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Chrosomus Erythrogaster Andc. Eos (Osteichthyes: Cyprinidae) Taxonomy, Distribution) Ecology a Thesis Submitted to the Faculty O

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Chrosomus Erythrogaster Andc. Eos (Osteichthyes: Cyprinidae) Taxonomy, Distribution) Ecology a Thesis Submitted to the Faculty O CHROSOMUS ERYTHROGASTER AND C. EOS (OSTEICHTHYES: CYPRINIDAE) TAXONOMY, DISTRIBUTION) ECOLOGY A THESIS SUBMITTED TO THE FACULTY OF THE GRADUATE SCHOOL OF THE UNIVERSITY OF MINNESOTA By Gary Lee Phillips IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY Pegree Granted June, 1968 FRONTISPIECE. Male Chrosomus erythrogaster in breeding color, headwaters of the Zumbro River, Dodge County, Minnesota, 4 June 1966. Photograph by Professor David J. Merrell of the University of Minnesota. 47?-a•4 V gir irck 4r4.4- 1,1! IL .1, 74ko2,4,944,40tgrAt skr#9 4.e4 riff4eotilired‘ ik tit "ital.:A-To 4-v.w.r*:ez••01.%. '.or 44# 14 46#41bie. "v1441t..4frw.P1)4iiriiitalAttt.44- Aiihr4titeec --N. 1 4r40•4-v,400..orioggit kf)f 4y 4:11 to_ r •ArPV .1 1 "11(4% tk eat'n'ik\Nthl haf ilif -7b111,6 10t 11*4 * A Aver44, wr. • 4‘4041:Nr 0141 -at 1,10,71mr--,• 4 TABLE OF CONTENTS INTRODUCTION ACKNOWLEDGMENTS SYNONYMY AND NOMENCLATURE METHODS AND MATERIALS DISTRIBUTION 23 Geographical Distribution ................... 23 Ecological Distribution ......,....•.....,.. 24 Distribution in Minnesota ............. 27 VARIATION 38 Reliability of Measurements •.*****••••••** 4 • * 38 Sexual Variation •.. 53 Ontogenetic Variation •••• • • • • • •••• 61 Geographical Variation .................. ft ft. 72 Interpopulation Mean Character Differences • • * 77 Anomalies 83 REPRODUCTION 86 Schooling BehaviOr, ....................- 86 , 000,. W.4,41 , 87 .Spawning ,Behavior. , ,10041.4100 .......„......... 00 90 Breeding Color .. Breeding Tubercle ......................,. 93 Sex Ratios ... ............... ..,. 97 Sexual cycles ' , .-•,................ ... 99 ft 99 Fecundity ... ft ft ft ft 0 ft S OlkOodt*40o,OWOOsoo•O*00-Ito , 41,* 111: Hybridization 40. 0.41400**************0.0 DIET 1.28 The Digestive Tract ...................... 128 Composition of Diet .• .. • •. • •.... • • •. • • •. • 130 Selection of Diet ftaftftftftftftftftftftft............... 142 Feeding Behavior .....• ........ ..... ....... 148 • • 151 Daily Cycle of Activity .• 6 * 000004611,44.10•••** Effects on Algal Growth .... 0000, 04,0. ..... 159 FAUNAL ASSOCIATIONS 165 Syntopic Fish Species ....................... 165 Predators and'Enemies ....................... 169 MO 0 170 Parasites ....... 0.**0•1100oWil ***W..** SUMMARY • 174 LITERATURE CITED 178 APPENDIX 193 INTRODUCTION Four species of minnows are generally recognized within the North American Genus Chrosomus (Osteichthyes: Cyprinidae): These are C. eos (northern redbelly dace), C. erythrogaster (southern redbelly dace C. neogaeus (finescale dace), and C. oreas (mountain redbelly dace). Chrosomus eos has been considered a subspecies of C. erythrogaster by some workers (see Jordan and Evermann, 1896:210; Fowler, 1909:520; Legendre, 1952:xi). All of these fishes except C. oreas occur in Minnesota. The taxonomic, distributional, and ecological relationships of the northern and southern redbelly dace were investigated in the present study. Relegation of these forms to two species is upheld. Their distributions in Minnesota are summarized, chiefly on the basis of collections available at the University of Minnesota. Diet and reproduction were emphasized in the ecological phase of the study, in which C. erythrogaster was studied in detail and compared, when possible, with C. eos. ACKNOWLEDGMENTS The contributions of the many persons whose assistance made this study possible are gratefully acknowledged. Professor James C. Underhill of the Zoology Department of the University of Minnesota served as my advisor, offered suggestions regarding all phases of the work, and participated in most collecting trips made in the course of the study. Professor William D. Schmid of the Zoology Department of the University of Minnesota made the statistical portion of the study possible by suggesting many of the analytical approaches used, arranging with the Computer Center at the University to have the complex analyses performed, helping with most of the calculations done by the author, and critically examining pertinent sections of the manuscript. The investigation of the algal diet of C. erythrogaster was done with the help of members of the Department of Ecology and Behavioral Biology of the University of Minnesota. Professor Alan J. Brook made identifications of algae, offered consultation, and critically examined the "DIET" section of the manuscript. Mr. Roscoe F. Colingsworth and Mr. Alden E. Hine generously made and checked identifications of algae. Photographs were taken by Professor Schmid, Professor David J. Merrell, and Mr. Dale W. Fishbeck. The individual acknowledgments are given in the text. Mts. Marilyn Steere and Mrs. Raymond G. Griffith provided invaluable assistance with the figures and tables. My father, Edward B. Phillips, and fellow graduate students Fishbeck, Raymond G. Griffith, John D. Hudson, and Richard H. Stasiak all took part in the field work. Professors Underhill Albert W. Erickson, and Lloyd L. Smith, Jr., critically examined the entire manuscript. Two fellowships awarded by the National Science Foundation (NSF Grant GB 3390 and an NSF Summer Traineeship), provided financial assistance while the study was in progress. 