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diff --git a/38425.txt b/38425.txt new file mode 100644 index 0000000..7a902ec --- /dev/null +++ b/38425.txt @@ -0,0 +1,1827 @@ +The Project Gutenberg EBook of Geographic Variation in the North American +Cyprinid Fish, Hybopsis gracilis, by Leonard J. Olund and Frank B. Cross + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Geographic Variation in the North American Cyprinid Fish, Hybopsis gracilis + +Author: Leonard J. Olund + Frank B. Cross + +Release Date: December 27, 2011 [EBook #38425] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK GEOGRAPHIC VARIATION *** + + + + +Produced by Chris Curnow, Joseph Cooper, Erica +Pfister-Altschul and the Online Distributed Proofreading +Team at http://www.pgdp.net + + + + + + + + UNIVERSITY OF KANSAS PUBLICATIONS + MUSEUM OF NATURAL HISTORY + + Volume 13, No. 7, pp. 323-348, pls. 21-24, 2 figs. + + February 10, 1961 + + + + + Geographic Variation + In the North American Cyprinid Fish, + Hybopsis gracilis + + BY + + LEONARD J. OLUND AND FRANK B. CROSS + + + UNIVERSITY OF KANSAS + LAWRENCE + 1961 + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, Henry S. Fitch, + Robert W. Wilson + + + Volume 13, No. 7, pp. 323-348, pls. 21-24, 2 figs. + Published February 10, 1961 + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + PRINTED IN + THE STATE PRINTING PLANT + TOPEKA, KANSAS + 1961 + + [Illustration] + + 28-5871 + + + + +Geographic Variation +In the North American Cyprinid Fish, +Hybopsis gracilis + +BY + +LEONARD J. OLUND AND FRANK B. CROSS + + + + +CONTENTS + + + PAGE + + INTRODUCTION 325 + + METHODS, MATERIALS, AND ACKNOWLEDGMENTS 326 + + DESCRIPTION OF THE SPECIES HYBOPSIS GRACILIS 327 + _Hybopsis gracilis gracilis_ 328 + _Hybopsis gracilis gulonella_ 330 + + INTRASPECIFIC VARIATION 333 + + NATURAL HISTORY 334 + _Habitat_ 334 + _Associated Species_ 336 + _Food_ 338 + _Spawning Season_ 338 + + DISCUSSION 340 + + LITERATURE CITED 343 + + + + +INTRODUCTION + + +The flathead chub, _Hybopsis gracilis_ (Richardson), occurs in the +Plains Region of Canada and the United States, in four major drainage +systems: Mackenzie River, which discharges into the Arctic Ocean; +Saskatchewan River, which discharges into Hudson Bay via Nelson River; +and Missouri-Mississippi System and Rio Grande, both draining into the +Gulf of Mexico. Each of these systems is occupied in part only. In the +Mackenzie Basin, _H. gracilis_ has been reported as far north as Fort +Good Hope (Walters, 1955:347). Flathead chubs occur in the Saskatchewan +Basin from Alberta eastward to Lake Winnipeg, Manitoba, but have not +been found in other streams that flow into Lake Winnipeg (Red River, +Brokenhead River and Whitemouth River) nor in Nelson River downstream +from Lake Winnipeg. In the Missouri Basin the species occurs more or +less continuously from the high plains adjacent to the Rocky Mountains +in Montana and Wyoming down the mainstream of the Missouri River to its +mouth, and down the mainstream of the Mississippi River as far as +Barfield, Arkansas, but not to the Gulf. The species probably attains +its greatest abundance in the Missouri Basin, but it is scarce or absent +in tributaries north and east of the Missouri mainstream, in the South +Platte Basin, and in the central part of the Platte River in Nebraska. +The flathead chub is unknown in the Mississippi Basin above the mouth of +the Missouri River, and in the Ohio River Basin above its mouth. In the +Arkansas River Basin, records are restricted to (1) the headwaters and +tributaries of the Arkansas River from eastern Colorado downstream as +far as Garden City, Kansas, (2) the Cimarron River at Kenton, Cimarron +County, Oklahoma, and (3) the South Canadian River and tributaries from +northeastern New Mexico eastward as far as Norman, McClain County, +Oklahoma, but rarely there. Thus, the range in the Arkansas Basin seems +to consist of three isolated segments. Likewise, isolated populations +exist in the Rio Grande System, where flathead chubs are confined to the +upper parts of the Rio Grande and Pecos basins, above the confluence of +the Rio Grande and Pecos Rivers. Records resulting from introductions +have been reported for the Gila River by Koster (1957:62) and from the +Snake River, Wyoming, by Simon (1946:72). + +Six names apply to the flathead chub, the earliest of which is _Cyprinus +gracilis_ Richardson (1836:120). Other names have sometimes been +accepted as applicable to valid species and/or subspecies, but usage, +diagnoses, and stated ranges have been confusingly inconsistent. For +most of the past 100 years, _Platygobio_ Gill has been recognized as the +appropriate generic name for the flathead chub, but Bailey (1951:192) +places _Platygobio_ and other nominal genera of barbeled minnows having +short guts, protractile premaxillae, and four teeth (primary row) in the +single genus _Hybopsis_ (Agassiz, 1854). Strangely, the orthotype of +_Hybopsis_, _H. gracilis_ Agassiz, is a junior synonym of _H. amblops_ +(Rafinesque) (Hubbs and Ortenburger, 1929b:66) and is a younger name +than _C. gracilis_ Richardson. + +The purpose of this paper is to redescribe the species and to make known +its pattern of geographic variation. Natural history will also be +considered, as will habitat, food habits, and breeding season. + + + + +METHODS, MATERIALS AND ACKNOWLEDGMENTS + + +Ten meristic characters and seventeen measurements of body-parts (the +latter expressed as proportions of standard length) have been analyzed. +They are: number of rays in the dorsal, anal, caudal, pectoral and +pelvic fins; number of scales in the lateral line, before the dorsal +fin, around the body and around the caudal peduncle; number of +vertebrae; body-depth, depth of caudal peduncle, length of caudal +peduncle, predorsal length, length of depressed anal and dorsal fins, +length of pectoral and pelvic fins, head-length, head-depth, +head-width, snout-length, postorbital length of head, length of orbit, +interorbital width, length of upper jaw and width of gape. + +Counts and measurements were made as described by Hubbs and Lagler +(1958), with the exception of scales before the dorsal fin, which were +counted as the number of vertical scale-rows between the upper margin of +the opercular cleft and the origin of the dorsal fin. Vertebral counts, +made from roentgenograms, excluded vertebrae in the Weberian complex +(presumably always four) but included the hypural vertebra. + +Counts and measurements were made on series (usually ten fish) from +localities throughout the range. To minimize effects of allometric +growth, the fish were