The Bantam Sunfish, lepomis symmetricus: Systematics and Distribution, and Life History in Wolf Lake, Illinois

Brooks M. Burr

STATE OF ILLINOIS DEPARTMENT OF REGISTRATION AND EDUCATION

NATURAL HISTORY SURVEY DIVISION URBANA, ILLINOIS

VOLUME 31, ARTICLE 10 SEPTEMBER, 1977 STATE OF ILLINOIS DEPARTMENT OF REGISTRATION AND EDUCATION BOARD OF NATURAL RESOURCES AND CONSERVATION

JOAN G. ANDERSON, B.S. Chairman; THOMAS PARK, Ph.D., Biology; L. L. SLoss, Ph.D., Geology; H. S. GUTOWSKY, Ph.D., Chemistry; ROBERT H. ANDERSON, B.S.C.E., Engineering; STANLEY K. SNAPino, Ph.D., Forestry; W. L. EvEsirr, E.E., Ph.D., Representing the President of the University of Illinois; JOHN C. GUYON, Ph.D., Representing the President of Southern Illinois University. NATURAL HISTORY SURVEY DIVISION, Urbana, Illinois SCIENTIFIC AND TECHNICAL STAFF GEORGE SPRUGEL, JR., Ph.D., Chief ALICE K. ADAMS, Secretary to the Chief

Section of Economic Entomology Section of Aquatic Biology WILLIAM H. LUCKMANN, Ph.D., Entomologist and D. HOMER BITCH, Ph.D., Aquatic Biologist Head WILLIAM F. CHILDERS, Ph.D., Aquatic Biologist JAMES E. APPLEBY, Ph.D., Entomologist R. WELDON LARIMORE, Ph.D., Aquatic Biologist /VIARCOS KOGAN, Ph.D., Entomologist ROBERT C. HILTIBRAN, Ph.D., Biochemist RONALD II. MEYER, Ph.D., Entomologist ALLISON BRIGHAM, Ph.D., Associate Aquatic Biol- STEVENSON MOORE, III, Ph.D., Entomologist, Ex- ogist tension WARREN U. BRIGHAM, Ph.D., Associate Aquatic Bi- EDWARD J. ARMBRUST, Ph.D., Associate Entomolo- ologist gist RICHARD E. SPARKS, PhD., Associate Aquatic Biologist JOSEPH V. MADDOX, Ph.D., Associate Entomologist TED W. STORCK, Ph.D., Assistant Aquatic Biologist ROBERT D. PAUSCH, Ph.D., Associate Entomologist JoHN TRANQUILLI, Ph.D., Assistant Aquatic Biolo- RALPH E. SECHRIEST, Ph.D., Associate Entomolo- gist gist RICHARD J. BAUR, M.S., Junior Professional Scientist JOHN K. BOUSEMAN, M.S., Assistant Entomologist CARL NI. THOMPSON, B.S., Junior Professional Sci- CHARLES D. BREMER, M.S., Assistant Entomologist, entist Extension t 9t LEE WAITE, M.S., Junior Professional Eden- MICHAEL E. IRWIN, Ph.D., Assistant Entomologist JANiA DONALD E. KUHLMAN, Ph.D., Associate Professor, DONALD W. DUFFORD, M.S., Research Associate Extension JOHN M. MCNURNEY, M.S., Research Associate ROSCOE RANDELL, Ph.D., Associate Professor, Ex- DAVID P. PHILIPP, Ph.D., Research Associate tension HARRY W. BERGMANN, B.S., Research Assistant WILLIAM G. RuEsiNx, Ph.D., Assistant Entomolo- WARNER D. BRIGHAM, B.S., Research Assistant gist KURT T. CLEMENT, B.S., Research Assistant DOUGLAS K. SELL, Ph.D., Assistant Entomologist LARRY W. COUTANT, M.S., Research Assistant JoHN L. WEDBERG, Ph.D., Assistant Entomologist, DONALD R. HALFFIELD, M.S., Research Assistant Extension EARL THOMAS JOY, JR., M.S., Research Assistant CLARENCE E. WHITE, B.S., Assistant Entomologist ROBERT MORAN, M.S., Research Assistant KEVIN D. BLACK, M.S., Assistant Specialist, Exten- MICHAEL J. STJLE, M.S., Research Assistant sion STEPHEN 0. SWADENER, M.S., Research Assistant DAVID A. GENTRY, M.S., Assistant Specialist, Ex- STEPHEN W. WAITE, M.S., Research Assistant tension CARL ALDE, B.S., Technical Assistant STEVEN TROESTER, ME., Assistant Systems Engineer PAUL BEATY, M.S., Technical Assistant JEAN G. WILSON, B.A., Superviaory Assistant KATHRYN EWING, B.S., Technical Assistant CATHERINE EASTMAN, Ph.D., Assistant Professional JEFF HurroN, B.S., Technical Assistant Scientist GEORGE LEWIS, M.S., Technical Assistant JOHN T. SHAW, B.S., Assistant Professional Scientist SUE MORAN, Technical Assistant DANIEL SHERROD, M.S., Assistant Professional Scientist DON MYRICK, B.S., Technical Assistant LESTER WEI, Ph.D., Assistant Professional Scientist JENS SANDBERGER, M.S., Technical Assistant CHARLES G. HELM, M.S., Junior Professional Sci- JOHN J. SULOWAY, B.S., Technical Assistant entist LIANN SULOWAY, M.S., Technical Assistant LINDA ISENHOWER, Junior Professional Scientist ROBERT THOMAS, B.S., Technical Assistant STEPHEN ROBERTS, B.S., Junior Professional Sci- GARY L. WARREN, B.S., Technical Assistant entist LOWELL DAVIS, Field Assistant LI-CHuN OHIO, Ph.D., Research Assistant C. RUSSELL ROSE, Field Assistant B.S., Research Assistant ELIZABETH ALLISON, Section of Faunistic Surveys and MARGARET ANDERSON, B.S., Research Assistant Insect Identification ROBERT J. BARNEY, B.S., Research Assistant Txu-Susx City, M.S., Research Assistant PHILIP W. SMITH, Ph.D., Taxonomist and Head MARION FARRIS, M.S., Research Assistant WALLACE E. LABERGE, Ph.D., Taxonomist JANET HARRY, B.S., Research Assistant GEORGE L. GODFREY, Ph.D., Associate Taxonomist BONNIE IRWIN, M.S., Research Assistant JOHN D. UNZICKER, Ph.D., Associate Taxonomist LOUIS JACKAI, M.S., Research Assistant DONALD W. WEBB, M.S., Associate Taxonomist JENNY KOGAN, M.S., Research Assistant LARRY M. PAGE, Ph.D., Associate Taxonomist PATRICIA MACKEY, B.S., Research Assistant BERNICE P. SWEENEY, Junior Professional Scientist BRIAN MELIN, B.S., Research Assistant Section of Wildlife Research MARY MILBRATH, M.S., Research Assistant JUDY MOLLETT, B.S., Research Assistant GLEN C. SANDERSON, Ph.D., Wildlife Specialist and LYNN PAUTLER, B.S., Research Assistant Head CELIA SHIN, M.S., Research Assistant FRANK C. BELLROSE, Sc.D., Wildlife Specialist BARBARA STANGER, B.S., Research Assistant WILLIAM R. EDWARDS, Ph.D. L Wildlife Specialist LEE ANNE TURNER, M.S., Research Assistant JEAN W. GRABER, Ph.D., Wildlife Specialist JO ANN AUBLE, Technical Assistant RICHARD R. GRABER, Ph.D., Wildlife Specialist CHARLOTTE JOHNSON, B.S., Technical Assistant HAROLD C. HANsoN, Ph.D., Wildlife Specialist W. W. COCHRAN, JR., B.S., Associate Wildlife Specialist Section of Botany and Plant Pathology CHARLES M. NIXON, M.S., Associate Wildlife CLAUS GRUNWALD, Ph.D., Plant Physiologist and Specialist Head KENNETH E. SMITH, Ph.D., Associate Chemist EUGENE B. HIMELICK, Ph.D., Plant Pathologist RONALD L. WEATEmEnn, M.S., Associate Wildlife DAN NEELY, Ph.D., Plant Pathologist Specialist D. F. ScEoENEwElss, Ph.D., Plant Pathologist LONNIE P. HANSEN, Ph.D., Assistant Wildlife Specialist J. LELAND CRANE, Ph.D., Associate Mycologist STEPHEN P. HAVERA, M.S., Assistant Wildlife KENNETH R. ROBERTSON, Ph.D., Assistant Taxono- Specialist mist DAVID R. VANCE, M.S., Assistant Wildlife Specialist BETTY S. NELSON, Junior Professional Scientist RICHARD E. WARNER, M.S., Assistant Wildlife Specialist GENE E. REID, Junior Professional Scientist RONALD E. DUZAN, Junior Professional Scientist JAMES E. SERGENT, Greenhouse Superintendent HELEN C. Scrcuurz, M.A., Junior Professional RICHARD WILSON, Technical Assistant Scientist

