-orientation; visual reception on the part of females 
seems to be secondary. Contrariwise, males apparently become aware of the 
proximity of females by seeing them; once a male sees a female he usually 
tries to clasp her. Possibly the males receive stimuli by means of chemo- 
reception, but in each observed instance the male obviously looked at the 
female. 

Amplexus is axillary in all members of the genus. Normally amplexing 
males hunch their backs and press their chins to the females' backs. Clasp- 
ing pairs are usually found at the edge of the water, but sometimes amplexus 
takes place in trees or bushes. 

Egg deposition. — Oviposition has been observed only in Smilisca baudini. 
On the night of June 28, 1961, at Chinaja, Alta Verapaz, Guatemala, a clasp- 
ing pair was observed at the edge of a shallow rain p>ool. After sitting for 
several minutes in shallow water, the female (with male on her back) swam 
part way across the pool and grasped an emergent stick with one hand. The 
female's body was nearly level with the surface of the water, and her hind 
legs were outstretched as deposition commenced; eggs were extruded rapidly. 
After a few seconds the female moved slowly to another twig a few centi- 
meters away and deposited more eggs. This process was repeated until the 
female was spent. The spawn resulted in a surface film covering roughly one 
square meter. It is doubtful if this type of egg deposition occurs in any other 
species in the genus, especially those that lay their eggs in streams. 

Breeding Call 

The breeding calls of the six species of Smilisca are alike in their explosive 
nature. Calls are emitted quickly vdth a short burst of air filling the vocal 
sac, which immediately deflates. Phonetically the calls can be described as a 
single "wonk" or series of such notes in S. baudini and S. cyanosticta, a low 
growl in S. phaeota, a relatively high pitched rattle in S. sordida, and a low 



352 



University of Kansas Publs., Mus. Nat. Hist. 



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Neotropical Hylid Frogs, Genus Smilisca 353 

squawk usually followed by one or more rattling secondary notes in S. puma 
and S. sila. Quantitatively, the calls of the six species differ in number of 
notes, duration of notes, and in pitch (Table 8, Pis. 10 and 11). Although 
no measurements were taken on the intensity of the calls, we observed in the 
field that each of the species has a loud voice. The call of S. baudini seems 
to carry farther than any of the others. 

Call rate. — The rate at which call-groups are produced varies from one 
every few seconds to one in several minutes. In S. baudini, cyanosticta, phae- 
ota, and sordida, call-groups are produced as frequently as every 12 seconds, 
but usually more time elapses between call groups. In S. sordida, five or more 
minutes sometimes elapse between call-groups. The interval is somewhat less 
in S. phaeota. Calls are repeated at much shorter intervals in S. puma (5-55 
seconds) and S. sila (4-20 seconds). 

Notes per call-group. — Except for S. puma and S. sila, the series of notes 
produced in any given call of a species of Smilisca is essentially the same; 
there is no differentiation into primary and secondary notes. Smilisca cya- 
nosticta and S. phaeota emit only one or two relatively long notes per call- 
group, whereas S. baudini and S. sordida produce as many as 15 and 6 notes, 
respectively. Males of S. puma and S. sila often produce only the primary 
note; sometimes this is done several times before the secondary notes are pro- 
duced. For example, one S. puma (KU 91711; tape No. 379) produced the 
following number of notes in consecutive call-groups: 1, 1, 1, 1, 2, 2, 3, 1, 4; 
secondary notes are present in only four of the nine call-groups. A typical 
series of consecutive call-groups in S. sHa (KU 91852; Tape No. 385) has 
1, 1, 1, 2, 4, 2 notes per call-group; secondary notes are present in only half 
of the call-groups. Smilisca puma apparently always produces at least two 
primary notes before emitting secondary notes; sometimes only primary notes 
are produced in one series of calls. The number of secondary notes following 
a given primary varies from one to nine; the modal number is one, and the 
mean is three in 27 call-groups. Smilisca sila frequently begins a series of calls 
with two or more primary notes, but sometimes the first primary note is fol- 
lowed immediately by two or more secondary notes. The number of secondary 
notes following a given primary varies from one to five; the modal number is 
one, and the average is two in 13 call-groups. 

Duration. — The average duration of call-groups consisting of two or more 
notes is 1,18 seconds in S. baudini; 1.02 in cyanosticta, 0.91 in plxaeota, 1.32 
in puma, 1.48 in stki, and 1.29 in sordida. Although there is considerable 
variation ia the lengths of the notes (only primary notes in S. puma and sila 
are considered here), S. cyanosticta, phaeota, and sordida have noticeably 
longer notes than do the other species (Table 8). The secondary notes are 
longer than the primary notes in S. puma (average 0.27 sees, as compared with 
0.13 sees.) and in S. sila (average 0.25 sees., as compared with 0.16 sees.). 

Note repetition rate. — The rate at which notes in call-groups containing two 
or more notes are produced varies in S. baudini from 2.5 to 7.1 (average, 3.7) 
calls per second; cyanosticta, 1.8-2.1 (1.9); phaeota, 2.0-2.4 (2.2); puma, 
1.9-2.9 (2.2); sila, 1.3-2.4 (1.8); and sordida, 1.5-2.6 (2.1). Smilisca baudini, 
which has notes of short duration (0.09 to 0.13 seconds), has the fastest note- 
repetition rate. Although the individual notes of S. cyanosticta and S. phaeota 
are relatively long (average, 0.38 and 0.31 seconds, respectively), the intervals 



354 University of Kansas Publs., Mus. Nat. Hist. 

between the notes is short; consequently, their note-repetition rates do not 
differ greatly from those of S. puma and S. sila, which have shorter notes (av- 
erage, 0.13 and 0.16 seconds, respectively) but longer intervals between notes. 
Pulse rate. — Pulses vary in frequency from 78 to 240 per second in the calls 
analyzed (only primary notes in S. puma and S. sila), but the variation in any 
given species is much less than that in the entire genus (Table 8). Smilisca 
puma is outstanding in having a high pulse rate, which is approached only by 
that of S. baudini. Even in the species having the lowest pulse rates, the 
pulsations are not audible. The secondary notes produced by S. puma and 
S. sila have a slower pulse rate than the primary notes; often the pulses are 
audible. In S. puma the pulse rate of secondary notes is sometimes as low as 
48 pulses per second, and in S. sila still lower (as low as 40 pulses per second). 
The upper limits of pulse rate in the secondary notes in these species merge 
imperceptibly with the rates of the primary note; consequently, on the basis 
of pulse rate alone it is not always possible to distinguish primary from sec- 
ondary notes. 

Frequency. — Smilisca produces noisy (as opposed to more musical) calls, 
and the energy is distributed throughout the frequency spectrum; the calls are 
poorly modulated, except in S. sordida, in which two usually discrete bands of 
frequency are present (Pi. IIC). For the most part the calls of Smilisca con- 
sist of little modified energy of the fundamental frequency and of its har- 
monics, some of which are emphasized. 

The upper frequency range varies within each species and even within the 
calls of one individual. Smilisca phaeota has the lowest upper frequencies; 
no calls ranged above 4400 cycles per second (cps. ), and half of the calls 
never exceeded 3000 cps. Smilisca cyanosticta produces calls in which the 
upper frequency is below 7000 cps. and usually below 6000 cps. Likewise, 
S. puma produces calls that are below 7000 cps., whereas S. sila has frequen- 
cies of up to 8400 cps. In both S. baudini and S. sordida, the highest fre- 
quencies attained are about 9100 cps. Variation in the highest frequencies in 
a series of consecutive calls by one individual frog was noted in all species. 
Such variation is especially prevalent in S. puma; for example one individual 
(KU 87771; Tape No. 376) recorded at a temperature of 24° C. at 7.5 kilome- 
ters west of Puerto Viejo, Heredia Province, Costa Rica, on July 31, 1964, 
produced three consecutive primary notes having upper frequencies of about 
6000, 4000, and 4000 cps., respectively. Apparently in a given species the 
production of the higher frequencies in some notes and not in others is cor- 
related with the amount of distention of the vocal sac and is not dependent 
upon the structure or tension of the vocal cords. 

Although the dominant frequency in S. sordida is lower than that in S. 
baudini and S. cyanosticta, the call of the former is audibly higher-pitched. 
This is due primarily to the emphasis on certain harmonics at a high frequency 
(sometimes as high as 9000 cps.) in S. sordida, whereas in S. baudini and 
other species, if harmonics are present at those frequencies, they are not em- 
phasized. 

The fundamental frequencies are as low as 90 cps. in S. sila and S. sordida 
and as high as 200 cps. in S. puma (Table 8). The fundamental frequency 
seemingly is relatively unimportant in determining the general pitch of the call, 
a characteristic most dependent on the dominant frequency and emphasized 
harmonics in the higher-frequency spectrum. In none of the species is the 



Neotropical Hylid Frogs, Genus Smilisca 355 

fundamental the dominant frequency. In the low-pitched call of S. phaeota 
the dominant frequency is the third harmonic (the second harmonic above the 
fundamental frequency, which is the first harmonic). In all other species a 
much higher harmonic is dominant; for examples, in S. cyanosticta harmonics 
from 10 to 15 are dominant; in S. baudini, 15-19; and S. sUa, 20-30. 

A glance at the audiospectrographs and their accompanying sections (Pis. 
10 and 11) reveals the presence of two emphasized bands of frequency in all 
species except S. phaeota, in which only the lower band is present. These two 
bands of emphasized harmonics are part of a continuous, or nearly continuous, 
spread of energy throughout the frequency spectrum, except in S. sordida in 
which the bands are usually distinct. As shown in the sections, certain har- 
monics in each of the bands are emphasized with nearly equal intensity. 
Therefore, with the exception of S. pJmeota, the calls of Smilisca are character- 
ized by two major frequencies, one of which is the dominant frequency and 
the other is a subdominant frequency (Table 8). The upper major frequency 
is dominant in all calls in S. baudini and S. cyanosticta, but either major fre- 
quency may be dominant in other species. The upper major frequency is 
dominant in 65 per cent of calls by S. puma, 87 per cent in S. sila, and 68 per 
cent in S. sordida. Individuals of these three species sometimes produce a 
series of calls in which the dominant frequency changes from one of the major 
frequencies to the other. Four consecutive notes emitted by an individual of 
S. sordida recorded 13 kilometers east-northeast of Golfito, Puntarenas Prov- 
ince, Costa Rica, had dominant frequencies of 910, 1950, and 750 cps., re- 
spectively. In each case, an alternation of major frequencies took place in 
respect to dominance. An individual of S. puma from 7.5 kilometers west of 
Puerto Viejo, Costa Rica, produced a primary note followed by one secondary 
note; each note had major frequencies at 600 and 1800 cps.; the dominant 
frequency of the primary note was at 1800 cps., whereas in the secondary note 
the dominant frequency was at 600 cps. The difference in emphasis on the 
major frequencies is so slight that shift in dominance is not audible. 

Effect of temperature on calls. — The present data are insufficient to test 
statistically the correlation between temperature and variation within certain 
components of the calls in Smilisca, but even a crude graph shows some gen- 
eral correlations. The widest range of temperatures is associated with the 
recordings of S. baudini. Three individuals recorded at a temperature of 30° 
C. at Tehuantepec, Oaxaca, had pulse rates of 180 pulses per second and fun- 
damental frequencies of 160-180 cps., as compared with an individual recorded 
at a temperature of 17° C, which had a pulse rate of 140 and a fundamental 
frequency of 135 cps. All individuals of S. baudini recorded at higher tem- 
peratxu-es had faster pulse rates and higher fundamental frequencies. Pulse 
rates differ in the other species in the genus but less strikingly ( probably owing 
to narrower ranges of temperatures at which recordings were made). In five 
recordings of S. sordida made at 20° C. the pulse rate is 80-90, as compared 
with four recordings made at 25° C. having pulse rates of 120-135. Thirteen 
recordings of S. sila made at 17° C. have pulse rates of 97-112 (average 105); 
one individual recorded at 26° C. has 120 pulses per second. Seemingly no 
correlation exists between temperature and other characteristics of the calls, 
such as duration and rate of note-repetition. 

The breeding call as an isolating mechanism. — Blair ( 1958), Bogert ( 1960), 
Duellman (1963a), Fouquett (1960), Johnson (1959), and others have pro- 



356 University of Kansas Publs., Mus. Nat. Hist. 

vided evidence that the breeding calls of male hylids (and other anurans) 
serve as isolating mechanisms in sympatric species. In summarizing this dis- 
cussion of the breeding calls of Smilisca we w^ant to point out what seem to 
be important differences in the calls that may prevent interspecific hybridiza- 
tion in sympatric species of Smilisca. 

The genus is readily divided into two species-groups on morphological 
characters; this division is supported by the breeding calls. In the species of 
the baudini group the calls are unmodulated and lack secondary notes. In the 
sordida group the calls either have secondary notes or are modulated. 

Smilisca baudini occurs sympatrically with S. cyanosticta and S. phaeota; 
where they occur together, both species sometimes breed in like places at the 
same time. We are not aware of these species breeding synchronously at ex- 
actly the same site, although S. baudini and S. cyanosticta were calling on the 
same nights and less than 100 meters apart in Oaxaca in June, 1964. Regard- 
less of their respective breeding habits, sympatric species have calls that differ 
notably. Except for the higher fundamental and dominant frequencies, the 
calls of S. cyanosticta and S. phaeota closely resemble one another, but the 
calls of both species differ markedly from that of S. baudini. The geographic 
ranges of S. cyanosticta and S. phaeota are widely separated. 

The calls of the allopatric species S. puma and S. sila are not greatly differ- 
ent. Smilisca sordida has a distinctive call and occurs sympatrically with S. 
puma and S. sila. In the streams in southern Costa Rica S. sordida and S. sila 
breed synchronously, but the high-pitched modulated call of the former is 
notably different from the lower, unmodulated call of S. sila. 

The data indicate that the calls of related sympatric species differ more 
than the calls of related allopatric species. We postulate that these differences 
evolved to support the reproductive isolation of the sympatric species. The 
data are insufficient to determine geographic variation in the calls and to de- 
termine if differences in the calls are enhanced in areas of sympatry as com- 
pared with the allopatric parts of the ranges. 

Other calls. — As stated previously, there is no direct evidence of territoriality 
in Smilisca; we have heard no calls that can be definitely identified as terri- 
torial. Single notes of S. baudini, phaeota, and sQa have been heard by day, 
just prior to rains, or during, or immediately after rains. Such calls can be 
interpreted as "rain calls," which are well knovni in Hyla eximia and Hyla 
squirella. Distress calls are known in several species of Rana and in Lepto- 
dactylus pentadactylus; such calls result from the rapid expulsion of air over 
the vocal cords and with the mouth open. Distress calls have been heard 
from S. baudini. At Charapendo, Michoacan, Mexico, a male that had one hind 
limb engulfed by a Leptodeira maculata emitted several long, high-pitched 
cries. A clasping pair of S. baudini was found in a bush at the edge of a 
marshy stream 2 kilometers northeast of Las Canas, Guanacaste Province, 
Costa Rica. When the pair was grasped, the female emitted a distress call. 

Eggs 

Eggs of S. baudini, cyanosticta, and phaeota have been found in the field, 
and eggs of S. sila have been observed in the laboratory. The eggs of S. puma 
and sordida are unknown. Insofar as known, Smilisca baudini is unique in 
the genus in depositing the eggs in a surface film. Each egg is encased in a 



Neotropical Hylid Frogs, Genus Smilisca 357 

vitelline membrane, but individual outer envelopes are lacking. The eggs are 
small; the diameter of recently-deposited eggs is about 1.3 mm. and that of 
the vitelline membrane is about 1.5 mm. The eggs of S. ctjanosticta and phae- 
ota are deposited in climips, and the eggs are larger than those of S. baudini. 
Diameters of eggs of S. cyanosticta are about 2.3 mm., and those of the outer 
envelopes are about 4.0 mm. Artificially fertilized eggs of S. sila raised in 
the laboratory have diameters of about 2.4 mm.; the diameter of the outer 
envelopes is about 4.9 mm. 