divided into several length-groups prior to +analysis of proportional measurements: 30-50mm, 50-70mm, 70-100mm, +100-150mm, 150-200mm and 200mm standard length and over. The majority of +specimens examined were 70-100mm in standard length. + +Specimens were obtained from the following institutions: University of +Alberta (abbreviated UA in the text); Museum of Zoology, University of +Michigan (UMMZ); University of Missouri (UM); Montana State College +(MSC); University of Oklahoma Museum of Zoology (UOMZ); University of +Saskatchewan; Royal Ontario Museum, Division of Zoology, Toronto (ROMZ); +University of Wyoming (WU); Museum of Natural History, University of +Kansas (KU). Specimens examined are listed in the accounts of the +subspecies. + +We are grateful to D. A. Boag, Reeve M. Bailey, Arthur L. Witt, C. J. D. +Brown, Carl Riggs, F. M. Atton, W. B. Scott, and George Baxter, all +staff-members of the institutions listed in the immediately preceding +paragraph, for placing specimens at our disposal. Mr. William Peters +analyzed the contents of stomachs of specimens that were used for study +of the food habits. Mr. Artie L. Metcalf assisted in collecting +specimens. Drs. Kenneth B. Armitage and E. Raymond Hall offered valued +suggestions in connection with the preparation of the manuscript. + + + + +DESCRIPTION OF THE SPECIES + + +=Hybopsis gracilis= (Richardson) + +Flathead Chub + +(Synonymy under accounts of subspecies) + +_Description._--Pharyngeal teeth 2,4-4,2, hooked; dorsal fin of +moderate size, falcate, first principal ray longest, extending beyond +posterior rays in depressed fin, its origin usually slightly in front +of insertion of pelvic fin, approximately equidistant from tip of snout +and base of caudal fin, rays 8, rarely 9; pectoral fin strongly +falcate, rays 14-20, usually 16-18; pelvic rays 8, rarely 9; anal fin +falcate, rays 8, rarely 9; caudal rays 19, rarely 20. + +Body slightly compressed, nearly terete; head-length 23.1-28.8 per cent +of standard length; head broad and flattened, snout subconical, +premaxillae protractile, upper lip not medially expanded; mouth +subterminal, nearly horizontal, large; a single pair of terminal +maxillary barbels; orbit usually 5-7 per cent of standard length; +lateral line slightly decurved; intestine short, peritoneum silvery. + +Color brown or olivaceous dorsally, silver or creamy white ventrally, +without distinctive markings; dusky lateral band evident in preserved +specimens. + +Taste-buds present on membrane between first and second principal rays +of all fins, and on first to sixth interradial membranes of pectoral +fin. On the caudal fin, taste buds between first and second principal +rays of upper and lower lobes, though present, are less well developed +than on other fins. Moore (1950:88) states that taste buds are numerous +on the barbels, cheeks, lips, chin, snout, opercles and branchial +membranes, and are present in decreasing numbers over the body. + +Nuptial tubercles of male minute and densely scattered over top of head +and snout; usually present on pectoral rays 1-8, weak when present on +rays beyond the eighth, never found beyond the eleventh ray; minute +tubercles usually found on dorsal, pelvic and anal fins, rarely on lower +scales of caudal peduncle; predorsal scales have a fine peripheral row +of tubercles. + + +=Hybopsis gracilis gracilis= (Richardson) + +(Plate 22) + + _Cyprinus (Leuciscus) gracilis_ Richardson, 1836:120 and Pl. 78 + (original description; Saskatchewan R. at Carlton House). + + _Coregonus angusticeps_ Cuvier and Valenciennes, 1848:534 + (original description; Saskatchewan R.). + + _Pogonichthys communis_ Girard, 1856:188 (in part; original + description); Girard, 1858:247 and plate 55 (in part; + characters; synonymy); Suckley, 1860:361 (Milk R.); Cope, + 1879:440 (Fort Benton, Mo. R.; Judith R.). + + _Platygobio gracilis_, Jordan and Gilbert, 1882:219 (in part; + characters; synonymy); Graham, 1885:74 (Kansas R.; synonymy); + Jordan, 1885:29 (records); Jordan and Meek, 1886:13 (Mo. R., St. + Joseph, Mo.); Meek, 1892:245 (characters; Mo. R., Sioux City, + Iowa); Eigenmann, 1895:111 (Craig; Poplar; Brandon; Medicine + Hat); Meek, 1895:137 (Platte R., Fremont, Neb.); Evermann and + Cox, 1896:412 (in part; habitat; synonymy); Jordan and Evermann, + 1896:326 (in part; characters; synonymy); Thompson, 1898:214 + (Brandon; Saskatchewan R.); Evermann and Goldsborough, 1907:98 + (records from Canada); Forbes and Richardson, 1920:170 + (characters; habitat; synonymy; records from Illinois; but Fig. + 45 is _Hybopsis meeki_ Jordan and Evermann, not _H. gracilis_); + Hankinson, 1929:446 (records from North Dakota); Jordan, 1929:76 + (in part; characters); Jordan, Evermann and Clark, 1930:136 (in + part; synonymy); Churchill and Over, 1933:45 (characters; food; + habitat; spawning; records from South Dakota); O'Donnel, + 1935:481 (Ohio R., Cairo, Ill.; Miss. R., Chester, Ill.); Hinks, + 1943:57 (records from Canada); Clemens, _et al._, 1947:17 + (records from Saskatchewan); Dymond, 1947:19 (distribution in + Canada); Rawson, 1951:208 (Great Slave Lake; Mackenzie R.); + Shoemaker, Pickering and Durham, 1951:84 (Miss. R., Cates, Tenn.; + Miss. R., between Hickman and Barfield, Ark.); Wynne-Edwards, + 1952:18 (distribution in Canada); Miller and Paetz, 1953:47 + (Peace R. at town of Peace River); Walters, 1955:347 + (distribution in Canada; dispersal into Canada); Keleher, + 1956:265 (Saskatchewan R., Manitoba); Lindsey, 1956:771 + (distribution in Canada); Keleher and Kooyman, 1957:110 (Kelsey + Lake, Manitoba); Lindsey, 1957:657 (Laird and Peace drainages, + British Columbia); Scott, 1958:16 (distribution in Canada); + Slastenenko, 1958:7 (distribution in Canada). + + _Platygobio pallidus_ Jordan and Gilbert, 1882:220 (original + description; Ohio R., Cairo, Ill.); Jordan and Evermann, + 1896:326 (characters; synonymy; Ohio R., Cairo, Ill.); Jordan, + Evermann and Clark, 1930:136 (Ohio R., Cairo, Ill.; synonymy). + + _Platygobio gracilis communis_, Simon, 1946:71 (in part; + characters; food; habitat; spawning); Moore, 1950:87 (habitat; + sense organs). + + _Hybopsis gracilis communis_, Bailey, 1951:192 (record from Iowa; + key); Harlan and Speaker, 1951:75 (characters; distribution in + Iowa); Hubbs, 1951:9 (habitat; Miss. R.); Harrison and Speaker, + 1954:516 (habitat); Personius and Eddy, 1955:42 (habitat; Little + Mo. R.). + + _Hybopsis gracilis_, Cleary, 1956:271 (record from Iowa; + distributional map); Bailey, 1956:332 (record from Iowa; key); + Harlan and Speaker, 1956:90 (characters; distribution in Iowa); + Eddy, 1957:111 (in part; characters; key); Moore, 1957:110 (in + part; key); Underhill, 1959:100 (Vermillion R., South Dakota). + +_Diagnosis._--Post-Weberian vertebrae 40-42, usually 41-42; lateral +line scales 50-56; pectoral rays 15-20, usually 17 or more; head-depth +12.3-15.1 per cent of standard length, usually 14.7 per cent or less. +See Figs. 1 and 2. + +_Other characters._--Circumference scale-rows 31-42; predorsal +scale-rows 20-29; size large, as much as 246 mm standard length (see +Fig. 1 of Pl. 24); head-length 23.4-27.4 per cent of standard length, +usually 25.5 per cent or less; postorbital length of head 10.9-13.9 per +cent of standard length, usually 12.5 per cent or less; predorsal length +46.0-51.7 per cent of standard length; orbit 5.1-6.8 per cent of +standard length; prepelvic length 46.6-52.2 per cent of standard length; +caudal peduncle length 17.2-22.1 per cent of standard length. + +_Range_ (Plate 21).