Continued on page 466

CONTENTS ACKNOWLEDGMENTS 437

METHODS AND MATERIALS 438

SYSTEMATICS 439 Synonymy 439 Types 440 Diagnosis 442 Description 443 Variation 445 Sexual 445 Allometric ...... 445 Geographic 445 Relationships 446 Specimens Studied 446 Ohio River Drainage 446 Mississippi River Drainage 446 Gulf Coast Drainage 447

DISTRIBUTION 447

CONSERVATION STATUS 448

LIFE HISTORY IN WOLF LAKE 449 Study Area 449 Habitat 449 Reproduction 450 Reproductive Cycle of the Male 450 Reproductive Cycle of the Female 451 Spawning 452 Development and Growth 453 Demography 454 Density 454 Composition 455 Survival 455 Diet 455 Interaction with Other Organisms 457 Competition 457 Predation 458 Hybridization 459 Parasitism 459 Summary 460

LITERATURE CITED 461

INDEX 465

This report is printed by authority of the State of Illinois, IRS Ch. 127, Par. 58.12. It is a contribution from the Section of Faunistic Surveys and Insect Identification of the Illinois Natural History Survey. Brooks M. Burr is a former Research Assistant, Illinois Natural History Survey. He is presently an assistant professor, Department of Zoology, Southern Illinois University, Carbondale.

dea&s, 2 (01284-2M-8-77) Fig. 1—Distribution of Lepomis symmetricus in relation to the Coastal Boundary (solid black line). Solid circles represent recent localities (1938 to the present) ; large open circles represent old records (pre-1900) where the species is presumably extinct. The most northern open circle also represents the type-locality. The life-history study area is enclosed within the square. The Bantam Sunfish, Lepomis symmetricus: Systematics and Distribution, and Life History in Wolf Lake, Illinois Brooks M. Burr

The bantam sunfish, described as lke; Garrett S. Glodek; Tomio Lepomis symmetricus by Stephen A. Iwamoto, California Academy of Sci- Forbes in 1883, is one of the least ences, including the Stanford Univer- known species in the genus, probably sity (SU) collection; Robert K. John- because of its small size, rarity over son; Craig E. Nelson; Donn E. Rosen, parts of its range, occurrence in rather Robert Schoknecht, Museum of Com- inaccessible swamp habitats, and drab parative Zoology (MCZ) at Harvard and nondescript appearance. This ef- University; and Keith S. Thompson. fort to remedy the gaps in our knowl- For providing laboratory space and/or edge of the species reviews all published the loan of specimens I thank Reeve references to L. symmetricus. To sup- M. Bailey, University of Michigan Mu- plement the meager information avail- seum of Zoology (UMMZ) ; Thomas able, this report includes an analysis M. Buchanan, Westark Community of morphological variation based on College at Fort Smith, Arkansas the study of museum specimens, an (ARP) ; Neil H. Douglas, Northeast assessment of the species' distribution, Louisiana University (NLU) ; David and a life-history study'based on peri- A. Etnier, University of Tennessee odic collections made at a study site in (UT) ; Robert K. Johnson, Field Mu- southern Illinois. seum of Natural History (FMNH) ; Ernest A. Lachner, U.S. National Mu- ACKNOWLEDGMENTS seum of Natural History (USNM) ; For aid in the literature search, I am Robert F. Martin, Texas Natural His- indebted to Philip W. Smith and Illi- tory Collection at Austin (TNHC) ; nois Natural History Survey librarian John D. McEachran, Texas Cooper- Doris L. Sublette; for help in collecting ative Wildlife Collection at College specimens, to present and former asso- Station (TCWC) ; William D. Pearson, ciates John A. Boyd, Lloyd R. Davis, University of Louisville, Kentucky Larry M. Page, Philip W. Smith, and (UL) ; Henry W. Robison, Southern Roger D. Wrisberg. Larry M. Page State College at Magnolia, Arkansas and Philip W. Smith provided counsel (HWR) ; and Royal D. Suttkus and on numerous matters. For identifying Michael M. Stevenson, Tulane Univer- trematode and acanthocephalan par- sity, Louisiana (TU) . For providing asites, I am grateful to David F. lists of locality records and/or distri- Oetinger and for advice on other par- bution maps I thank Thomas M. Bu- asitological problems, to Mary H. chanan; B. E. Gandy, Mississippi Mu- Pritchard, both of the Nebraska State seum of Natural Science; Henry W. Museum. E. Donald McKay III of the Robison; Morgan E. Sisk, Murray Illinois State Geological Survey sup- State University, Kentucky; Neil H. plied information on the age of Wolf Douglas; and Glenn H. Clemmer, Mis- Lake. sissippi State University. For information about the location Unless stated otherwise, the majority of syntypes and other pertinent study of the specimens used in this study are material, I am indebted to museum deposited at the Illinois Natural His- officials Reeve M. Bailey; James E. tory Survey (INHS) . 437

