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 0 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. CO s =i "o 4 bC i CD "o "o 1—4 o "o o O no" c3 O O O O •N •« Ul c3 Si^^ ^ ^ ^ ^ ^3 -o p 4.3 KH -o T3 -o rs tn C c ^'^ u *^ P„ o o o o ^ a a 3 "cl a a a a a a a "m • > o a a a a i -u «4-l o o 1 OS CO ^3 OS TJ TJ -a I CO -3 o CO 1— ( a; Tfl 0) ICI o3 a u ^ > >> a)pH o3 03 tn C 03 « 02 03 i 1 >> * OS OQ QQ "oj "tn bJ3 C a ■A 0? §03- D, Ph Ph 3 03 S 03 j CQ QQ OQ oo m ai *3 ., a? bb 1^ H o o c o3 03 03 a o 7-1 - - - - - m CO 00 CO '^ s c CO as CO 00 ^ g CO .*» .2 s S s <» 2 ^ *(S> oo ;s '-3 s s S i 8 S o <3 S ".S a "•+3 55, CO s o s s § i CO CO 00 00 (—1 h s~ h O •(S» ?i. a, O !5, ?i. s S S "co o o o .'^ O O 1 S ^ 1 3 4J o +3 ■+3 o s^ +3 ^ < 1 i 1 CO .4.3 +3 o < O < 8 ■Si O -Q Xi < 1 o O 01 G, O o oo 3 '?^ !-. c o oo +3 a o CO s a) a, 1 CO ! CO s s s S s i s -1 o a. o s m ^ ^ CO S 00 CO 00 "co •2 § o o i~ O •r«i §• S O a. ^ ^ 00 ■? 5~ ■^ p s s s "a s o ^ s .o ^ ^ s a. ^ ?s CO "^^ ^ <» ^ ^ Fishes of Kansas River System 93 01 (U (U c 6 •4^ -4^ •4^ S^.^ rt ^ rt ^P= E s E ^fi a o a a p a a o a i"i kl aj &-( CD u *H -4-J t« ^ >> 3 CO jaCQ 3 to e3 ki c3 U c3 t-l "^ t- V. Xi 0) JO 0) ^ a jD CU M O > O > O > 0 > G tH Ui 1^ Lri .^ •■-■ f^rt ;^P5 £rt £tf a >> ^ 3 3 "ro 'm in m O lO O /-^>^ «' 0! CO CL, 00 Ch CO « lO 05 c 05 1 (M 00 ^ S o 1 -C s -OiC cog t>. t^ -d 00 00 OJ T— ( «J 0) > 2 t-, & 03 i >>* > >> > >> o s > .2 'a > a hf) 02 1 a; -4^ 1 •4^ s 1 m MM :^0 S ^1 ^ "* "• "* ** •^ s ., o ^ X 2 ^ e s "-0 •2 ■'^ •r» "cj C3 !"§ J3 CO Q5 1-^- CO 1 t ^^2 CO t3 e -« CO >> 1 00 00 •2| »- 1 i •g 1 "3 1 s 0 ^ § s § -* 6 ^ f-^ s a 0) a; >> e H-l CO CO V i e z3 o o o c a O CO S e O O 00 H e CO a o O CO s fa. o i C4 0 CO «3 s ■Jo .00 p. CO 1 S CO > 5 s G5 s s "—J 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 ^ II ^ .. CO uT eg ^^ ^ s o z z w U b O^ « 2 < S O fa Z < J H < K ■ oQ P (« b ^ SOS ^ -J zS CO S en " * , bC •* ■* ■* CO c C T— t »-H 1— ( I— I d. 1 i tJ. Q (M (N CO 1— 1 »-H 1—4 I— < ?-H 5 bC C 05 00 05 1-H <— 1 t3 ^ e3 -3 ^ L-"^ o CO 00 s t^ 00 00 «o -3 * Tf< c^ ■"— ' CO c^ ^ V (h D <: t- ^ 0) a a > a h > 0 SS s o: 0 5" S^ c ^5? ^ -1^ 3 o £ c O 3 s 0 o u a , g^tC « i 'or fiOi 'gd bO c 31M 0 ^ i. cj _ a, 3 £ c ►i-Ol S 0 0 'S 3 O tZ) CO 1 1-=- (N CO (N (N (M (N (N O io CO ■^ 00 ▼— 1 in (M »-H ■* CO CD in CO (N •* (M (N (N (N (N (M (M (N t^ C^ C^ s CO CD CO T}H in lO CD CO "■J* in 10 5 -3 o 1 1 (N 1-H .J. 1-H ci 1-H 1 1-H 1-H 1 CO 1— < fO 1-H 1-H 1-H 1-H 00 1-H CO »-H o CO ?; CO CD f^ CO ^ 1-H *-H lO 1-H in 1-H CO 1-H 1-H 1-H c ^H t^ in CO - S" 1-H 00 OJ t^ o o t>i 00 05 J (N (M Ol c^ CO (N C<1 c< ■e, "O 1 1 1 1 1 i 1 Sf's "^ i OC t~. tr tr 00 t^ Widi of orbi C I 1 1 CO 1 ^ 1 1 1-H cc CC t^ CD CO CO t^ t^ •' *■>— ■^ ^«— . ■^ v^. N-^ cc b- l> C C5 »— ^ in CC cc t^ l> CO t^ CO t^ C 0 z 0 1-H H ^ "2 ^_x CO rM Number and Number of Specim c i o: ts; c 2 a o ^_ ^ a s > p: N 1-H (-C > ID "3- u 0) > C o t-. O i m ro 1 T— ( S3 e2 (N 0 in 1-H CS3 be > C c3 > a ^ j> .