--Mackenzie Basin south from Fort Good Hope; +Saskatchewan Basin east to Lake Winnipeg; mainstream of Missouri River +and Mississippi River south to Barfield, Arkansas; intergrading with _H. +g. gulonella_ in upper Missouri Basin and lower parts of major +tributaries to Missouri River in Nebraska and Kansas. + +_Specimens examined._--Below are listed museum numbers, number of +specimens (in parentheses), localities, and year of collection. +Collections marked with asterisk (*) are intergrades more closely +resembling _H. g. gracilis_ than _H. g. gulonella_. Records from +literature are cited in the synonymy. + +ALBERTA: UA (6), Milk R. at town of Milk River, 1950; UA (3), Athabasca +R. at Fort McMurray, 1955; UA (1), Red Deer R. at Steveville, 1952; UA +(2), Peace R. at town of Peace River, 1952; UA (11), Peace R. at +Dunvegan, 1956; UA (2), Simonette R. tributary to Smoky R., date +unknown; ROMZ 17704 (1), Milk R. W town of Milk River, 1955. + +ARKANSAS: UMMZ 128573 (5), Mississippi Co., Mississippi R., 1939. + +ILLINOIS: UMMZ 134799 (146), Mississippi R. at Grand Tower, 1936; UMMZ +147045 (8), Mississippi R. at Cairo, 1944. + +KANSAS: KU 1234 (173), Leavenworth Co., backwater of Missouri R. near +Corral Cr., 1940; * KU 1814 (1), Douglas Co., floodpool of Kansas R., +below Lakeview, 1951; * KU 1825 (1), Douglas Co., floodpool of Kansas +R., 1951; * KU 1841 (56), Douglas Co., Kansas R. at Lawrence, 1951; * KU +1898 (6), Douglas Co., floodpool of Kansas R., 1951; * KU 1911 (5), +Douglas Co., floodpool of Kansas R., 1951; * KU 1928 (2), Jefferson Co., +floodpool of Kansas R., 1951; KU 3850 (30), Atchison Co., Missouri R., +1957; * KU 4377 (2), Douglas Co., Kansas R. at Lawrence, 1958; * KU 4655 +(2), Douglas Co., Kansas R. at Lawrence, 1959. + +MANITOBA: ROMZ 13834 (1), Kelsey Lake, 25 miles east of the Pas, no +date; ROMZ 14500 (25), Saskatchewan R. at the Pas, 1947; ROMZ 16325 (1), +Lake Winnipeg, no date. + +MISSOURI: UMMZ 147126 (130), Mississippi R. at Cliff Cave, 1944. + +MONTANA: * MSC 1878 (36), Carbon Co., Elbow Cr., 1957; * MSC 1943 (11), +Phillips Co., Frenchman Cr., 1957; * MSC 2021 (10), Pondora Co., Marias +R., 1955; * MSC 2022 (4), Lewis and Clark Co., Missouri R. below Holter +Dam, 1948; * MSC 2052 (6), Gallatin Co., Missouri R. near Trident, 1948; +* MSC 3074 (3), Custer Co., Hardy Reservoir, 1952; UMMZ 94146 (34), near +mouth of Powder R., 1926. + +NEBRASKA: * KU 4158 (9), Holt Co., Niobrara R. 6 mi. N Midway, 1958; * +UM (field no. 59-81) (56), Butler Co.-Colfax Co. line, Platte R. 1.5 mi. +S Schuyler, 1959; * UM (field no. 59-74) (5), Dodge Co., Platte R. 1 mi. +S North Bend, 1959; UMMZ 134826 (46), Otoe Co., Missouri R. 1.5 mi. E +Minersville, 1940; UMMZ 134799 (67), Cass Co., Missouri R., 1940; UMMZ +135341 (43), Knox Co., Missouri R. 2 mi. NE Niobrara, 1940; UMMZ 135818 +(95), Thurston Co., Missouri R. NE Macy, 1941. + +NORTHWEST TERRITORY: ROMZ 13627 (1), Great Slave Lake, no date; ROMZ +13628 (1), Great Slave Lake, no date. + +SASKATCHEWAN: * ROMZ 3885 (2), Sucker Cr., trib. Cypress Lake, 1927; +ROMZ 14368 (2), South Saskatchewan R. at Yorath Island, 1941; ROMZ 16620 +(5), South Saskatchewan R. at Saskatoon, 1953; KU 5126 (5), South +Saskatchewan R. at Birson Ferry, 1957; KU 5127 (3), South Saskatchewan +R. at Leader, 1957; KU 5128 (2), North Saskatchewan R. at Cecil Ferry, +1957; KU 5129 (1), South Saskatchewan R. at Clarkboro Ferry, 1957. + +SOUTH DAKOTA: * KU 4961 (9), Haakon Co., Bad R. at Midland, 1959; * KU +4963 (17), Washabaugh Co., White R. 6 mi. SW Belvidere, 1959; * UMMZ +120362 (168), White R. 6.5 mi. S Kadoka, 1934; * UMMZ 127484 (11), Todd +Co., Little White R., 1934; UMMZ 127488 (29), Charles Mix Co., Missouri +R., 1934; * UMMZ 127678 (32), Cheyenne R., E Wasta, 1939; UMMZ 166762 +(21), Hughes Co., Missouri R. 3 mi. NE Pierre, 1952; * UMMZ 166803 (91), +Harding Co., Little Missouri R. at Camp Crook, 1952; UMMZ 166845 (121), +Carson Co.-Walworth Co. line, Missouri R. 2.5 mi. N Mobridge, 1952; UMMZ +166985 (61), Yankton Co., Missouri R. at Yankton, 1952. + +WYOMING: * WU 2073 (6), Washakie Co., Big Horn R. at Worland, 1956. + + +=Hybopsis gracilis gulonella= (Cope) + +(Plate 23) + + _Pogonichthys communis_ Girard, 1856:188 (in part; original + description); Girard, 1858:247 (in part; characters; synonymy); + Cope and Yarrow, 1875:653 (characters; Pueblo, Colo.). + + _Pogonichthys (Platygobio) gulonellus_ Cope, 1864:277 (original + description; near Bridger's Pass, Wyo.). + + _Platygobio gulonellus_ Cope, 1865:85 ("Platte R., near Fort + Riley" [Fort Riley is on Kansas R., not Platte R.; Cope's + specimens probably are from Platte drainage, on basis of known + distributions of other species reported]). + + _Ceratichthys physignathus_ Cope and Yarrow, 1875:651 (original + description; Arkansas R., Pueblo, Colo.). + + _Platygobio communis_, Gill, 1876:408 (characters; Platte Valley; + Green River, Utah [the latter probably erroneous]). + + _Couesius physignathus_, Jordan and Gilbert, 1882:219 (characters; + synonymy; Arkansas R., Pueblo, Colo.); Jordan, 1885:29 + (records). + + _Platygobio gracilis_, Jordan and Gilbert, 1882:219 (in part; + characters; synonymy); Cragin, 1885:109 (Garden City, Kans.); + Gilbert, 1885:98 (Garden City, Kans.); Jordan, 1885:29 + (records); Evermann and Cox, 1896:412 (in part; habitat; + synonymy); Jordan and Evermann, 1896:326 (in part; characters; + synonymy); Ortenburger and Hubbs, 1927:125 (Canadian R., Norman, + Okla.); Hubbs, 1927:75 (parasites; teratology; records from New + Mexico); Hubbs and Ortenburger, 1929a:28 (S. Canadian R., + Durham, Okla.); Jordan, 1929:76 (in part; characters); Jordan, + Evermann and Clark, 1930:136 (in part; synonymy). + + _Platygobio physignathus_, Jordan and Evermann, 1896:325 + (characters; synonymy; records from upper Arkansas R.); Ellis, + 1914:62 (characters; synonymy; records from Colorado); + Cockerell, 1927:123 (distribution in Colorado); Jordan, Evermann + and Clark, 1930:136 (synonymy; records from upper Arkansas R.). + + _Platygobio gracilis communis_, Simon, 1946:71 (in part; + characters; food; habitat; spawning). + + _Platygobio gracilis gulonellus_, Simon, 1946:72 (characters; + records from Wyoming; Arkansas R.). + + _Platygobio gracilis:_ _communis_ x _gulonellus_, Simon, 1946:92 + (North Platte R., Neb.