438 ILLINOIS NATURAL HISTORY S URVEY BULLETIN Vol. 31, Art. 10 Most of the illustrations for this Observations and minnow-seine col- paper were prepared by Larry Farlow, lections were made in Wolf Lake in Technical Photographer; Lloyd Le- Union County, Illinois, at approx- Mere, Technical Illustrator; and Craig imately 1-month intervals, except dur- Ronto, all of the Illinois Natural His- ing the spawning season, when more tory Survey; the drawing of the sub- frequent observations were needed. adult was done by Alice A. Prickett of The life-history study began 2 June the University of Illinois School of Life 1973 and ended 27 May 1975. Sciences. Computer analysis of some of Specimens were preserved in 10-per- the data was undertaken by Stephen D. cent form alin and were returned to Cowan of the Survey. The manuscript the laboratory for study. In all, 233 was edited by Shirley McClellan, As- specimens from Wolf Lake were pre- sistant Technical Editor at the Survey, served and examined. Because the spe- and Dr. Neil H. Douglas, Northeast cies is protected by the Illinois Fish Louisiana University, served as guest Code, usually no more than 20 spec- reviewer. Partial support for the field imens were taken on one visit even work was provided by the U.S. Depart- when the species was commonly en- ment of Agriculture Forest Service; the countered, so as not to seriously dec- Illinois Natural History Survey ren- imate the population. Collecting was dered the other support. Special per- done by bag seine; minnow seine; dip mission to collect specimens of the net; and, in one instance, by electro- bantam sunfish, which is protected by fishing. Potential predators of the ban- the Illinois Fish Code, was given by tam sunfish were occasionally collected the Department of Conservation. Per- for examination of stomach contents. mits to take specimens in the National Field notes were routinely taken. In Park were issued by Joe L. Newcomb the laboratory, specimens were sexed, of the Forest Service: Paul Brown of measured, and aged, and their gonads the Trojan Powder Plant granted per- and stomachs were excised and studied. mission to collect on powder plant During the spawning season, breeding property. adults were brought to the laboratory and placed in observation tanks. Aging to year class was done by METHODS AND MATERIALS counting scale annuli removed from An attempt was made to compile as the dorsum. Aging to month was done complete a synonymy as possible for by using May, the month of greatest Lepomis symmetricus, and it is be- breeding activity in Wolf Lake, as lieved that virtually all published ref- month zero. Thus, a sunfish collected erences to it have been examined. in October with one scale annulus was Morphological data were taken on se- estimated to have lived 1 year and 5 lected series that could be expected to months. For certain comparisons sun- show geographical variation, allomet- fishes were divided into young (through ric variation, or sexual dimorphism in 12 months) and adult (over 12 months) the species. Meristic and morphomet- age groups. ric data were taken in the conventional Weights of the ovaries of 30 females manner of Hubbs gc Lagler (1964: 19— were obtained and recorded as a pro- 26) . One-way analysis of variance tests portion of the adjusted body weight were run to determine significant dif- (the specimen minus the ovaries, stom- ferences in means of samples deter- ach, intestine, and liver) of the female. mined by sex. Unless stated otherwise, Mature ova from 14 preserved breed- measurements are standard lengths ing females were counted. Indicators (SL). used for ascertaining probable spawn- Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 439 ing periods in other localities and in Indiana) ; Seamster 1948: 165, 168 other years were that males exhibited (trematode parasite cited) ; Baugh- breeding color patterns and that fe- man 1950: 247 (Texas list) ; Moore males were heavy with ova. & Cross 1950: 146 (recorded from The relative survival of each year Oklahoma) ; Reeves 8c Moore 1951: class of the study population was cal- 112 (Oklahoma Coastal Plain) ; culated by expressing the number of Biililke 1953: 71 (SU syntypes individuals in that year class as a pro- listed) ; Moore 1952: n.p. (Oklahoma portion of the number of individuals list) ; Jurgens & Hubbs 1953: 15 in a younger year class. (Texas list) ; Knapp 1953: 115 (key, Texas range) ; Gunning & Lewis SYSTEMATICS 1955: 556 (habitat, food habits in SYNONYMY Illinois) ; Gunning 8c Lewis 1956: 24 (Wolf Lake and Pine Hills, Illinois) ; Leporms symmetricus Forbes Eddy 1957: 191 (key, range) ; Hubbs Lepomis symmetricus McKay 1882: 88 1957a: 97 (Texas range) ; Hubbs (nomen nudum); Forbes in Jordan 1957b: 9 (Texas list) ; Moore in 8c Gilbert 1883: 473-474 (original de- Blair et al. 1957: 170 (key, range) ; scription, Illinois River [at Pekin] Hubbs 1958: 10 (Texas list) ; Bou- Illinois) ; Forbes 1884: 68 (Illinois dreaux et al. 1959: 8, 10 (Sour Lake, range) ; Jordan 1884: 320-321 (rede- Hardin County, Texas) ; Cook 1959: scription, museum specimens cited) ; 180 (redescription, ecology, Missis- Jordan 1888: 117 (redescription) ; sippi range) ; Bailey et al. 1960: 27 'Bollman 1892: 566, 571 (key, range) ; (list) ; Smith 8c Bridges 1960: 254 Evermann 8c Kendall 1894: 84, 93, (INHS syntypes) ; Hubbs 1961: 10 111 (redescription, Texas records) ; (Texas range) ; Branson & Moore Hay 1894: 255, 261 (redescription, 1962: 9, 15, 24, 27, 29, 31, 33, 41, 48, key, not taken in Indiana) ; Richard- 65, 72, 91, 99 (relationships, acous- son 1904: 31, 33 (relationships, key, tico-lateralis system) ; Clay 1962: 119 Illinois range) ; Forbes 8c Richard- (Kentucky range) ; Collette 1962: son 1908: 251-252 (redescription, 146, 177 (associate of slough darter key, Illinois range) ; Forbes 1909: and swamp darter) ; Lambou 1962: 388 (Illinois range) ; Pratt 1923: 118 78 (Lake Bistineau, Louisiana) ; (key, range) ; Greene 1927: 309 (not Walker 1962: 40 (Jackson, Lincoln, in Wisconsin) ; Hildebrand 8c and Bienville parishes, Louisiana) ; Towers 1927: 133—134. (Greenwood, Walker 1963: 48 (Choudrant Bayou, Mississippi, food habits in Missis- Louisiana) ; Sharma 1964: 533 (mu- sippi); Summers 1937: 434 (new cus cells in canal linings) ; Burton & trematode parasite, Baton Rouge, Douglas 1965: 94 (Bayou De Siard, Louisiana) ; Mizelle 1938: 160 (tre- Louisiana) ; Smith 1965: 9 (Illinois matode transferred to new genus) ; range) ; Pflieger 1966: 53 (Missouri Mizelle & Hughes 1938: 351 (tre- key) ; Breder & Rosen 1966: 413 matode parasite cited) ; Summers & (breeding habits unknown) ; Chil- Bennett 1938: 248 (trematode para- ders 1967: 160 (tribe Lepomini) ; site cited) ; Kuhne 1939: 110 (Ten- Douglas & Davis 1967: 23 (Louisi- nessee list, redescription, sexes fig- ana list) ; Hoffman 1967: 340 ured) ; Lamb 1941: 45 (Willow (known parasites) ; Pflieger 1968: 54 Creek, San Jacinto drainage, Texas) ; (Missouri key) ; Moore in Blair Fowler 1945: 364, 370 (Louisiana et al. 1968: 128-129 (key, range) ; and Mississippi records, figure erro- Whitaker 1968: 96-97 (key, range) ; neous) ; Gerking 1945: 115 (possible Eddy 1969: 217 (key, range) ; 440 ILLINOIS NATURAL HISTORY S URVEY BULLETIN Vol. 31, Art. 10 Smith 8c Sisk 1969: 66 (Obion spawning) ; Webb 8c Sisk 1975: 63, Creek, Kentucky) ; Bailey et al. 67, 69 (Bayou de Chien, Kentucky, 1970: 36 (list) ; Jenkins et al. endangered in Kentucky) ; Hubbs 1971: 74 (possibly present in lower 1976: 6 (Texas list) ; Hubbs & Pigg Tennessee or Cumberland rivers) ; 1976: 116 (indeterminate status in Pflieger 1971: 413-414 (habitat, zoo- Oklahoma); Seehorn 1976: 21 (South- geography, Missouri range) ; Smith eastern National Forest list) . et al. 1971: 10 (not in upper Missis- Apomotis symmetricus: Boulenger sippi River) ; Hubbs 1972: 6 (Texas 1895: 21 (redescription) ; Jordan & range) ; Miller 1972: 244 (threatened Evermann 1896: 998-999 (redescrip- in Illinois and Missouri) ; Rozenburg tion) ; Evermann 1899: 310 (Lake et al. 1972: iii, 22, 28, 30, 32, 33, 36, Lapourde, Louisiana) ; Large 1903: 40, 45, 51, 82, I 1 1 (Navasota River, 24 (Illinois range) ; Jordan et al. Texas) ; Buchanan 1973a: 29 (Ar- 1930: 299 (list, range) ; Gowanloch kansas list) ; Buchanan 1973b: 51 1933: 348, 351 (Louisiana range) ; (key, Arkansas range) ; Miller 8c Schlaikjer 1937: 12 (phylogeny) ; Robison 1973: 184-185 (key, rede- Schrenkeisen 1938: 243-244 (rede- scription, ecology, Oklahoma range) ; scription, range) . Moore 1973: 6 (McCurtain County, Lethogrammus symmetricus: Hubbs in Oklahoma) ; Smith 1973: 33 (Illinois Jordan 1929: 147 (transfer to new key) ; Lopinot 8c Smith 1973: 46-47 genus erected by C. L. Hubbs) ; (status in Illinois) ; Buchanan 1974: Greene 1935: 220 (not in Wiscon- 89 (status undetermined in Arkan- sin) ; O'Donnell 1935: 486 (Illinois sas) ; Douglas 1974: 312-313 (rede- range) ; Breder 1936: 28 (breeding scription, Louisiana range) ; Pflieger habits unknown) ; Baker 1937: 48 in Holt et al. 1974: n.p. (rare in (redescription, rare at Reelfoot Missouri) ; Ackerman 1975: 10 (en- Lake) ; Baker 8c Parker 1938: 162 dangered in Illinois) ; Boyd et al. (Reelfoot Lake list) ; Baker 1939a: 1975: 11, 21 (status in Illinois) ; Clay 34 (redescription, sexes figured, com- 1975: 267, 276, 280 (redescription, mon at Reelfoot Lake) ; Baker key, Kentucky range) ; Douglas & 1939b: 45 (Reelfoot Lake key) . Davis 1975: 23 (Louisiana list) ; Mc- Reynolds 1975: 253 (LaRue Swamp, TYPES Illinois) ; Pflieger 1975: 254, 265 (fig- Lepomis symmetricus was described ure, key, redescription, Missouri by Forbes in Jordan 8c Gilbert (1883: range) ; Robison 1975: 54, 56 (Saline 473-474) from a syntypic series consist- River, Arkansas, evidence of recent ing of 15 specimens collected 16 April