3 SYNONYMY AND NOMENCLATURE Rafinesque (1820:48) listed "Chrosomus" as a subgenus of the Genus Luxilus when he described C. erythrogaster, butstated: "It might probably form a • s peculiar genus and be called Chrosomus erythrogaster or Kentucky Red belly." The type specimen(s) came from the Kentucky River, with no one locality specified. Cope, who brought the name "Chrosomus" into general use, distinguished C. eos from C. erythrogaster (1862: 523) on the basis of specimens from Meshoppen Creek, Susquehanna County) Pennsylvania. He listed several features that he considered diagnostic, perhaps the most significant of which are the comparatively more oblique mouth and shorter snout in C. eos. Cope later again discussed differences between C. erythrogaster and "C. eos" (1864:281), but confused C. eos with the yet-undescribed C. neogaeus and actually. 0.rom.IMme 101.0•0.0 — gave characteristics pertinent to the latter. He subsequently acknowledged this error (1869a:375), and, on the basis of specimens taken at New Hudson, Livingston County, Michigan, named C. neogaeus as Phoxinus neogaeus (p. 374). In the same year, he named Chrosomus oreas from specimens apparently taken in Montgomery County, Virginia (1869:233-4). Jordan in the first edition of his "Manual of the vertebrates of the United States ... " (1876:284), -4.. recognized three species of Chrosomus: C. pyrrhter o C. erythrogaster, and C. cos. Of these three, only C. cos was in synonymy with its original description. 11,11.1..1.10.111.1 Jordan named C. asEL5222Ela as a new species and stated that it was not the C. erythrogaster of Rafinesque. can find nothing in his diagnostic description to support this assertion. Jordan considered his C. erythrogaster to be the same fish as Cope's C. oreas (loc. cit.). Thus it seems clear that Jordan's C. pyrrhogaster is referable to C. erythrogaster, and his C. erythrogaster is referable to C. oreas. In the second edition of "Manual of the vertebrates ...", Jordan (1878:302) listed only C. erythrogaster under ”Chrosomus" and stated: "There seems to be but one well-defined species." However, Jordan and Gilbert (1882:153-4) listed C. erythrogaster, C. eos, and C. oreas as separate species. This arrangement remained rather stable although C. eos has been regarded as a subspecies of C. erythrogaster by certain workers and C.- necgaeus was added to the genus later. The finescale dace was called Phoxinus neogaeus until Jordan (1924:71) assigned it to a 'new' genus, Pirille (which he stated Floc. cit.] "is the German name of Phoxinus phoxinus"). The circumstances under which “Pfrille” was placed in the Genus Chrosomus are obscure. Regarding this action, Professor Reeve M. Bailey, curator of fishes at -5- the University of Michigan stated (personal communication, 29 August 1967): "1 am not sure who first assigned neogaeus to the genus Chrosomus but for want of an earlier date I may refer you to its positioning in that genus by C. L. Hubbs (1955, Systematic Zoology, Vol. 4, No. 1, p. 10). It was my conclusion after examination of the generic status of American freshwater fishes in 1949 that this action should be taken and Dr. Hubbs was aware of this belief. However, I do not recall having referred to the species in question during the interval between 1949 and 1955. It is entirely possible that someone else made the shift earlier. Many people have adopted the shift in generic names (Pfrille to Chrosomus). Most recently Banarescu in a book on the fishes of Roummia has gone one step further and synonymized both Chrosomus and Pfrille in the genus Phoxinus which genus had formerly been thought to be confined to the Old World. To my knowledge no American worker has yet followed this latter action but that is perhaps due to the fact that few are aware of it." The generic name Chrosomus will be retained in the ensuing discussion of the North American members of the ”Phoxinus-Chrosomus complex". METHODS AND MATERIALS The collections of Chrosomus erythrogaster and C. eos upon which the present study was based were either taken in the course of the investigation by seining or were available in the James Ford Bell Museum of Natural History at the University of Minnesota. Unless otherwise identified collection numbers refer to those housed in the fish collection at the Museum. All specimens in the available collections were
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