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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/38425-8.txt b/38425-8.txt new file mode 100644 index 0000000..a9bb533 --- /dev/null +++ b/38425-8.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: ISO-8859-1 + +*** 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_ × _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 Böhlke 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° +F.) but not at Mora (temperatures below 70° F.). In the Purgatoire +River, a thriving population was found where the water temperature was +92° 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), Tåning (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 Tåning (1952:181-182), who states: +"Shock treatment produced by especially great changes in temperature +(_c._ 10-14° 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 Tåning'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° N to 66° N, and +longitude 89° W to 123° 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° N to 48° N, and longitude 97° W to 100° 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. 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Soc., 27(2):169-193. + +THOMPSON, ERNEST SETON. + + 1898. A list of fishes known to occur in Manitoba. Forest and Stream. + 51(11):214. + +UNDERHILL, JAMES C. + + 1959. Fishes of the Vermillion River, South Dakota. Proc. South + Dakota Acad. Sci., 38:96-102. + +VLADYKOV, VADIM D. + + 1934. Environmental and taxonomic characters of fishes. Trans. Royal + Canadian Inst., 20(43):99-140. + +WALTERS, VLADIMIR. + + 1955. Fishes of western Arctic America and eastern Arctic Siberia. + Bull. Amer. Mus. Nat. Hist., 106(5):255-368. + +WEISEL, GEORGE F. + + 1955. Variation in the number of fin rays of two cyprinid fishes + correlated with natural water temperatures. Ecology. 36(1):1-6. + +WYNNE-EDWARDS, V. C. + + 1952. 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 "Société Géologique"] + + + + + +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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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: ISO-8859-1 + +*** 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 + + + + + + +</pre> + + +<div class="spacer"></div> + +<div> + <span class="pagenum invisible"> + <a name="Page_323" id="Page_323">[pg 323]</a> + </span> +</div> + +<h4><span class="smcap">University of Kansas Publications<br /> +Museum of Natural History</span></h4> + +<h4>Volume 13, No. 7, pp. 323-348, pls. 21-24, 2 figs.<br /> +February 10, 1961</h4> + +<h1>Geographic Variation<br /> +In the North American Cyprinid Fish,<br /> +Hybopsis gracilis</h1> + +<h2><small>BY</small><br /> +LEONARD J. OLUND AND FRANK B. CROSS</h2> + +<h4><span class="smcap">University of Kansas<br /> +Lawrence</span><br /> +1961</h4> + +<hr class="cb" /> + +<div> + <span class="pagenum invisible"> + <a name="Page_324" id="Page_324">[pg 324]</a> + </span> +</div> + +<h4><span class="smcap">University of Kansas Publications, +Museum of Natural History</span></h4> + +<h4>Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br /> +Robert W. Wilson</h4> + +<h4>Volume 13, No. 7, pp. 323-348, pls. 21-24, 2 figs.<br /> +Published February 10, 1961</h4> + + +<h4><span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas</h4> + +<h5>PRINTED IN<br /> +THE STATE PRINTING PLANT<br /> +TOPEKA, KANSAS<br /> +1961</h5> + +<p class="center"> +<img src="images/ukp-logo.png" width="100" height="35" +alt="Allied Printing Trades Council Topeka" title="" /> +</p> + +<h5>28-5871</h5> + + +<div> + <span class="pagenum"> + <a name="Page_325" id="Page_325">[pg 325]</a> + </span> +</div> + +<hr class="cb" /> + +<h2>Geographic Variation +In the North American Cyprinid Fish, +Hybopsis gracilis</h2> + +<h3>BY<br /> +LEONARD J. OLUND AND FRANK B. CROSS</h3> + +<h2>CONTENTS</h2> + +<ul class="TOC"> + <li> + <span class="ralign">PAGE</span></li> + <li><span class="smcap"><a href="#INT">Introduction</a></span> + <span class="ralign"> + <a href="#Page_325">325</a></span></li> + <li><span class="smcap"><a href="#MET_MAT_ACK">Methods, Materials, and Acknowledgments</a></span> + <span class="ralign"> + <a href="#Page_326">326</a></span></li> + <li><span class="smcap"><a href="#DES_SPE">Description of the Species Hybopsis gracilis</a></span> + <span class="ralign"> + <a href="#Page_327">327</a></span></li> + <li class="section"><a href="#H_G_GRA"><i>Hybopsis gracilis gracilis</i></a> + <span class="ralign"> + <a href="#Page_328">328</a></span></li> + <li class="section"><a href="#H_G_GUL"><i>Hybopsis gracilis gulonella</i></a> + <span class="ralign"> + <a href="#Page_330">330</a></span></li> + <li><span class="smcap"><a href="#INT_VAR">Intraspecific Variation</a></span> + <span class="ralign"> + <a href="#Page_333">333</a></span></li> + <li><span class="smcap"><a href="#NAT_HIS">Natural History</a></span> + <span class="ralign"> + <a href="#Page_334">334</a></span></li> + <li class="section"><a href="#HAB"><i>Habitat</i></a> + <span class="ralign"> + <a href="#Page_334">334</a></span></li> + <li class="section"><a href="#AS_SPE"><i>Associated Species</i></a> + <span class="ralign"> + <a href="#Page_336">336</a></span></li> + <li class="section"><a href="#FOO"><i>Food</i></a> + <span class="ralign"> + <a href="#Page_338">338</a></span></li> + <li class="section"><a href="#SPA_SEA"><i>Spawning Season</i></a> + <span class="ralign"> + <a href="#Page_338">338</a></span></li> + <li><span class="smcap"><a href="#DIS">Discussion</a></span> + <span class="ralign"> + <a href="#Page_340">340</a></span></li> + <li><span class="smcap"><a href="#LIT_CIT">Literature Cited</a></span> + <span class="ralign"> + <a href="#Page_343">343</a></span></li> +</ul> + + + + +<h2><a name="INT" id="INT"></a>INTRODUCTION</h2> + + +<p>The flathead chub, <i>Hybopsis gracilis</i> (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, <i>H. gracilis</i> 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 +prob<span class="pagenum"><a name="Page_326" id="Page_326">[pg 326]</a></span>ably +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).</p> + +<p>Six names apply to the flathead chub, the earliest of which is +<i>Cyprinus gracilis</i> 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, <i>Platygobio</i> Gill +has been recognized as the appropriate generic name for the flathead +chub, but Bailey (1951:192) places <i>Platygobio</i> and other +nominal genera of barbeled minnows having short guts, protractile +premaxillae, and four teeth (primary row) in the single genus +<i>Hybopsis</i> (Agassiz, 1854). Strangely, the orthotype of <i>Hybopsis</i>, +<i>H. gracilis</i> Agassiz, is a junior synonym of <i>H. amblops</i> (Rafinesque) +(Hubbs and Ortenburger, 1929b:66) and is a younger name than +<i>C. gracilis</i> Richardson.