Table 1.—Frequency distribution for number of caudal peduncle scales in selected pop- ulations of Lepomis symmetricus. Coefficient Number of Scales Standard Drainage Mean of 17 18 19 20 21 22 Deviation Variation Illinois R., Ill. 6 4 1 11 19.5 0.69 3.5 Wabash R., Ill. .. 3 7 2 12 20.9 0.67 3.2 Mississippi R., Ill., Mo., Ky. 3 13 13 17 5 51 20.2 1.21 6.0 Mississippi R., Tenn. 12 13 25 19.5 0.51 2.6 Mississippi R., Ark., La. 1 3 10 9 2 25 19.3 0.95 4.9 Ouachita R., Ark., La. 4 5 11 4 2 26 18.8 1.13 6.0 Red R., Okla., Tex., Ark., La. 5 12 10 5 1 33 18.5 1.03 5.6 Gulf Slope, Tex., La. 7 7 10 9 6 39 19.0 1.34 7.1 Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 441

Table 2.-Frequency distribution for number of lateral line scales in selected populations of Lepomis symmetricus. Number of Scales Coefficient Drainage Mean Standard of Deviation 30 31 32 33 34 35 36 37 38 Variation Illinois R., III. 7 2 2 11 32.5 0.82 2.5 Wabash R., Ill 3 5 1 2 1 12 35.4 1.51 4.3 Mississippi R., III., Mo., Ky. 2 12 15 9 5 3 4 1 51 33.6 2.87 8.5 Mississippi R., Tenn. 2 4 6 7 4 25 33.5 1.39 4.2 Mississippi R., Ark., La. 1 1 5 7 4 4 25 33.4 1.58 4.7 Ouachita R., Ark., La. 2 3 5 4 4 4 3 1 .. 26 33.3 1.95 5.9 Red R., Okla., Tex., Ark., La. 6 4 10 7 6 .. .. 33 32.0 1.39 4.3 Gulf Slope, Tex., La. 5 4 10 8 8 3 1 .. . 39 32.6 1.57 4.8

Table 3.-Frequency distribution for number of dorsal soft rays in selected populations of Lepomis symmetricus. Coefficient Number of Rays Standard Drainage N Mean of Deviation 9 10 11 12 Variation Illinois R., Ill. .. 7 3 1 11 10.5 0.69 6.6 Wabash R., III. 2 9 1 .. 12 9.9 0.51 5.2 Mississippi R., Ill., Mo., Ky. .. 23 24 4 51 10.6 0.40 3.8 Mississippi R., Tenn. . . 17 6 2 25 10.4 0.65 6.3 Mississippi R., Ark., La. 3 18 4 .. 25 10.0 0.54 5.4 Ouachita R., Ark., La. 3 19 3 1 26 10.1 0.63 6.2 Red R., Okla., Tex., Ark., La. 4 17 12 .. 33 10.2 0.66 6.5 Gulf Slope, Tex., La. 4 22 10 3 39 10.3 0.77 7.5

Table 4.-Frequency distribution for number of anal soft rays in selected populations of Lepomis symmetricus. Coefficient Number of Rays Standard Drainage N Mean of Deviation 9 10 11 12 Variation Illinois R., Ill. .. 10 .. 1 11 10.2 0.60 5.9 Wabash R.,111. 1 9 2 12 10.1 0.51 5.0 Mississippi R., Ill., Mo., Ky. 11 22 17 1 51 10.2 0.61 6.0 Mississippi R., Tenn. 7 17 1 25 9.8 0.52 5.3 Mississippi R., Ark., La. 8 15 2 25 9.8 0.60 6.1 Ouachita R., Ark., La. 8 17 1 26 9.7 0.53 5.5 Red R., Okla., Tex., Ark., La. 3 21 9 33 10.2 0.58 5.7 Gulf Slope, Tex., La. 9 27 3 39 9.8 0.54 5.5 442 ILLINOIS NATURAL HISTORY SURVEY BULLETIN Vol. 31, Art. 10 and 2 June 1880 from the Illinois INHS 220 and 226 are paralectotypes, River (Mississippi drainage) at Pekin, now INHS 75005 and INHS 75006, re- Tazewell County, Illinois (Fig. 1) . All spectively. The USNM, SU, and MCZ 15 of the original syntypes are extant: syntypes also became paralectotypes, INHS 220 (8, 32.7-39.5 mm SL) ; keeping their original catalogue num- INHS 226 (2, 50.1-51.2 mm SL) ; MCZ bers. 25014 (I, 49.5 mm SL) ; SU 1276 (3, It is unlikely that the original mate- 49.8-56.9 mm SL) ; USNM 29864 (1, rial of L. symmetricus collected by 51.0 mm SL) . All 15 are in a good state Forbes and associates was captured of preservation. To preserve customary from the Illinois River proper. Al- nomenclature and in accordance with though the Illinois River has changed the International Code of Zoological rather drastically since Forbes's era, it Nomenclature Article 74, recommenda- probably never maintained habitat tion 74D, a lectotype of L. symmetricus suitable for L. symmetricus. More Forbes is herewith designated (INHS likely the specimens came from one of 75004, 39.5 mm SL) . The specimen, the natural floodplain lakes in the a juvenile, conforms to the character- Pekin area, where favorable habitat ization of the species given under De- has been present in past years. scription and in Tables 1-4. The in- complete lateral line has 34 scales with DIAGNOSIS 6 scales above and 13 scales below the The most diminutive species of Le- lateral line. There are 19 caudal pe- porn is (the largest specimen measured duncle scales, 5 cheek row scales, and is 75.5 mm SL) is distinguished from 6 branchiostegal rays. Fin ray counts other members of the genus by this are: dorsal spines, 10; anal spines, 3; combination of characters: Lateral pectoral rays, 12-12; dorsal soft rays, line incomplete (1-18 scales unpored) 10; anal soft rays, 10. The nine other or interrupted (as many as 6 times) . specimens originally accessioned as Gill rakers long (longest in the genus,