^ N &H •« p m > ; p^s > -4- ^ i ^ S2 ^ c 3 O c 'SE C 3 3x_ ' g^ : s § ^ =s^ c o Oo- &c ^^ c i 3'? c ^ ' ^^ <1)(M CCC > Ch 0 bX) o; a c- 3c .22 •-3 O o o: r cT t OcC 1 ^N : is .2 H-i -is: ^ a T S^ oT = J ! 35 ' CO ^ > 'o 3 4^ X ! J2^ C ^ " .S C a ^ o ■^ HH W 1 122 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 University of Kansas Publs., Mus. Nat. Hist. o u 2 to 5. o o q C/5 Z o o V @ o g M go S w » H h O z o i CQ K H O z u p o u cq (N n 9) CO CO IM CO 05 CO co c CO C<1 o b- 6 SS2 gS:2§§§22 § 02 8 c a PQ CO CO CO CO g n I— ( 00 (M N f— i CO CO r^ lO rt CO ?-H o 6 O o o m 6 !^ w . pi V 03 C o § d O (U tji CO o 6 o (O -3 V s s o d = IS d^ oc O 3 O d « O d CO as O d > c a; hH d OC cc I. % Q d O c • 1 — . t- ci a: *; 6 O 1 1— V 1— ' o tc a i d g c 'c o p cc Q d o < ' i ir. •-, <<- d O 3 S PQ oc fe : CO : o • .-2 '■ is -a; - ^^ 12; Fishes of Kansas Riveb System 127 IC CO N «o IC OC w eo o o CO >* «-H r-H IC CC b- b- b- h- b- CD t- 00 b- 00 2P t^ M (N (N cs Cl (N (N M (N w C^ C ;■ eS « t> • a ^ "^^ "* Of •^ ,^- J3 03 a ■« c • 1— ; PS ! 1 •<* : 6\ ^ ^ ) P: 1 ""^ •^ '^ ' c ; ^ < 03 I « i r « o ; o p: i |- ' ^ t— 1 «3 2. < o: : P: o t. n (— c ) -1- ) p: : t c . (4-1 O .-2 H d OS -4^ b£ 03 e i .1 ^ P5 c a b C c c > N ) c ! ^ 1 -1- 5 n ij 1 -s > ; ^ i 1 r c ) I. I 1 ; p: 1 p: r ^■ ) c ; C : a s J iver Drainage (east C ) 1 ! C 0 I c c r c ) ^ 1 c P a ) it > K * c i c ; & J c bawnee Co., Shung; U 1955) 1. ^ 1. : s. .1 b 2 -C : X. 3 _c 5 oC ^ 03 a ) *- > V- : ^-S V c ) < > a ) c > < J c > o ^ 3 a 3 0 \ >' 1 go ^ i 12 5 ^ ; ^ ^ ^ ; ;2 ^ L ) U '■ C^ ; ;i i \i. 1 U \ "A 03 03 P^ w 128 University of Kansas Publs., Mus. Nat. Hist. S (N n < c ■ i> b- b- oc b- or b- oc b- OS § iM (M 00 IN or t^ cc oc IN o- CO 10 ^ lO ;^ (N 1 t^ CC c: rt Tt 't cc c^ ■t -t ^H 3 (N ^ V O iC Tf Tf ,— (N ^ r- cs CS • >. (M -§ « CO Tj- IN ■* 1— ^" ■ 1— CC • '^ f— t (N • IM CO (N (N IM 1-H (N ^ ^„^ OC t— 1 c< t^ c^ 10 6m ."i" || o £ ID o c3 CO (N oc ^ 3^ OC Pi 5 a 3 oc 0. 2 y / — b- CO c b- ,1 i cc 0 AND L .in Pa -1-3 a> 03 u 6 0 c c > -1^ r 0 a 0 ^ c -1-= -»^ 0 p: a Dbainage (Museum No 00 03 .2 c c c C c 0 bDir. coc Is 0 ^ is Sec fL|OC ^^ 0 s a a Ph c % - C c a, - -Q 0 0 .s 0; c ~ d 0 c 0 1 0 cj EC CK cc m cn a, X a • • '-' IM ec -< CO lO lO M O IM lO O CI IC M •* • »o Tj. CO Tf* rt COO5C00QC5 (N rH IC IN -H -H o ^H lOioeC'— iio cj t^ (M (N (N IN -H IN .-1 —1 IN ><}< n IN CO (N >0 I-H IN ^ IN IN Tt •* IC -^ IC lO CO !>• LC i c C a c p: c a c. O ! ^•c ; SI s ) • -gp: O c M «■ > t Si m C " 1 O -►J o a o a ) cd c s ^^ 1 ^"s ! « -3 'S, o O . CO OS "^ ^ CO 00 OS CO oc g OS ■«j5 rji -<}* CO CO «d § CO CO CO CO CO CO CO CO 6 o o o CO to 00 CO 00 o ^ CO CO CO T-H CO CO ■* eo w CO c^ 1-H 1-H CO ^^ ^H CO 1— 1 CO o iC « ■^ 61 eo 1^ oo § Lt iC rt< 1-H V u C 00 K •* CO 1 IM t* 05 ^H CD S (N 'o 1 CO CO Pq -H C5 e^ — C5 3 T3 O •».» u « C<1 — l^ cc — 1 CO M — K cc a u CO a CQ < CO CO H CO 1-H eo ^■ a ^ ■«* a 3 a • I:? ii ID s a ' ? (5 !