</p> + +<p>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.</p> + + + + +<h2><a name="MET_MAT_ACK" id="MET_MAT_ACK"></a> +METHODS, MATERIALS AND ACKNOWLEDGMENTS</h2> + + +<div class="descrip"> +<p>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, +<span class="pagenum"><a name="Page_327" id="Page_327">[pg 327]</a></span> +head-width, snout-length, postorbital length of head, length of orbit, interorbital +width, length of upper jaw and width of gape.</p> + +<p>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.</p> + +<p>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.</p> + +<p>Specimens were obtained from the following institutions: University of +Alberta (abbreviated AU 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.</p> + +<p>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.</p> +</div> + + + + +<h2><a name="DES_SPE" id="DES_SPE"></a> +DESCRIPTION OF THE SPECIES</h2> + + +<p class="section"><b>Hybopsis gracilis</b> (Richardson)</p> + +<p class="center">Flathead Chub</p> + +<p class="center">(Synonymy under accounts of subspecies)</p> + +<div class="descrip"> +<p><i>Description.</i>--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.</p> + +<p>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.</p> + +<p>Color brown or olivaceous dorsally, silver or creamy white ventrally, without +distinctive markings; dusky lateral band evident in preserved specimens.</p> + +<p><span class="pagenum"><a name="Page_328" id="Page_328">[pg 328]</a></span> +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.</p> + +<p>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.</p> +</div> + + +<p class="section"><a name="H_G_GRA" id="H_G_GRA"></a> +<b>Hybopsis gracilis gracilis</b> (Richardson)</p> + +<p class="center">(<a href="#PL22">Plate 22</a>)</p> + +<div class="species"> +<p><i>Cyprinus (Leuciscus) gracilis</i> Richardson, 1836:120 and Pl. 78 (original +description; Saskatchewan R. at Carlton House).</p> + +<p><i>Coregonus angusticeps</i> Cuvier and Valenciennes, 1848:534 (original description; +Saskatchewan R.).</p> + +<p><i>Pogonichthys communis</i> 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.).</p> + +<p><i>Platygobio gracilis</i>, 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 <i>Hybopsis meeki</i> Jordan and Evermann, not <i>H. gracilis</i>); +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, <i>et al.</i>, 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).</p> + +<p><i>Platygobio pallidus</i> 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).</p> + +<p><i>Platygobio gracilis communis</i>, Simon, 1946:71 (in part; characters; food; +habitat; spawning); Moore, 1950:87 (habitat; sense organs).</p> + +<p><span class="pagenum"><a name="Page_329" id="Page_329">[pg 329]</a></span> +<i>Hybopsis gracilis communis</i>, 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.).</p> + +<p><i>Hybopsis gracilis</i>, 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).</p> +</div> + +<div class="descrip"> +<p><i>Diagnosis.</i>--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.</p> + +<p><i>Other characters.</i>--Circumference scale-rows 31-42; predorsal scale-rows 20-29; +size large, as much as 246 mm standard length (see Fig. 1 of <a href="#PL24">Pl. 24</a>); 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.</p> + +<p><i>Range</i> (<a href="#PL21">Plate 21</a>).--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 <i>H. g. gulonella</i> in upper +Missouri Basin and lower parts of major tributaries to Missouri River in Nebraska +and Kansas.</p> + +<p><i>Specimens examined.</i>--Below are listed museum numbers, number of specimens +(in parentheses), localities, and year of collection. Collections marked +with asterisk (*) are intergrades more closely resembling <i>H. g. gracilis</i> than +<i>H. g. gulonella</i>. Records from literature are cited in the synonymy.</p> + +<p><span class="smcap">Alberta:</span> 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.</p> + +<p><span class="smcap">Arkansas:</span> UMMZ 128573 (5), Mississippi Co., Mississippi R., 1939.</p> + +<p><span class="smcap">Illinois:</span> UMMZ 134799 (146), Mississippi R. at Grand Tower, 1936; +UMMZ 147045 (8), Mississippi R. at Cairo, 1944.</p> + +<p><span class="smcap">Kansas:</span> 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.</p> + +<p><span class="smcap">Manitoba:</span> 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.</p> + +<p><span class="smcap">Missouri:</span> UMMZ 147126 (130), Mississippi R. at Cliff Cave, 1944.</p> + +<p><span class="smcap">Montana:</span> * 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 +<span class="pagenum"><a name="Page_330" id="Page_330">[pg 330]</a></span> +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.</p> + +<p><span class="smcap">Nebraska:</span> * 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.</p> + +<p><span class="smcap">Northwest Territory:</span> ROMZ 13627 (1), Great Slave Lake, no date; +ROMZ 13628 (1), Great Slave Lake, no date.</p> + +<p><span class="smcap">Saskatchewan:</span> * 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.</p> + +<p><span class="smcap">South Dakota:</span> * 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.</p> + +<p><span class="smcap">Wyoming:</span> * WU 2073 (6), Washakie Co., Big Horn R. at Worland, 1956.</p> +</div> + + +<p class="section"><a name="H_G_GUL" id="H_G_GUL"></a> +<b>Hybopsis gracilis gulonella</b> (Cope)</p> + +<p class="center">(<a href="#PL23">Plate 23</a>)</p> + +<div class="species"> +<p><i>Pogonichthys communis</i> Girard, 1856:188 (in part; original description); +Girard, 1858:247 (in part; characters; synonymy); Cope and Yarrow, +1875:653 (characters; Pueblo, Colo.).</p> + +<p><i>Pogonichthys (Platygobio) gulonellus</i> Cope, 1864:277 (original description; +near Bridger's Pass, Wyo.).</p> + +<p><i>Platygobio gulonellus</i> 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]).</p> + +<p><i>Ceratichthys physignathus</i> Cope and Yarrow, 1875:651 (original description; +Arkansas R., Pueblo, Colo.).</p> + +<p><i>Platygobio communis</i>, Gill, 1876:408 (characters; Platte Valley; Green +River, Utah [the latter probably erroneous]).</p> + +<p><i>Couesius physignathus</i>, Jordan and Gilbert, 1882:219 (characters; synonymy; +Arkansas R., Pueblo, Colo.); Jordan, 1885:29 (records).</p> + +<p><i>Platygobio gracilis</i>, 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).