In order to determine the reproductive potential of the six species, ovulated 
eggs were removed from females and counted. The numbers of eggs recorded 
are: 3 S. baudini— 2620, 2940, 3320; 1 S. cyanosticta— 910; 3 S. phaeota^ 
1665, 1870, 2010; 1 S. puma— 518; 3 S. yt/<j— 369, 390, 473; 3 S. sordida— 
524, 702, 856. These limited data indicate that the large species (S. baudini, 
cyanosticta, and pliaeota) have more eggs than do the smaller species. The 
stream-breeding species (S. sila and sordida) have relatively iew eggs by com- 
parison with the pond-breeders. Possibly this is a function of size of eggs 
rather than a correlation with the site of egg-deposition. 

Tadpoles 

The acquisition of tadpoles of all of the species of Smilisca has made pos- 
sible the use of larval characters in erecting a classification and in estimating 
the phylogenetic relations of the several species. Furthermore, developmental 
series of tadpoles of four species allow a comparison of the growth and de- 
velopment in these species. Throughout the discussion of tadpoles we have 
referred to the various developmental stages by the Stage Numbers proposed 
by Gosner (1960). 

General Structure 

Tadpoles of the genus Smilisca are of a generalized hylid type, having % 
tooth-rows, unspeciaUzed beaks, mouth partly or completely bordered by papil- 
lae, lateral fold present in the lips, spiracle sinistral, anal tube dextral, and 
caudal musculature extending nearly to tip of caudal fin. Although minor 
differences exist in coloration, proportions, and mouthparts, no great modifica- 
tions of the basic structure are present. 

Comparison of Species 

The larval characters of the species of Smilisca are compared below and 
illustrated in Figures 11-15. 

Shape and Proportions. — The bodies of S. baudini, ctjanosticta, phaeota, and 
puma are rounded and about as wide as deep; the eyes are moderately large 
and directed dorsolaterally, and the nostrils are about midway between the 
bluntly rounded snout and the eyes. The mouths are medium-sized and di- 
rected anteroventrally. The bodies of tadpoles of S. sila and sordida are 
slightly compressed dorso-ventrally. The snout is moderately long and sloping; 
the eyes are larger and directed more dorsally than in the other species, and 
the nostrils are closer to the eyes than the snout. The mouths are moderately 
large and directed ventrally. 

The tail is about half again as long as the body in S. baudini, cyanosticta. 



358 



University of Kansas Publs., Mus. Nat. Hist. 



phaeota, and puma; in these species the caudal musculature is moderately 
heavy, and the caudal fins are deep. The caudal musculature is upturned 
distally in S. baudini and phaeota, and the dorsal fin extends anteriorly onto 
the body in these two species and in S. puma. The tail is about twice as long 
as the body in S. sila and sordida. In both species the caudal fins are shallow 
in comparison with the depth of the caudal musculature, especially in S. sor- 
dida (Fig. 14); in neither species does the dorsal fin extend anteriorly onto the 
body. 

Mouthparts. — The mouth of S. sordida is completely bordered by two rows 
of papillae, whereas in the other species the median part of the upper lip is 
devoid of papillae. Sm^ilisca baudini and puma have two rows of papillae; S. 
sila has one complete row (except medially on the upper hp) and one incom- 
plete row, and S. cyanosticta and phaeota have only one row (Fig. 15). All 
species have numerous papillae in the lateral fold; the fewest lateral papillae 






Fig. 11. Tadpoles of Smilisca baudini: (A) Stage 21 (KU 62155) xlO; (B) 
Stage 25 (KU 68467) x5; (C) Stage 30 (KU 60018) x4; (D) Stage 41 

(KU 60018) x3. 



Neotropical Hylid Frogs, Genus Smilisca 



359 







Fig. 12. Tadpoles of Smilisca cyanosticta: (A) Stage 21 (KU 87648) (B) 
Stage 25 (KU 87651) X5; (C) Stage 30 (KU 87652) X4; (D) Stage 40 

(KU 87650) x3. 

are found in S. cyanosticta and phaeota. Although all species have two rows 
of teeth in the upper jaw and three rows in the lower jaw, specific diflFerences 
in the nature of the rows exist between certain species. The second upper 
tooth-row is narrowly interrupted medially in S. sila and sordida and broadly 
interrupted in the other species. The first upper row is strongly arched in S. 
puma, moderately arched in S. baudini and sila, and weakly arched in the 
other species. In all species the third lower tooth-row is the shortest, only 
slightly so in S. sila and sordida, but only about half the length of the second 
lower row in S. puma. 

The beaks are well developed and finely serrate in all species. The lower, 
broadly V-shaped, beak is slender in S. puma, rather robust in S. baudini and 



360 



University of Kansas Publs., Mus. Nat. Hist. 



sila, and moderately heavy in the other species. The lateral processes of the 
upper beak are shortest in S. puma and longest in S. baudini and sordida. In 
the latter the inner margin of the upper beak and lateral process have the form 
of a shallow S, whereas in the other species the inner margin of the upper 
beak forms a continuous arch with the lateral processes (Fig. 15). 

Coloration. — The tadpoles of Smilisca lack the bright colors or bold mark- 
ings characteristic of some hylid tadpoles; even so, the subdued colors and 
arrangement of pigments provide some distinctive markings by which the 
species can be distinguished from one another. The species comprising the 
baudini group (S. baudini, cyanosticta, and phaeota) are alike in having the 
body brown or grayish brown dorsally and transparent with scattered brown 
pigment ventrally. A cream-colored, crescent-shaped mark is present on the 
posterior edge of the body; this mark is usually most noticeable in S. baudini 
and least so in S. cyanosticta. Other differences in coloration in members of 
the baudini group are relative and subtle. Smilisca phaeota usually is more 







Fig. 13. Tadpoles of Smilisca phaeota: (A) Stage 21 (KU 68479) Xl4; 
(B) Stage 25 (KU 68480) x5; (C) Stage 30 (KU 68482) X4; (D) Stage 

40 (KU 68483) x3. 



Neotropical Hylid Frogs, Genus Smilisca 



361 



pallid than baudini, and cyanosticta usually is darker than baudini; both spe- 
cies have larger dark markings on the tail than does S. pJiaeota. Smilisca 
baudini has a dark streak on the middle of the anterior one-fourth of the tail 
(Figs. 11-13). 

Smilisca puma is distinctive in having a grayish brown body and dark gray 
reticulations on the tail. Smilisca sila and sordida are distinctive in having 
pairs (sometimes interconnected) of dark marks on the dorsal surfaces of the 
caudal musculature, and in dorsal viev^' the tail appears to be marked with 
dark and pale creamy-tan transverse bars. These dark marks, as well as the 
small flecks on the tail, are brown in S. sila and red in sordida. Smilisca sila 
has dark browTi flecks on the dorsal surface of the body and small white flecks 
laterally; these markings are absent in S. sordida (Fig. 14). 

Descriptions of the coloration of living tadpoles are given in the accounts 
of the species. 

Growth and Development 

Information on the growth and development of Middle American hylids is 
scanty. Adequate descriptions have been published for Phyllomedusa annae 
(Duellman, 1963b), Phrynohyas venulosa (Zweifel, 1964), and Triprion pe- 
tasatus (Duellman and Klaas, 1964). Material is available for adequate de- 
scriptions of the developmental stages of fomr species of Smilisca (Tables 9-12, 
Figs. 11-13). Because none of the tadpoles was raised from hatching to meta- 
morphosis, the rate of growth and duration of the larval stages are unknown. 






Fig. 14. Tadpoles of Smilisca: (A) S. puma. Stage 30 (KU 91807); (B) 
S. sila. Stage 25 (KU 80260); S. sordida. Stage 30 (KU 68475). All X3.5. 



6—3430 



362 



University of Kansas Publs., Mus. Nat. Hist. 




•T.'-JLA/^^'rAV^^-^'^-^-'t^A'.ViWt^'Wi^*^^^^^^^ 








'S^<L^r I riiuiiiiimif uuU Miiniiii ^<•f l\ n ^^^ ^ ,^^^ f„„lm nu iii i iiiii mu iiii i iijiiiii m uumu,. 




^,.ii n i i uu h Uiiiiu i ii iu<)». | ,„m„)|p || nm, , „mmnmi)«nBi 

Fig. 15. Mouthparts of tadpoles of Smilisca; (A) S. baudini (KU 60018); 
(B) S- P«"ia (KU 91807); (C) S. cyanosticta (KU 87625); (D) S. sila 
(KU 80620); (E) S, p/iaeofa (KU 68482); (F) S. sordtda (KU 68475). 

All Xl7. 

Table 9. — Growth and Development of Tadpoles of Smilisca baudini. 
( Means Are Given in Parentheses After the Observed Ranges. ) 



Stage 


N 


21 


10 
10 
10 
10 
10 

3 
10 

2 
10 

3 

6 
23 


24 


25 


27 


29 


37 


38 


40 


41 


42 


45 


46 





Total length 



5.1-5.4 
6.0-6.5 
7.2-8.3 
18.5-21.5 
21.5-24.5 
28.5-31.0 
35.0-37.5 
34.0-37.0 
34.0-37.0 
24.0-30.0 
14.0-24.0 



(5.22) 
(6.20) 
(7.78) 
(20.22) 
(22.60) 
(30.00) 
(35.50) 
(35.50) 
(35.50) 
(27.00) 
(17.58) 



Body length 



2.6-2.7 

2.3-2.6 

3.0-3.3 

8.0-9.0 

8.5-10.0 

11.0-12.5 

12.0-13.5 

12.5-13.5 

12.5-13.5 

12.5-13.0 

12.5-14.0 

12.0-15.5 



(2.54) 

(2.45) 

(3.14) 

(8.38) 

(9.25) 

(11.67) 

(12.80) 

(13.00) 

(13.00) 

(12.67) 

(13.37) 

(13.34) 



Tail length 



2.5-2.7 (2.58) 

3.5-3.9 (3.69) 

4.2-5.0 (4.64) 

10.4-13.0 (11.84) 

12.5-14.5 (13.35) 

17.5-19.0 (18.00) 

21.5-24.0 (22.70) 

21.5-23.5 (22.50) 

21.5-23.5 (22.50) 

11.5-17.0 (14.33) 

1.5-10.0 (4.17) 



Neotropical Hylid Frogs, Genus Smilisca 



363 



Table 10. — Growth and Development of Tadpoles of Smilisca cya- 
NOSTicTA. ( Means Are Gin^en in Parentheses After the Observed Ranges. ) 



Stage 


N 


Total length 


Body length 


Tail length 


21 


10 

10 

7 

10 

2 

2 


5.8-6.5 (6.28) 

7.9-9.2 (8.44) 

22.5-25.0 (23.50) 

27.0-30.0 (28.75) 

26.0-27.0 (26.50) 


2.8-3.1 (3.00) 
2.7-3.2 (2.96) 
8.5-9.5 (9.00) 
9.5-11.5 (10.80) 

10.00 

14.00 


3.0-3.5 (3.28) 


25 


4.8-6.0 (5.48) 


30 


14.0-15.5 (14.57) 


36 

42 

46 


17.0-18.5 (17.95) 
16.0-17.0 (16.50) 









Hatchlings of three species (S. baudini, cyanosticta, and pJiaeoia) are avail- 
able. These larvae have non-functional eyes and large oral suckers. By the 
time the larvae have developed to stage 21, external gills are present, the 
caudal musculatiu-e and caudal fin have been differentiated, and the head is 
distinguishable from the body. In stage 21 oral suckers and a large amount of 
yolk are still present. 

The developmental data on the four species show no significant variations; 
consequently, we will describe the development of only one species, Smilisca 
phaeota (Table 11, Figs. 13 and 16). 

Stage 21. — Bulging cream-colored yolk mass, transparent cornea, and mod- 
erately long, unbranched filamentous gills, and oral suckers present; mouth 
having irregular papillae on lower lip; teeth and beaks absent; caudal myo- 
meres distinct; pigmentation uniform over body and caudal musculature; cau- 
dal fin transparent with scattered small flecks. 

Stage 25. — Operculum complete; gills absent; sinistral spiracle apparently 
functional; cloacal tail-piece, nasal capsules, and external nares present; gut 



o 

z 

UJ 



40 
35 
30 
25 
20 
15 
10 
5 



TOTAL LENGTH 
BODY LENGTH 
TAIL LENGTH 



Mefotorsol Tubercle. 



Mouthports Degenerating 



L = 2 XD 




Moulhports Connplete 
LM/2D^ 

Mouthparts Appear a . — ' 

Orel Suckers Lost "^ / ,'' 

V y 

Holchling 



Resorption of Toil 



Metamorphosis 
Complete 



15 17 19 2! 23 25 27 29 31 33 35 37 39 41 43 45 
DEVELOPMENTAL STAGES 

Fig. 16. Relative rate of growth in tadpoles of Smilisca phaeota as correlated 
with developmental stages. Formulas for the limb bud refer to its length (L) 

in relation to basal diameter (D). 



364 



University of Kansas Publs., Mus. Nat. Hist. 



partly formed; mouth bordered by single row of papillae, except medially; 
small papillae present in lateral fold of lips; two upper and three lower tooth- 
rows present, but not fully developed; beaks apparently fully developed; depth 
of dorsal and ventral fins less than depth of caudal musculatmre; tip of tail 
upturned; pigment on body most dense on dorsvun and sides; faint, nearly 
pigmentless crescent-shaped mark on posterior edge of body; concentrations of 
pigment forming small spots on tail. 

Stage 28. — Mouthparts complete; limb bud about half as long as thick; 
other structural features and coloration closely resemble those in stage 25. 

Stage 30. — Limb bud approximately twice as long as thick; body as deep 
as wide; dorsal fin deepest just posterior to body; ventral fin deeper than cau- 
dal musculature; tail sharply uptmrned distally; anal tube dextral; brown pig- 
ment sparse on flanks. 



Table 11. — Growth and Development of Tadpoles of Smilisca phaeota. 
( Means Ajeie GrvEN in Parentheses After the Observed Ranges. ) 



Stage 


N 


Total length 


Body length 


Tail length 


15 


10 

8 




1.9-2.1 (1.97) 
2.0-2.2 (2.07) 




16 






18 


4 




2.2-2.6 (2.31) 




21 


3 


7.9-8.6 (8.21) 


4.1-4.5 (4.31) 


3.8-4.1 (3.92) 

4.3-6.0 (5.05) 

6.7-9.8 (8.41) 

7.7-10.5 (8.88) 

8.5-12.6 (9.85) 

11.5-14.0 (12.60) 

10.5-15.5 (13.53) 

14.7-17.3 (16.02) 

15.6-18.5 (16.80) 

18.9-20.0 (19.44) 


25 


10 


8.7-10.6 (9.69) 


4.5^.8 (4.64) 


26 


11 


12.3-16.1 (14.01) 


4 2-6.3 (5 60) 


27 


10 


13.0-15.7 (14 28) 


4 9-6 2 (5 40) 


28 


13 


13 9-20 9 (15 62) 


5 2-8 3 (5 75) 


29 


8 


17 8-22 3 (19 79) 


6 3-8 4 (7 19) 


30 


9 


20.3-24.8 (22.85) 


8.1-10.5 (9.32) 


31 


5 


24.1-28.5 (26.61) 


9.4-11.2 (10.59) 


34 


5 


24.8-29.4 (27.31) 


9.2-11.6 (10.73) 


36 


3 


30.0-30.1 (30.07) 


10.1-12.2 (11.15) 


37 


4 


28.9-34.1 (31.75) 


11.5-12.4 (11.88) 


17.4-22.5 (19.88) 


38 


1 
2 


28.98 
35.6-36.9 (36.25) 


12.88 
14 00 


16 10 


39 


21.6-22.9 (22.25) 


40 


2 


32.3-39.8 (36.05) 


14.00 


18.3-21.8 (20.05) 


43 


2 


21.5-23.0 (22.25) 


14.2-14.8 (14.45) 


6.8-8.8 (7.80) 


44 


4 


14.5-15.6 (15.08) 
12.7-16.7 (14.26) 




46 


11 













Neotropical Hylid Frogs, Genus Smilisca 



365 



Table 12. — Growth and Development of Tadpoles of Smilisca sordida. 
(Means Abe Given in Parentheses After the Observed Ranges.) 