■ c : ^ ; c c west): r Boggy R. < : c : p= c . p : tf 1 6S > IS < ) ) ) ) < < RAINAGE AND L seum No. in Pa c Q Km a < If i C ! s ' -^ < I- ;^ < ) C 5 c ■ainage (east to itotoc Co., Cleai a nston Co., Blue rray Co., Washi 9998 99fi4 .ti.^4 i IS r Op: 1 %z kham Co., Tim . Red R. Dr.) ( > ■ S 1 Pl \ I 1 ..^ 1 5)^ 1 s<-* 1 PQ^ 1 1 c 1 c ; % I . i .fS'^ a "c 3 "c Ph-« ^ J s ^ ^ M L ) C \ :z ; ^o c ) o o o o ki 4^ < Ph Fishes of Kansas River System 131 m CD 00 r^ 1 lO CO CO ec (N C^l (N (M O 00 ■* 05 (N i-H — (N — > : ^ 13 a 1 O : a • ^^ re H 1 o '. -u s •^ CO U^ 03 ..2 o - ■3 O ©2 River Drainage: Hays Co., Blanc TT f;QQR^ : *3^ ico, Coahuila, San imenez (KU 3006, r e 3 O ShS i a.^u -a O 2 -S g- o o O s 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 171 C/3 < fa O o I— I < 0) C/3 o .S 8 in iH tf.S <2 O Wt3 fc — 6 Si! - ^ s.j . ^ 8-» " =^3 <;^^ =^i 10 o A, To ^ o - -s 0> en \ i »'"' :^ o ^7!i fP^I ^^ •. k . h^ V ^ \^ktl3i If) * « a. to Z3 i ) Q.- ' 1 o o c o ■a o z| SI*' i •q. ^^ »r X ^ ^)M,|f/ni (P \\\ /?1 I 1 / "IK^ \\\\ //' If/ ~ - ^ ^/// 4\ wk\ ' s. k _ r^f/z -Ja^pm " = 1 o u t w ■ L - (. a — - ~~ t. b J, 4m °^ ■' 10 « to 5^ x\ ^^s ?:>;;7L,\ ■ ffi i ^^m ;^ o ^ o . Q. Y^x*^ fl ^M^ #^P ( - . ^s T3 CD Q. 1 ^r- fffmi ^ fl> 3 ^cl - 'l\ 1/ ( Q." " O) i "^t ' /-^ I f 3 O c ^M^ # 1 r O o £ 1 T a. o fffi) • o Q i i s ■(>^ 1 -'- i ■ f // Wf ■ 0 ^.S\ w -1 Q^*^"^ ^ ^WiW ■J 1 o o ^^\^ I j_ !j -I' 172 University of Kansas Publs., Mus. Nat, Hist. ' ^ ^ m I n i ^ LD ^f HiA^l^' 3 /y / V^ r^ L Y^ 4) CD - in w \ / f/1 Q. LT) & ^^\ -. o z. W\\ ^r A s ^ J3 ^^ vT* vs. V\ 1 >s ^v \V, ■ /f //[*: X ,<^ ^M 1 Ss wwr nwi - -^ s r - o4 % 1 /^ ^3h f »1 . feJ.;^ % OJ 1 isaL/^i, Tf h ^'i'/^s' 7^ \ 'co < ):^iV (| \ — - Q_ r o O SI ilu^ ^' ^ sS >, ^i i^vW / z 'S X pffl t [at i ^ - <>ViSK¥ 8-8 c3^ _ ^m ^ t: \^^\'^aV) ''^ CO h-^ -Q O tSw '"nl 1* O c O) I /'"'/p/x ' V v\\v\ w r /TT Ik jr 1 Q- -^ i (IN. 0-,5 ^ S-s - "3i^^'' / °-s ~/-7 '?=cs:;vr _£Z jaJ?R.^ a CO to 3 iMi\\\f'1(* L^ E ^M y \i o r^^hvW ii/ CO V^A^JVM'W O f^^Xx {^ i/y Wl -C ^•V/'K^ Y\V'/^ Ijl . O o ^ ? ■ Fishes of Kansas Ri\ter System 173 ■ /^^ Sis - - ^^m f ^Tf ^ ^\ (iIt ^ ■ — Vi\ r u/7w55WVt? J? 2 nAw iv -Q \ ^tj U^'Rs^ 3^ - E Z3 < s /<*i ii;f^j \ Q- cn 5 AS^K'U -H o CL £/ l^M^^^'f ,»> o h ^ \l u / ^^ ^V\V\ i X 5 o ^OT ^ \ Vv, m IC 2 '!^ x\iM 1 . 4 Jk luV-^v / - n /''' I ^^tvU^ 1 s %4W o , s- . j^ifj^%g »l3 3 8-S (? ?r^ ^x^iC^^v^or^ y o {YfitS^ ^?/A^ ?? ^i: i O) ft]^ ii7=^;^*X £ CL l{27C*3/6 ^ - O «*^ mbr/-l CM. cn O piwf 1 o « ^ ^U f^P// " 1 . \{d p ft u —k ' fis <3 OT 8-s ~ W tn V. ^ \U)Ni*^ f^ ^ 3 N WiHft ^j/y L>y O) J^-^ W = ^n^^ ' --y ^ )/y^ (fri Cl" in "1* &.\\' 1 \ ) / / o Q- ^\t\ ii 'Al O ^ u \ \ \ V ^- \ My 3 o \\ \\AA 2 z ^Ai^ i 1 i ^^^ ^^^ w ! b b k at I " - s ^t j^^T^A^ftJ* "• ^ s=f^^^^^^ tn ftr fr /oyvJ "^^rff jO- :Sg^ 9 tTaQ^^JcI!"