</p> + +<p><span class="pagenum"><a name="Page_331" id="Page_331">[pg 331]</a></span> +<i>Platygobio physignathus</i>, 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.).</p> + +<p><i>Platygobio gracilis communis</i>, Simon, 1946:71 (in part; characters; food; +habitat; spawning).</p> + +<p><i>Platygobio gracilis gulonellus</i>, Simon, 1946:72 (characters; records from +Wyoming; Arkansas R.).</p> + +<p><i>Platygobio gracilis:</i> <i>communis</i> × <i>gulonellus</i>, Simon, 1946:92 (North Platte +R., Neb.-Wyo. line).</p> + +<p><i>Platygobio gracilis physignathus</i>, Moore, 1950:87 (habitat; sense organs).</p> + +<p><i>Hybopsis gracilis communis</i>, Beckman, 1952:50 (characters; food; habitat); +Cross, Dalquest and Lewis, 1955:222 (records from Texas).</p> + +<p><i>Hybopsis gracilis physignathus</i>, Beckman, 1952:50 (characters; habitat).</p> + +<p><i>Hybopsis gracilis</i>, 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.).</p> +</div> + +<div class="descrip"> +<p><i>Diagnosis.</i>--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.</p> + +<p><i>Other characters.</i>--Circumference scale-rows 30-40, slightly fewer than in +<i>H. g. gracilis</i>; 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 <a href="#PL24">Pl. 24</a>); +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 <i>H. g. gulonella</i>); +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 <i>H. g. gracilis</i>; +caudal peduncle length 17.1-22.7 per cent of standard length, essentially the +same in both subspecies.</p> + +<p>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 Böhlke 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.</p> + +<p>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 +<span class="pagenum"><a name="Page_332" id="Page_332">[pg 332]</a></span> +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.</p> + +<p>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 <i>P. gulonellus</i>, because +the flathead chub has not since been found in either of these streams. The +most likely collection site for <i>P. gulonellus</i> 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.</p> + +<p><i>Range</i> (<a href="#PL21">Plate 21</a>).--Upper mainstream and tributaries of Rio Grande, Pecos, +Arkansas and North Platte Rivers; isolated populations in tributaries of the +upper Missouri River.</p> + +<p><i>Specimens examined.</i>--Below are listed museum numbers, number of specimens +(in parentheses), localities and year of collection. Series marked by asterisks +(*) are intergrades tending toward <i>H. g. gulonella</i>. Literature reports +are cited in the synonymy.</p> + +<p><span class="smcap">Colorado:</span> 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.</p> + +<p><span class="smcap">Kansas:</span> 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.</p> + +<p><span class="smcap">Montana:</span> * MSC 1960 (8), Powder River Co., E. Fork of Powder R., +1957; MSC 2010 (64), Dawson Co., Redwater R., 1957.</p> + +<p><span class="smcap">Nebraska:</span> * 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.</p> + +<p><span class="smcap">New Mexico:</span> 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 +<span class="pagenum"><a name="Page_333" id="Page_333">[pg 333]</a></span> +Cr. at Sapello, 1937; UMMZ 133131 (7), Pecos R. 0.5 mi. N Santa Rosa, 1940; +UMMZ 133136 (1), Rio Grande at Albuquerque, 1940.</p> + +<p><span class="smcap">Oklahoma:</span> 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.</p> + +<p><span class="smcap">Texas:</span> KU 3409 (18), Hemphill Co., Canadian R. at town of Canadian, +1955.</p> + +<p><span class="smcap">Wyoming:</span> 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.</p> +</div> + + + + +<h2><a name="INT_VAR" id="INT_VAR"></a> +INTRASPECIFIC VARIATION</h2> + + +<p>Two subspecies of <i>H. gracilis</i> are recognized by us: one northern +and eastern, characteristically inhabiting large rivers (<i>H. g. gracilis</i>), +and one southern and western, characteristically inhabiting small +streams (<i>H. g. gulonella</i>). Other scientific names that have been +applied to this fish in the past are listed in the synonymy.</p> + +<p><i>H. g. gulonella</i> is a chubby, deep-bodied fish, whereas <i>H. g. gracilis</i> +is long and slender. The head of the creek subspecies is deeper +and longer than that of <i>H. g. gracilis</i>, being rounded anteriorly when +seen in sideview. The head of the large-river subspecies is acutely +wedge-shaped in profile. <i>H. g. gracilis</i> has a larger orbit than <i>H. g. +gulonella</i>. Fins of <i>H. g. gracilis</i> are more strongly falcate than those +of the other subspecies. <i>H. g. gracilis</i> 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 (<a href="#PL24">Plate 24</a>). 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 <i>H. g. gracilis</i> from the Saskatchewan River (several +localities) and in <i>H. g. gulonella</i> from Beaver Creek, Arkansas River +Drainage (KU 4769) are shown in <a href="#PL24">Plate 24</a>. Although values for +the largest specimens of <i>H. g. gracilis</i> are omitted from <a href="#PL24">Plate 24</a>, 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.</p> + + + + +<div><span class="pagenum"> +<a name="Page_334" id="Page_334">[pg 334]</a> +</span></div> +<h2><a name="NAT_HIS" id="NAT_HIS"></a> +NATURAL HISTORY</h2> + + +<p class="section"><a name="HAB" id="HAB"></a> +<i>Habitat</i></p> + +<p>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).</p> + +<div class="figcenter"> + <a href="images/fig1-full.png"> + <img src="images/fig1-th.png" width="800" height="474" alt="" title="" /> + </a> +</div> + +<div class="caption"> + <p><span class="smcap">Figure 1.</span> + Graphic analysis of lateral line scales, pectoral rays and + post-Weberian vertebrae in <i>Hybopsis gracilis</i>. 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. + 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.</p> +</div> + +<p><i>H. g. gracilis</i> is found in large rivers throughout its range, +occa<span class="pagenum"><a name="Page_335" id="Page_335">[pg 335]</a></span>sionally +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.</p> + +<p><i>H. g. gulonella</i> 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 +<span class="pagenum"><a name="Page_336" id="Page_336">[pg 336]</a></span> +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.