-Wyo. line). + + _Platygobio gracilis physignathus_, Moore, 1950:87 (habitat; sense + organs). + + _Hybopsis gracilis communis_, Beckman, 1952:50 (characters; food; + habitat); Cross, Dalquest and Lewis, 1955:222 (records from + Texas). + + _Hybopsis gracilis physignathus_, Beckman, 1952:50 (characters; + habitat). + + _Hybopsis gracilis_, Eddy, 1957:111 (in part; characters; key); + Koster, 1957:61 (characters; habitat; spawning; food); Moore, + 1957:110 (in part; key); Smith, 1958:177 (fossil record; Doby + Springs, Okla.). + +_Diagnosis._--Post-Weberian vertebrae 36-38, rarely 39; lateral line +scales 42-54, usually less than 50; pectoral rays 14-19, usually fewer +than 17; head-depth 13.5-18.0 per cent of standard length, usually 14.8 +per cent or more. See Figures 1 and 2. + +_Other characters._--Circumference scale-rows 30-40, slightly fewer than +in _H. g. gracilis_; predorsal scale-rows 17-27, somewhat fewer than in +specimens from Canada, but much the same as specimens from the +Missouri-Mississippi system; size small, rarely as much as 130 mm +standard length (Fig. 1 of Pl. 24); head-length 24.0-28.0 per cent of +standard length, usually more than 25.5 per cent; postorbital length of +head 11.2-14.4 per cent of standard length, usually more than 12.5 per +cent (both characters illustrate the larger head of _H. g. gulonella_); +predorsal length 46.4-52.7 per cent of standard length, longer than in +the other subspecies; orbit 5.0-6.6 per cent of standard length; +prepelvic length 47.4-53.7 per cent of standard length, longer than in +_H. g. gracilis_; caudal peduncle length 17.1-22.7 per cent of standard +length, essentially the same in both subspecies. + +The label on types of this subspecies, in the Academy of Natural +Sciences of Philadelphia, states merely "near Bridger's Pass, Wyo., +Expedition of 1856, Dr. W. A. Hammond" (letter from Dr. James Boehlke to +Cross, dated Jan. 27, 1960). Dr. Hammond was a surgeon who also +collected scientific specimens, assigned to an expedition under the +command of Lt. F. T. Bryant. Bryant's log is recorded in the Proceedings +of the 35th Congress (1858:455-481). The site at which these specimens +were taken cannot be ascertained from the log, but study of it is +helpful in indicating the probable locations. + +The expedition left Fort Riley on June 21, 1856, on the following route: +up Republican River; across to Fort Kearney on Platte River; west along +Platte River to S. Platte River; up S. Platte River to Pole (Lodgepole) +Creek; Pine-Bluffs (Neb.-Wyo. line); across East Fork to West Fork of +Laramie River; Cooper's Creek; West Fork of Medicine Bow; Pass Creek and +down canyon of Pass Creek; across N. Platte River; up Sage Creek; on +August 15, camped on Muddy Creek, tributary to Green River (first record +of fish, trout); back to Sage Creek; August 19-21, camped on island in +North Platte River; to Pass Creek; Elk Creek; west branch of Medicine +Bow; Aspen Creek; West Fork of Laramie River; August 29, to East Laramie +River where a large supply of fish was caught; tributary of Cache la +Poudre then downstream to mouth of this river; down South Platte River +past mouth of Crow Creek and Beaver Creek; left South Platte River 14 +miles below mouth of Beaver Creek, toward Republican River; down Rock +Creek to Arikaree; down Arikaree to Republican River and down the +Republican to Fort Riley. + +Mention is made of fish only twice in the entire log. We doubt that +Muddy Creek or the East Laramie River is the type locality of _P. +gulonellus_, because the flathead chub has not since been found in +either of these streams. The most likely collection site for _P. +gulonellus_ is the North Platte River near the mouth of Sage Creek, in +what is now Carbon County, Wyoming, where the expedition was camped for +three days. This species is known to occur in the North Platte River, +and since the type locality is reported as "near Bridger's Pass" this is +the probable location. + +_Range_ (Plate 21).--Upper mainstream and tributaries of Rio Grande, +Pecos, Arkansas and North Platte Rivers; isolated populations in +tributaries of the upper Missouri River. + +_Specimens examined._--Below are listed museum numbers, number of +specimens (in parentheses), localities and year of collection. Series +marked by asterisks (*) are intergrades tending toward _H. g. +gulonella_. Literature reports are cited in the synonymy. + +COLORADO: KU 4742 (162), Bent Co., Purgatoire R. at Las Animas, 1959; KU +4748 (105), Pueblo Co., Arkansas R. at west edge of Pueblo, 1959; KU +4758 (50), Fremont Co., Arkansas R. at Florence, 1959; KU 4769 (64), +Fremont Co., Beaver Cr., 1959. + +KANSAS: KU 2648 (2), Finney Co., Arkansas R., 1958; KU 2858 (13), Finney +Co., Arkansas R. at Garden City, 1951; KU 3964 (12), Kearney Co., +Arkansas R., 1958; * KU 4041 (2), Cheyenne Co., Republican R., 1958; KU +4732 (30), Hamilton Co., Arkansas R. at Kendall, 1959; * KU 4868 (1), +Kansas-Nebraska line, Republican R. 1.5 mi. S. Hardy, 1959. + +MONTANA: * MSC 1960 (8), Powder River Co., E. Fork of Powder R., 1957; +MSC 2010 (64), Dawson Co., Redwater R., 1957. + +NEBRASKA: * KU 2140 (2), Dawson Co., Platte R., at Gothenburg, 1931; * +KU 4863 (20), Furnas Co., Republican R. at Cambridge, 1959; * UM (field +no. 59-49) (74), Scotts Bluff Co., North Platte R. at Morrill, 1959; * +UMMZ 133918 (17), Dixon Co., Logan Cr., 1939; * UMMZ 134813 (31), North +Platte R., Neb.