Table 5.-Proportional measurements of Lepomis symmetricus from throughout the range, expressed in thousandths of standard length.' 10 Males (54-64 mm SL) 10 Females (50-64 mm SL) Coeffi- Coeffi- Stan- Stan- cient cient Measurement dard Range Mean dard of Range Mean of Devi- . Devi- Vari- Vari- ation ation atiorl a tiOfl Head length 375-423 396 013 3.5 361-403 390 012 3.2 Body depth 471-531 491 018 3.6 468-527 494 017 3.4 Caudal-peduncle depth 150-169 160 006 3.6 142-192 163 014 8.8 Pectoral fin length 245-285 263 014 5.4 248-291 263 012 4.7 Pelvic fin length 227-255 238 009 3.9 212-243 225 010 4.6 Longest dorsal spine 116-153 139 010 7.2 126-164 140 014 9.7 Head width 180-219 203 013 6.5 191-234 213 013 6.0 Bony interorbital width 078-096 087 006 6.6 074-093 084 007 7.8 Snout length 071-086 078 005 6.7 074-088 081 006 7.4 Upper jaw length 123-151 140 009 6.2 124-158 133 010 7.8 Predorsal length 439-480 459 014 3.0 445-483 462 014 2.9 Base dorsal fin length 461-508 478 015 3.1 455-517 480 017 3.6 Longest anal spine 120-151 137 010 6.9 124-157 139 012 8.7 Base anal fin length 216-297 244 022 8.8 206-257 234 015 6.6 Orbit length 087-105 095 005 5.7 083-105 095 002 7.8 Based on NLU 29918, 12804, 1954; UT 90.116, 90.140; TCWC 3643; HWR 74-8; INHS 75020, 75021, 75022, 75023, 18151, 18143, 17547. Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 443 longest rakers 2.3-2.9 mm) , and slen- 473-474) and Forbes & Richardson der (0.3-0.5 mm wide, 7-9 times longer (1908: 251-252) adequately described than wide) , numbering 12-15, modally the specimens available to them. The 13. Opercle stiff to its bony margin, following description is an amplifica- the dark opercular spot slightly diffuse tion, which includes additional meris- on narrow, bordering membrane. Dor- tic and morphometric data, and a more sal coloration dusky with dark coffee- comprehensive description of colora- colored spots on body, spots occasion- tion. Body proportion values are ally forming irregular vertical bands. presented in Table 5. When no geo- Head and cheeks darkened and with- graphic variation was noted, the var- out patterns. Juveniles often more ver- iation data from throughout the range tically barred than adults and have a of the species are merely summarized. prominent black blotch in the poste- When geographic variation was noted, rior rays of the soft dorsal fin, becom- the ranges and modes are given in ing less intense with age. Branson 8c the description, but their frequencies Moore (1962) showed these additional are discussed under Variation. Counts characters to be distinctive: only one of lateral-line scales, caudal-peduncle posterior pore on the post-temporal, scales, dorsal soft rays, and anal soft lateralis ending under the soft dorsal rays, all of which show slight clinal fin, preopercle angle 110° to 115°, variation, are presented in Tables 1-4. lachrymal bone nearly twice as tall as General physiognomy and pigmenta- wide, supramaxilla shorter than max- tion of adults and juveniles are shown illa, and no teeth on tongue or ptery- in Fig. 2 and 3. goids. Lateral line scales 30-38, modally 32 (Table 2) . Bailey (1938) reported one DESCRIPTION specimen with 40 lateral line scales. Forbes (in Jordan 8c Gilbert 1883: Scales above the lateral line 5 (in 7

Fig. 2—Breeding male Lepomis symmetricus 53.6 mm in standard length collected in Wolf Lake on 27 May 1975. Pigmentation in the fins is somewhat subdued by preservation. 444 ILLINOIS NATURAL HISTORY SURVEY BULLETIN Vol. 31, Art. 10 specimens) , 6 (92) , 7 (11) , = 6.1. Scales below the lateral line 12 (in 48 specimens) , 13 (52) , 14 (15) , i = 12.7. Caudal peduncle scales 17-22, modally 19 (Table 1) . Scales on cheek 4-6, modally 5. Scales well developed on preopercle, subopercle, interopercle, and opercle, all such scales about the same size and shape. No scales on top of head. Dorsal spines 9 (in 22 specimens) , 10 (133) , 11 (6) , I = 9.9. Dorsal soft rays 9-12, modally 10 (Table 3) . Anal spines 2 (in 1 specimen) , 4 (2) , 3 in all others. Anal soft rays 9-12, modally 10 (Table 4) . Pectoral rays 11 (in 8 specimens) , 12 (66) , 13 (32) , I = 12.2. All pelvic fins counted had 1 spine. Pelvic rays 4-4 (in 1 specimen) , 4-5 (2) , 5-5 (42) . Principal caudal rays Fig. 3—Lepcmis symmetricus prejuvenile 17 (in 41 specimens) , 18 (1) . 12.0 mm in standard length (above) and ju- Gill rakers on first arch (all rudi- venile 30.0 mm in standard length (below). ments counted) 12 (in 12 specimens) , 13 (36) , 14 (22) , 15 (6) , 1 = 13.3. imens are almost solid black on the Rakers long and slender (see Diagno- midbody with discrete black punctate sis). Rudimentary rakers (usually 3-5) marks on the cheeks. The fins, except are shorter and more blunt. The lat- the pectorals, are dusky overall with eral line on the body is incomplete or the soft dorsal and anal usually having interrupted (see Diagnosis and Fig. 3). several light spots. The pectoral fin The cephalic lateral-line system was rays are outlined by melanophores but described in detail by Branson are otherwise clear. The cheeks and Moore (1962) . Caudal fin slightly head are very dark and have no pat- emarginate. No teeth on tongue and terns. The dark opercular spot is pterygoids. Teeth present on vomers usually bordered with a light area on and palatines. Pharyngeal arches nar- its posterior margin. row with many small, blunt subconical Juveniles contrast with adults in teeth present (Richardson 1904) . Peri- generally having more distinct vertical toneal color is usually fleshy with many bands, in always having a black spot scattered melanophores, but occasional in the soft dorsal, and in having some specimens have a more silvery ground red-orange pigmentation in both the color with melanophores scattered soft dorsal and soft anal fins. Juveniles throughout. are lighter overall than adults and gen- Dorsal coloration is dusky olive- erally have seven to nine rather distinct brown or, in life, dark green with a vertical bands that are darker (brown) somewhat lighter venter of yellowish than the overall light greenish ground brown. Many dark coffee-colored spots color. The vertical barring is occasion- occur over the body, often one spot per ally obscured by flecks of darker pig- scale, creating vague, irregular vertical ment over the body, giving it a spotted bands or longitudinal rows. The belly, appearance. The juveniles of both breast, throat, and chin have many sexes have a distinct black blotch on tiny, dark melanophores. Some spec- the last five to eight rays of the soft Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 445 dorsal fin; the pigment is distributed sex organs was evident. Pelvic fin both on the radial and interradial length is significantly greater at the membranes. Rarely, there is a black 0.05 level (F = 8.29) in the male than spot in the soft anal fin (NLU 2907, in the female (Table 5). The urogen- 2 of 21 specimens; INHS 18151, 1 of ital papilla of the adult female is en- 45 specimens) on the last three rays, larged and protruding during the and again the pigment is both on the spawning season, whereas that of the radial and interradial membranes. Red- adult male is only slightly enlarged orange pigmentation is also present in (Fig. 4) . The male is not appreciably the soft dorsal and soft anal fins of larger than the female. The largest both sexes, on the radial and inter- individuals from the study area were radial membranes, and is very prom- females (61 and 63 mm SL), the inent in specimens collected during the largest specimen examined from fall and winter months. The belly, throughout the range was a female breast, throat, and chin sometimes are (75.5 mm SL) . The largest male was marked with discrete, tiny brown mel- 73.5 mm SL. anophores. Jordan (1884:320-321) re- marked that small specimens from New Allometric Orleans had faint blue spots on the No allometric variation in meristic sides of their heads. Breeding color- characters was found. Although allo- ation is discussed under Reproductive metric variation in morphometric char- Cycles of both sexes. acters was not investigated, adults are more robust than juveniles, as in other VARIATION sunfishes. Moreover, it is the adult that Sexual is symmetrical in shape and thus is re- No sexual variation in meristic char- sponsible for the trivial name of the acters was noted, but some dimorphism species. Juveniles have body propor- in one proportional character and in tions similar to those of other juvenile sunfishes. The number of vertical bands, if present at all, is the same in the juvenile and adult. The most no- table allometric variation is the ten- dency for the black spot in the soft dor- sal fin to become more diffuse and weak with age. It is prominent in the smallest young and absent in the adult, except in an occasional female. The soft anal and soft dorsal fins have a red-orange coloration that disappears when the fish becomes adult.