Stage 


N 


Total length 


Bodj' length 


Tail length 


25 


6 
2 
8 
7 
3 
1 
9 


25.5-28.0 (26.1) 
28.5-30.0 (29.3) 
29.5-34.5 (32.3) 
31.6-37.5 (34.6) 
33.0-37.2 (35.2) 


9.0-9.5 (9.3) 
10.2-10.5 (10.4) 
10.2-11.7 (10.8) 
11.0-12.5 (11.5) 
11.6-12.2 (11.9) 

12.4 
13.1-15.7 (14.9) 


16.2-18.5 (16.7) 


33 


18.0-19.8 (18.9) 


36 


19.3-23.0 (21.5) 


37 


21.6-25.0 (23.2) 


41 


21.4-25.2 (23.2) 


43 




46 













Stages 34, 36, 37, and 38. — Stage 34, foot paddle-shaped wiih four toe 
buds; stage 36, five toe buds; stages 37 and 38, lengthening of toes. In all 
four stages, spiracle persistent, and pigmentation resembling that of early 
stages. 

Stage 39. — Metatarsal tubercle present; greatest total length (36.9 mm.) 
attained. 

Stage 40. — Subarticular tubercles prominent; skin over forelimbs transpar- 
ent; cloacal tail-piece and spiracle absent; outer tooth-rows degenerating; cau- 
dal fins shallower than in preceding stages; distal part of tail nearly straight; 
size of dark markings on tail decreased; pigment present on hind Hmb. 

Stage 43. — Forelimbs erupted; larval mouthparts absent; comer of mouth 
between nostril and eye; transverse bands present on hind limbs; tail greatly 
reduced (about 8 mm. in length). 

Stage 44. — Sacral hump barely noticeable; tail reduced to a stub; comer 
of mouth at level of pupil of eye; dorsal surfaces pale oHve-green; venter 
white. 

Changes proceed in a definite pattern during the growth and development 
of tadpoles. Larval teeth are absent in hatchlings; the inner tooth-rows de- 
velop first, and the third lower row last. At metamorphosis the third lower 
row is the first to be lost. The tail increases gradually in length relative to 
the body. In stage 25 the tail is 52.1 per cent of the total length, and in stage 
36, 64.6 per cent. In later stages the tail becomes relatively shorter through 
resorption. Duellman and Klaas (1964:320) noted a great size-variation in 
Triprion tadpoles in stage 25. No such variation is apparent in any stage of 
any of the species of Smilisca studied. 

The growth and development of the other species of Smilisca do not difiFer 
significantly from that of S. phaeota. The tadpoles of S. sila and sordida from 
streams have relatively longer tails at hatching. For example, in tadpoles of 
S. sordida the average length of tail is 64.0 per cent of the body-length in stage 
25, and in stage 37, 67.0 per cent. 



Behavior 

The tadpoles of S. baudini, cyanosticta, phaeota, and puma are pelagic in- 
habitants of shallow ponds. Early stages of S. baudini in which external gills 



366 University of Kansas Publs., Mus. Nat, Hist. 

are present have been observed to hang vertically with the gills spread out at 
the surface of the water, a behavior noted by Zweifel (1964:206) in tadpoles 
of Phrynohyas venulosa, which also develop in warm, standing water having a 
relatively low oxygen-tension. When disturbed the pelagic tadpoles usually 
dive and seek shelter amidst vegetation or in mud on the bottom. This be- 
havior was observed in S. baudini, cijanosticta, and phaeota by day and at 
night. No tadpoles of S. puma were observed by day; those seen at night 
were near the surface of small water-filled depressions in a grassy marsh; they 
responded to light by taking refuge in the dense grass. Perhaps tadpoles of 
this species are negatively phototactic and remain hidden by day. 

The stream-inhabiting tadpoles of S. siln and sordida live in clear pools in 
rocky streams, where they were observed to cling by their mouths to rocks in 
the stream and to seek shelter amidst pebbles or beneath rocks and leaves on 
the bottom. These tadpoles are not found in shallow riflBes. 

We have not found tadpoles of two species of Smilisca in the same body 
of water and therefore cannot offer observations on ecological relationships in 
sympatric situations. 

PHYLOGENETIC RELATIONSHIPS 

Identifiable hylid remains are known from the Miocene to the Recent, but 
these fossils are mostly fragmentary and provide little useful infonnation re- 
garding the phylogenetic relationships of living genera. Frogs of the genus 
Smilisca are generalized and show no striking adaptations, either in their struc- 
ture or in their modes of life history. 

Interspecific Relationships 

In attempting to understand the relationships of the species of Smilisca we 
have emphasized osteological characters. The phylogeny suggested by these 
characters is supported by other lines of evidence, including external morphol- 
ogy, tadpoles, and breeding calls. 

Our concept of the prototype of the genus Smilisca is a moderate-sized 
hylid having: (1) a well-developed frontoparietal fontanelle, (2) frontoparietal 
lacking lateral processes, (3) no bony squamosal-maxillary arch, (4) a fully 
ossified ethmoid, (5) paired subgular vocal sac, (6) moderately webbed 
fingers and toes, (7) relatively few supernumerary tubercles on the digits, (8) 
eggs deposited in clumps in ponds, (9) anteroventral mouth in tadpoles 
bordered by one row of labial papillae, but median part of upper lip bare, 
( 10 ) tail relatively short and deep in tadpoles, and (11) a breeding call con- 
sisting of a series of like notes. 

Two phyletic lines evolved from this prototype. The first of these was the 
stock that gave rise to the baudini group. The evolutionary changes that 
took place in this line included increase in size, development of a lateral 
curvature of the maxillary, and an increased amount of cranial ossification, 
especially in the dermal roofing bones. This phyletic line retained the larval 
characters and breeding call of the prototype. The second phyletic line gave 
rise to the sordida group and diverged from the prototype in the development 
of an angular maxillary and a breeding call consisting of a primary note fol- 
lowed by secondary notes. The frogs in this phyletic line retained the moderate 



Neotropical Hylid Frogs, Gexus Smilisca 367 

size of the protoh'pe and did not develop additional dermal bone. Our con- 
cept of the phylogenetic relationships is shown graphically in Figure 17. 

Within the baudini group one stock retained separate nasals and did not 
develop a bony squamosal-maxillary arch, but broad lateral processes developed 
on the frontoparietals. The tadpoles remained unchanged from the primitive 
t>'pe. This stock evolved into S. phaeota. In the other stock the nasals be- 
came fully ossified and a bony squamosal-maxillary arch developed. One 
branch of this second stock retained tadpoles having only one row of labial 
papillae and did not develop lateral processes on the frontoparietals; this 
branch evolved into S. cyanosticta. The other branch diverged and gave rise 
to S. baudini by developing relatively shorter hind legs, large lateral processes 
on the frontoparietals, and tadpoles having two rows of labial papillae. 

Within the sordida group the cranial features remained unchanged in one 
line, which gave rise to S. sila, whereas in a second hne the nasals were re- 
duced, and their long axes shifted with the result that they are not parallel to 
the maxillaries; the amount of ossification of the ethmoid was reduced, and 
the tadpoles developed two rows of labial papillae. In this second line one 
branch retained the pond-breeding habits and gave rise to S. pinna, whereas 
a second branch became adapted to stream-breeding and gave rise to S. 
sordida. 

Certain aspects of this proposed phylogeny warrant further comment. 
Features such as the deposition of additional bone that roofs the skull or 
that forms lateral projections from the frontoparietals, like those in S. baudini 
and phaeota, are minor alterations of dennal elements and not basic modifica- 

baudini 

cyanosticta 

phaeota sordida 




Prototype 

Fig. 17. Hypothesized phylogenetic relationships of the species of Smilisca. 



368 University of Kansas Publs., Mus. Nat. Hist. 

tions of the architecture of the skull. Consequently, we hypothesize the inde- 
pendent development of these dermal changes in S. baudini and phaeoia. 
Similar kinds of dermal modifications have evolved independently in many 
diverse groups of frogs. 

Likewise, we propose the parallel development of stream-adapted tadpoles 
in S. sordida and sila; in both cases the tadpoles adapted to changing environ- 
mental conditions (see following section on evolutionary history). Tadpoles of 
S. sordida already had two rows of labial papillae before entering the streams; 
subsequently the tadpoles developed complete rows of papillae, ventral mouths 
and long tails having low fins. Possibly the tadpoles of S. sUa had two rows 
of labial papillae prior to their adapting to stream conditions; in the process 
of adapting they developed ventral mouths and long tails having low fins. 
Similar modifications in tadpoles have occurred in many diverse groups of 
Middle American hylids, such as Plectrohyla, Ftychohyla, the HyJa uranochroa 
group, and the Hyla taeniopus group. 

Our lack of concern about coloration is due to the fact that, with the excep- 
tion of the blue spots on the flanks and posterior sxuiaces of the thighs in some 
species, the coloration of Smilisca, consisting of a pattern of irregular dark 
marks on a paler dorsum and dark transverse bars on the limbs, is not much 
different from that of many other Neotropical hylids. Blue is a structural color, 
rare among Amphibia, which is achieved by the absence of lipophores above 
the guanophores. Thus, the incident hght rays at the blue end of the spectrum 
are reflected by the guanophores without interference by an overlying yellow 
lipophore screen. According to Noble (1931), lipophores are capable of 
amoeboid movement that permits shifts in their positions, between or beneath 
the guanophores. We do not know whether this behavior of lipophores is 
widespread and is efi^ected in response to environmental changes, or whether 
it is a genetically controlled attribute that is restricted in appearance. If the 
latter is the case we must assume that the prototype of Smilisca possessed such 
an attribute which was lost in S. baudini, phaeota, and puma. The develop- 
ment of blue spots is not constant in S. sordida and S. sila; in S. cyanosticta 
the spots range in color from blue to pale green. 

The coloration of the tadpoles is not distinctive, except for the presence of 
dorsal blotches on the tails of S. sila and sordida. However, the similarity in 
pattern cannot be interpreted as indicating close relationships because nearly 
identical patterns are present in Hyla legleri and some species of Prostherapis. 
This disruptive coloration seems to be directly associated with the pebble- 
bottom, stream-inhabiting tadpoles. 

In the baudini group, S. phaeota and cyanosticta are allopatric, whereas S. 
baudini occurs sympatrically with both of those species. The call of S. baudini 
differs notably from the calls of S. phaeota and cyanosticta, which are more 
nearly alike. Although in the phylogenetic scheme proposed here S. sila is 
considered to be more distantly related to S. puma than is S. sordida, the calls 
of S. sila and puma more closely resemble one another than either resembles 
that of S. sordida. Smilisca sila and puma are allopatric, whereas S. sordida 
is broadly sympatric with both of those species. We assume that in their 
respective phyletic lines the difFerentiation of both S. baudini and sordida was 
the result of genetic changes in geographically isolated populations. Sub- 
sequently, each species dispersed into areas inhabited by other members of their 
respective groups. Selection for differences in the breeding calls helped to 



Neotropical Hylid Frogs, Genus Smilisca 369 

reinforce other diflferences in the populations and thereby aided in maintaining 
specificity. 

Evolutionary History 

With respect to temporal and spatial aspects of evolution in Smilisca, we 
have tried to correlate the phylogenetic evidence on Smilisca with the geologic 
data on Middle America presented by Lloyd (1963), Vinson and Brineman 
(1963), Guzman and Csema (1963), Maldonado-Koerdell (1964), and Whit- 
more and Stew^art ( 1965). Likewise, we have borne in mind the evidence for, 
and ideas about, the evolution of the Middle American herpetofauna given by 
Dunn (1931b), Schmidt (1943), Stuart (1950, 1964) Duellman (1958, MS), 
and Savage (MS). 

According to Stuart's (1950) historical arrangement of the herpetofauna, 
Smilisca is a member of the Autochthonous Middle American Favmal Element, 
and according to Savage's ( MS ) arrangement the genus belongs to the Middle 
American Element, a fauna which was derived from a generalized tropical 
American unit that was isolated in tropical North America by the inundation 
of the Isthmian Link in early Tertiary, that developed in situ in tropical North 
America, and that was restricted to Middle America by climatic change in the 
late Cenozoic. 

Savage (MS) relied on the paleogeographic maps of Lloyd (1963) to 
hypothesize the extent and centers of differentiation of the Middle American 
Faunal Element. According to Lloyd's concept, Middle America in the Miocene 
consisted of a broad peninsula extending southeastward to about central 
Nicaragua, separated from the Panamanian Spur of continental South America 
by shallow seas. A large island, the Talamanca Range, and remnants of the 
Guanarivas Ridge formed an archipelago in the shallow sea. The recent dis- 
covery of remains of mammals having definite North American affinities in the 
Miocene of the Canal Zone (Whitmore and Stewart, 1965) provides substantial 
evidence that at least a peninsula was continuous southeastward from Nuclear 
Central America to the area of the present Canal Zone in early mid-Miocene 
time. South America was isolated from Central America by the Bolivar Trough 
until late mid-Pliocene. 

Thus, in the mid-Tertiary the broad peninsula of Nuclear Central America, 
which consisted of low and moderately uplifted regions having a tropical 
mesic climate, provided the site for the evolution of Smilisca. It is not pos- 
sible to determine when the genus evolved, but to explain the differentiation of 
the species it is unnecessary to have the ancestral Smilisca present prior to the 
Miocene. 

We view the Miocene Smilisca as the prototype described in the preceding 
section, and suppose that it lived in the mesic tropical environment of the 
eastern part of the Central American Peninsula (in what is now Costa Rica 
and western Panama). Two stocks differentiated, probably in middle Miocene 
times; one of these, the ancestral stock of the haudini group, was widespread 
on the Caribbean lowlands from the Nicaraguan Depression to the Bolivar 
Trough, and the other, the ancestral stock of the sordida group, was restricted 
to the Pacific lowlands of the same region. In late Miocene time the ancestral 
stock of the baudini group dispersed northwestward around the deep embay- 
ment in the Nicaraguan depression into upper Central America (in what is 
now Honduras and Guatemala) and thence into southern Mexico. Apparently 



370 University of Kansas Publs., Mus. Nat. Hist. 

differentiation took place on each side of the Nicaraguan Depression; the frogs 
to the south of the depression evolved into S. phacota, whereas those to the 
north of the depression represented the stock from which S. baudini and 
cyanosticta arose. Prior to the uphft of the mountains in the late Miocene 
and the Pliocene the baudini-cyanosticta stock probably was widespread in 
northwestern Central America. The elevation of the mountains resulted in 
notable climatic changes, principally the development of sub-humid environ- 
ments on the Pacific lowlands. The frogs living on the Pacific lowlands be- 
came adapted to sub-humid conditions and developed into S. baudini. The 
stock on the Caribbean lowlands remained in mesic environments and evolved 
into S. cyanosticta. 