-Wyo. line, 1941; * UMMZ 135084 (14), Harlan Co., Beaver +Cr. 0.25 mi. S Stamford, 1940; * UMMZ 135200 (41), Scotts Bluff Co., +North Platte R. 1 mi. SE Henry, 1940; * UMMZ 135280 (59), Cherry Co., +Niobrara R. 3 mi. SE Valentine, 1940; * UMMZ 135700 (25), Buffalo Co., +South Loup R. 8 mi. N Miller, 1941; * UMMZ 135778 (54), Thurston Co., +Logan Cr. 2.5 mi. W Pender, 1941; * UMMZ 135786 (25), Dixon Co., Logan +Cr. 0.5 mi. NW Wakefield, 1941. + +NEW MEXICO: KU 4219 (50), Colfax Co., Cimarron Cr. at Springer, 1958; KU +4235 (19), Mora Co., Sapello Cr. near Sapello, 1958; KU 4245 (157), +Bernalillo Co., Rio Grande 12 mi. S Bernalillo, 1958; KU 4255 (22), Rio +Arriba Co., Rio Grande at Velarde, 1958; KU 4266 (53), Sandoval Co., Rio +Grande 2 mi. N Cochiti Pueblo, Marcelino Baca bridge, 1958; KU 4269 +(91), San Miguel Co., Pecos R., 3 mi. S Pecos, 1958; KU 4274 (25), +Sandoval Co., Jemez R. at Jemez Canyon Dam, 1958; KU 4294 (113), +Guadalupe Co., Pecos R. 3 mi. N. Dilia, 1958; UMMZ 94897 (146), Pecos R. +at San Tuan, 1926; UMMZ 94898 (1), Pecos R. at San Juan, 1926; UMMZ +118209 (68), Sapello Cr. at Sapello, 1937; UMMZ 133131 (7), Pecos R. +0.5 mi. N Santa Rosa, 1940; UMMZ 133136 (1), Rio Grande at Albuquerque, +1940. + +OKLAHOMA: KU 2329 (1), Cleveland Co.-McClain Co. line, S. Canadian R., +1952; UOMZ 26355 (10), Cimarron Co., Cimarron R. 2 mi. N. Kenton, 1957; +UOMZ 5917 (2), Cleveland Co., S. Canadian R. S Norman, 1925. + +TEXAS: KU 3409 (18), Hemphill Co., Canadian R. at town of Canadian, +1955. + +WYOMING: WU 2084 (4), Platte Co., N. Platte R. at Glendo, 1956; WU 2095 +(3), Converse Co., N. Platte R. at Douglas, 1956; UMMZ 104064 (58), N. +Platte R. below Guernsey Dam, 1937; * UMMZ 114642 (7), drainage ditch in +Wind R. drainage, 1936; * UMMZ 114644 (20), drainage ditch at Riverton, +1936; * UMMZ 127518 (63), Weston Co., Beaver Cr., 1934; * UMMZ 127681 +(20), Big Horn Co., Big Horn R. tributary, 1939; * UMMZ 136488 (9), +Crook Co., Belle Fourche R. 15 mi. N Devil's Tower, 1941; * WU 2122 and +two uncatalogued series at WU (13), Belle Fourche R., no precise +locality or date; UMMZ 159969 (14), Natrona Co., N. Platte R. 2 mi. E +Casper, 1950. + + + + +INTRASPECIFIC VARIATION + + +Two subspecies of _H. gracilis_ are recognized by us: one northern and +eastern, characteristically inhabiting large rivers (_H. g. gracilis_), +and one southern and western, characteristically inhabiting small +streams (_H. g. gulonella_). Other scientific names that have been +applied to this fish in the past are listed in the synonymy. + +_H. g. gulonella_ is a chubby, deep-bodied fish, whereas _H. g. +gracilis_ is long and slender. The head of the creek subspecies is +deeper and longer than that of _H. g. gracilis_, being rounded +anteriorly when seen in sideview. The head of the large-river subspecies +is acutely wedge-shaped in profile. _H. g. gracilis_ has a larger orbit +than _H. g. gulonella_. Fins of _H. g. gracilis_ are more strongly +falcate than those of the other subspecies. _H. g. gracilis_ has a +greater number of lateral line scales, pectoral rays and post-Weberian +vertebrae than the creek subspecies. The large-river subspecies attains +much larger size than does the creek subspecies (Plate 24). Except in +areas of intergradation, complete separation of the two subspecies can +be made on the basis of lateral line scales, pectoral rays, +post-Weberian vertebrae and head-depth. The regressions of head-depth on +standard length in _H. g. gracilis_ from the Saskatchewan River (several +localities) and in _H. g. gulonella_ from Beaver Creek, Arkansas River +Drainage (KU 4769) are shown in Plate 24. Although values for the +largest specimens of _H. g. gracilis_ are omitted from Plate 24, the +regression remains essentially linear to standard lengths of +approximately 250 mm. On the basis of head-depth alone, separation of +the two subspecies is possible in specimens larger than 40 mm. Similar +results were obtained by using the regression of postorbital length on +standard length, and could have been obtained by using other +proportional measurements. + + + + +NATURAL HISTORY + + +_Habitat_ + +The species inhabits alkaline streams with shifting sand bottoms where +the waterlevel fluctuates considerably with heavy rains and melting +snow. The flathead chub is found in silty water and often is the +predominant species in streams that have high turbidity. The remarkable +ability of this fish to withstand exceedingly high turbidity is +illustrated by its predominance in the Little Missouri River, which has +an average concentration of suspended silt two and one-half times that +of the Missouri River at Kansas City (Personius and Eddy, 1955:42). + +[Illustration: FIGURE 1. Graphic analysis of lateral line scales, +pectoral rays and post-Weberian vertebrae in _Hybopsis gracilis_. +In each symbol, horizontal line = range, vertical line = mean, +open rectangle = one standard deviation on each side of mean, black +rectangle = twice the standard error on each side of mean. + +_H. g. gracilis_ is found in large rivers throughout its range, +occasionally migrating into smaller streams, especially in the spawning +season. It prefers the main channel of rivers in moderate to strong +current. All series examined are from elevations lower than 3,000 feet. + +Numbers to left of symbols = number of specimens examined from that +locality; combined collections indicated by brackets. The dash-lines +represent drainage patterns of rivers in which this species occurs.] + +_H. g. gulonella_ occupies small rivers and creeks, preferring pools +with moderate currents. In fall, dense concentrations of this subspecies +have been found in small pools, where brush, driftwood or other debris +deflects the current and prevents filling with drifting sand. Hundreds +of flathead chubs were collected in such pools in the Purgatoire and +Arkansas rivers. Specimens were also collected with ease in Beaver +Creek, Colorado, from pools with murky water and slight flow, over +bottoms of gravel and bedrock. No brush or other debris was near the +pools. In each case the streams carried little water, although they +undoubtedly carry greater volumes of water in spring and early summer +after rains and spring thaws. The preferred bottom-type for this +subspecies seems to be gently shifting sand. + +_H. g. gulonella_ is found in warm-water streams, whereas _H. g. +gracilis_ occurs in cooler water. The southwestern subspecies was taken +in August in the Mora River drainage at Sapello (temperatures above 80 deg. +F.) but not at Mora (temperatures below 70 deg. F.). In the Purgatoire +River, a thriving population was found where the water temperature was +92 deg. F., on September 6, 1959. In the Arkansas and Pecos rivers and the +Rio Grande this subspecies is most abundant below the mountainous parts +of the stream-courses, but at elevations higher than 4,000 feet on the +plains. + + +_Associated Species_ + +[Illustration: FIGURE 2. Graphic analysis of head-depth, postorbital +length of head and predorsal length of _Hybopsis gracilis_, expressed +as thousandths of standard length. Numbers in parenthesis = number +of specimens examined from each locality. In each symbol, horizontal +line = range, vertical line = mean, open