^ 3v ^'J^^ O) ( ( WCSi^ IJ^/ en V i ifvr/JtfwW^ ^^7^^ TD Y, vM^n ^V)l Z5 vAl nWluWc? /'/ ) v// •o ^ vW imriMh L-f/ ;3 X nW^ ]M5l3J^f/ H< fy'r- "teyfiV) U c \Af' \ y ^B^ g % . 0) 5^ i 1^ mv^ M ^-S 01 - O o -y^i^riT^ — ■" o . '^ - CO S ^/i /I) r J z' Al ^ Q. o s/ SMuKif 3^ O ^fO^ \\o\i] f^/ 1 2 Z fmi f 3 c\w ^ 3 174 University of Kansas Publs., Mus. Nat. Hist. ^ <^ '1? - -^■v * i ^ '" J tTA #il^-^ ^^^ 3 P yy,'^ I ^^2 ^^^' o c ^^\1^ *^i "^ 1 vi %i '^ 'r a] ^ " o 1 'S t\\ \ ^Jt/f 8" ^1 ^ \\\ ^i( ■" 2 D. ^-n Va \s (A § CD i/ '5 E ^ ^<\\\v\ Q- '/il f^\ vm - r wM o i^ii f u i b 1 ' \ s-i - ^!!!^r^V Ol v v)^-/ 'Ji- C i - g lO s „ D siy 1^ -im' |J CM. I/) i«K 1' H/ A) ^ g ^V\ v\ 1 / ill cn '^^J^.'W \\\ Id/ O J3 ^^■l^ miTj it >< w^ J) s . J- v/wf •'> f 1 Vf"A m,4 l^^w ^ t b 1 ^ i' i^rt^ s-s - ^ r^w] '- ■~^\ o-^S ^ ik i^^ cn p iV. „ D. N^? -^7 ■i^ i CO >N K \^ ^ ' / 5\> Cl" Ol § /'^-'w / T 1 (/) * j!^Sn \ ■- ( r "^ =J ^ 1^3 \\ \ < l I ^ "^^ J3 ^^k 111 \ , ' O j^^^n V \\\ / >*^-^^ r\ V\\ 4 7 1^ JJ ^^^OT t{ jj %. . (jj^ i\Ss uf s: " 1 f ' b ' iN ^ - ^%^^jM\ F / ^^^ X ^i "rf^ v3j(rj3vf>'rfe ^ O V r/*VA=^ ^i^V^ ?H s- jM'Jm^jJ IS- H o J^' ' A^^i^vr ^ e c o V ^tll^Vf/i i CO i (y3 i^^ 1 r/Vv "^ CO - D F i\il M Q. i> o o S/: s 2 01 O Im" 3 Q. / V>IH J i- ^^ ^1 {«vS M 71 o < y 'L ^>> yAWi. JJ . O l-^/ m ^ ■ /!( ^*\^ ^ ? f^ii T' f> rO . O) ^/)r / A\ (^ o & dMK V ] Y D E ^/ iWr/ ' a ^ o ^ rnvvW 7 J s ld /^ ^s\\s\ ■ (1/ n J it O /, ^^vM'f/ frl x; (^^ yf J 1 t<>!;N>\> . L O '^- Jj . S .% A Jv; IVly-^* 1 1 £Ajxj \ 1^^^ t Fishes of Kansas River System 175 b » A. yO/fT* bj V • ^"VC^/L \ •• ■'^I—^Jj^ri ^^jS r^;^:^^^ :sv in i ^£(a ^ 3 \ 1 S ttM^ _^/ a M.\ \ W Sj/ o \ J v^ fy 3 \^ 1 ^ 0> - CL s V ^i^tv^ f> to l/^ yk/fti u /I '^ 3 /Alvir-ii 13 5^ ^;v s O ^^ \\L r ^ o ^ tw s \IfO r 9 iwV M^^ fit Nlp^ o 1 1 tf IjO 5T^ '4' s o 2 i i 1 KANSOS Map 39 1 £ i - ? w //A! '\^^ m m ) o // ^ 1 ^ m s 3 O % h ■ !> 1 (K J^ H - ,<5i?ft a-> " ^M ^\ t - £ f - '1^ i^^^^ a "• R|^ D N(\ ^ i; - ^ ii M "l o /''-y[/ V" A cl -jr^ o 3 !^i B \\\\ ^7 a ^ cn ^ ^A iR \\ // c J^^ /"ilA \\. W - / § < 1 1" 9 O o 6; mWP^ o b t 1 b b 1 /V 0) J D F ij 8-° " r CO u> U9 5 W^ s KANSAS Map 3 1 V &< o Mi 01 ffl o ■I s 1 B KANSAS Map 4 *1 'y^7sy^^,'i§'i* k Q- ^/mff/ VT vi^^m ill K - '-iff) . JL/^ V Y ^ nr - o O - 'til o o %^« b b V * ' 4s^ f s-s " \ g 1 CO n}\ 1) jMq^Tj }> D \ ) %^ "^A >s' ^ O Vj %%v^\^^ ; = \ 1^ - C ?< f^'%U/ H ) K) J to k^A A V 1 / i o. 3 s i a b ^k in\\^' .2 D ^g tZA l\ \l 1 f _o ^ iw i r 'i'^ i 1 1 - 176 University of Kansas Publs., Mus, Nat. Hist. b 1 ^ i3 f « S-S " t ^ ^. 9> o S )fj^j\ /\\ I" /' h CL £/^vi\ U "'" // l> U ':3 2^^ \ ft \ \ J 1 / ^ ^2 o ,^^M V\ \ ^ 'A i o 2 o cr ri^^WII'TV r 1 - V^A^^ ) O \t "mJ^W §^#^(\1W s o - 7 *"■ S < a- 5 i^ (( lif ^^ O) \> i>V ) o - O o p IMK ki o .^ I \ . 