Possibly in the middle Miocene before the Talamanca Range in Costa Rica 
and western Panama was greatly uplifted, the ancestral stock of the sordida 
group invaded the Caribbean lowlands of what is now Costa Rica. The sub- 
sequent elevation of the Talamanca Range in the Pliocene effectively isolated 
the ancestral stock of S. sila on the Pacific lowlands from the puma-sordida 
stock on the Caribbean lowlands. The former was subjected to the sub-humid 
conditions which developed on the Pacific lowlands when the Talamanca 
Range was uplifted. It adapted to the sub-humid environment by living 
along streams and evolving stream-adapted tadpoles. On the Caribbean side 
of the Talamanca Range the puma-sordida stock inhabited mesic environments. 
The stock that evolved into S. puma remained in the lowlands as a pond- 
breeding frog, whereas those frogs living on the slopes of the newly elevated 
mountains became adapted for their montane existence by developing stream- 
adapted tadpoles and thus differentiated into S. sordida. 

Probably the six species of Smilisca were in existence by the end of the 
Pliocene; at that time a continuous land connection existed from Central Amer- 
ica to South America. The climatic fluctuations in the Pleistocene, and the 
post-Wisconsin development of present climatic and vegetational patterns in 
Middle America, brought about the present patterns of distribution of the 
species. From its place of origin on the Caribbean lowlands of lower Central 
America, S. phaeota dispersed northward into Nicaragua and southward along 
the Pacific slopes of northwestern South America. Perhaps in the late Pleis- 
tocene or in post-Wisconsin time when mesic conditions were more widespread 
than now, S. phaeota moved onto the Pacific lowlands of Costa Rica. Its route 
could have been through the Arenal Depression. Subsequent aridity restricted 
its range on the Pacific lowlands to the Golfo Dulce region. Climatic fluctua- 
tion in northern Central America restricted the distribution of S. cyanosticta 
to mesic habitats on the slopes of the Mexican and Guatemalan highlands and 
to certain humid areas on the lowlands. Smilisca baudini was well adapted 
to sub-humid conditions, and the species dispersed northward to the Rio 
Grande Embayment and to the edge of the Sonoran Desert and southward into 
Costa Rica. In southern Mexico and Central America the species invaded 
mesic habitats. Consequently, in some areas it is sympatric with S. cyanosticta 
and phaeota. 

Smilisca puma dispersed northward onto the Caribbean lowlands of southern 
Nicaragua. Its southward movements probably were limited by the ridges of 
the Talamanca Range that extend to the Caribbean coast in the area of Punta 
Cahuita in Costa Rica. Smilisca sila dispersed along the Pacific lowlands and 
slopes of the mountains from eastern Costa Rica and western Panama through 



Neotropical Hylid Frogs, Genus Smilisca 371 

eastern Panama to northern Colombia. Climatic fluctuation in the Pleistocene 
evidently provided sufficient altitudinal shifts in environments in the Talamanca 
Range to permit S. sordida to move onto tlie Pacific slopes. From its upland 
distribution the species followed streams down to both the Caribbean and 
Pacific lowlands, where it is sympatric with S. puma on the Caribbean low- 
lands and S. sila on the Pacific lowlands. 

The evolution of the species-groups of Smilisca was effected through isola- 
tion by physical barriers in the Cenozoic; the differentiation of the species was 
initiated by further isolation of populations by changes in physiography and 
climate. Present patterns of distribution resulted from Pleistocene and post- 
Wisconsin climatic changes. Today, sympatric species have different breeding 
habits and breeding calls wliich reinforce the differences in morphology. 

SUMMARY AND CONCLUSIONS 

The genus Smilisca is composed of six species of tree frogs; each species is 
defined on the basis of adult morphology, larval characters, and breeding be- 
havior. Keys are provided to aid in the identification of adults and of tadpoles. 

Analysis of the characters and examination of type specimens indicates that 
several currently-recognized taxa are synonymous, as follows: 

1. Hijla heltrani Taylor, 1942 = Smilisca haudini. 

2. Hyla gabbi Cope, 1876 = Smilisca sordida. 

3. Htjla manisorum Taylor, 1954 = Smilisca baiidini. 

4. Hyla nigripes Cope, 1876 = Smilisca sordida. 

5. Hijla wellmanorum Taylor, 1952 = Smilisca puma. 

Smilisca phaeota cyanosticta Smith, 1953 is elevated to specific rank, and 
one new species, Smiluica sila, is named and described. 

The skeletal system of developmental stages and the adult of Smilisca baudini 
is described, and the skull is compared with that of other members of the 
genus. 

The tadpoles are described, compared, and illustrated; the larval develop- 
ment of Smilisca phaeota is described. 

Breeding behavior and breeding calls are described and compared. Some 
species of Smilisca have breeding choruses. Two species, S. sila and sordida, 
breed in streams, whereas the others breed in ponds. 

The genus is considered to be part of the Middle American Faunal Element: 
the species are thought to have differentiated in response to ecological diversity 
and historical opportunities provided by Cenozoic changes in physiography and 
climate. 



372 University of Kansas Publs., Mus. Nat. Hist. 

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Neotropical Hylid Frogs, Genus Smilisca 375 

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Transmitted March 14, 1966. 



n 

31-3430 



(Continued from inside of front cover) 

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

2. A distributional studv of tlie amphibians of the Isthmus of Tehuantepec, 
Mexico. By William' E. DucUman. Pp. 19-72, pis. 1-8, 3 figures in text. 
August 16, 1960. 50 cents. 

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

"4. Autecolog> of the copperhead. By Henry S. Fitch. Pp. 85-288, pis. 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 Hemy 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, pis. 21-24, 2 figures 
in text. February 10, 1961. 

8. Descriptions of two species of frogs, genus Ptychohvla; studies of American 
hylid frogs, V. By William E. Duellman. Pp. 349-357, pi. 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, pis. 26-30, 3 fig- 
ures in text. August 11, 1961. 73 cents. 

10. North American recent soft-shelled trutles (Family Trionychidae ). By 
Robert G. Webb. Pp. 429-611, pis. 31-54, 24 figures in text. February 
16, 1962. $2.00. 

Index. Pp. 613-624. 

Vol. 14. 1. Neotropical bats from western Mexico. By Sydnev Anderson. Pp. 1-8. 
October 24, 1960. 

2. Geographical variation in the harvest mouse, Reithrodontomys inegalotis, on 
the cenbal Great Plains and in adjacent regions. Bv J. Knox Jones, Jr., and 
B. Mursaloglu. Pp. 9-27, 1 figiu-e in text. July 24', 1961. 

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

4. A new subspecies of the black myotis (bat) from eastern Mexico. By E. 
Raymond Hall and Ticul Alvarez. Pp. 69-72, 1 figure in text. December 
29, 1961. 

5. North American yellow bats, "Dasypterus," and a list of the named kinds 
of the genus Lasiurus Gray. By E. Ravmond Hall and J. Knox Jones, Jr. 
Pp. 73-98, 4 figures in text. December 29, 1961. 

6. Natural historv' of the brush mouse (Peromyscus boylii) in Kansas with 
description of a new subspecies. Bv Charles A. Long. Pp. 99-110, 1 figure 
in text. December 29, 1961. 

7. Taxonomic status of some mice of the Peromyscus boylii group in eastern 
Mexico, with description of a new subspecies. By Ticul Alvarez. Pp. 111- 
120, 1 figure in tex-t. December 29, 1961. 

8. A new subspecies of ground squirrel ( Spermophilus spilosoma) from Ta- 
maulipas, Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962. 

9. Taxonomic status of the free-tailed bat, Tadarida yucatanica Miller. By J. 
Knox Jones, Jr., and Ticul Alvarez. Pp. 125-133, 1 figure in text. March 7, 
1962. 

10. A new doglike carnivore, genus Cynarctus, from the Clarendonian, Pliocene, 
of Texas. By E. Ravmond Hall and Walter W. Dalquest. Pp. 133-138, 
2 figures in text. April 30, 1962. 

11. A new subspecies of wood rat (Neotoma) from northeastern Mexico. By 
Ticul Alvarez. Pp. 139-143. April 30, 1962. 

12. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones, Jr., Ticul 
Alvarez, and M. Ra>anond Lee. Pp. 145-159, 1 figure in text. May 18, 
1962. 

13. A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, 
1 figure in text. May 21, 1962. 

"14. The mammals of Veracruz. By E. Raymond Hall and Walter W. Dalquest. 
Pp. 165-362, 2 figures in text. May 20, 1963. $2.00. 

15. The Recent mammals of Tamaulipas, Mexico. By Ticul Alvarez. Pp. 363- 
473, 5 figures in text. May 20, 1963. $1.00. 

16. A new subspecies of the fruit-eating bat, Stiunira ludovici, from western 
Mexico. By J. Knox Jones, Jr., and Gary L. PhiUips. Pp. 475-481, 1 figure 
in text. March 2, 1964. 

17. Records of the fossil mammal Sinclairella, Family Apatemyidae, from the 
Chadronian and Orellan. By William A. Clemens, Jr. Pp. 483-491, 2 figures 
in text. March 2, 1964. 

18. The mammals of Wyoming. By Charles A. Long. Pp. 493-758, 82 figures 
in text. July 6, 1965. $3.00. 

Index. Pp. 759-784. 

(Continued on outside of back cover) 



(Continued from inside of back cover) 

Vol. 15. 1. The amphibians and reptiles of Michoacan, Mexico. By William E. Duell- 
man. Pp. 1-148, pis. 1-6, 11 figures in text. December 20, 1961. $1.50. 

2. Some reptiles and amphibians from Korea. By Bobert G. Webb, J. Knox 
Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962. 

3. A new species of frog (genus Tomodactylus ) from western Mexico. By 
Bobert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962. 

4. Type specimens of amphibians and reptiles in the Museum of Natural His- 
tory, The University of Kansas. By William E. Duellman and Barbara Berg. 
Pp. 183-204. October 26, 1962. 

5. Amphibians and reptiles of the rainforests of southern El Peten, Guatemala. 
By William E. Duellman. Pp. 205-249, pis. 7-10, 6 figures in text. October 
4, 1963. 

6. A revision of snakes of the genus Conophis (Family Colubridae, from Middle 
America). By John Wellman. Pp. 251-295, 9 figures in text. October 4, 
1963. 

7. A review of the Middle American tree frogs of the genus Ptychohyla. By 
William E. Duellman. Pp. 297-349, pis. 11-18, 7 figures in text. October 
18, 1963. 50 cents. 

8. Natural historv of the racer Coluber constrictor. By Henry S. Fitch. Pp. 
351-468, pis. 19-22, 20 figures in text. December 30, 1963. $1.00. 

9. A review of the frogs of the Hyla bistincta group. By William E. Duellman. 
Pp. 469-491, 4 figures in text. March 2, 1964. 

10. An ecological study of the garter snake, Thamnophis sirtalis. By Henry S. 
Fitch. Pp. 493-564, pis. 23-25, 14 figures in text. May 17, 1965. 

11. Breeding cycle in the ground skink, Lygosoma laterale. By Henry S. Fitch 
and Harry W. Greene. Pp. 565-575, 3 figures in text. May 17, 1965. 

12. Amphibians and reptiles from the Yucatan Peninsula, Mexico. By William 
E. Duellman. Pp. 577-614, 1 figure in text. June 22, 1965. 

13. A new species of tvutle, genus Kinosternon, from Central America. By John 
M. Legler. Pp. 615-625, pis. 26-28, 2 figures in text. June 20, 1965. 

14. A biogeographic account of the herpetofauna of Michoacan, Mexico. By 
William E. Duellman. Pp. 627-709, pis. 29-36, 5 figures in text. December 
30, 1965. 

15. Amphibians and reptiles of Mesa Verde National Park, Colorado. By Charles 
L. Douglas. Pp. 711-744, pis. 37 and 38, 6 figures in text. March 7, 1966. 
Index in preparation. 

Vol. 16. 1. Distribution and taxonomv of mammals of Nebraska. By J. Knox Jones, Jr. 
Pp. 1-356, plates 1-4, 82 figures in text. October 1, 1964. $3.50. 

2. Synopsis of the lagomorphs and rodents of Korea. By J. Knox Jones, Jr., 
and David H. Johnson. Pp. 357-407. February 12, 1965. 

3. Mammals from Isla Cozumel, Mexico, with description of a new species of 
harvest mouse. By J. Knox Jones, Jr., and Timothy E. Lawlor. Pp. 409-419, 
1 figure in text. April 13, 1965. 

4. The Yucatan deer mouse, Peromyscus yucatanicus. By Timothy E. Lawlor. 
Pp. 421-438, 2 figures in text. July 20, 1965. 

5. Bats from Gautemala. By J. Knox Jones, Jr. Pp. 439-472. April 18, 1966. 
More numbers will appear in volume 16. 

Vol. 17. 1. Localities of fossil vertebrates obtained from the Niobrara Formation (Cre- 
taceous) of Kansas. By David Bardack. Pp. 1-14. January 22, 1965. 

2. Chorda tympani branch of the facial nerve in the middle ear of tetrapods. 
By Richard C. Fox. Pp. 15-21. June 22, 1965. 

3. Fishes of the Kansas River System in relation to zoogeography of the Great 
Plains. By Artie L. Metcalf. Pp. 23-189, 4 figures in text, 51 maps. March 
24, 1966. 

4. Factors affjcting growth and production of channel catfish, Ictalurus punctatus. 
By Bill A. Simco and Frank B. Cross. Pp. 191-256, 13 figures in text. 
June 6, 1966. 

5. A new species of fringe-limbed tree frog, genus Hyla, from Darien, PanamS. 
By William E. Duellman. Pp. 257-262, 1 figure in text. June 17, 1966. 

6. Taxonomic notes on some Mexican and Central American hylid frogs. By 
William E. Duellman. Pp. 263-279. June 17, 1966. 

7. .Neotropical hylid frogs, genus Smilisca. By William E. Duellman and Linda 

Trueb. Pp. 281-375, pis. 1-12, 17 figures in text. July 14, 1966. 
More numbers will appear in volume 17. 



u 



University of Kansas Publications 

Museum of Natural History 



Volume 17, No. 8, pp. 377-433, 1 fig. 
October 27, 1966 



Birds From North Borneo 



BY 



MAX C. THOMPSON 



'■■-■" jVI 

rt. 
UNiVBRSlTV 



University of Kansas 

Lavi^rence 

1966 



UNIVERSITY OF KANSAS PUBLICATIONS 
MUSEUM OF NATURAL HISTORY 

Institutional libraries interested in publications exchange may obtain this 
series by addressing the Exchange Librarian, University of Kansas Library, 
Lawrence, Kansas. Copies for individuals, persons working in a particular 
field of study, may be obtained by addressing instead the Museum of Natural 
History, University of Kansas, Lawrence, Kansas. When copies are requested 
from the Museiun, 25 cents should be included (for each 100 pages or part 
thereof) for the pvirpose of defraying the costs of wrapping and mailing. For 
certain longer papers an additional amount indicated below, toward the cost 
of production, is to be included. Materials published to date in this series 
are as follows. 

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

Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950. 
•Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 
figures in text. April 9, 1948. 