Geographic Geographic variation in some meris- tic characters was evident when samples Fig. 4—Genital papillae of Lepomis sym- were grouped according to major river metricus. A, nonbreeding male; B, breeding systems and arranged in a north-to- male; C, nonbreeding female; D, breeding south order from the Mississippi drain- female. The nonbreeding specimens were 1+ age of Illinois; through the Ouachita years old, collected on 19 October 1973; the breeding specimens were 2 years old, collected and Red river drainages of Arkansas, on 27 April 1974. Oklahoma, Louisiana, and Texas; to 446 ILLINOIS NATURAL HISTORY S URVEY BULLETIN Vol. 31, Art. 10 the Gulf Coast drainages of Texas and be most closely related to L. cyanellus Louisiana (Tables 1-4) . (Bailey 1938; Branson 8c Moore 1962) No significant geographic variation as a highly specialized congener with was found in any of the body propor- several unique characters. Hubbs (in tions measured (Table 5) . In fact, in Jordan 1929) considered L. symmetricus this respect L. symmetricus is remark- 'distinctive enough to warrant place- ably conservative for a species with a ment in a new monotypic genus, Le- rather long north-to-south distribution thogrammus, and Bailey (1938), adopt- (Fig. 1) . These meristic characters ing the use of subgenera, placed L. varied clinally: numbers of caudal- symmetricus in the subgenus Lethog- peduncle scales, lateral-line scales, and rammus. anal soft rays. The number of dorsal More recent studies on species of soft rays showed a slight but somewhat Lepomis using the techniques of elec- irregular trend toward more ray ele- trophoresis (Avise 8c Smith 1974) , hy- ments in the north (Table 3) . The bridization (Hester 1970) , and chro- Mississippi drainage samples from Ar- mosome analysis (Roberts 1964) have kansas, Louisiana, Tennessee, Missouri, not included specimens of L. symmetri- Illinois; and Kentucky were intermedi- cus. Thus, it is not known where the ate in caudal-peduncle and lateral-line species would be placed in the classi- scale counts between Red River-Gulf fication schemes presented by these Coast samples and those from the Wa- authors. bash drainage of Illinois. In these counts (Tables 1 and 2) the samples SPECIMENS STUDIED showed a gradual increase toward the The following list includes only north, whereas the soft-ray counts (Ta- those collections of L. symmetricus that bles 3 and 4) were more discordant, were used for meristic and morpho- with specimens from the Red River- metric features. Others were used for Gulf Coast samples having means close the assessment of distribution, descrip- to that of the Illinois River specimens. tive features, and life-history data. Col- The most aberrant samples are those lections are listed generally from north that formerly occurred in oxbow ponds to south. The number of specimens ex- along the Wabash River in White amined is given in parentheses follow- County, Illinois. They have a slightly ing the catalog number. Specific local- higher mean number of lateral-line ity data may be obtained upon request scales and slightly higher mean num- from the author. ber of caudal-peduncle scales, but they have lower means for the soft fin ray Ohio River Drainage counts than samples from the upper Mississippi drainage (Tables 1-4) . WABASH RIVER SYSTEM.—ILLINOIS, No apparent geographic trends in White County: 2 October 1882, INHS coloration or pattern could be per- 75008 (1) ; 1 October 1882, INHS ceived. Individual variation occurs in 75009 (1) ; 3 October 1882, INHS the prominence of the vertical bars and 75007 (10) . overall darkness, due perhaps in part to the strength of the preservative and Mississippi River Drainage age of the individuals. ILLINOIS RIVER SYSTEM.—ILLINOIS, Tazewell County: 16 April 1880, RELATIONSHIPS I NHS 75004 (1) , INHS 75005 (7) , Because of various features of mor- USNM 29864 (1) ; 2 June 1880, INHS phology, cytology, and paleontology, 75006 (2) . L. symmetricus has been considered to CLEAR CREEK SYSTEM.—ILLINOIS, Sept., 1977 BURR: THE BANTAM S UNFISH, LEPOMIS SYMMETRICUS 447 Union County: 18 July 1883, INHS ISIANA, Red River Parish: 22 June 75102 (5) ; 16 September 1959, 1965, NLU 1954 (7) . Winn Parish: INHS 17547 (6) ; 27 April 1963, INHS 23 June 1965, NLU 1989 (7) . Caddo 17566 (1) ; 27 May 1965, INHS 17583 Parish: 22 February 1969, NLU 12804 (1) ; 31 August 1970, INHS 17557 (1) ; (5) . TEXAS, Bowie County: 24 May 21 June 1973, INHS 18143 (5) ; 25 July 1957, TNHC 4984 (10) . Harrison 1913, INHS 18151 (2) ; 24 January County: 17 March 1972, TCWC 1974, INHS 75025 (6) ; 28 March 1974, 4068.14 (2) . INHS 75022 (1) ; 30 May 1974, INHS LAKE PONTCHARTRAIN.-LOUISI- 75021 (1) ; 27 May 1975, INHS 75020 ANA, Orleans Parish: 15 April 1974, (1) . NLU 29918 (5) . ORION CREEK SYSTEM.-KEN- TUCKY, Hickman County: 21 Jan- Gulf Coast Drainage uary 1964, INHS 75024 (1) ; no date, CALCASIEU RIVER SYSTEM.-LOUISI- UL 5617 (10) . Fulton County: 15 ANA, Calcasieu Parish: 10 August June 1948, UL 10691 (4) . 1965, NLU 2534 (6) . Allen Parish: 10 SAINT FRANCIS RIVER SYSTEM.-MIS- August 1965, NLU 2907 (5) . Jefferson SOURI, Stoddard County: 25 October Davis Parish: 10 August 1965, NLU 1973, INHS 75023 (10) . 2909 (4) . NATURAL LAKES AND BACKWATERS.- MERMENTAU-TECHE RIVER SYSTEM.- TENNESSEE, Lake County: 11-13 LOUISIANA, Avoyelles Parish: 20 March 1968, UT 90.27 (8) ; 8 April April 1975, NLU 31572 (3) . 1950, FMNH 80532 (2) . Lauderdale NECHES RIVER SYSTEM.-TEXAS, Jef- County: 9 October 1972, UT 90.102 ferson County: 2 May 1970, TCWC (2) . ARKANSAS, Chicot County: 17 3643 (14) . Hardin County: August August 1974, HWR 74-35 (8) . 1950, TNHC 585 (1). Newton County: FORKED DEER RIVER SYSTEM.