1 ^P, \\\\ A QJ l^iAhVWli^ /7 3 D- i f o ^ '-' 1 i ! ■j^ o 1 _, / 'n^ ' fc *^ J "" S,:^f ° ■ - s xS^S^Sir'i -=^'f^(^^?5^^^ (; V. to ^Iwl^^^l/ £ r -c: ^^?^M ^ . en E o Cl a) s- ^^y s "%^ b 5> ^5=^ yJ^SM g^ f vri ^^-V' ^ o \w yiK^^ ^ m yj O \ iO L cu ^ , ^ 2S Mt I'lf ( CL J a i- y 'a <( o ^^ ^v\v\^ 1 Cl /^ ^^K\\\\ X / %A^vM r^ I _J /I-- '- 1 '-^ / ON 9 %4l HM ft w - . ; ■-■ ' 6 1 J|, '-?- <^^ =^5 0) f ^^^ at r\f>m^^^ J3 ^ O \<*K*J )^^ s'^ o 1 CL - cn 2\^ %v'^4\ '^- m O ••' r-^y / ■ V <^A O 1= o ^yfeflUT/M S ^ cn ^si" O LlI ^ \4C0—. . . j[*y 5yvi:^ f^l O J, 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. 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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 0 to 22 per cent (aver- age of 5 per cent); mortalities in 1961-64 varied from 0 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. •-^ O c ^-s « bC 03 03 a « c ;-i ■1.3 bl ■3 >- • (.^ S. 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 > < 0 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 0 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 0 % PER CENT PROTEIN IN FEED 3 0 % 4 0 % 5 0 % ■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 0 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 0 213 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 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 0 694 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 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. u 13 1 > (MrH 05 10 t^O cot^ CO— 1 t^iO occo 00 u too TflO >Ct^ coco 00 lO OOO oco (N ca o O ■*iO (Nrt (NOS CO— 1 coo 1-H lOOO CO"* 'i^ h2 1— > fl . t^lM 00 1^ t^l^ (NO OQO IN 00 co- r^ o t^O CCiO (NO Tfo 00 CO coco (N G Catfish Ponds in s^ OO — IIM (NO (N Tj< IM^ CO(N CO 1-H r-H T— 1 lO (N (N T) (U en CO 00 00 00 COO CD>0 (NiO ^O ot- CO O COCO — OS 00 lO lOCD O^ 0
i ■*co d^' Tt o O <:D(N ^05 cr.— ( O^iO OO ■*(N oo o CO(N IC 1—1 1—) 1-H ■* (N 1— 1 T— ) OO lod (N CO^ (NCO (N ^r^ +s b C ^«4 fe « a . ^00 ■tf 1> cot^ ^00 0(N COCO Tt< 00 (N t o z w »— t 1—1 OCO r^ 4.1 (~ COCO 1— ( 1—1 lOO .-HO I>1> coo COt^ CO O fflPn (N(N coco Tt05 coo dio lO o ^H 1-H 1—1 cc -^f* 00 ^,^, > o OiiO io«o coco OO Ttico coo coco o <;U t>t^ , (NC 0(N 1—1 Pi Wco (TXT! lOO i>»o coco ooo ^00 I>iO CO l^o do .-KM ^(N eo(N C0(N (N— 1 (N(N kC ^s \N — 1 1-H (N .3 w z i-*\ 5 U2 M cc 00 OO t^o OO t^O CO t^i> o>a> t^O CO(N COTJH 1>.I> (N (N lO .g 9 o T-t 1—1 1—1 1—1 (NCO CO-* lO iC —1 1-H t^OO IC —1 —1 (N(N »o b WU 2 ^z. -t — wZ > o co-t-- lOO <3iOO Tto u; ^g t>co di-i OiOO —id t^CO dd M^ o (N •* (NIN 1-H 1-H (N 13 ^ U ^ IS q; Co ni 0=2 CC':** Oi(N (N^ (N —1 oco 0(» 00 Tt* (N o 3 do 1 1—1 1— ( 00 lO O— i (N(N coco coco (N— i .-n'd .-H 1-H (N (N (N es 3 o «4H tf H O 5 W C^ -73 . CD m ft,'-' coco Oico coco COO coco cot^ ■^ t^ ^ §1 O^ ug 00 00 dd — d coco 1—1 1—1 00 t^ —1 —1 (N(N 1-H 05 dio (N-«*< Tj'CO rH rH 1> (N Id 7^ a Or-I O— 1 O— 1 O— 1 O— 1 o^ O— 1 "•9 ^ 1— ( 1—1 1-H 1—1 1—1 r— 1 .-H 1-H 1-H 1-H .-H .