•Vol. 3. Nos. 1-4 and index. Pp. 1-681. 1951, 

•Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 
figiires in text. December 27, 1951. 
Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953. 
•Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. 
Durrant. Pp. 1-549, 91 figtures in text, 30 tables. August 10, 1952. 
Vol. 7. Nos. 1-15 and index. Pp. 1-651, 1952-1955. 
Vol, 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956. 
Vol. 9. Nos. 1-23 and index. Pp. 1-690, 1955-1960. 
Vol. 10. Nos. 1-10 and index. Pp. 1-626, 1956-1960. 
Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960. 

Vol. 12. •I. 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 modem 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 Sidney Anderson. Pp. 181-216, 49 
figiu-es 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. 

5. Natural history of the Bell Vireo. By Jon C, Barlow. Pp. 241-296, 6 
figures in text. March 7, 1962. 

6. Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C. 
Fox. Pp. 297-307, 6 figures in text. May 21, 1962. 

7. Vertebrates from the barrier island of Tamaulipas, Mexico. By Robert K. 
Selander, Richard F. Johnston, B. J. Wilks, and Gerald G. Raun. Pp. 309- 
345, pis. 5-8. June 18, 1962. 

8. Teeth of edestid sharks. By Theodore H. Eaton, Jr. Pp. 847-362, 10 figures 
in text. October 1, 1962. 

9. Variation in the muscles and nerves of the leg in two genera of grouse (Tym- 
panuchus and Pedioecetes). By E. Bruce Hohnes. Pp. 363-474, 20 figures. 
October 25, 1962. $1.00. 

10. A new genus of Pennsylvanian fish ( Crossopterygii, Coelacanthiformes ) from 
Kansas. By Joan Echols. Pp. 475-501, 7 figures. October 25, 1963. 

11. Observations on the Mississippi kite in southwestern Kansas. By Henry S. 
Fitch. Pp. 503-519. October 25, 1963. 

12. Jaw musculature of the Mourning and White-winged doves. By Robert L, 
Merz. Pp. 521-551, 22 figures. October 25, 1963. 

13. Thoracic and coracoid arteries in two families of birds, Cdlumbidae and 
Hirundinidae. By Marion Anne Jenkinson, Pp, 553-573, 7 figures in text. 
March 2 1964. 

14. The breeding birds of Kansas. By pfechard F. Johnston. Pp. 575-655, 10 
figures. May 18, 1964. 75 cents. 

15. The adductor muscles of the jaw in some primitive reptiles. By Richard C. 
Fox. Pp. 657-680, 11 figures in text. May 18, 1964. 

Index. Pp. 681-694. 

(Continued on inside of back cover) 



University of Kansas Publications 
Museum of Natural History 



Volume 17, No. 8, pp. 377-433, 1 fig. 
October 27, 1966 



Birds From North Borneo 



BY 



max C. THOMPSON 



University of Kansas 

Lawrence 

1966 



University of Kansas Publications, Museum of Natural History 

Editors: E. Raymond Hall, Chairman, Henry S. Fitch, 
Frank B. Cross 



Volume 17, No. 8, pp. 377-433, 1 fig. 
Published October 27, 1966 



University of Kansas 
Lawrence, Kansas 



MAR '^ 1967 

UNIVERSITY 



PRINTED BY 

ROBERT R. fBOB) SANDERS. STATE PRINTER 

TOPEKA. KANSAS 

1 966 



31-4627 



Birds From North Borneo 

BY 
MAX C. THOMPSON 

CONTENTS 

PAGB 

Introduction 379 

Acknowledgments 379 

Methods 380 

Notes on Zoogeography 380 

Collecting Localities and Collectors 381 

Ecology of the Collecting Localities 382 

Ecological Affinities of the Avifauna at Quoin Hill 385 

Seasonality of Breeding 387 

Accounts of Species 390 

Literature Cited 432 

INTRODUCTION 

The major part of this report is an account of birds collected by 
the expedition of the Bemice P. Bishop Museum of Honolulu, 
Hawaii, to North Borneo, from June 24, 1962, through January 14, 
1963. Most of the time spent in the then British Colony was devoted 
to collecting in lowland habitats. The chief collecting localities 
were in the vicinity of Quoin Hill on the Sempoma Peninsula, and 
near Kalabakan. Approximately two weeks were spent in surveying 
the Tenom area. Additional work was done by the North Borneo 
Department of Agriculture after my departure, mainly by Antonio 
D. Garcia. 

ACKNOWLEDGMENTS 

I am indebted to J. L. Gressitt of the Entomology Department of the Bishop 
Museum for providing the opportunity for me to work on the expedition and 
to examine and report on the material collected. Without the help of the 
North Borneo Department of Agriculture, the success of our expedition would 
have been restricted. The Entomologist of North Borneo, G. R. Conway, was 
of great help with our logistic problems as was the Director of the Depart- 
ment, Mr. E. J. H. Berwick, and the Agronomist of Cocoa Research Station, 
Ed Wyrley-Birch. The Bombay Burmah Trading Corporation, Ltd., pro- 
vided facilities and transportation at Kalabakan. Mr. Dai Rees of that cor- 
poration should be especially mentioned. Others who helped are: J. A. 
Comber, Ronnie Young, Mr. and Mrs. Horace Traulsen, Maureen Wyrley- 

(379) 



380 University of Kansas Publs., Mus. Nat. Hist. 

Birch, and the Resident, Tawau, Mr. Peter Edge. The Conservator of Forests 
kindly provided the necessary permits for collecting. 

Authorities of the United States National Museum and The American 
Museum of Natural History generously permitted me to work at those institu- 
tions, using their specimens for comparative studies. Other specimens were 
borrowed from the Museum of Comparative Zoology, Rijksmuseum Van 
Natuuriijke Historic, British Museum (Natural History), and the Yale Peabody 
Museum. Dr. Alexander Wetmore, Herbert Deignan, and Charles Vaurie 
helped with some of the more difficult taxonomic problems. Specimens cited 
in this report are in the Bemice P. Bishop Museum, The University of Kansas 
Museum of Natural History, The University of Michigan Museum of Zoology, 
and the U. S. National Museum. 

Richard F. Johnston and Robert M. Mengel kindly read the manuscript 
and made many helpfid suggestions. The latter re-read it and assisted with 
the editing. 

The most recent comprehensive work published previous to my preparation 
of manuscript for the present account was Smythies ( 1960 ) "The Birds of 
Borneo." 

This report is a partial result of field work supported by a grant from 
the United States Army Medical Research and Development Command, Depart- 
ment of the Army, to the Bemice P. Bishop Museum for research on ectopara- 
sites of vertebrates. The contract numbers were DA-MD-49-193-62-G47 and 
G65. The Chapman Fund of The American Museum of Natural History met 
part of the cost of transporting, to and from the United States, specimens from 
North Borneo collected after I left there. 

METHODS 

While collecting at Quoin Hill, we used only guns in taking birds. At an 
area 12 miles north of Kalabakan, we supplemented the guns with mist nets 
in the primary forest. This method was excellent for taking rarely seen 
species. For example the thrush Zoothera interpres was never seen in the 
field but was taken several times in mist nets. 

Another method of collecting was the use of native snares. Such snares 
were made of heavy nylon string tied to a sapling, held down by a nylon 
string attached to a treadle. When a bird stepped on the treadle, it tripped 
the snare and a loop closed about its feet, hoisting it aloft. To divert large 
groimd birds and mammals into the snare, natives placed brush barriers 
along the top of a ridge for one or two miles. Animals were diverted by 
these barriers until they came to an opening; if they went through they 
usually tripped the trap. Pheasants and the large ground cuckoo were taken 
in this manner. 

NOTES ON ZOOGEOGRAPHY 

The avifauna of Borneo is of Indo-Malayan aflBnities. The num- 
ber of birds endemic to Borneo is relatively small; most species 
are shared with the Asian mainland. Only 29 birds are known to 
be endemic to the island and 17 of these are montane. The large 
proportion of montane endemics is not surprising, because Borneo 



I 



I 



Birds From North Borneo 



381 



has been connected with the Asian continent in recent geological 
time; lowland isolation, and differentiation, has been less exten- 
sive than the montane. The Sunda Shelf, on which Borneo is 
situated, hes in a shallow sea generally less than 300 feet deep. 
Beaufort has shown that the Malay Peninsula, Sumatra, and Java 
were connected until early historic times (Darlington, 1957:488). 
The endemic species in Borneo are members of four, possibly 
five, genera that are also endemic. Four of these five genera are 
montane in distribution. The only endemic for which the geo- 
graphic history cannot be adequately explained is the monotypic 
Pityriasis gymnocephala. Its affinities seem to be with the Cracti- 
cidae of New Guinea and Australia. The species has been found 
throughout Borneo. Since Pityriasis is endemic to Borneo, it prob- 
ably was detached from the parent stock at an early period. The 
Australasian affinities of Pityriasis emphasize its zoogeographical 
pecuharities. A more detailed discussion of this species appears 
in the annotated hst below. 

COLLECTING LOCALITIES AND COLLECTORS 



1. Cocoa Research Station, Quoin Hill, elevation 750 feet, Tawau. Max 
C. Thompson (MCT) and Antonio D. Garcia (ADG). 

2. Tawau. Max C. Thompson. 

of Kala- 
Max C. 



116" 



NORTH BORNEO 



-v 



10 "(? 

HLOHeiERS 




3. Twelve miles north 
bakan, elevation 600 feet. 
Thompson. 

4. Kalabakan, elevation 50 feet. 
Max C. Thompson. 

5. Tiger Estate, 20 miles north- 
west of Tawau. Max C. Thompson, 
Antonio D. Garcia. 

6. Ulu Balung Cocoa Estate, Mile 
27, Quoin Hill, elevation 750 feet, 
Tawau. Antonio D. Garcia. 

7. Karindingen Island. Max C. 
Thompson. 

8. Siamil Island. Max C. Thomp- 
son. 

9. Lahad Datu. Antonio D. Gar- 
cia. 

10. Kuala Sumawang, 25 miles west of Sandakan. Antonio D. Garcia. 

11. Agricultural Station, Mile 17, Sandakan (Gum-Gum). Antonio D. 
Garcia. 

12. One-fourth mile east Gum-Gum, Sandakan. Antonio D. Garcia. 

13. Lamag, Kinabatangan River. Antonio D. Garcia. 

14. Pintasan Agriculture Station, Kinabatangan River. Antonio D. Garcia. 

15. Kampong Kuamut, Kinabatangan River. Antonio D. Garcia. 

16. Kampong Maluwa, Kinabatangan River. Antonio D. Garcia. 



Fig. 1. Localities from which collec- 
tors from the Department of Agricul- 
ture or I saved specimens in North 
Borneo. 



382 University of Kansas Publs., Mus. Nat. Hist. 

17. Ka-Karis, Kinabatangan River, elevation 200 feet. Antonio D. Garcia. 

18. Tongod, Kinabatangan River, elevation 300 feet. Antonio D. Garcia. 

19. Tuaran. Max C. Thompson, Antonio D. Garcia, S. F. W. Cheng 
(SFWC). 

20. Tehpok. Antonio D. Garcia, G. R. Conway. 

21. Mt. Rumas, 5 miles northwest of Tuaran, elevation 75 feet, Antonio D. 
Garcia. 

22. Five and one-half miles southwest of Tenom, elevation 4,000 feet. Max 
C. Thompson. 

23. Tenom, elevation 600 feet. Max C. Thompson. 

24. Kampong Banjar, Mile 29, Keningau. Antonio D. Garcia. 

25. Oil Palm Research Station, Mile 32, elevation 40 feet, Sandakan. 
Antonio D. Garcia. 

ECOLOGY OF THE COLLECTING LOCALITIES 

Quoin Hill. — At this locality I recognized five habitat types as 
follows : 

Primary forest. — We were fortunate to be able to work at Quoin 
Hill because it had been opened to cultivation (of Cocoa, Theo- 
broma cacao) for only a few years. Thus the primary forest here 
started at the edge of the Cocoa Research Station. This was in 
marked contrast to areas on the west coast, where one would need 
to travel many miles inland to find virgin forest. The forest at 
Quoin Hill was typical tropical rain-forest, composed mostly of 
dipterocarps (Dipterocarpaceae). These comprise an essentially 
Indo-Malayan family, members of which are so conspicuous that 
we commonly referred to it as Evergreen Dipterocarp Forest. The 
lowland forests of Borneo are composed of approximately 3,000 
species of trees (Browne, 1955). At Quoin Hill, as in most of the 
tropical rain-forest of Borneo, the forest canopy is stratified in three 
layers, a distinct and easily recognizable top story and less easily 
separable middle and lower stories. The top canopy is composed 
of fohage of giant trees that may tower to heights of 200 feet and 
have trunks three to seven feet in diameter. The trunk is usually 
unbranched for 50 to 100 feet and the whole tree is supported by 
buttresses jutting out from the main trunk. Some of the most im- 
portant plants in the tropical rain-forest are the strangler figs 
( Ficus sp. ) . These plants, when in fruit, draw birds in large flocks 
to feed upon them. Such figs were common about the edges of the 
research station and some birds taken from these trees were never 
taken elsewhere. The birds seemed to wait for a certain degree of 
ripeness of fruits; on one day the figs were unmolested and the 
next day the trees would be swarming with birds. Strangler fig 
trees reach tremendous size and help form the upper forest canopy. 



Birds From North Borneo 383 

The middle and lower forest canopies are not easily separable 
and I shall speak of them together. The trees forming these varied 
from 10 to 60 feet in height. The ground surface beneath the trees 
was usually bare except for leaf litter and dead branches. Sun- 
light penetrates only where the big trees have been removed or 
where the larger trees are otherwise widely spaced. At Quoin Hill 
the large trees of species affording lumber of commercial quality 
had been taken out, modifying somewhat the character of the 
forest. Such forest actually contained many of the animals charac- 
teristic of primary forest, and I refer to it as badly disturbed pri- 
mary forest. 

Secondary forest. — In some of the areas adjoining the research 
station, roads had been bulldozed for future expansion and trees 
had been cut. These areas were starting to grow dense stands of 
grass and shrubs and will be jungle in a few years unless cut back. 
Most of the trees in this area are saplings with some trees as large 
as a foot in diameter. 

Fluviatile waters. — There are numerous small streams in the 
Quoin Hill area, the largest being the Balung River and Apas 
River. Little work was done along these streams and only the 
thrushes of the genus Enictirus and some kingfishers seemed to be 
confined to them. 

Cocoa plantations. — Artificial plantings of cocoa, Theobroma 
cacao, formed a major habitat type at Quoin Hill, and provided a 
major source of food for birds. Cocoa planters have found it nec- 
essary to provide shade with trees of some other species. In some 
instances trees from the original primary forest were left standing 
to provide this shade, but more often exotic trees were planted. 
Most of the shade trees were of no use to birds save for providing 
resting places. Trema orientalis was the most important in pro- 
viding food for birds. Its fruit was used more by the frugivorous 
species of birds than that of any other tree in the cocoa plantings. 
Tree Cassava, an exotic, was constantly attended by the nectariniids, 
or honey creepers. 

Although the cocoa plantings did not provide much plant ma- 
terial for bird food, they did apparently nourish a horde of insects, 
which the birds fed upon. A Drongo-cuckoo, Sumiculus lugubris, 
had 50 caterpillars in its stomach. Healthy cocoa trees were 
sparsely inhabited by birds but areas that were obviously infested 
with insects literally swarmed with birds. Dead shade trees in the 
cocoa plantings also provided food for woodpeckers, with four 
species being found utilizing these dead trees. 



384 University of Kansas Publs., Mus. Nat. Hist. 

Abaca.— The last of the habitat types that I recognized at Quoin 
Hill was a small grove of Abaca, Musa textilis, and wild bananas, 
Musa sp. This habitat type was frequented by spiderhunters 
( Arachnothera sp. ) of the family Nectariniidae. 