rectangle = one standard +deviation on each side of mean, black rectangle = twice the standard +error on each side of mean. The dash-lines represent drainage patterns +of rivers in which this species occurs. All measurements are of +specimens 70 to 100 mm in standard length.] + +In the Pecos and Arkansas basins, species commonly taken with _H. g. +gulonella_ are _Catostomus commersonnii_, _Hybognathus placita_, +_Notropis lutrensis lutrensis_, _Notropis stramineus missuriensis_, +_Pimophales promelas_, and _Campostoma anomalum plumbeum_. The only +spiny-rayed fishes that we have found with _H. g. gulonella_ are +_Lepomis cyanellus_ and _L. humilis_, both of which are scarce. +Associates of _H. g. gracilis_ include the same species, plus other +ostariophysan fishes such as species of _Carpiodes_, _Ictiobus_, and +silt-adapted species of _Hybopsis_ and _Notropis_. + +We failed to find the flathead chub at any of 11 localities in the South +Platte drainage, where we collected in September, 1959. Dr. George +Baxter, of the Department of Zoology, University of Wyoming, told +us that he has never found _H. gracilis_ in that drainage. The fauna +of the South Platte includes _Catostomus catostomus_, _Semotilus +atromaculatus_, _Hybopsis biguttata_, _Hybognathus hankinsoni_, +_Notropis cornutus frontalis_, _Etheostoma nigrum_ and _E. +exile_--species rarely if ever found with _H. gracilis_. + +Ecologically, _H. g. gulonella_ seems to be the counterpart of +_Semotilus atromaculatus_ in streams where the latter species is absent. +Observations of _H. g. gulonella_ in the Purgatoire River indicated +that loosely-organized groups of flathead chubs congregated one to four +inches above the bottom of pools, and near or under protective cover +such as roots of vegetation or debris lodged against shore. Individuals +moved about independently within the group (rather than as schools), and +occasionally rose to the surface, perhaps for food. + + +_Food_ + +The flathead chub is chiefly carnivorous, but its food includes some +aquatic vegetation (Table 1). Most organisms found in specimens (both +subspecies) were terrestrial insects (Coleoptera, Diptera, Orthoptera); +all insects were adult stages, except those designated as larvae in +Table 1. Roundworms probably were parasites, rather than food. + +Hubbs (1927:76) states that the food of young flathead chubs that were +obtained from the Arkansas River System in New Mexico consisted "almost +entirely of crustaceans (small ostracods and cladocerans to the +exclusion of all else but an occasional larval or adult insect, etc.)." + + +_Spawning Season_ + +Specimens of _H. g. gulonella_ that have been examined reach sexual +maturity at approximately 65 mm standard length. Most specimens of _H. +g. gracilis_ less than 85 mm in standard length are immature, but larger +specimens probably are mature. + +The spawning season is in late summer, beginning in July and extending +into September. Specimens from the Peace River, collected on August 10, +1952, include females that were mostly spent and tuberculate males. +Males and females in spawning condition were collected in the Milk River +in August of 1955. A large prespawning female was obtained in Red Deer +River in June of 1952. A male from Fort McMurray had fairly well +developed tubercles on August 9, 1955. A prespawning female was taken +from the Saskatchewan River at Clarkboro Ferry on June 7, 1957. +Tuberculate males were collected in the Powder River on June 30, 1957. +Specimens from the White River in South Dakota, collected on July 7, +1934, include tuberculate males. The specimens discussed above are _H. +g. gracilis_ or intergrades tending toward that subspecies. + +Specimens of _H. g. gulonella_ collected in the Arkansas River at Pueblo +and Florence, Colorado, on September 7, 1959, include some tuberculate +males, although most females are spent. On August 8, 1957, a series of +flathead chubs that includes tuberculate males was collected in the +Redwater River, Montana. In the Pecos River on August 25, 1958, +spawning seemingly had been completed, although a few males still bore +tubercles. + +TABLE 1. ORGANISMS FOUND IN STOMACHS OF HYBOPSIS GRACILIS FROM VARIOUS +LOCATIONS, EXPRESSED AS PERCENTAGE OF TOTAL VOLUME. + + A: S. Saskatchewan R., Clarkboro Ferry, Sask. + B: Milk R., Alberta + C: Missouri R., S. D. + D: Missouri R., Neb. + E: Arkansas R., Fremont Co., Colo. + F: Arkansas R., Pueblo Co., Colo. + G: Pecos R., San Miguel Co., N. M. + + ==============================+=====+=====+=====+=====+=====+=====+===== + | A | B | C | D | E | F | G + ------------------------------+-----+-----+-----+-----+-----+-----+----- + No. specimens examined | 1 | 7 | 6 | 10 | 10 | 10 | 10 + | | | | | | | + No. specimens containing food | 1 | 6 | 1 | 2 | 1 | 3 | 7 + ------------------------------+-----+-----+-----+-----+-----+-----+----- + KIND OF ORGANISM | | | | | | | + | | | | | | | + Aphasmidia |10.0 |00.7 | |03.0 | | | + Arthropoda | | | | | | | + Araneae | | | | | | | + Argiopidae | | | | |04.0 | | + Theridiidae | | | | |04.0 | | + Insecta | | | | | | | + Ephemeroptera (nymph) | | | | | | | + Baetidae | |05.0 | | | | | + Heptagenidae | |08.0 | | | | | + Hemiptera | | | | | | | + Corixidae |35.0 |00.3 | | | | | + Hymenoptera | | | | | | | + Formicidae | |21.0 | | | | |60.0 + Coleoptera | | | | | | | + Staphylinidae | |01.7 |07.0 | | | | + Scolytidae | |13.3 |70.0 | | | | + Tenebrionidae | |05.7 | | |70.0 | | + Carabidae | |05.7 | | | |01.0 | + Curculionidae | |01.0 | | | | | + Coccinellidae | | | | | | |09.0 + Trichoptera (case) | |01.7 | | | | | + Diptera | | | | | | | + Mymaridae | |00.3 | | | | | + Empididae | |01.3 | | | | | + Cecidomyiidae | | | | |04.0 | | + Trachinidae | |00.7 | | | | | + Simulidae | |06.7 |20.0 | | | | + Tabanidae | | | | |06.0 | | + Chironomidae | | | | |06.0 | | + Not identified to family| |01.0 | | | | | + Orthoptera | | | | | | | + Locustidae | |07.7 | | | | | + Tettigoniidae | | |03.0 |70.0 | | |09.0 + Tetrigidae | | | | |06.0 | | + Homoptera | | | | | | | + Fulgoridae | |05.0 | | | | |01.0 + Insect egg | |00.7 | | | | | + Plants | | | | | | | + Cyanophyceae | | 09.0| | | | 99.0| 20.0 + Cyperaceae | | 02.0| | | | | 01.0 + Zannichellia palustris | | 00.3| | | | | + Vascular remains | 55.0| | | 27.0| | | + | | | | | | | + Miscellaneous | | | | | | | + Sand | | 00.7| | | | | + Pharyngeal tooth | | 00.3| | | | | + +-----+-----+-----+-----+-----+-----+----- + Total (%) |100.0| 99.8|100.0|100.0|100.0|100.0|100.0 + ------------------------------+-----+-----+-----+-----+-----+-----+----- + +Spawning apparently occurs when river levels recede to the