-H — ( — ( §9 1"^ cSfDS 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 0 66 0 70 0 81 1103 1245 1314 1515 1964 2000 1 2000 1 2000 32 32 32 0 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 0 074 0 076 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. o ^? ■i>rtir>coooiMi-i-^ ;^ ■1 • rti CD TJH lO CC CO 1-i (N • « 5 m •cofoeococccccoco • c : ■— 1 -o Oj . OOt^COCOCCOOOJOO • a 5 s OJ r-l 1^ O5Oif0t^00"*iOiOO3 •C£ 5 CD C^l "* o C^JrJHCDGO^^'— llOt>. • o 2 O 05 !^ T-H I— t »— ( 1-H H C^ >— 1 ^1 eoiot^o-^ooQOioos • c 3 r-lOO'-Hi— Ir- 1>— IC: ) d ^'P •-lOOiOCO-^i-HOSCCO eoo6cci0505'^05 05d ^ eocotccco^coiroirc T3 Tota wt. O-^CMCC—ICOIM-*^ CD O '^ r^ lOrfiOOiiOfNOOJCO M^ I-H lO 3 0, CC>— iCCiMCC-^iO-^C^l lO 00 05 I-H it IMCDOiOOOOlTtiOicO CCOCDl>CCO'>*Tj*(M ^ ^ ^ ^ rt C^ (^j' C^ jvj i? — -^ooirsooc^iocoo C0G0C0OO03IM— i-H (N COCClOOTtiTtiCOTtdO to I-H C^ (M ^ <1 ^ OC^ IC OOOiOOiOCO • O lO CO T— O rt iC to I> • _ ^' rt e^ (N (N (N c^ CC| ■ ^u. ooooooooo • -(— o 03 s=y OOOLQOiOOiQO • o iC tH o iOiOiO-rtTM?0C0 • -OCMTt* (l g c^'d fe feco o3 2 c: < iC 05 -(J -w C C (D --OJ 2 2 g g^ S-i m a s ■> '•+3 3 1^ hS ►S ^ ■^ Q> OJ o CO o (1h en HI e .s ' ^ ■>«' Tf T) -0 C a a n . >i: ^ Tl r > C3 O B s ■D o W z . fr! > a 4-* .c O V) (U w u. w o iH ■* w" nl •a a a in CO -a a lO a T) 13 M O (1, 13 n a o 4-* (^ g fi o o u * V n-a 01 rf > « i a kl 01 V h t- ^ «M O s ^ Pi 3 ,a rt ■d ca o o 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. ■^ ,'-.^ 30 A \.^ ./'■■ — •■^^^^>^. ^_^ .♦'' *• *■ * -25 - **-V^ < ""--.^ (T \ O \ 1^ 20 _ \ \ z LLl \ o '\, o lU tr 3 1- < 30 ■ ,"'' - Ll) Q- *"---— — ,_-- 5 ^ 25 • . . ■ " " "^^ .^'-^ ■•-....^ ^**H, B ^,...^ ^' 1- ' ~ — . ^"■■■■■•■. ". '•"'^^ ^~-^~~-.'-!^,^^ 20 - " *— 1 — .,. .. . 1 -■ 1 1 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. o > August 27- September 9 . 1— ( 00 00 CO m CO o o o «> o ■<1< 00 00 00 CO CO -* o o o 00 t> CO t- t>. 00 CO lO CO o o o 'T3 w a> (N S 00 t" 00 "W o< N CJ CO —I O) TJH ^^ o l^ o 1— t 1—1 tH CO ■* t^ '* tN. 1> t^ l> 00 Oi o ■>*< o o 1—1 »H lO CO 1> lO t^ t^ 1> t>. ■* Oi 00 o o o o 1—1 Tt* CO IC CO t^ i> l^ t^ CO o Oi o o I^H o 1—1 'W lO ■* lO CO <1> c^ 00 o t* t>^ f-o a 8 CO lO 00 CO ?; o o o o ~2 S, 3 1— t t^ t^ t^ 00 t^ t^ t* l^ t^ 00 t^ ■* CO o 05 o o I— ( o Oi o CO CO t^ 00 t^ l> CD CD b£ bC •*^ -4^ c c 1^ "S 3 T3 3 V O bC bC bO c c a «a «3 «3 »o »o lO r^ CM (N C) (N CO 1-1 05 1"^ 1-H CO 1 CM O >c lO lO C5 00 1-H o 1-H CO 1 CO 7 o o Th CO »-l 1-H r-i o 1— I lO CM 1-H 1 1 1 1 -« IC CO Oi t^ !U r^ o> 1—1 1-H >4) »— ( 1-H t*- C T 1-H 1 1 CM S a. R. 00 T-H lO O » 1—1 o '^ •f»a 1-H CO CO CO -* 1—^ 1—1 1-1 CO 1-H O ? o T - T—i »» CO o (N c< o OS 1-H 1— ( 1-H cn tn bO bO ■u -tj a c "C 'u 3 3 a> b O CO O CO CO 05 M H CO X 9 u P4 August 27- Septembcr 9 DIM bC 1 SCO <1^ July 30- August 12 July 16-29 >=2 3 1 June 18- July 1 H^CO Q 2 O CO 00 (N (M (N -^ ^ ^ ^ o o to CM CM O 1—* CM CM CO CO no CO O 1-H CM -* CD CM CO C^l Tt< •* ^ CM CM CM CM CM CM CO CM CD 05 CO iC CM 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 0 013 .