</p> + +<p><i>H. g. gulonella</i> is found in warm-water streams, whereas <i>H. g. +gracilis</i> occurs in cooler water. The southwestern subspecies was +taken in August in the Mora River drainage at Sapello (temperatures +above 80° F.) but not at Mora (temperatures below 70° F.). +In the Purgatoire River, a thriving population was found where the +water temperature was 92° 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.</p> + + +<p class="section"><a name="AS_SPE" id="AS_SPE"></a> +<i>Associated Species</i></p> + +<div class="figcenter"> + <a href="images/fig2-full.png"> + <img src="images/fig2-th.png" width="800" height="309" alt="" title="" /> + </a> +</div> + +<div class="caption"> + <p><span class="smcap">Figure 2.</span> + Graphic analysis of head-depth, postorbital length of head and + predorsal length of <i>Hybopsis gracilis</i>, 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.</p> +</div> + +<p>In the Pecos and Arkansas basins, species commonly taken with +<i>H. g. gulonella</i> are <i>Catostomus commersonnii</i>, <i>Hybognathus placita</i>, +<span class="pagenum"><a name="Page_337" id="Page_337">[pg 337]</a></span> +<i>Notropis lutrensis lutrensis</i>, <i>Notropis stramineus missuriensis</i>, <i>Pimophales +promelas</i>, and <i>Campostoma anomalum plumbeum</i>. The +only spiny-rayed fishes that we have found with <i>H. g. gulonella</i> are +<i>Lepomis cyanellus</i> and <i>L. humilis</i>, both of which are scarce. Associates +of <i>H. g. gracilis</i> include the same species, plus other ostariophysan +fishes such as species of <i>Carpiodes</i>, <i>Ictiobus</i>, and silt-adapted +species of <i>Hybopsis</i> and <i>Notropis</i>.</p> + +<p>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 <i>H. gracilis</i> in that drainage. +The fauna of the South Platte includes <i>Catostomus catostomus</i>, <i>Semotilus +atromaculatus</i>, <i>Hybopsis biguttata</i>, <i>Hybognathus hankinsoni</i>, +<i>Notropis cornutus frontalis</i>, <i>Etheostoma nigrum</i> and <i>E. exile</i>--species +rarely if ever found with <i>H. gracilis</i>.</p> + +<p>Ecologically, <i>H. g. gulonella</i> seems to be the counterpart of <i>Semotilus +atromaculatus</i> in streams where the latter species is absent. +Observations of <i>H. g. gulonella</i> in the Purgatoire River indicated +<span class="pagenum"><a name="Page_338" id="Page_338">[pg 338]</a></span> +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.</p> + + +<p class="section"><a name="FOO" id="FOO"></a> +<i>Food</i></p> + +<p>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.</p> + +<p>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.)."</p> + + +<p class="section"><a name="SPA_SEA" id="SPA_SEA"></a> +<i>Spawning Season</i></p> + +<p>Specimens of <i>H. g. gulonella</i> that have been examined reach +sexual maturity at approximately 65 mm standard length. Most +specimens of <i>H. g. gracilis</i> less than 85 mm in standard length are +immature, but larger specimens probably are mature.</p> + +<p>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 <i>H. g. gracilis</i> or intergrades tending toward that subspecies.</p> + +<p>Specimens of <i>H. g. gulonella</i> 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 +<span class="pagenum"><a name="Page_340" id="Page_340">[pg 340]</a></span> +August 25, 1958, spawning seemingly had been completed, although +a few males still bore tubercles.</p> + +<p><span class="pagenum invisible"> +<a name="Page_339" id="Page_339">[pg 339]</a></span> +<span class="smcap">Table 1. Organisms Found in Stomachs of Hybopsis gracilis From +Various Locations, Expressed as Percentage of Total Volume.</span></p> + +<table summary="Table 1"> + <tr> + <th class="dbt bb"> </th> + <th class="dbt bb bl">S. Saskatchewan R., Clarkboro Ferry, Sask.</th> + <th class="dbt bb bl">B: Milk R., Alberta</th> + <th class="dbt bb bl">Missouri R., S. D.</th> + <th class="dbt bb bl">Missouri R., Neb.</th> + <th class="dbt bb bl">Arkansas R., Fremont Co., Colo.</th> + <th class="dbt bb bl">Arkansas R., Pueblo Co., Colo.</th> + <th class="dbt bb bl">Pecos R., San Miguel Co., N. M.</th></tr> + <tr> + <td class="id1">No. specimens examined</td> + <td class="bl">1</td> + <td class="bl">7</td> + <td class="bl">6</td> + <td class="bl">10</td> + <td class="bl">10</td> + <td class="bl">10</td> + <td class="bl">10</td></tr> + <tr> + <td class="bb id1">No. specimens containing food</td> + <td class="bb bl">1</td> + <td class="bb bl">6</td> + <td class="bb bl">1</td> + <td class="bb bl">2</td> + <td class="bb bl">1</td> + <td class="bb bl">3</td> + <td class="bb bl">7</td></tr> + <tr> + <td class="id1"><span class="smcap">Kind of Organism</span></td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id1">Aphasmidia</td> + <td class="bl">10.0</td> + <td class="bl">00.7</td> + <td class="bl">.....</td> + <td class="bl">03.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id1">Arthropoda</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id2">Araneae</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id3">Argiopidae</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">04.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id3">Theridiidae</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">04.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id2">Insecta</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id3">Ephemeroptera (nymph)</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id4">Baetidae</td> + <td class="bl">.....</td> + <td class="bl">05.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Heptagenidae</td> + <td class="bl">.....</td> + <td class="bl">08.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id3">Hemiptera</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id4">Corixidae</td> + <td class="bl">35.0</td> + <td class="bl">00.3</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id3">Hymenoptera</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id4">Formicidae</td> + <td class="bl">.....</td> + <td class="bl">21.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">60.0</td></tr> + <tr> + <td class="id3">Coleoptera</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id4">Staphylinidae</td> + <td class="bl">.....</td> + <td class="bl">01.7</td> + <td class="bl">07.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Scolytidae</td> + <td class="bl">.....</td> + <td class="bl">13.3</td> + <td class="bl">70.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Tenebrionidae</td> + <td class="bl">.....</td> + <td class="bl">05.7</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">70.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Carabidae</td> + <td class="bl">.....</td> + <td class="bl">05.7</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">01.0</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Curculionidae</td> + <td class="bl">.....