Kalabakan. — We worked at three localities in this area: 12 miles 
north of Kalabakan, Brantian Estate, and Kalabakan. 

Primary forest. — We were fortunate in being able to work on the 
very edge of the primary forest 12 miles north of Kalabakan. The 
composition of the primary forest was much like that at Quoin Hill 
and will not be discussed further. About a month after we arrived 
at our forest camp, logging crews moved in and cut the commercial 
timber near our area. The only immediately noticeable difference 
in the makeup of the avifauna after destruction of the forest canopy 
was the appearance of the drongo Dicrurus aeneus. This drongo 
was seen in areas where the trees had been cut, sitting on limbs and 
darting out after insects. 

Secondary forest. — The area around Kalabakan proper was in 
secondary forest, which was almost impossible to penetrate. At 
Kalabakan, Cymbirhynchus macrorhynchus, Cecropsia striohta, and 
Macronous ptilosus were taken and I did not see them elsewhere. 
Kalabakan is situated on the Kalabakan River at the upper tidal 
limit. The Nipa-Mangrove association, not investigated, lies im- 
mediately below Kalabakan. 

The Brantian Estate area was mostly in secondary forest and was 
situated on the Brantian River. There were some fairly large areas 
of grass with water buffalo wallows in them. These grassy areas 
were favorite haunts of the Painted Quail, Coturnix chinensis. 

Tenom. — The first locality that we investigated was 5.5 miles 
southwest of Tenom, approximately 4,000 feet elevation, in moss 
forest. A few days were spent collecting in the area of Tenom 
itself. 

Moss forest. — The lower altitudinal limit of the moss forest was 
about 3,600 feet. The trees on top of the mountain were mostly 
oaks (Quercus) and were festooned with ferns, orchids, and other 
epiphytes. The area had been used as a triangulation station by a 
survey team and a small area on top of the mountain had been 
cleared earlier. At the time of our visit this small area had grown 
to secondary vegetation, mostly Fandanus. The oaks in the pri- 
mary forest surrounding this disturbed area were generally about 
30 to 50 feet high and there was little undergrowth in virgin stands. 
This area was usually swathed in fog from three o'clock in the 



Birds From North Borneo 385 

afternoon until eleven o'clock the next morning. One morning of 
our fourteen there was clear. 

Paddy. — The area visited at Tenom itself consisted mostly of old 
paddy grown to grass and scrub. Forest did occur but was of 
secondary nature in the immediate vicinity of Tenom. 

SiAMiL Island. — This island is about one mile in circumference 
and the highest point is about 300 feet above sea level. The island 
has high bluffs on three sides but slopes gently to the sea on the 
other. There were patches of forest left on the island, one on the 
north side and one on the south. The sheer bluffs on the east side 
of the island were covered with Pandanus sp. The undergrowth 
of the north forest had been cut, leaving extensive bare areas. The 
principal undergrowth was rattan. The natives are clearing and 
planting more of the island to coconuts and hope eventually to 
clear it completely. 

Karindingen Island. — This island, about half a mile in circum- 
ference and between 10 and 20 feet above sea level at its highest 
point, was surrounded by extensive coral reefs and sand; the prin- 
cipal vegetation was mangroves. 

ECOLOGICAL AFFINITIES OF THE AVIFAUNA AT 

QUOIN HILL 

More time was spent at Quoin Hill than at any other locality. 
Fifty five of the more common and hence best-known birds are 
hsted in Table 1 together with their primary and secondary prefer- 
ences of habitat. The habitat distribution of the birds shows the 
amount of secondary utilization of habitats by birds that occurred 
predominantly in one habitat. Cocoa was utilized by 6.2 per cent 
of the birds of the primary forest, and 88.8 per cent of birds of the 
secondary forest. This indicates that cocoa is an effective sub- 
stitute for secondary forest for some birds. Of the species of the 
primary forest, 18.7 per cent occurred also in secondary forest; 
thus, three times as many species of primary forest utilized sec- 
ondary forest as utilized cocoa. This too might be expected, since 
"secondary" forest is of frequent natural occurrence and an ancient 
feature while the comparatively simple cocoa plantings are new and 
artificial. 

The avifauna at Quoin Hill was a mixture of montane, submon- 
tane, and lowland species. Smythies (1957:527) defines four 
altitudinal areas of distribution: Higher Montane, Montane, Sub- 
montane, and Lowland. Higher Montane birds have not been 



386 University of Kansas Publs., Mus. Nat. Hist. 

TABLE 1. — Habitat preferences of 55 Quoin Hill birds. 



X=Primary 
O^Secondary 

Species 



Treron curvirostra . . . . 
Cacomantis merulinus . 
Chalcites malayanus . . 



Phaenicophaeus 
chlorophaeus . 



Harpactes diardi 

Uarpactes duvauceli 

Alcedo euTyzona 

Ceyx erithacus 

Eurystomus orientalis 

Calorhamphus fuliginosua . 
Megalaima chrysopogon . . . 

Megalaima henrici 

Sasja abnormis 

Meiglyptes tukki 

Dryocopus javensis 

Chrysocolaptes validus . . . . 
Eurylaimus ochromalus . . 

Pitta guajana 

Coracina fimbriata 

Aegithina viridissima. . . . 
Chloropsis cyanopogon . . . 

Irena puella 

Pycnonotus brunneua. . . . 

Criniger bres 

Criniger phaeocephalus . . . 

Criniger finschii 

Hypsipetes criniger 

Copsychus pyrrhopygus . . 
Copsychus stricklandi . . . . 



X 



o 



X 



X 



X 



X 



X 



X 



X 



o 



X=Primary 
0=Secondary 
Species 



Enicurus ruficapillus . . . . 
Pellorneum capistratus . . . 
Trichastoma malaccense . . 
Triehastoma sepiarium. . . 
Malacopteron magnum .... 
Malacopteron magnirostre. . 

Kenopia striata 

Stachyris poliocephala 

Alcippe brunneicaiida 

OrthotomuB atrogularis . . . . 

Orthotomux sepium 

Rhipidura perlata 

Muscicapa dumetoria 

Rhinomyias umbratilis. . . . 

Hypothymis azurea 

Antkreptes simplex 

Anthreptes rhodolaema . . . . 
Nectarinia hypogrammica. . 
Arachnothera longirostris . . 
Arachnothera flavigaster . . . 
Arachnothera chrysogenys . . 

Arachnothera affinis 

Ziosterops everetti 

Lonchura fuscans 

Oriolus xanthonotus 

Platysmurus leucopterus . . , 

Total Primary 

Total Secondary 



X 



tn 



o 



o 



X 



recorded on mountains the summits of which are lower than 5,000 
feet, although on higher peaks the actual lower limit of occurrence 
may be considerably below 5,000 feet. Montane birds have not 
been recorded on mountains the summits of which are lower than 



Birds From North Borneo 387 

3,000 feet, although specimens may have been taken below that 
altitude on higher peaks. Submontane, as defined by Smythies, is 
a comprehensive term appHed to birds occurring from sea level to 
an elevation of 5,000 feet but ordinarily not found away from 
mountainous country. The Lowland birds normally range from 
sea level to 3,000 feet. Of the 125 species of birds observed at 
Quoin Hill, 1.6 per cent were Montane, 14.4 per cent were Sub- 
montane and 84 per cent were Lowland species. The distribution of 
birds 12 miles north of Kalabakan closely resembled that at Quoin 
Hill except for the total absence of Montane species and an increase 
of Submontane species to 25 per cent. The observation of fewer 
species (48) can be attributed to the nearly uniform habitat. 

The avifauna in the moss forest 5.5 miles southwest of Tenom 
was unusual in that 45.4 per cent consisted of Lowland species; 
this locality hes 4,000 feet above sea level, yet only 27.3 per cent of 
its species were Submontane and 27.3 per cent Montane. If one 
looks at these figures from the standpoint of the actual importance 
of the three groups at this place, however, a different picture 
emerges. Some of the Lowland species were seen only once while 
I was there and few were common, while all of the Submontane 
and most of the Montane forms were more or less common. 

SEASONALITY OF BREEDING 

The breeding season in North Borneo. — Birds in the Quoin Hill 
area of eastern Borneo seem to breed most commonly in June, July, 
and August. Table 2 lists 34 of the more common species at Quoin 
Hill for which evidence on breeding was available. The actual 
evidence was provided by females with active brood patches or 
active ovaries, males with enlarged testes, birds in juvenal plumage, 
or birds actively in annual molt. From such data dates of presumed 
breeding were extrapolated. In Table 2, the solid black lines 
indicate dates for which both male and female were in breeding 
condition. The dotted lines indicate enlarged testes but no evidence 
of breeding in females. In the bottom line of Table 2, the figures 
indicate the percentage of the population breeding in any one 
month. For instance, 2.9 per cent of the birds were breeding in 
March, but 73 per cent were breeding in June. Rainfall records 
from the Cocoa Research Station from April, 1959, to December, 
1964, were available to me. These data, along with the average 
for each month, are given in Table 3. There appears to be little 
correlation between rainfall and breeding season at Quoin Hill. 



388 



University of KIaj^sas Publs., Mus. Nat. Hist. 



TABLE 2. — Seasonality of common breeding birds at Quoin Hill. Solid lines 
indicate times of occurrence of known breeding; dotted lines represent times 

of presumed breeding. 



Species 


J 


F 


M 


A 


M 


J 


J 


A 


S 





N 


D 


Treron curvirostra 


J 












• 












Cacomantix merulinus 








Chnlrite.s Tnalatianus 


- 


• 








Phaenicophaeus curvirostris 

Collocalia fuciphaga 

Chaetura leucowQialis 




















— 




Harvactes diardi 








Euriistojnus orientalis 










Calorhamphus fuliginosus 

Megalaima chrysopogon 

Megalaima myslacophanes 

Siafsia ahnormis 






























— 




Micropternus brachyurus 

Dryocopits javensis 


- 







Pycnonotus cyaniventris 

Pvcnonotus atricevs 














Purnnnntiift brnnne.us 










Pycnonotus erythrophthalmus . . . . 
Zoothera intervres 






— 


- 








— 




Malacopteron magnirostre 

Piilocichla leucogrammica 

Kp.nnrtia striata, 




— 















Stachyris niaculata 








Orthntnmus alrooularis .... 










Orthntnmiis serice.us 














Orthotomus seviuni 












Rhividura verlata 






- 








Prionichilus xavthopygiixs 

A nthreTotps rhodolaema 


- 














Arachnothera flaviqaster 

Pityriasis gymnocephala 

Oriolus xanthonotus 




















Platysmurus leucopterus 
















F 


M 


A 


M 


J 


J 


A 


S 





N 


D 


The percentage of breeding in 
any one month is as follows: 








2.9 


8.8 


38 


73 


58 


50 


35 


17 


11 


8 



A true dry season in the Quoin Hill area does not occur, but 
monthly rainfall has varied from 0.57 inches to 21.27 inches in a 
single year. 

Birds in the moss forest near Tenom appeared to be breeding in 
January, paralleling the trend found by Voous (1950a) for the 
lowlands of Borneo. 

Other Bornean observations. — Voous (1950a) summarized data 
assembled by Coomans de Ruiter on the breeding of birds in the 
lov^'land of western Borneo near Pontianak, It appears that the 



Birds From North Borneo 



389 



TABLE 3. — Monthly rainfall records. Cocoa Research Station, Quoin Hill. 



Year 


Jan. 


Feb. 


Mar. 


April 


May 


June 


July 


Aug. 


Sept. 


Oct. 


Nov. 


Dec. 


1959.. . 








6.49 
10.65 


12.16 
8.84 


11.11 
11.00 


7.64 
6.31 


12.11 
11.25 


4.75 
8.56 


8.33 
5.49 


12.10 
8.39 


13.81 


I960.... 


9.24 


8.17 


3.76 


11.81 


1961... . 


6.68 


8.06 


4.35 


4.74 


7.55 


7.25 


5.93 


2.40 


7.47 


5.58 


4.38 


10.73 


1962.... 


3.82 


6.76 


13.72 


9.68 


6.82 


7.49 


6.59 


5.82 


7.81 


9.47 


19.80 


9.28 


1963.... 


21.27 


8.18 


7.64 


0.57 


5.83 


4.62 


0.64 


12.49 


5.24 


8.75 


7.43 


11.05 


1964.... 


4.17 


7.92 


4.40 


11.20 


11.82 


8.04 


2.42 


7.52 


5.69 


13.15 


8.82 


9.88 


Average 


9.03 


7.81 


6.77 


7.22 


8.83 


8.25 


4.92 


8.59 


6.58 


8.46 


10.15 


11.09 



breeding season in that part of Borneo, and indeed in all of western 
Borneo (Banks, 1950), starts in December and reaches a peak in 
March. 

Gibson-Hill (1952) has questioned Banks' (1950) interpretation 
of data from the egg collection of V. W. Ryves. Gibson-Hill has 
shown that the data collected by Ryves covered two widely 
separated localities, one at Kiau near Kota Belud and the other 
near Sandakan. The former locality is on the west coast of North 
Borneo and the latter on the east coast of North Borneo. Gibson- 
Hill points out, and rightly so, that the timing of the rainfall in 
different parts of Borneo must be taken into account because of 
the large regional variation. The nesting data from the Ryves 
egg collection are scant and when used alone possibly yield a 
distorted view of the actual breeding season. Ryves did no col- 
lecting in the Sandakan area between September and March, and 
in the Kiau area between May and January. Although the breed- 
ing data from North Borneo accumulated by both Ryves and myself 
are limited, and records of rainfall are scant, there appears to be 
a trend toward breeding after the heavy rains have fallen. 

Seasonality of breeding in tropical birds. — Possibly Bomean birds 
breed mostly in the "driest" part of the year. If so, this is in con- 
trast with the time of breeding of birds of other tropical areas. 
Moreau (1950) found that in the Congo there was no distinct 
breeding season for most groups of birds, but that in East Africa 
there was a double breeding season; the peaks coincided with the 
two rainy seasons. Lack (1950) found that the Geospizinae of the 
Galapagos breed only when it rains and that rainfall causes a 
flurry of nest building and singing. If the rains stop, then the 
courtship activities stop until the next rains. Miller (1963) found 
that in birds of a western Andean cloud forest the breeding season 



390 University of Kansas Publs., Mus. Nat. Hist. 

was spread over the year and that breeding could not be correlated 
with rainfall. 

Obviously more study is required on breeding of birds in Borneo 
before the timing of the annual cycle can be ascertained. 

ACCOUNTS OF SPECIES 

The EngHsh names used in this report follow Smythies (1960) 
where possible. If the bird has not been recorded from Borneo 
previously, then I have resorted to Delacours and Mayr's "Birds of 
the Philippines" ( 1946). The taxonomy is that of Smythies (1960), 
except where current American or my own opinion differs and 
where new evidence has warranted a change. The sequence of 
families is that of Wetmore ( I960 ) . 

Two species listed beyond that had not previously been recorded 
from the island of Borneo are: Red-footed Booby, Sida stila; and 
Whitehead's Thick-head, PachycepJiala whiteheadi. Six additional 
kinds listed beyond had not previously been recorded from North 
Borneo. They are: Chinese Egret, Egretta eidophotes; Knot, Cali- 
dris canutus; Ground Cuckoo, Carpococcijx radiceus; Stachyris 
nigriceps hartleyi; Finch's Bulbul, Criniger finschii; and Pale Blue 
Flycatcher, Muscicapa imicolor. 