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. X CO en X CD lO CO (N O 00 HKi CO uO ^ CO (N (MOOcooooa)C^ico©i-Heo'-Hr^-^ O-^^COfNlMfMOi^icStOrHlcsC^cO^ II III III I I ^SO'^fNOOi—iOOOOO'-'COiOi.OOi CO CO ^^Co CO (N CO C O^ CO 1— I O-— (N O O ^^rt lO CO iCi t>. t^ lO CO rJH CO t^ ©* O CO Oo ■^(N O O ^^1— I CO IQ CO CO 1— I CO Co C^ 1— I 00 COCjOS'-H'— i^C'^iOOoCiLO'— 1-^ (NCo(NC^)— ie^,-iO»i5CoO(NCO I I 111 ■— iCoOCOt'-J^O^^CoCO'-i'JS II III I T lO C lO (M(M O I I I ■<-H lC5 .— I Co ^<^co o»ci COC5COCD<^>C5C)lQiCO ^*Oi CO O Co LO ~<^ (M (M t-H ^ »Q ^* Co 00 Co Co f^ O IM ^O (M Co ■-^ o r^ ^ CO lo ?^ — I O O O ■— ' II II C5 lO CO CO '+ <55 t^ 00 0<^(M(NCOCOOCO 0CIMC0'*CO(Nr-ITt< II III Ci(NI> O^CO f- (M OC^(M ^CO ICI I I I r-l C^> lO IC CO O ^ (N (M C^ --ilM I III CO (M CO ©} GO CO ~*C0 O o o o e^ *<(-'-H oc (N J^ III I '#lO-*C0^00t^>,'5OG0 ^ooi^o^cacooco III II I I OOCoCQiO^^ lOCOt^Clt^-fNCJi^^COO-* OCo(MCO<^ >— iCOOC-ir-}i>(>}e^coOoo iQi »ij ^^ IM Co Co CO oo Coo 00 to Co O >o III III III I '0>~< l>0C)C005COOr-(^c|Oi C^ Ot)'C)0'^CO'^~^T-'-'5(MCO(MrtCO'-i-sr II III II III l-OOOCiCO>~(<3-Jf^COCOCOlOCOC5®:)^00 (MO00C0e^C0G0(N(NC0OCc»CiC0O'O I III III I i-i ^ y. bC+i a> c > o gsgsgsss fcD c c bJD CO 'i. o a c o 03 Production of Channel Catfish 249 CO 9« lO (N 05 00 o iC CO (N I III III ■^0500(NTtiOO'rtH050'*»0®5»Cirti(M O"OO'*"OCCiO00'*^^ — CO ' ' ' ' I II I " I I I I (M I Oc 05 O <^ i^ ^ f^ ^ '3'^ -^J 05 O to CD O OiCO(MCitC5to?--'— ''Otof^lM lot^i— (lO I III I I I I I ■-tcsoiMOeo^toc-i^to Ttii-HtoiNco III III I oos-^-o-iOiMtoi-ot^oo e^ ^ ^ f ^ CO •* <^ (N " (M I I I ■<* C5 ^ "50 to O O "^ 00 iCi O to II I I C5LOI>lOOOOooOOOOo i:^^cof^O'^'*'^<3i III I CO 05 1^ >-H CO ^ ~^ --1 »Q --1 to c^ " "^ I II I ®^ lO CO Oi ^- O lO >-< ?^Tj.cotoOio-*»o I I I I I ?-rt^00lO^Tti00 I I I I I -* ^ 05 CO 00 LO --I C^ -^ (M "-H C^4 C^ ^ I I I >i^C0C0COQ^lClC500O ■* ^ T-H t^ i^ o 00 o <^ III I to (M ■— I <5.> CO CO to IM O J^ C ■— I I I lO --^ CO >-. iC rfi CO ?^ -^ CO OCOfMCoCfN-T^iqiCO^ III III ^^00 r^ 9^ "^ ir^ ^ CO <- 'Ci I I I Ot^OOlM Tt^ ^ O Tt" I I I OCO'-hO^OOOOQ^iCOO T-i TfH CO CO O lO f ^ to •* O CO I I I I I I I I C>OO(NC;C5»-0>-iQO'-iT-i III I I CO I COOOI>C^OOC5LOt^OOCOOi-OCO CO-— Q'-HCOOiMCO^'-^O'-'rHC^IO Till II I I t>.cDeocotooo>-Q0C5O'-H(MC0T} S e £ £ £ £ £ £ ■53, & O hC C fcC CO f-> ~' v— ^-^ ■ JlHr44U966 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 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 individuals request copies from the 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 some of the costs of production, is to be included. ' An asterisk designates those numbers 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 figures 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. 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. 