</td> + <td class="bl">01.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Coccinellidae</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">09.0</td></tr> + <tr> + <td class="id3">Trichoptera (case)</td> + <td class="bl">.....</td> + <td class="bl">01.7</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id3">Diptera</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id4">Mymaridae</td> + <td class="bl">.....</td> + <td class="bl">00.3</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Empididae</td> + <td class="bl">.....</td> + <td class="bl">01.3</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Cecidomyiidae</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">04.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Trachinidae</td> + <td class="bl">.....</td> + <td class="bl">00.7</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Simulidae</td> + <td class="bl">.....</td> + <td class="bl">06.7</td> + <td class="bl">20.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Tabanidae</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">06.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Chironomidae</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">06.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Not identified to family</td> + <td class="bl">.....</td> + <td class="bl">01.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id3">Orthoptera</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id4">Locustidae</td> + <td class="bl">.....</td> + <td class="bl">07.7</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id4">Tettigoniidae</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">03.0</td> + <td class="bl">70.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">09.0</td></tr> + <tr> + <td class="id4">Tetrigidae</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">06.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id3">Homoptera</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id4">Fulgoridae</td> + <td class="bl">.....</td> + <td class="bl">05.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">01.0</td></tr> + <tr> + <td class="id3">Insect egg</td> + <td class="bl">.....</td> + <td class="bl">00.7</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id1">Plants</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id2">Cyanophyceae</td> + <td class="bl">.....</td> + <td class="bl">09.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">99.0</td> + <td class="bl">20.0</td></tr> + <tr> + <td class="id2">Cyperaceae</td> + <td class="bl">.....</td> + <td class="bl">02.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">01.0</td></tr> + <tr> + <td class="id2">Zannichellia palustris</td> + <td class="bl">.....</td> + <td class="bl">00.3</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id2">Vascular remains</td> + <td class="bl">55.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">27.0</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id1">Miscellaneous</td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td> + <td class="bl"> </td></tr> + <tr> + <td class="id2">Sand</td> + <td class="bl">.....</td> + <td class="bl">00.7</td> + <td class="bl"> </td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td> + <td class="bl">.....</td></tr> + <tr> + <td class="id2"> Pharyngeal tooth</td> + <td class="bb bl">.....</td> + <td class="bb bl">00.3</td> + <td class="bb bl">.....</td> + <td class="bb bl">.....</td> + <td class="bb bl">.....</td> + <td class="bb bl">.....</td> + <td class="bb bl">.....</td></tr> + <tr> + <td class="bb">Total (%)</td> + <td class="bb bl">100.0</td> + <td class="bb bl">99.8</td> + <td class="bb bl">100.0</td> + <td class="bb bl">100.0</td> + <td class="bb bl">100.0</td> + <td class="bb bl">100.0</td> + <td class="bb bl">100.0</td></tr> +</table> + +<p>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.</p> + +<p class="center"><a name="PL21" id="PL21"></a>PLATE 21</p> + +<div class="figcenter"> + <a href="images/plate-21-full.png"> + <img src="images/plate-21-th.png" width="600" height="772" alt="" title="" /></a> +</div> + +<div class="caption"> + <p>Distribution of collections examined.</p> +</div> + +<p class="center"><a name="PL22" id="PL22"></a>PLATE 22</p> + +<div class="figcenter"> + <a href="images/plate-22-full.jpg"> + <img src="images/plate-22-th.jpg" width="600" height="608" alt="" title="" /></a> +</div> + +<div class="caption"> + <p><i>Hybopsis gracilis gracilis.</i> Missouri River, Thurston County, northeast of Macy, + Nebraska. Largest specimen 87.5 mm standard length.</p> +</div> + +<p class="center"><a name="PL23" id="PL23"></a>PLATE 23</p> + +<div class="figcenter"> + <a href="images/plate-23-full.jpg"> + <img src="images/plate-23-th.jpg" width="600" height="785" alt="" title="" /></a> +</div> + +<div class="caption"> + <p><i>Hybopsis gracilis gulonella.</i> Pecos River, San Miguel County, 3 miles south + of town of Pecos, New Mexico. Largest specimen 91 mm standard length.</p> +</div> + +<p class="center"><a name="PL24" id="PL24"></a>PLATE 24</p> + +<div class="figcenter"> + <a href="images/plate-24-1-full.jpg"> + <img src="images/plate-24-1-th.jpg" width="600" height="291" alt="" title="" /></a> +</div> + +<div class="caption"> + <p><span class="smcap">Fig. 1.</span> Top: <i>Hybopsis gracilis gracilis</i>, + 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.</p> + + <p>Bottom: <i>Hybopsis gracilis gulonella</i>, 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.</p> +</div> + +<div class="figcenter"> + <a href="images/plate-24-2-full.png"> + <img src="images/plate-24-2-th.png" width="600" height="414" alt="" title="" /></a> +</div> + +<div class="caption"> + <p><span class="smcap">Fig. 2.</span> Regression of head-depth on + standard length in <i>Hybopsis gracilis gracilis</i> + from the Saskatchewan River, and in <i>H. g. gulonella</i> from Beaver Creek, + Arkansas River Drainage (KU 4769).</p> +</div> + + + + +<h2><a name="DIS" id="DIS"></a>DISCUSSION</h2> + + +<p><i>Hybopsis gracilis</i> 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, <i>H. g. gracilis</i>, 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 <i>H. g. gulonella</i> is +the oldest applicable name, is small, and has 1) a deep, chubby +body, 2) head convex in dorsal contour (less depressed than in <i>H. g. +gracilis</i>), 3) fins less falcate than in the latter subspecies, with the +dorsal and pelvic fins originating more posteriorly, and 4) fewer +<span class="pagenum"><a name="Page_341" id="Page_341">[pg 341]</a></span> +scales, vertebrae, and pectoral fin-rays than <i>H. g. gracilis</i>. 