Sula sula (Linnaeus): Red-footed Booby. — A captive seen in the Tawau 
police station was said to have been caught when it landed at night on a 
pohce launch anchored off Siamil Island in or near May, 1962. The bird was 
in first-year or second-year plumage, seemed to be tame, and was thriving on a 
diet of fish. Identification was made by Alexander Wetmore from a photo- 
graphic print. This is the first seemingly substantiated record of the species 
from Borneo, although it probably occurs there regularly. The species breeds 
on Bankoran Island and on Tubbataha Reef in the Sulu Sea (Smythies, 1960: 
113). 

Fregata ariel (G. R. Gray): Lesser Frigatebird. — This conspicuous bird on 
the waters around Tawau is occasionally seen in Cowie Harbor but more often 
along the coast outside the Harbor. On September 20 an estimated 300-500 
birds circled over the shore northeast of Tawau. 

Ardea sumatrana sumatrana Raffles: Dusky-gray Heron. — Specimens, 2. 
Karindingen Island: $ testis 12 X 23 mm., November 24, 1962, MCT 3308; 
$ , November 24, 1962, MCT 3309. 

The specimens were taken on the coral sand beach surrounding Karindingen 
Island. At least 10 were seen feeding on the beach and on a reef. One ( MCT 
3308 ) had testes of a size suggesting that it was in breeding condition. 

Casmerodius alba tnodestus (Gray): Common Egret. — Specimen, 1. Karin- 
dingen Island: 9 , November 24, 1962, MCT 3310. 

This species was seen on three dates: August 17 at Karindingen Island, 
where 30 were observed feeding along the mangroves bordering the island; on 
November 24, on the same island, where a specimen was taken from a flock 



Birds From North Borneo 391 

of 30 birds that had been feeding on the coral sand and reef; and on November 
30 when a single bird landed at the small reservoir at the Cocoa Research 
Station and began to feed in the shallow water. Only four specimens appear 
to have been recorded earlier (Smythies, 1957:561), although Smythies (1963: 
270) lists additional sight records. 

Egretta garzetta ( Linnaeus ) : Little Egret. — On November 24 at Karin- 
dingen Island, I observed several mixing freely with individuals of E. sacra 
on the coral sand beaches and reefs. 

Egretta eulophotes (Swinhoe): Chinese Egret. — Specimen, 1: Ka-Karis, 
Kinabatangan River, 200 feet: $ , October 20, 1963, ADC 326. 

This is the only record known to me of tliis species from North Borneo. 
Smythies ( 1960:126) lists 11 specimens from other parts of Borneo. 

Egretta sacra ( Gmelin ) : Reef Egret. — The species was observed at Karin- 
dingen Island on August 17 and November 24, 1962, and was the most com- 
mon egret on the island. One compact flock of 50 was seen at high tide. No 
white-phased birds were in the flock. 

Bubulcus ibis coromandus (Boddaert): Cattle Egret. — Specimen, 1: Ton- 
god: 9 , October 21, 1963, ADC 328. 

Individuals were observed daily at Tuaran with cattle at the Agricultural 
Research Centre. I saw them on December 9 when I arrived at Tuaran and 
again on January 13, when I departed. 

Ardeola Bacchus (Bonaparte): Chinese Pond Heron. — Specimen, 1: 12 
mi. N Kalabakan: 5, October 26, 1962, MCT 3151. 

The specimen was brought to our camp by a local boy who obtained it on a 
stream that ran through his kampong (village) near our sulap (hut). The 
stream was a small one that ran out of the primary forest, through the village, 
and back into secondary forest. 

Butorides striatus (Linnaeus): Little Green Heron. — Specimens, 2: Teli- 
pok: 9, March 10, 1963, TM 67; Sex?, December 13, 1962, TM 6. 

This heron was seen at Karindingen Island on August 17, 1962, in a small 
swamp near the Tawau airport on September 16, and on the reef at Siamil 
Island on September 18. The specimens collected at Telipok were not identi- 
fied to subspecies. 

Ixobrychus cinnamomeus cinnamomeus ( Gmelin ) : Chestnut Bittern. — 
Specimen, 1: Tuaran: Sex?, December 24, 1963. SCFC 32. 

This specimen was probably taken in the paddy around the Agricultural 
Research Centre, where I saw Chestnut Bitterns in January 1963. 

Leptoptilos javanicus (Horsfield): Lesser Adjutant Stork. — Specimen, 1: 
Karindingen Island: $ testis small, November 24, 1962, MCT 3311, 

My first observation of this species was at Karindingen Island on August 17, 
when approximately 100 birds were seen feeding on the coral sand and reefs. 
I saw them also within a mile of Sempoma along the ship channel, usually 
in the vicinity of fish traps. On August 31 two birds flew high overhead at 
Quoin Hill, proceeding in the direction of Cowie Harbor. On November 24, 
large numbers at Karindingen Island were sitting in mangroves and feeding 
on the reefs around the island. Two birds were caught by the rising tide 
while feeding on the reef and were unable to take off. One of these was shot 
in approximately six to seven feet of water; its feathers were completely water- 



392 University of Kansas Publs,, Mus. Nat. Hist. 

logged. Interestingly tlie flaky skin of the crown had blue-green algae growing 
on it. These birds were heard calling many times on the reef and in the 
mangroves; the call is much like the deep growl of a dog. 

Sims, Banks, and Harrison have found the storks common in this area 
(Smythies, 1957:569). Although I could find no evidence of it, possibly 
Karindingen Island is a nesting locality. 

Anas querquedula Linnaeus: Garganey. — Specimen, 1: Kg. Banjar: sex? 
November 13, 1963, ADG 329. 

This teal obtained by Garcia is our only record of any duck from North 
Borneo. 

Haliastur indus (Boddaert): Brahminy Kite. — This common resident of 
the Cocoa Research Station was observed almost daily while I was in the 
field. It seemed to prefer the cleared areas replanted to cocoa and oil palm 
and was common also at Tawau, Sempoma, and Karindingen Island. 

Accipiier trivirgatus microstictus Mayr: Crested Goshawk. — Specimens, 2: 
Pintasan Agriculture Station: $ , October 17, 1963, ADG 314. Cocoa Re- 
search Station: $ , April 30, 1963, ADG 89. 

Spizaetus cirrhatus limnaetus ( Horsfield ) : Changeable Hawk-eagle. — 
Specimen, 1: Tiger Estate: 5 , April 28, 1963, ADG 74 (Specimen in black 
phase ) . 

Hieraetus kienerii formosus (E. Geoffroy): Rufous-bellied Hawk-eagle. — 
Specimens, 2: Cocoa Research Station: 9 imm., July 10, 1962, MCT 2615. 
Tiger Estate: 2 , July 13, 1962, MCT 2621. 

The presence of an immature bird of this species in North Borneo lends 
support to tlie suggestion of Smythies (1957:580) that this species breeds in 
Borneo. 

Haliaeetus leucogaster ( Gmehn ) : White-bellied Sea Eagle. — This eagle 
appears to be fairly common along the coast in the Tawau-Darvel Bay area. 
I saw it around Cowie Harbor, Tawau, Sempoma, and Siamil Island. 

SpUornis cheela Latham: Crested Serpent Eagle. — I saw this eagle but 
once, circling overhead at the Cocoa Research Station on September 26, 1962. 

Microhierax latifrons Sharpe: White-fronted Falconet. — Specimens, 2: 

Cocoa Research Station: $ testis 2x1 mm., 41.2 gm., July 6, 1962, MCT 

2600. Tiger Estate: $ oviduct regressing, old brood patch, December 1, 
1962, MCT 3418. 

This species was first seen at the Cocoa Research Station. The specimen 
taken there bobbed its head in the manner of various other falcons. On two 
occasions individuals were observed sitting in a low tree in the front yard of 
a home in Kalabakan. 

These records extend the known range of this species south from Darvel 
Bay (Smythies, 1960:161) to Cowie Harbor. 

Falco peregrinus japonensis Gmelin: Peregrine Falcon. — Specimen, 1: 
Kampong Kuamut: S , October 20, 1963, ADG 327. 

Coturnix chinensis ( Linnaeus ) : Painted Quail. — Specimen, 1 : Cocoa Re- 
search Station: $ testis 8x5 mm., September 6, 1962, MCT 2881. 

This specimen is intermediate between C. c. lineata and C. c. caerulescens, 
tending shghtly toward the latter in having more rufous coloring on the tertials. 



Birds From North Borneo 393 

R. E. Kuntz took a male at Ranau (USNM 472504) that was also intermediate 
in its characters but was referable to C. c. lineata. These two specimens, when 
compared with series of specimens from the Philippines and Sumatra, fitted 
into a clinal progression of increasingly rufous tertials toward Sumatra. Peters 
(1934:96) united C. c. caerulescens with C. c. palmeri, but Amadon (in litt.) 
retains C. c. caerulescens (Smythies, 1957:588). No clear-cut distributional 
pattern is yet discernible in North Borneo and the arrangement of Amadon 
(MS) should probably be reviewed again when more specimens become 
available. 

These birds were observed several times at the Cocoa Research Station but 
were difficult to collect. They were abundant in the grasslands on the 
Brantian Estate. 

RoUulus roulrotd (Scopoli): Crested Green Wood Partridge. — It was ob- 
served once along the Apas River at the Cocoa Research Station, in primary 
forest. 

Haetnctortyx sanguiniceps Sharpe: Crimson-headed Wood Partridge. — 
Frederick Durm saw one fly across the padang at the Cocoa Research Station 
rest house on September 9. 

Lophura ignita (Shaw and Nodder): Crested Fireback Pheasant. — Speci- 
mens, 3: Cocoa Research Station: 9, molting, July 20, 1962, MCT 2624. 
12 mi. N Kalabakan: 2 , November 4, 1962, MCT 3206; $ , November 7, 
1962, MCT 3216. 

This pheasant was taken in native snares from primary forest. 

Lophura btdweri (Sharpe): Bulwer's Pheasant. — Specimens, 3: 5.5 mi. 
SW Tenom: $ , December 25, 1962, MCT 3534; $ , December 27, 1962, 
MCT 3539; 9 , December 27, 1962, MCT 3540. 

This bird was first taken in a snare 12 miles north of Kalabakan. Un- 
fortunately, the specimen was stolen and only its tail feathers were brought 
in by the trappers. Mr. Comber of Sapong Estates in Tenom said this species 
comes in numbers with the wild-pig migration and that he had observed this 
at least three times. He has also seen three-quarters grown young at Tenom, 
so they seemingly nest there. They were found only in primary forest. 

These specimens constitute the first records of the species for the west coast 
of North Borneo (cf. Smythies, 1957:593). 

Argusianus argus grayi (Elliott): Great Argus Pheasant. — Specimens, 6: 
12 mi. N Kalabakan: $, October 14, 1962, MCT 3034; $, October 22, 1962, 
MCT 3121; S , October 23, 1962, MCT 3130; $ , October 27, 1962, MCT 
3166; $ ?, October 29, 1962, MCT 3307. 5.5 mi. SW Tenom: $ , December 
19, 1962, MCT 3464. 

This species is a common resident of the primary forest at Kalabakan and 
in the lower areas around Tenom (J. A. Comber, pers. comm. ). It is found 
only in primary forest. 

Rallina fasciata (RaflBes): Malaysian Banded Crake. — Specimen, 1: Tiger 
Estate: $ , July 11, 1963, ADG 193. 

Amauromis phoenicurus javanicus (Horsfield): White-breasted Water- 
hen. — Specimens, 3: Tuaran: sex?, March 1, 1963, ADG 5. Telipok: $, 
February 2, 1963, TM 23; $ , February 2, 1963, TM 24. 

This species was commonly observed in marshes near Tawau and on the 
Brantian Estate. 

2—4627 



394 University of Kansas Publs., Mus. Nat. Hist. 

Gallicrex cinerea (Gmelin): Watercock. — Specimen, 1; Tiger Estate: 
sex?, December 17, 1962 (taken on dry grassland). 

Squatarola squatarola (Linneaus): Black-bellied Plover. — On September 2 
I saw a flock of 12 plovers on the Tawau golf course. They were in a mottled 
plumage indicating extensive molt and feather growth. Later in the day a 
bird in almost complete breeding plumage flew overhead. 

Charadrius peroni Schlegel: Malay Sand Plover. — On September 15, one 
was sitting on the Tawau Airport runway. Another was observed on November 
20, feeding near a water buffalo wallow on tlie Brantian Estate. Smythies 
(1960:191) lists sandy beaches as the only habitat. 

Charadrius leschenaulti Lesson: Large Sand Plover. — On September 16 one 
was sitting on the Tawau Airport runway. 

Numenius phaeopus variegatus ( Scopoli ) : Whimbrel. — Specimen, 1 : Kar- 
indingen Island: $ , November 24, 1962, MCT 3315. 

This was the most common curlew around Karindingen Island on November 
24. Flocks of 10 to 20 individuals were feeding on the coral sand around the 
island, and a few individuals were sitting in the tops of dead mangroves at 
low tide. 

Numenius arquata ( Linnaeus ) : Common Curlew. — On November 24, a 
Common Curlew flew from Karindingen Island toward the mainland. 

Numenius madagascariensis ( Linnaeus ) : Eastern Cm-lew. — I first observed 
this curlew on August 17 at Karindingen Island, where 50 were feeding on 
the coral sand. At that time it was the most abundant shorebird. When I 
returned to the island on November 24, several were seen around the island, 
but the species was not so abundant as N. phaeopus. 

Limosa lapponica ( Linnaeus ) : Bar-tailed Godwit. — A flock of five was ob- 
served at Karindingen Island on November 24. 

Tringa totanus eurhinus ( Oberholser ) : Redshank. — Specimens, 2: Karin- 
dingen Island: ?, November 24, 1962, MCT 3312; ?, November 24, 1962, 
MCT 3313. 

On August 17, this bird was common and feeding on the coral sand at 
Karindingen Island. When 1 revisited the island on November 24 the Redshank 
seemed to prefer the mangrove areas for feeding and was the most common 
wader. 

Tringa ochropus Linnaeus: Green Sandpiper. — Specimen, 1: Brantian Es- 
tate: $ , November 19, 1962, MCT 3305. 

The specimen, one of three or four birds seen, was taken near a grassland 
pond. 

Heteroscelus incanum (Gmelin): Wandering Tattler. — One was feeding 
along a sandy beach and later on rocks on Siamil Island on September 18, 1962. 

Capella megala (Swinhoe): Swinhoe's Snipe. — Specimens, 3: Tiger Es- 
tate: sex?, December 9, 1962; sex?, December 9, 1962. Pintasan Agriculture 
Station: $ , October 17, 1963, ADG 317. 

These three specimens lend support to the theory of Smythies (1960:206) 
that this species is the common snipe of North Borneo. 

Calidris canutus ( Linnaeus ) : Knot. — On August 17, I saw 20 Knots feed- 
ing on the coral sand at Karindingen Island. They were stUl partly in breeding 
feather, showing rusty color here and there on the breast. I saw no Knots on 



Birds From North Borneo 395 

November 24 at Karindingen Island. There is but one prior record for 
Borneo, from the North Natuna Islands (Chasen, 1935:39). 

Erolia ruficollis (Pallas): Red-necked Stint. — Specimen, 1: Karindingen 
Island: $ , November 24, 1962, MCT 3314. 

The species was common on November 24 around Karindingen Island, 
usually in flocks of 5 to 10 birds. 

Glareola pratincola ( Linnaeus ) : Collared Pratincole. — Specimen, 1 : Tiger 
Estate: S > April 28, 1963, ADG 75. 

Chlidonias hybrida (FaWas) : Whiskered Tern. — Specimen,!: Kuala Suma- 
wang: se.\?, September 18, 1962, ADG 280. 