-TEN- 7 June 1952, TNHC 2889 (3) . NESSEE, Haywood County: 3 Novem- TRINITY RIVER SYSTEM.-TEXAS, ber 1973, UT 90.138 (1) ; 27 April Chambers County: 14 July 1953, 1974, UT 90.140 (6) . Gibson County: TNHC 3873 (2) . 19 October 1973, UT 90.139 (10) . SAN JACINTO RIVER SYSTEM.-TEXAS, L'ANGVILLE RIVER SYSTEM.-AR- Montgomery County: 23 March 1951, KANSAS, St. Francis County: 7 Au- TNHC 1211 (1) . gust 1939, UMMZ 128537 (2) . ARKANSAS RIVER SYSTEM.-ARKAN- SAS, Arkansas County: 13 August DISTRIBUTION 1974, ARP-79 (10) . All known locality records for L. OUACHITA RIVER SYSTEM.-ARKAN- symmetricus are plotted in Fig. 1. SAS, Bradley County: 23 May 1974, Along the Gulf Coast the species ex- UT 90.116 (1) , HWR 74-8 (1) ; 10 Au- tends from Eagle Lake (Colorado River gust 1974, HWR 74-26 (7) . Calhoun drainage, UMMZ 129793) in Texas County: 6 October 1974, JLS 74-14 east to marshes of the Jordan River sys- (2) . Union County: 25 April 1975, tem in Mississippi. In the Mississippi NLU 31455 (10) . LOUISIANA, Oua- Valley it presently extends north to the chita Parish: 17 October 1964, NLU bottomland oxbow lakes and swamps 894 (5) . of southern Illinois. A published rec- RED RIVER SYSTEM.-ARKANSAS, ord for the St. Joseph River of Mich- Little River County: 13 September igan (Dolley 1933) is clearly based on 1940, UMMZ 170879 (1) . OKLA- a misidentification, as Michigan is far HOMA, McCurtain County: 20 Au- out of the range of the bantam sunfish. gust 1948, UMMZ 155830 (1) . LOU- L. symmetricus is now almost en- 448 ILLINOIS NATURAL HISTORY SURVEY BULLETIN Vol. 31, Art. 10 tirely restricted to the Coastal Plain. It The distribution of L. symmetricus formerly traversed the Coastal Plain suggests that it is autochthonous to the boundary far northward to the Illi- lower Mississippi River valley (Pflieger nois River (at Pekin) and backwater 1971:413-414). It apparently dispersed ponds and sloughs of the Wabash River through oxbow lakes, swamps, and system in White County, Illinois (Fig. sloughs, created by varying water levels 1) . The species has not been collected during the history of the Mississippi from the type-locality since 1880, a fact River. (Pflieger (1971:414) suggested which Richardson (1904) noted only that L. symmetricus may have had its 24 years after its original description. origin in the lower Mississippi valley, Indeed, it was collected only twice from dispersed northward to central Illinois Pekin. It has not been collected from during the postglacial Climatic Opti- the Wabash valley since 1882, whence mum, and become disjunct in its north- it was known from three localities and ern distribution subsequently. 12 specimens (INHS 75007, 75008, 75009) . The distribution of L. sym- metricus has thus changed rather dra- CONSERVATION STATUS matically in Illinois, the decimation Miller (1972) listed L. symmetricus probably being the result of radical as rare in both Illinois and Missouri in changes brought on by human modifi- a compilation of threatened fishes of cations, notably the stocking of non- the United States. At that time it was native sunfishes, a reduction in aquatic known in those states from only two vegetation, draining of lowland swamps localities: the LaRue-Pine Hills area and sloughs, and various forms of ag- of southwestern Illinois (Union County) ricultural and industrial pollution and the Duck Creek Wildlife Area of (Smith 1971) . Mills et al. (1966) southeastern Missouri (Bollinger clearly demonstrated the effects of hu- County) , where it has been reported to man modification on the fauna and be common (Pflieger 1971:413) . It has flora of the Illinois River, and the fac- since been found to be common in tors listed above almost surely caused Wolf Lake, Illinois, and Mingo Na- the extirpation of the species from the tional Wildlife Refuge, Missouri (Pflie- Pekin area. It is also possible that the ger 1975:265) . The species is on the relatively short life span of the species protected list of both states but not (3+ years) is somehow associated with presently endangered in either because its fairly rapid extirpation from dis- its habitat is now rigidly protected in turbed or polluted areas in the Missis- refuges. Recently, Webb 8c Sisk (1975: L. symmetricus sippi Valley of Illinois, Missouri, and 69) recommended that be placed on Kentucky's rare and en- Kentucky. dangered species list in view of its rar- The species is virtually absent east of ity in Kentucky. the Mississippi River in Mississippi. In Oklahoma the species is found Perhaps the Mississippi River has been only in the swamps of McCurtain an effective barrier to dispersal in this County in the southeastern corner of region, or the species' apparent absence the state (Fig. 1). L. symmetricus was there may be because collectors tend to not considered threatened by Robison avoid swamps, sloughs, and lowland et al. (1974) in their list of threatened streams. The species is statewide in Oklahoma fishes, but it may presently occurrence in Louisiana, where it is be reduced in numbers according to common, and it is rather common in Hubbs 8c Pigg (1976:116) . In Arkan- eastern Texas, southern Arkansas, and sas the status of the species was listed parts of western Tennessee (Fig. 1) . as indeterminate by Buchanan (1974) , Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 449 but L. symmetricus was not cited by Powder Plant. Most observations and Robison (1974) in his list of threat- collections in Wolf Lake were made ened Arkansas fishes. The species is near the powder plant bridge, where apparently in no danger in southern access to the lake was easy although Arkansas (Fig. 1) , where it is known other portions of the lake were from many localities. sampled.