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.</p> + +<p>Variation in <i>H. gracilis</i>, as shown in the graphic analysis (Figs. 1 +and 2) and distribution map (<a href="#PL21">Plate 21</a>), presents two clines: a north-south +cline and a large-river to small-river (mainly east-west) cline. +The absence of <i>H. gracilis</i> 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 <i>H. g. gulonella</i> seems to be the ecological +equivalent of <i>Semotilus atromaculatus</i> in streams in which +<i>S. atromaculatus</i> is not found. <i>S. atromaculatus</i> 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 <i>H. gracilis</i> +tend to limit gene flow.</p> + +<p>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), Tåning (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 <i>H. gracilis</i>, 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 +<i>H. g. gracilis</i> and <i>H. g. gulonella</i>. To our knowledge, the most extreme +differences that have been induced by modification of temperature +are those reported for <i>Salmo trutta</i> by Tåning (1952:181-182), +who states: "Shock treatment produced by especially great +changes in temperature (<i>c.</i> 10-14° C), especially during the +super-<span class="pagenum"><a name="Page_342" id="Page_342">[pg 342]</a></span>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 <i>H. g. gracilis</i> and +<i>H. g. gulonella</i>. Although we cannot assume that the sensitivity of +the brown trout is the same as that of the flathead chub, the causative +conditions in Tåning'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 <i>H. gracilis</i>, 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.</p> + +<p>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 <i>H. g. gracilis</i> from <i>H. g. gulonella</i> 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 <i>H. g. gracilis</i>, 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.</p> + +<p>A remarkable effect of extreme parasitism in <i>H. gracilis</i> has been +described by Hubbs (1927). Very young chubs that harbored numerous +tapeworms (<i>Proteocephalus</i>) 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.</p> + +<p><span class="pagenum"><a name="Page_343" id="Page_343">[pg 343]</a></span> +<i>H. g. gracilis</i> occurs in three separate river systems (Mackenzie, +Saskatchewan, Missouri-Mississippi) from latitude 36° N to 66° N, +and longitude 89° W to 123° W. <i>H. g. gulonella</i> 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° N to 48° N, and longitude 97° W to +100° W.</p> + +<p>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 <i>H. g. gulonella</i> (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 <i>H. g. gulonella</i> 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. <i>H. g. gracilis</i> undoubtedly entered the Saskatchewan +and Mackenzie basins from the upper Missouri Basin, following +glacial retreat (Walters, 1955:347).</p> + + + + +<h2><a name="LIT_CIT" id="LIT_CIT"></a> +LITERATURE CITED</h2> + + +<p><span class="smcap">Agassiz, L.</span></p> + +<div class="biblio"> +<p>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.</p> +</div> + +<p><span class="smcap">Bailey, Reeve M.</span></p> + +<div class="biblio"> +<p>1951. A check-list of the fishes of Iowa, with keys for identification. <i>In</i> +Iowa fish and fishing, by James R. Harlan and Everett B. Speaker. +1st edition. Iowa Cons. Comm. 185-237.</p> + +<span class="pagenum"><a name="Page_344" id="Page_344">[pg 344]</a></span> +<p>1956. A check-list of the fishes of Iowa, with keys for identification. <i>In</i> +Iowa fish and fishing, by James R. Harlan and Everett B. Speaker. +3rd edition. Iowa Cons. Comm. 325-377.</p> +</div> + +<p><span class="smcap">Beckman, William C.</span></p> + +<div class="biblio"> +<p>1952. Guide to the fishes of Colorado. Univ. Colorado Mus., 11:1-110.</p> +</div> + +<p><span class="smcap">Bryant, Francis T.</span></p> + +<div class="biblio"> +<p>1858. Report of Francis T. Bryant to Col. J. J. Abert, Chief, Corps +Top. Engs., USA. <i>In</i> Executive Documents of the Senate of the +U. S., First Session, 35th Congress, and Special Session of 1858. +3:455-481.</p> +</div> + +<p><span class="smcap">Churchill, Edward P.</span>, and <span class="smcap">William H. Over</span>.</p> + +<div class="biblio"> +<p>1933. Fishes of South Dakota. South Dakota Dept. Game and Fish. +1-87.</p> +</div> + +<p><span class="smcap">Cleary, Robert E.</span></p> + +<div class="biblio"> +<p>1956. The distribution of the fishes of Iowa. <i>In</i> Iowa fish and fishing, by +James R. Harlan and Everett B. Speaker. 3rd edition. Iowa +Cons. Comm. 267-324.</p> +</div> + +<p><span class="smcap">Clemens, W. A.</span>, <span class="smcap">A. MacDonald</span>, <span class="smcap">H. McAllister</span>, <span class="smcap">A. Mansfield</span> and <span class="smcap">D. Rawson</span>.</p> + +<div class="biblio"> +<p>1947. The fishes of Saskatchewan. <i>In</i> Report of the Royal Commission +on the fisheries of the province of Saskatchewan. 1-131.</p> +</div> + +<p><span class="smcap">Cockerell, Theodora D. A.</span></p> + +<div class="biblio"> +<p>1927. Zoology of Colorado. Univ. Colorado, Boulder. 1-262.</p> +</div> + +<p><span class="smcap">Cope, E. D.</span></p> + +<div class="biblio"> +<p>1864. Partial catalogue of the cold-blooded Vertebrata of Michigan. Proc. +Acad. Nat. Sci. Philadelphia, part 1:276-285.</p> + +<p>1865. Partial catalogue of the cold-blooded Vertebrata of Michigan. Proc. +Acad. Nat. Sci. Philadelphia, part 2:78-88.</p> + +<p>1879. A contribution to the zoology of Montana. Amer. Nat., 13:432-441.</p> +</div> + +<p><span class="smcap">Cope, E. D.</span>, and <span class="smcap">H. C. Yarrow</span>.</p> + +<div class="biblio"> +<p>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. <i>In</i> 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.</p> +</div> + +<p><span class="smcap">Cragin, F. W.</span></p> + +<div class="biblio"> +<p>1885. Preliminary list of Kansas fishes. Bull. Washburn Lab. Nat. Hist., +1(3): 105-111.</p> +</div> + +<p><span class="smcap">Cross, Frank B.</span>, <span class="smcap">Walter W. Dalquest</span> and <span class="smcap">Leo Lewis</span>.</p> + +<div class="biblio"> +<p>1955. First records from Texas of <i>Hybopsis gracilis</i> and <i>Notropis girardi</i>, +with comments on geographic variation of the latter. Texas Jour. +Sci., 7(2):222-226.</p> +</div> + +<p><span class="smcap">Cuvier, M. le B<sup>on</sup></span> and <span class="smcap">M. A. Valenciennes</span>.