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, 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 figin-es in text. March 7, 1962. 6. Two new pelvcosaurs from the lower Pennian 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. 347-362, 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). Bv 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 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. 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. 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 0 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 ilWl ii^miiiiitllijaiiu awflllfailllWllii ¥>^^^ 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 0 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 CO CJ3 a 5 cs . a tic V « o a» H-5 ,2 (7-. o i; a S S « a-£o Sum ■Vtc a C3 CO iT o o S 5 a*' i; " a ..05 -Is C c~ 01.- X .--O OS = c c a i> - S« o oj - ca 1) fc- \M < o d h O >< u o •-I o u H CO o < < U < < :§ o u w n < 60 os.5'x 4) ° 4) lc'o'3) c g oj C' — — ^ ca OS 0) T3 -" o o a u c B o .. V j3 ca AJ3 V o .. o bi Q o .. >>* — U3 5^i cc £ a.2 0) 5 £ c (1) .. BJ'S 9 1-5 c M M 01 — 0) c o -•J — -*j C3 oj . (U L. ei a 3 CO oi V bfi — - - 0-5 CO »- ir..S .. O ^ 3 S- «' 2 .s M 03 O M 3-3 O . O ,« o 2 *> — a a 4) a o =i C3 O w ♦- 03 sis ttl w 3 o; bil " a 4>!n aj a a a o 03 «:^ § i' >. "3 4)'S oi 4) a o a. CO ■7 a u ** 2 S :►. a c3 ux^ .a o c , "a " §.S 5 O 4)_ C • a 2 I" 2 E C a o3 -*^ !" h "^ ff; O ^ , te-d a o O t. 4> I- o O M - £ 4) .— *^ -w o a o » a tr .. O 4) 1-9 ca a *i a o 03 4J.— .-__—. (nX! ^^ O JD t. 5 '" 03 O 03 = 4) — '- nT - 4) 3 a, 4) " *^ !> a M.5 >> a tiC j'3t3= S >- »3 «,^-5 a — ..'■^ X 4) S 05 =3 X o CO .« -c a hj OS £.£.- J & >. ** OS >.: fe <" M a ti oj a ° M— a o 4P O QJ ^ d-d - ..'S hJ O 4> 4< ^tli) 01 J3 M O a o o a te «3 g a; i;j -£fc^£ *^ • -— S3 a 2 o — . 3 e CO < K ■< K o ^ ^ k — a2 *« **- CJ _ o-g in M ^ bfi 83 g.2 .. ■5 — ^2 X t; '^ 3 > 3 4) 5 - O -*^ 03 - - 05 a = f^ « 2.2 a 'E'fc- a O OI — ^ -^ -t^ e tc 4) ti g'S O bfl 4) CJ o £ ■ - 01 1-3 IS ■3 B lilt .— =* a ^- s > a .S 4; o ® a S; VOL. - 1- a a* — a" a.2j:. as w a ° ol a o e p "^ a .2T. 41 S 'a *j OS o cs .^ oJ 1= ..• t. u i; 03 -iJ o £ 03 3 K 340 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 0 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. (A a t/2 ^:s 6 lO 1^ c f^ o ^,_^ . (M o -* o S. ss o^^ "7 S^ S8 2o 2=^ -+ CO •* 05 CO (M 1— ( T-H ^H CM C 3 ~-^ ^^ "-— • ^^ D" flj u <«-i ^^ /■ — s ,— N y— -s ^ o ^ lO LO o o qc ■* o 05 t> 05 00 *«l & IST — lOi (N'* eoci Oi '■^ 'c? ■^ 1 COO ■* 1 05 1 Si VH o co>o QOO t-iO 00 lO o 1 eo 1 lO 1 coT coV 1—1 3^ «0 lO TJ< IC ^o ■^lO 2g ^eo -H CO -hC^ 3 cr >„_• s^ ^_^ v_^ "*— ^ ^-^ r^. Oi '^ «- U-i &.I o oeo o§ ^g « g '*< 1 t^ 1 coT 00 1 oo7 ^7 .2 ^ S 00 1-H 0 O (N ^H (N .— t ^H cc U e" u ^j 60 ^ ^ M •*»a o "c Q ^ ts s 2 CO CO 1 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/