Smythies (1960:217) lists no record for North Borneo. 

Sterna bergii Lichtenstein: Greater Crested Tern. — Specimens, 2: Kuala 
Sumawang: 2 , September 18, 1963, ADG 278; $ , September 18, 1963, 
ADG 279. 

This tern was observed several times off the coast of North Borneo near 
Tawau. 

Treron curvirostra curvirostra (Gmehn): Thick-billed Pigeon. — Specimens, 
9: Cocoa Research Station: $ testis 9x5 mm., 166.7 gm., August 1, 1962, 
MCT 2693; S testis 5x4 nmi., 167.2 gm., August 1, 1962, MCT 2694; 
S testis 13 X 6 mm., 167.8 gm., August 1, 1962, MCT 2695; $ testis 
9x5 mm., 155.5 gm., August 1, 1962, MCT 2700; 9 , 112.9 gm., August 2, 
1962, MCT 2712; $ , 185.8 gm., August 2, 1962, MCT 2713; 2 , 135.4 gm., 
growing new 5th primary, August 25, 1962, MCT 2806; 9 , 132.2 gm., August 
31, 1962, MCT 2842; $ , 112.5 gm., August 31, 1962, MCT 2843. 

This pigeon was the most common bird eating wild figs (Ficus) in the com- 
munal feeding trees, where there were as many as 30 gathered in one tree 
to feed. While resting, individual birds commonly dipped their tails. 

Treron olax olax (Temminck): Little Green Pigeon. — Specimens, 2: Cocoa 
Research Station: $ testis 11 X 5 mm., September 1, 1962, MCT 2844. 
Pintasan Agriculture Station: $ , October 14, 1963, ADG 306. 

The species was seen only once. At the Cocoa Research Station, one bird 
sat on a dead tree and fed on a red berry from a vine. The call resembled the 
crying of a child. Others called in the cocoa. The bird taken had testes of a 
size indicating possible breeding condition. 

Treron vernans purpurea ( Gmelin ) : Pink-necked Green Pigeon. — Speci- 
mens, 10: Tiger Estate: $, November 25, 1962, MCT 3323; 9, November 
25, 1962, MCT 3325. Telipok: 9 , January 31, 1963, TM 22; 9 , January 
31, 1963, TM 20; $ , January 31, 1963, TM 19; 9 , January 31, 1963, TM 
21; $ , February 10, 1963, TM 35; $ , February 2, 1963, TM 25. Mt. Rumas: 
9 , March 6, 1963, ADG 11. Tuaran: 9 , November 29, 1963, SCFC 34. 

The species was confined to the lowlands around Tawau, as at the Tawau 
Airport in the scrub growth. Flocks of 50 to 100 were observed at Tuaran. 
None of the specimens taken in November was in breeding condition. 

Ducula bicolor (Scopoli): Pied Imperial Pigeon. — I observed this pigeon 
on September 18 and 19 at Siamil Island. On the first day two were seen in 
the few remaining trees on the island and on September 19 a flock of 12 flew 
southwest over the island at about 8:30 A. M. 

Streptopelia bitorquata ( Temminck ) : Javanese Turtle Dove. — On Siamil 
Island on September 18 and 19, two were seen at close range feeding with 



396 University of Kansas Publs., Mus. Nat. Hist. 

10 S. chinensis. There is only one other record from Borneo; Pryer took one 
at Sandakan in the 1880's (Everett, 1889:193) and it has not been recorded 
since. Chasen (1935:22) speculated that the Javanese Turtle Dove was in- 
troduced to Borneo as a cage bird. But, Borneo is seemingly well within the 
normal range of the species and probably it is a resident of North Borneo. 
The Javanese Turtle Dove and the Spotted-necked Dove, S. chinensis, closely 
resemble each other; this resemblance may help to account for the lack of 
records of S. bitorquata. 

Streptopelia chinensis (Scopoli): Spotted-necked Dove. — Specimens, 2: 
Tiger Estate: $ , June 19, 1963, ADG 149. Telipok: $ , February 10, 1963, 
TM34. 

This is a common bird of the coconut groves aroimd Tawau and on Siamil 
Island. 

Chalcophaps indica ( Linnaeus ) : Emerald Dove. — Specimen, 1 : Cocoa Re- 
search Station: $ , June 17, 1963, ADG 146. 

Birds, always solitary, of this species often were seen in the cocoa groves at 
the Cocoa Research Station. 

Tanygnathus lucionensis lucionensis ( Lirmaeus ) : Blue-naped Parrot. — 
Specimens, 3: Siamil Island: $ , September 19, 1962, MCT 2928; 9 , Sep- 
tember 19, 1962, MCT 2929; 9 , September 19, 1962, MCT 2930. 

These birds were seen on September 18 and 19. I saw flocks of 10 to 20 
in the remnant of forest on the north side of the island. The birds were almost 
entirely inhabitants of the forest and were rarely seen in the coconut groves. 
I estimated the entire island population to be between 30 and 100 birds. The 
Japanese residents knew nothing of the birds, although they were aware of a 
cockatoo (Cacatua galerita) that had lived at large on the island for several 
years. The Blue-naped Parrot has been found only on the Maratuas and on 
Mantanani Island. Smythies (1960:242) surmised that the Mantanani popula- 
tion was introduced by sailing craft from the Sulu Sea. In the light of the 
present discovery, I think the species is a naturally-established resident of the 
North Bornean islands. 

Psittinus cyanurus cyanurus ( Forester ) : Little Malay Parrot. — Specimens, 
2: Ticcer Estate: 9, October 11, 1962, MCT 2998; $, October 11, 1962, 
MCT '2997. 

Smythies (1963:277) was the first to record this species from North Borneo. 
However, the Harvard Primate Expedition in 1938 took three specimens as 
follows: $ , Sandakan, June 6, 1937, MCZ 197123; $ , Morutai Besar, June 
27, 1937, MCZ 197124; $ , Kalabakan River, July 16, 1937, MCZ 197125. 
The Harvard collection of birds from North Borneo appears to have been over- 
looked, although it was mentioned in passing by Smythies (1960:526). The 
specimens in my collection were taken in the same general area where H. G. 
Deignan took the Harvard specimens. 

Loriculus galgulus ( Lirmaeus ) : Malay Lorikeet. — Specimen, 1 : Cocoa Re- 
search Station: 9 , May 1, 1963, ADG 103. 

This lorikeet was rare at all of our collecting localities. 

Cuculus fugax fugax Horsfield: Malayan Hawk-Cuckoo. — Specimens, 3: 
Cocoa Research Station: $ , 86.0 gm., August 28, 1963, MCT 2825; $ , 
79.0 gm., September 11, 1962, MCT 2899; 9 , September 28, 1962, MCT 2977. 



Birds From North Borneo 397 

This species was first observed on August 28 in primary forest and was 
seen regidarly from then until September 28 in secondary forest, primary forest, 
and in cocoa shade trees. 

Cacomantis sonnerati (Latham): Banded Bay Cuckoo. — Birds that may 
have been of this species were observed on several occasions. E. J, H. Berwick 
(pers. comm. ) claimed he had heard C. sonnerati at the Cocoa Research Sta- 
tion. I have heard many times a call sometimes ascribed to this species but I 
have not actually seen the bird making the sound. If the call note 1 heard is 
actually of this species it is not rare in the Quoin Hill area. 

Cacomantis merulinus threnodes Cabanis and Heine: Plaintive Cuckoo. — 
Specimens, 5: Cocoa Research Station: $ , 25.0 gm., September 8, 1962, 
MCT 2891; $ imm., 27.0 gm., September 8, 1962, MCT 2892; ^ testis 
4X3 mm., November 29, 1962, MCT 3382. Tenom: $ , January 1, 1963, 
MCT 3563. Ulu Balung: $ , July 15, 1963, ADO 199. 

This cuckoo was common in all habitats examined at all of our collecting 
stations, except the moss forest near Tenom. 

Cacomantis variolosus sepulchralis (S. Muller): Fantailed Cuckoo. — Speci- 
mens, 2: Cocoa Research Station: $ , 30.8 gm., August 28, 1962, MCT 2824. 
Ulu Balung: $ , July 10, 1963, ADG 183. 

The specimens were collected in primary forest. There are only five earlier 
records for all of Borneo (Smythies, 1960:253-254). Probably this species 
nests in Borneo; it is unlikely that specimens taken in August and July are 
migrants. 

Chalcites xanthorhynchus xanthorhynchus (Horsfield): Violet Cuckoo. — 
Specimens, 1: Cocoa Research Station: 9 largest ovum 1 mm., September 
26, 1962, MCT 2964. 

This species was seen twice, both times in the research station cocoa plant- 
ings. The specimen taken was from a shade tree, Trema orientalis. 

Chalcites malayanus aheneus Junge: Malaysian Green Cuckoo. — Specimens, 
9: Cocoa Research Station: 2 definite brood patch, 17.5 gm., body molt, 
July 4, 1962, MCT 2587; $ testis 4x4 mm., 19.1 gm., August 8, 1962, MCT 
2736; 5 , 17.5 gm., August 8, 1962, MCT 2737; $ , 17.5 gm., August 8, 1962, 
MCT 2738; $ testis 4x3 nam., 18.2 gm., August 8, 1962, MCT 2739; 9 , 
August 25, 1962, MCT 2809; $ , 21.1 gm., September 11, 1962, MCT 2900; 
$ , October 2, 1962, MCT 2984. Tiger Estate: 2 oviduct enlarged, brood 
patch, November 25, 1962, MCT 3318. 

This cuckoo was common in the cocoa planting at the Cocoa Research 
Station and not found in any other type of habitat. Smythies (1960:255) 
thought that possibly two species of Chalcites were represented in the series of 
Chalcites malayanus from Borneo. I have assembled all 20 known specimens, 
however, including seven in the Museum of Comparative Zoology at Harvard 
unreported by Smythies (1957:638) and find that the variation in the colora- 
tion of the head and upperparts is due to the difi^erence in sexes, the males 
being darker than the females. There is much variation in the length of the 
wing, but the meaning of this variation is not yet clear. 

Surniculus luguhris barussarum ( Oberholser ) : Drongo-cuckoo. — Specimens, 
2: Cocoa Research Station: 2 imm., July 28, 1962, MCT 2672, discarded; 
9 imm., 28.6 gm., August 25, 1962, MCT 2810. 

The first specimen was taken in secondary forest; it had been sitting in a 



398 University of Kansas Publs., Mus. Nat. Hist. 

dead tree, occasionally darting out after insects. The second specimen was 
taken in cocoa; its stomach contained 50 caterpillars. 

Eudynamys scolopacea ( Linnaeus ) : Koel. — This species was observed at 
Tawau and on Siamil Island, on August 30 and September 18, respectively. 

Clamator coromandus ( Linnaeus ) : Red-vraiged Crested Cuckoo. — Speci- 
men, 1: Telipok: 5 , February 10, 1963, TM 33. 

Phaenicophaeus chlorophaeus fuscigularis ( Baker ) : RaflBes Malcoha. — 
Specimen, 1: Cocoa Research Station: S, August 26, 1962, MCT 2813. 

Flocks of three and four were seen in the cocoa. At Kalabakan the species 
was feeding about 40 feet up in the second canopy layer of the primary forest. 

Phaenicophaeus diardi borneensis (Salvadori): Lesser Green-billed Mal- 
coha. — Specimens, 3: Cocoa Research Station: $, 58.2 gm., July 22, 1962, 
MCT 2636; 9 , 55.8 gm., September 8, 1962, MCT 2890; $ , September 13, 

1962, MCT 2918. 

This malcoha was seen only three times; it was the second most common 
malcoha. 

Phaenicophaeus javanicus pallidus ( Robinson and Kloss ) : Red-billed Mal- 
coha. — Specimens, 2: Cocoa Research Station: 9 , 97.0 gm., August 31, 1962, 
MCT 2841; s , 98.0 gm., September 8, 1962, MCT 2889. 

The two specimens were taken in cocoa. On October 2, 1962, I saw one 
about 100 feet up in the top canopy layer of the primary forest at the Cocoa 
Research Station and, on October 3, two more hopping from branch to branch 
about 150 feet up in a tree. 

Phaenicophaeus curvirostris borneensis (Blasius and Nehrkom): Chestnut- 
breasted Malcoha. — Specimens, 7: Cocoa Research Station: 9 old brood 
patch, 121.8 gm., July 6, 1962, MCT 2602; 9 , September 4, 1962, MCT 2864; 
$ testis 3X2 mm., 143.6 gm., wing molt, July 7, 1962, MCT 2611; $ testis 
6X2 mm., 111.0 gm., August 11, 1962, MCT 2763; 9 , May 25, 1963, ADG 
110. Ulu Balung: sex?, July 24, 1963, ADG 216. Tiger Estate: $ , June 22, 

1963, ADG 156. 

This was the most numerous of the malcohas at the Cocoa Research Station. 
It was observed in primary forest, secondary forest, citrus trees, and cocoa trees. 
In the primary forest it ranged in the upper canopy from 100 to 150 feet up. 

As Peters has indicated (1940:56), the name P. c. borneensis (Blasius and 
Nehrkom) 1881 has priority over P. c. microrhinus Berlepsch 1895 (used by 
Smythies ) . 

Centropus sinensis (Stephens): Common Coucal. — Specimen, 1: Cocoa 
Research Station: 9 , May 15, 1963, ADG 108. 

The finding of a coucal at the Cocoa Research Station on May 15, 1963, 
came as something of a surprise, since none had been seen there earlier by our 
group. Coucals were seen at Tawau but were not collected or identified to 
species. 

Centropus bengalensis (Gmelin): Lesser Coucal. — Specimens, 2: Tuaran: 
9 , April 1, 1963, ADG 46; sex?, December 3, 1963, SCFC 17. 

Carpococctfx radiceus radiceus (Temminck): Ground Cuckoo. — Specimen, 
1: 12 mi. N Kalabakan: $ imm., November 7, 1962, MCT 3217. 

This male was taken in a native snare in primary forest and provides our 
only record. The specimen is in the postjuvenal (first prebasic) molt. 



Birds From North Borneo 399 

This record is the first for this species from North Borneo (Smythies, 1957: 
643 ) ; others are known from Sarawak and Indonesian Borneo. 

Otus hakkamoena lemurum Deignan. Collared Scops Owl. — Specimens, 3: 
Tiger Estate: 9 , November 25, 1962, MCT 3319. Tenom: $ testis 6x5 
mm., body molt, January 1, 1963, MCT 3552. Agricultural Oil Pabn Station: 
$ , October 6, 1963, ADO 299. 

The specimen from Tenom was taken in a bird net set in a grass-scrubland 
situation; the testes were regressing. Harrison (Smythies, 1957:645) found 
this species breeding in tlie Kelabit uplands in January. 

Glaucidium brodiei bomeense Sharpe: Pygmy Owlet. — Specimen, 1: Ulu 
Balung: $ , July 19, 1963, ADC 210. 

The specimen taken by Garcia is the sixth known (Smythies, 1957:646) 
from Borneo and the first from the east coast. Specimens were collected in 
1956 in North Borneo by the Cambridge Expedition. 

Ninox scutulata borneensis (Bonaparte): Hawk-owl. — Specimen,!: Tenom: 
9 largest ovum 2 mm., oviduct evident, January 6, 1963, MCT 3583. 

This female, taken in a bird net in a grass-scrubland association, was coming 
into breeding condition. 

Strix leptogrammica Temminck: Malaysian Wood Owl. — Specimens, 2: 
Cocoa Research Station: sex?, July 9, 1963, ADC 182; $, May 25, 1963, ADG 
111. 

On September 7, 1962, I flushed two