</p> + +<div class="biblio"> +<p>1848. Histoire naturelle des poissons. Libraire de la Société Géologique +de France, Paris, 21:1-536.</p> +</div> + +<p><span class="smcap">Dymond, J. R.</span></p> + +<div class="biblio"> +<p>1947. A list of the freshwater fish of Canada east of the Rocky Mountains, +with keys. Royal Ontario Mus. 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Govnt. +Printing Office, Washington. 325-429.</p> +</div> + +<p><span class="smcap">Evermann, Barton Warren</span>, and <span class="smcap">Edmund Lee Goldsborough</span>.</p> + +<div class="biblio"> +<p>1907. A check list of the freshwater fishes of Canada. Proc. Biol. Soc. +Washington. 20:89-120.</p> +</div> + +<p><span class="smcap">Forbes, Stephen Alfred</span>, and <span class="smcap">Robert Earl Richardson</span>.</p> + +<div class="biblio"> +<p>1920. The fishes of Illinois. Illinois Nat. Hist. Survey. 1-357.</p> +</div> + +<p><span class="smcap">Frye, J. C.</span>, and <span class="smcap">A. B. Leonard</span>.</p> + +<div class="biblio"> +<p>1952. Pleistocene geology of Kansas. Kansas Geol. Survey. 99:1-230.</p> +</div> + +<p><span class="smcap">Gilbert, Chas. H.</span></p> + +<div class="biblio"> +<p>1885. Second series of notes on the fishes of Kansas. Bull. Washburn +Lab. Nat. Hist., 1(3):97-99.</p> +</div> + +<p><span class="smcap">Gill, Theodore.</span></p> + +<div class="biblio"> +<p>1876. Report on ichthyology. <i>In</i> report of explorations across the Great +Basin of the territory of Utah in 1859, by Captain J. H. Simpson. +Appendix L. 385-431.</p> +</div> + +<p><span class="smcap">Gerard, Charles.</span></p> + +<div class="biblio"> +<p>1856. Researches upon the cyprinoid fishes inhabitating the fresh waters +of the United States of America, west of the Mississippi Valley, +from specimens in the museum of the Smithsonian Institution. +Proc. Acad. Nat. Sci. Philadelphia, 8:165-213.</p> + +<p>1858. Fishes. <i>In</i> Explorations and surveys for a railroad route from the +Mississippi River to the Pacific Ocean. War Dept., Washington. +1-368.</p> +</div> + +<p><span class="smcap">Graham, I. D.</span></p> + +<div class="biblio"> +<p>1885. Preliminary list of Kansas fishes. Trans. Kansas Acad. 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Sci., Arts and Letters, 26(1940):229-239. + +1951. <i>Notropis amnis</i>, a new cyprinid fish of the Mississippi fauna, with +two subspecies. Occ. Papers Mus. Zool., Univ. Michigan. 532:1-30.</p> +</div> + +<p><span class="smcap">Hubbs, Carl L.</span>, and <span class="smcap">Karl F. Lagler</span>.</p> + +<div class="biblio"> +<p>1958. Fishes of the Great Lakes region. Cranbrook Inst. Sci., 26:1-213.</p> +</div> + +<p><span class="smcap">Hubbs, Carl L.</span>, and <span class="smcap">A. I. Ortenburger</span>.</p> + +<div class="biblio"> +<p>1929a. Further notes on the fishes of Oklahoma, with descriptions of new +species of Cyprinidae. Publ. Univ. Oklahoma Biol. Sur., 1(2):17-44. + +1929b. Fishes collected in Oklahoma and Arkansas in 1927. Publ. Univ. +Oklahoma Biol. Sur., 1(3):45-112.</p> +</div> + +<p><span class="smcap">Jordan, David Starr.</span></p> + +<div class="biblio"> +<p>1885. A catalogue of the fishes known to inhabit the waters of North +America, north of the Tropic of Cancer. Annual report of the Commissioner +of Fish and Fisheries for 1884. Govnt. Printing Office, +Washington. 1-185. + +1929. Manual of the vertebrate animals of the northeastern United States, +inclusive of marine species. World Book Company, N. Y. 1-446.</p> +</div> + +<p><span class="smcap">Jordan, David Starr</span>, and <span class="smcap">Barton Warren Evermann</span>.</p> + +<div class="biblio"> +<p>1896. The fishes of North and Middle America: a descriptive catalogue +of the species of fish-like vertebrates found in the waters of North +America, north of the Isthmus of Panama. Part 1. Bull. U. S. Nat. +Mus., 47:1-1240.</p> +</div> + +<p><span class="smcap">Jordan, David Starr</span>, <span class="smcap">Barton Warren Evermann</span>, and <span class="smcap">Howard Walton +Clark</span>.</p> + +<div class="biblio"> +<p>1930. Check list of the fishes and fish-like vertebrates of North America +north of the northern boundary of Venezuela and Colombia. Report +of the United States Commissioner of Fisheries for the fiscal year +1928, with appendices. Part 2. Govnt. Printing Office, Washington. +1-670.</p> +</div> + +<p><span class="smcap">Jordan, David Starr</span>, and <span class="smcap">Charles H. Gilbert</span>.</p> + +<div class="biblio"> +<p>1882. Synopsis of the fishes of North America. Bull. U. S. Nat. Mus., +16:1-1018.</p> +</div> + +<p><span class="smcap">Jordan, David Starr</span>, and <span class="smcap">Seth E. Meek</span>.</p> + +<div class="biblio"> +<p>1886. List of fishes collected in Iowa and Missouri in August, 1884, with +descriptions of three new species. Proc. U. S. Nat. Mus., 8:1-17.</p> +</div> + +<p><span class="pagenum"><a name="Page_347" id="Page_347">[pg 347]</a></span> +<span class="smcap">Keleher, J. J.</span></p> + +<div class="biblio"> +<p>1956. The northern limits of distribution in Manitoba for cyprinid fishes. +Canadian Jour. Zool., 34(4):263-266.</p> +</div> + +<p><span class="smcap">Keleher, J. J.</span>, and <span class="smcap">B. Kooyman</span>.</p> + +<div class="biblio"> +<p>1957. Supplement to Hinks' "The fishes of Manitoba." Dept. Mines and +Nat. Res., Province of Manitoba. 103-117.</p> +</div> + +<p><span class="smcap">Koster, William J.</span></p> + +<div class="biblio"> +<p>1957. Guide to the fishes of New Mexico. Univ. New Mexico Press, in +cooperation with the New Mexico Dept. of Game and Fish, Albuquerque, +New Mexico. 1-116.</p> +</div> + +<p><span class="smcap">Lindsey, C. C.</span></p> + +<div class="biblio"> +<p>1956. Distribution and taxonomy of fishes in the Mackenzie drainage of +British Columbia. Jour. Fish. Res. Bd. Canada. 13(6):759-789. + +1957. Possible effects of water diversions on fish distribution in British +Columbia. Jour. Fish. Res. Bd. Canada. 14(4):651-668.</p> +</div> + +<p><span class="smcap">Martin, W. R.</span></p> + +<div class="biblio"> +<p>1949. The mechanics of environmental control of body form in fishes. Pub. +Ontario Fish. Res. Lab., 70:1-91.</p> +</div> + +<p><span class="smcap">Meek, Seth E.</span></p> + +<div class="biblio"> +<p>1892. A report upon fishes of Iowa, based upon observations and collections +made during 1889, 1890, and 1891. Bull. U. S. Fish Comm., +10:217-248. + +1895. Notes on the fishes of western Iowa and eastern Nebraska. Bull. +U. S. Fish Comm., 14:133-138.</p> +</div> + +<p><span class="smcap">Miller, R. B.</span>, and <span class="smcap">M. J. Paetz</span>.