seasonal +lows. In late summer, temperatures of these rivers probably are maximal, +their turbidities are reduced, and their sandy bottoms are stable. +Underhill (1959) reports that this species is rare in the Vermillion +River, a northeastern tributary of the Missouri River, except in autumn +when large numbers occur near the mouth of the river. We suspect that +this is associated with spawning. + +[Illustration: PLATE 21 + +Distribution of collections examined.] + +[Illustration: PLATE 22 + +_Hybopsis gracilis gracilis._ Missouri River, Thurston County, northeast +of Macy, Nebraska. Largest specimen 87.5 mm standard length.] + +[Illustration: PLATE 23 + +_Hybopsis gracilis gulonella._ Pecos River, San Miguel County, 3 miles +south of town of Pecos, New Mexico. Largest specimen 91 mm standard +length.] + +[Illustration: PLATE 24 + +FIG. 1. Top: _Hybopsis gracilis gracilis_, 230.0 mm standard length, +one of the largest specimens examined. Missouri River, Carson +County-Walworth County line, 3 miles northeast of Mobridge, South +Dakota, at mouth of Grand River. + +Bottom: _Hybopsis gracilis gulonella_, 121.6 mm standard length, the +largest specimen examined of this subspecies. Beaver Creek, Fremont +County, 10 miles northeast of Florence, Colorado, on Highway 115.] + +[Illustration: FIG. 2. Regression of head-depth on standard length in +_Hybopsis gracilis gracilis_ from the Saskatchewan River, and in _H. g. +gulonella_ from Beaver Creek, Arkansas River Drainage (KU 4769).] + + + + +DISCUSSION + + +_Hybopsis gracilis_ is highly variable in several morphological +characteristics, including size and shape of head, body, and fins, and +number of scales, vertebrae, and fin-rays. The variations are correlated +in a way that indicates the existence of two subspecies. One of these, +_H. g. gracilis_, attains large size, and has 1) a slender, streamlined +body, 2) a depressed head that is acutely wedge-shaped in profile, 3) +strongly falcate fins with the dorsal and pelvic fins originating +anteriorly, and 4) many scales, vertebrae, and pectoral fin-rays. The +second subspecies, for which _H. g. gulonella_ is the oldest applicable +name, is small, and has 1) a deep, chubby body, 2) head convex in dorsal +contour (less depressed than in _H. g. gracilis_), 3) fins less falcate +than in the latter subspecies, with the dorsal and pelvic fins +originating more posteriorly, and 4) fewer scales, vertebrae, and +pectoral fin-rays than _H. g. gracilis_. These differences are +consistently expressed throughout the size-ranges of the subspecies, and +in series collected at the same or nearby localities in several +different years. Considerable variability was found in features other +than those mentioned above, but individual variation among specimens +from the same locality and adjacent localities is so great that none is +diagnostic of subspecies. For example, orbital size and length of fins +(but not their falcate shape) are variables that have little diagnostic +value, although both features seem to vary in clinal fashion, with the +higher values in the north. + +Variation in _H. gracilis_, as shown in the graphic analysis (Figs. 1 +and 2) and distribution map (Plate 21), presents two clines: a +north-south cline and a large-river to small-river (mainly east-west) +cline. The absence of _H. gracilis_ from certain portions of river +systems is a matter of concern. The species has not been found in the +lower Arkansas River and the Rio Grande, nor in sandy tributary creeks +in eastern Kansas and Missouri that appear to provide suitable habitat. +It has already been noted that _H. g. gulonella_ seems to be the +ecological equivalent of _Semotilus atromaculatus_ in streams in which +_S. atromaculatus_ is not found. _S. atromaculatus_ occurs in creeks of +eastern Kansas and Missouri, and may provide interspecific competition +that prevents establishment of the flathead chub in these creeks. +Regardless of cause, the gaps in distribution of _H. gracilis_ tend to +limit gene flow. + +Many characters used in the separation of the two subspecies are known +to be influenced by environmental conditions, especially temperature. +Hubbs (1922, 1926, 1941), Schultz (1927), Vladykov (1934), Taning (1952) +and Weisel (1955), among others, have pointed out a correlation between +temperature (or developmental rate of fish) and the number of vertebrae, +scales, and fin-rays. Likewise, Martin (1949) and Hart (1952) have shown +that the proportions of some body-parts vary in response to temperature +during early development. In _H. gracilis_, the general nature of the +clines found in a majority of characters (but not all characters) +suggests a temperature influence. However, temperature-dependent +variability that has so far been demonstrated experimentally in fishes +is generally of lesser magnitude than the differences distinguishing _H. +g. gracilis_ and _H. g. gulonella_. To our knowledge, the most extreme +differences that have been induced by modification of temperature are +those reported for _Salmo trutta_ by Taning (1952:181-182), who states: +"Shock treatment produced by especially great changes in temperature +(_c._ 10-14 deg. C), especially during the super-sensitive period [of +somatic differentiation that fixes vertebral number] may produce ... a +difference of 3-4 vertebrae ... in offspring of the same parents." The +difference cited approximates that which distinguishes natural +populations of _H. g. gracilis_ and _H. g. gulonella_. Although we +cannot assume that the sensitivity of the brown trout is the same as +that of the flathead chub, the causative conditions in Taning's study +could scarcely be expected in nature; furthermore, it seems significant +that extremely high (as well as extremely low) mean numbers of scales +and vertebrae were found at southern localities, and that low mean +numbers of scales and vertebrae were found as far north as Wyoming and +Montana. We think it likely that temperature does influence the +expression of characters in _H. gracilis_, directly in individual +development, and indirectly as a selective mechanism in the evolutionary +process. The extent to which each kind of influence exists can be proved +only by experimental work with both subspecies, which we hope to +undertake at a later date. + +Other environmental factors that may have selective influence in this +species are rate of current, volume of flow, and turbidity. Interaction +of these environmental factors could result in genetic fixation of +morphological characters through natural selection. The characters that +distinguish _H. g. gracilis_ from _H. g. gulonella_ seem adaptive to +life in large rivers and small streams. Evidence that these characters +are under limited, direct environmental influence is found among +populations in the Arkansas River System. Although populations in the +Arkansas River have no continuity with populations of _H. g. gracilis_, +upstream-downstream variations like those found in other river systems +are apparent, but in lesser degree. The direction of variation in the +Arkansas River is the reverse of that in the Platte and other +tributaries of the Missouri River. For example, the populations farthest +upstream (Florence, Pueblo) have slightly higher mean numbers of lateral +line scales than do populations from Kansas, downstream. + +A remarkable effect of extreme parasitism in _H. gracilis_ has been +described by Hubbs (1927). Very young chubs that harbored numerous +tapeworms (_Proteocephalus_) had unusually large numbers of lateral-line +scales, large eyes, short snouts, small fins, small mouths lacking +barbels, and coalescent nares (internarial bridge weak or absent). Some +of these abnormalities presumably resulted from retention of larval +characteristics of the fish, correlated with the degree of infestation +by tapeworms. No teratological adults were found, indicating that severe +infections prevent survival to maturity. + +_H. g. gracilis_ occurs in three separate river systems (Mackenzie, +Saskatchewan, Missouri-Mississippi) from latitude 36 deg. N to 66 deg. N, and +longitude 89 deg. W to 123 deg. W. _H. g. gulonella_ exists as several +seemingly-isolated populations in the upper parts of the Rio Grande, +Pecos, South Canadian, Cimarron, Arkansas, Platte, and upper Missouri +basins, from latitude 35 deg. N to 48 deg. N, and longitude 97 deg. W to 100 deg. W. + +There is evidence of high mobility on the part of both subspecies, based +on irregularity of their occurrence in certain localities. Many +collections have been made in the Cimarron River in the vicinity of +Kenton, Oklahoma, from 1925 to the present, but only one of these (in +1957) contained flathead chubs. Bait dealers who seine the South +Canadian River in Dewey County, Oklahoma, have taken flathead chubs in +abundance in some seasons, but not at all in others. Seasonal variation +in abundance in the lower Vermillion River, South Dakota (Underhill, +1959:100) has been cited, and the number collected in the lower Kansas +River near Lawrence has varied similarly. Many rivers occupied by _H. g. +gulonella_ (and by intergrades) are intermittent, and in some years +their sand-filled channels become wholly dry for many miles. These +factors probably promote mixing of the two subspecies, and may account, +over long periods of time, for the wide dispersal of _H. g. gulonella_ +in the Missouri Basin. Flathead chubs are known from Pleistocene beds at +Doby Springs, Oklahoma (the Doby Springs local fauna) (Smith, 1958:177). +Drainage connections between the Arkansas, Kansas and Platte river +systems existed in Kansan and Nebraskan times (Frye and Leonard, +1952:189-190). Populations that have subsequently become isolated in +those rivers could be accounted for in this way. Flathead chubs could +have entered the Rio Grande-Pecos system by stream-capture from the +Arkansas System, in northeastern New Mexico or southern Colorado. _H. g. +gracilis_ undoubtedly entered the Saskatchewan and Mackenzie basins from +the upper Missouri Basin, following glacial retreat (Walters, 1955:347). + + + + +LITERATURE CITED + + +AGASSIZ, L. + + 1854. Notice of a collection of fishes from the southern bend of the + Tennessee River, in the state of Alabama. Amer. Jour. Sci. and + Arts, 17(50):297-308 and 353-365. + +BAILEY, REEVE M. + + 1951. A check-list of the fishes of Iowa, with keys for + identification. _In_ Iowa fish and fishing, by James R. Harlan + and Everett B. Speaker. 1st edition. Iowa Cons. Comm. 185-237. + + 1956. A check-list of the fishes of Iowa, with keys for + identification. _In_ Iowa fish and fishing, by James R. Harlan + and Everett B. Speaker. 3rd edition. Iowa Cons. Comm. 325-377. + +BECKMAN, WILLIAM C. + + 1952. Guide to the fishes of Colorado. Univ. Colorado Mus., 11:1-110. + +BRYANT, FRANCIS T. + + 1858. Report of Francis T. Bryant to Col. J. J. Abert, Chief, Corps + Top. Engs., USA. _In_ Executive Documents of the Senate of the + U. S., First Session, 35th Congress, and Special Session of + 1858. 3:455-481. + +CHURCHILL, EDWARD P., and WILLIAM H. OVER. + + 1933. Fishes of South Dakota. South Dakota Dept. Game and Fish. 1-87. + +CLEARY, ROBERT E. + + 1956. The distribution of the fishes of Iowa. _In_ Iowa fish and + fishing, by James R. Harlan and Everett B. Speaker. 3rd + edition. Iowa Cons. Comm. 267-324. + +CLEMENS, W. A., A. MACDONALD, H. MCALLISTER, A. MANSFIELD and D. RAWSON. + + 1947. The fishes of Saskatchewan. _In_ Report of the Royal Commission + on the fisheries of the province of Saskatchewan. 1-131. + +COCKERELL, THEODORA D. A. + + 1927. Zoology of Colorado. Univ. Colorado, Boulder. 1-262. + +COPE, E. D. + + 1864. Partial catalogue of the cold-blooded Vertebrata of Michigan. + Proc. Acad. Nat. Sci. Philadelphia, part 1:276-285. + + 1865. Partial catalogue of the cold-blooded Vertebrata of Michigan. + Proc. Acad. Nat. Sci. Philadelphia, part 2:78-88. + + 1879. A contribution to the zoology of Montana. Amer. Nat., + 13:432-441. + +COPE, E. D., and H. C. YARROW. + + 1875. Report upon the collections of fishes made in portions of + Nevada, Utah, California, Colorado, New Mexico, and Arizona, + during the years 1871, 1872, 1873, and 1874. _In_ Report upon + geographical and geological explorations and surveys, west of + the one hundredth meridian, in charge of First Lieut. Geo. M. + Wheeler. Volume V, Zoology. 635-703. + +CRAGIN, F. W. + + 1885. Preliminary list of Kansas fishes. Bull. Washburn Lab. Nat. + Hist., 1(3): 105-111. + +CROSS, FRANK B., WALTER W. DALQUEST and LEO LEWIS. + + 1955. 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Freshwater vertebrates of the Arctic and Subarctic. Bull. Fish. + Res. Bd. Canada. 94:1-28. + + +_Transmitted November 8, 1960._ + + +28-5871 + + +[Transcriber's Note: + +The following changes have been made to the original text: + + Table of Contents: page number of "Food" and "Spawning Season" + changed from 339 to 338 + Page 327: "abbreviated AU" changed to "abbreviated UA" + Page 344: "Societe Geologique" changed to "Societe Geologique"] + + + + + +End of the Project Gutenberg EBook of Geographic Variation in the North +American Cyprinid Fish, Hybopsis gracilis, by Leonard J. Olund and Frank B. 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