</p> + +<div class="biblio"> +<p>1953. Preliminary biological surveys of Alberta watersheds. 1950-1952. +Govmt. Province of Alberta, Edmonton, Alberta. 2:1-114.</p> +</div> + +<p><span class="smcap">Moore, George A.</span></p> + +<div class="biblio"> +<p>1950. The cutaneous sense organs of barbeled minnows adapted to life in +the muddy waters of the Great Plains region. Trans. Amer. Micro. +Soc., 69(1):69-95. + +1957. Fishes. <i>In</i> Vertebrates of the United States, by W. F. Blair, A. P. +Blair, P. Brodkorb, F. R. Cagle and G. A. Moore. McGraw-Hill +Book Company, Inc., New York, New York, 30-210.</p> +</div> + +<p><span class="smcap">O'Donnel, D. John.</span></p> + +<div class="biblio"> +<p>1935. Annotated list of the fishes of Illinois. Illinois Nat. Hist. Sur. Bull., +20(5):473-500.</p> +</div> + +<p><span class="smcap">Ortenburger, A. I.</span>, and <span class="smcap">Carl L. Hubbs</span>.</p> + +<div class="biblio"> +<p>1927. A report on the fishes of Oklahoma, with descriptions of new genera +and species. Oklahoma Acad. Sci., 6(1926):123-141.</p> +</div> + +<p><span class="smcap">Personius, Robert G.</span>, and <span class="smcap">Samuel Eddy</span>.</p> + +<div class="biblio"> +<p>1955. Fishes of the Little Missouri River. Copeia. 1:41-43.</p> +</div> + +<p><span class="smcap">Rawson, D. S.</span></p> + +<div class="biblio"> +<p>1951. Studies of the fish of Great Slave Lake. Jour. Fish. Res. Bd. +Canada. 8(4):207-240.</p> +</div> + +<p><span class="smcap">Richardson, John.</span></p> + +<div class="biblio"> +<p>1836. Fauna Boreali-Americana. Part 3, the fish. Richard Bentley, New +Burlington Street, London. 1-327.</p> +</div> + +<p><span class="pagenum"><a name="Page_348" id="Page_348">[pg 348]</a></span> +<span class="smcap">Schultz, Leonard P.</span></p> + +<div class="biblio"> +<p>1927. Temperature-controlled variation in the golden shiner, <i>Notemigonus +crysoleucas</i>. Proc. Michigan Acad. Sci., Arts and Letters. 6:417-432.</p> +</div> + +<p><span class="smcap">Scott, W. B.</span></p> + +<div class="biblio"> +<p>1958. A checklist of the freshwater fishes of Canada and Alaska. Royal +Ontario Mus., 1-30.</p> +</div> + +<p><span class="smcap">Shoemaker, Hurst H.</span>, <span class="smcap">Quentin H. Pickering</span> and <span class="smcap">Leonard Durham</span>.</p> + +<div class="biblio"> +<p>1951. The occurrence of the flathead chub, <i>Platygobio gracilis</i>, in Tennessee. +Jour. Tennessee Acad. Sci., 24(1):84.</p> +</div> + +<p><span class="smcap">Simon, James R.</span></p> + +<div class="biblio"> +<p>1946. Wyoming fishes. Wyoming Game and Fish Dept., 1-129.</p> +</div> + +<p><span class="smcap">Slastenenko, E. P.</span></p> + +<div class="biblio"> +<p>1958. The distribution of freshwater fishes in the provinces and main +water basins of Canada. Bull. Shevchenko Sci. Soc., 1(6):1-11.</p> +</div> + +<p><span class="smcap">Smith, C. Lavett.</span></p> + +<div class="biblio"> +<p>1958. Additional Pleistocene fishes from Kansas and Oklahoma. Copeia. +3:176-180.</p> +</div> + +<p><span class="smcap">Suckley, George.</span></p> + +<div class="biblio"> +<p>1860. Report upon the fishes collected on the Pacific Railroad Survey. +Pacific Railroad Report. 12(3):307-368.</p> +</div> + +<p><span class="smcap">Tåning, A. Vedel.</span></p> + +<div class="biblio"> +<p>1952. Experimental study of meristic characters in fishes. Biol. Rev. Cambridge +Phil. Soc., 27(2):169-193.</p> +</div> + +<p><span class="smcap">Thompson, Ernest Seton.</span></p> + +<div class="biblio"> +<p>1898. A list of fishes known to occur in Manitoba. Forest and Stream. +51(11):214.</p> +</div> + +<p><span class="smcap">Underhill, James C.</span></p> + +<div class="biblio"> +<p>1959. Fishes of the Vermillion River, South Dakota. Proc. South Dakota +Acad. Sci., 38:96-102.</p> +</div> + +<p><span class="smcap">Vladykov, Vadim D.</span></p> + +<div class="biblio"> +<p>1934. Environmental and taxonomic characters of fishes. Trans. Royal +Canadian Inst., 20(43):99-140.</p> +</div> + +<p><span class="smcap">Walters, Vladimir.</span></p> + +<div class="biblio"> +<p>1955. Fishes of western Arctic America and eastern Arctic Siberia. Bull. +Amer. Mus. Nat. Hist., 106(5):255-368.</p> +</div> + +<p><span class="smcap">Weisel, George F.</span></p> + +<div class="biblio"> +<p>1955. Variation in the number of fin rays of two cyprinid fishes correlated +with natural water temperatures. Ecology. 36(1):1-6.</p> +</div> + +<p><span class="smcap">Wynne-Edwards, V. C.</span></p> + +<div class="biblio"> +<p>1952. Freshwater vertebrates of the Arctic and Subarctic. Bull. Fish. +Res. Bd. Canada. 94:1-28.</p> +</div> + + +<p><i>Transmitted November 8, 1960.</i></p> + +<p class="center">□<br />28-5871</p> + +<div class="tnote"> + <h3>Transcriber's Note</h3> + + <p>The following changes have been made to the original text:</p> + + <ul> + <li><a href="#Page_325">Table of Contents:</a> + page number of "Food" and "Spawning Season" changed from 339 to 338</li> + <li><a href="#Page_327">Page 327:</a> + "abbreviated AU" changed to "abbreviated UA"</li> + <li><a href="#Page_344">Page 344:</a> + "Societe Geologique" changed to "Société Géologique"</li> + </ul> +</div> + +<div class="spacer"></div> + + + + + + + + +<pre> + + + + + +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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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. 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Philadelphia, 8:165-213. + + 1858. Fishes. _In_ Explorations and surveys for a railroad route from + the Mississippi River to the Pacific Ocean. War Dept., + Washington. 1-368. + +GRAHAM, I. D. + + 1885. Preliminary list of Kansas fishes. Trans. Kansas Acad. Sci., + 9:69-78. + +HANKINSON, THOMAS L. + + 1929. Fishes of North Dakota. Papers of the Michigan Acad. Sci., Arts + and Letters., 10:439-460. + +HARLAN, JAMES R., and EVERETT B. SPEAKER. + + 1951. Iowa fish and fishing. Iowa Cons. Comm. 1st edition. 1-237. + + 1956. Iowa fish and fishing. Iowa Cons. Comm. 3rd edition. 1-377. + +HARRISON, HARRY M., and EVERETT B. SPEAKER. + + 1954. An annotated list of the fishes in the streams tributary to the + Missouri River in Iowa. Proc. Iowa Acad. Sci., 61:511-523. + +HART, J. S. + + 1952. Geographic variations of some physiological and morphological + characters in certain freshwater fishes. Pub. Ontario Fish. + Res. Lab., 72:1-79. + +HINKS, DAVID. + + 1943. The fishes of Manitoba. Dept. 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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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