LIFE HISTORY IN WOLF LAKE HABITAT STUDY AREA Wolf Lake is characterized by two Wolf Lake is a long (ca. 1.9 km) , predominant habitats: a heavily veg- narrow (ca. 0.1 km) , and ancient ox- etated shoreline with many submerged bow of the Big Muddy River (Missis- logs and stumps (Fig. 5) and an open sippi drainage) situated south of the deepwater area in the center of the lake LaRue-Pine Hills Ecological Area to free from vegetation and submerged ob- which it is connected by bottomland jects. The lake is not shaded and the swamp. The lake is apparently still in water is usually turbid. The vegetated a fairly natural, undisturbed condition shoreline, where L. symmetricus occurs and is estimated to be at least 2,000 (Fig. 5) , is dominated by spatterdock years old (E. Donald McKay III, per- (Nymphaea advena), American lotus sonal communication) . The northern (Nelumbo lutea), common arrowhead portion of the lake was recently ac- (Sagittaria latifolia), coontail (Gera- quired by the U. S. Forest Service, tophyllum demersum) and duckweed whereas the southern portion of the (Lemna spp., Wolffia spp.) . The bot- lake is privately owned by the Trojan tom consists mostly of decomposed veg-

Fig. 5—Vegetated margin of Wolf Lake, Union County, Illinois, illustrating the preferred habitat of Lepomis symmetricus. Photo taken in May 1974. 450 ILLINOIS NATURAL HISTORY S URVEY BULLETIN Vol. 31, Art. 10 etation, silt, and mud, with some sand. REPRODUCTION Water depth ranges from 300 mm to 18 Reproductive Cycle of the Male meters. Dissolved oxygen averages 9.0 The genital papilla (Fig. 4) of ripe L. ppm; temperatures range from 4° to symmetricus males enlarged slightly as 8° C from December to February and the spawning season approached. The are as high as 29° C in July and August. testes, normally small, translucent, and L. symmetricus was found in similar elongate, became large, opaque white, habitat during a 1—year study of fishes and thickened. in the adjacent LaRue-Pine Hills Breeding males (Fig. 2) , in contrast swamp (Boyd et al. 1975) . During the to non-breeding males and females fall and winter months L. symmetricus (which were nearly identical in color was characteristically found at a depth and pattern) , became very dark on the of 150-300 mm usually near the shore- head, and the irregular vertical cross line in Wolf Lake. During the summer bars grew subdued. The venter from months the species could be found at the cltin and throat to the anterior rays depths of 600-1200 mm but still within of the anal fin became grayish black. the vegetated periphery of the lake. Many small greenish flecks were present Elsewhere in its range L. symmetri- on the head and opercle, and the dark cus is invariably found in lentic waters opercular spot was outlined by a silvery- characterized by standing timber, sub- cream color with a hint of suffused red. merged logs and stumps, and rich vege- The pectoral fins were relatively dusky tation. Sloughs, oxbows, ponds, back- overall but with no definite patterns. waters, lakes, and swamps typical of The posterior edges of the pelvic fins the undisturbed portions of the Coastal were almost solid black with the re- Plain are optimal habitat. L. symmet- mainder of the fins cream color. The riots is found in greatest numbers over dorsal fin had many light spots sur- substrates consisting of mud, detritus, rounded by dusky brown or black areas. and decayed plant material. The iris of the eye was brilliant red Although L. symmetricus is syntopic with a distinct black transverse bar with several other species of Lepomis through it. in Wolf Lake, it was almost always col- Because of the silty darkly-stained lected by itself in the areas mentioned. water of Wolf Lake, no nests of L. sym- The other Lepomis were usually taken metricus could be observed in nature, in more open areas and generally in and nothing is known of territory size. deeper water. In Wolf Lake the fishes However, Robison (1975:56) reported most often found with L. symmetricus that on 23 May 1974 in a roadside pool, in descending order of association were Saline County, Arkansas, L. symmetri- L. macrochirus, L. gulosus, Pomoxis cus had recently spawned, inasmuch as nigromaculatus, Notemigonus crysoleu- "depressions in the mud and leaf litter cas, Gambusia affinis, Micropterus sal- substrate were filled with numerous moides, Elassoma zonatum, and Etheos- eggs." Since males were observed to be toma gracile. Other inhabitants of the habitat of L. symmetricus occurring in highly aggressive toward females and less frequent numbers are Lepisosteus other sunfishes are known to be terri- oculatzts, L. platostomus, Dorosoma ce- torial (Larimore 1957) , it is assumed pedianum, Umbra limi, Cyprinus car- that L. symmetricus defends an area in pio, Ictiobus cyprinellus, Ictalurus nat- nature. An aquarium-held male col- alis, I. nebulosus, Fundulus dispar, lected in May was seen on several occa- Aphredoderus sayanus, Centrarchus sions to form a shallow nest by rapidly macro pterus, Lepomis microlophus, swimming forward, then turning his and L. punctatus. body straight up in a vertical position Sept., 1977 BURR: THE BANTAM S UNFISH, LEPOMIS SYMMETRICUS 451 and descending, sweeping his tail vig- Small white ova were present in fe- orously back and forth until a nest de- males 1+ years of age and 35 mm long pression was formed. Such nests were as early as September but were difficult formed over both sand and gravel sub- to distinguish in younger and smaller strates. These nests were approximately females. By January and February 90-120 mm in diameter. It is likely larger yellowish ova were found in 1+- that L. symmetricus males build shal- year females of 40 mm or longer. Large, low depressions in the mud bottom of coarse, maturing orange ova were pres- Wolf Lake along the shallow edges ent from March to May in larger and close to the vegetation where egg at- older females and in some smaller fe- tachment may take place. This behav- males over 34 mm and approaching 1 ior has been described for L. cyanellus year of age. Just prior to spawning (Hankinson 1908:210-211) . time, the mature ova became a translu- Only large males developed the cent orange. breeding patterns, the slightly enlarged The largest and oldest females pro- genital papilla, and the enlargement of duced the largest number of mature the testes. Only males of at least 1+ ova. In 14 ripe females collected in years and 40 mm or longer appeared to April and May the number of ova var- be sexually mature, according to color- ied from 219 to approximately 1,600 ation and condition of the testes. The (Table 6) . For these females the rela- largest males probably do most or all tionship between the number of ma- of the spawning. ture ova (F) and the adjusted body weight (W) was F = —50.94 + 210.70W, Reproductive Cycle of the Female with r = 0.818, and between the num- ber of mature ova and the standard Generally the largest females devel- length (L) was log F = —2.785 + 3.383 oped the earliest mature ova and prob- log L, with r = 0.663. ably contributed most to the spawning Ovaries of postspawning females col- effort. Females as short as or shorter lected in June were smaller than those than 34 mm and 1 year of age devel- of females collected in April and May. oped mature ova and were potential They averaged slightly heavier than spawners. ovaries from females collected in Females underwent some changes in March. Ovaries from females taken in coloration associated with the breeding July and August were small. A relative season. In contrast to males and non- increase in ovary size was evident by breeding individuals, the breeding fe- late fall and continued to the spawning male had 9 or 10 distinct vertical bars period the following spring (Fig. 6) . of a dark bluish-purple color with light For the females examined, the relation- greenish flecks in the spaces between ship between the weight of the ovaries the bars. The cheek and opercle con- divided by the adjusted body weight tained bright spots of golden green, but (Y) and the month (X) , with July = the fins were relatively clear and not 1 and May = 11, was log Y = 0.699 + dusky. Some females retained a diffuse 0.099X, with r = 0.782 (Fig. 6) . The ocellus in the posterior rays of the dor- proportionally largest ovaries (equaling sal fin. As in males, the iris was bright 30.8 percent of the adjusted body red. Other marked morphological weight) were found in a 51-mm, 2- changes were the distended belly caused year-old female collected on 27 April by the maturing ova and the enlarge- 1974 (UT 90.140) . In the 14 females ment of the genital papilla (Fig. 4). represented in Table 6, overy-weight- Enlargement of the papilla was notice- to-adjusted-body-weight ratios ranged able only in ripe females. from 0.070 to 0.308 and averaged 0.107. 452 ILLINOIS NATURAL HISTORY SURVEY BULLETIN Vol. 31, Art. 10 Table 6.—Relationship between size, age, and ovary weight of Lepomis symmetricus fe- males and the number of mature ova produced. An age of 1 year = 1 1-1 3 months, 2 years -= 23-25 months. Data from TCWC 3643, UT 90.140, and INHS 1 7 5 8 3, as well as that from Wolf Lake, are included. Number of Adjusted Mature Ovary Standard Body Age (orange or Weight Length Weight in translucent, in Grams in mm in Grams' Years 0.6-0.9 mm) Ova 34 1.42 0.10 326 34 1.49 0.12 219 36 1.78 1 0.13 491 37 1.65 1 0.20 368 37 2.06 1 0.18 403 38 2.32 1 0.20 330 39 2.11 1 0.21 432 40 2.43 1 0.18 417 42 2.44 1 0.26 421 43 2.75 1 0.20 374 45 3.22 1 0.33 364 45 3.34 1 0.31 378 51 4.51 2 1.39 ca 1600 52 7.57 2 0.88 ca 1400 Adjusted body weight is the specimen's weight after removal of the ovaries, stomach, intestine, and liver.

peratures ranged from 18° to 22° C. 500— Field observations and examination of LOG Y= 0.699+0.099X museum specimens collected during all 100- months of the year indicated that mid- 50- April to early June was the typical spawning period for the species throughout its range (Table 7) . Although spawning was not observed 10- in the study area, ripe aquarium-held individuals collected 27 May 1975 en- 5- gaged in prespawning activity for 7 days at water temperatures varying from 24° to 26° C. After presumed

WEIGHT OF OVARIES X 1000/ADJUSTED BODY 1 l l l l l l l l l l stimulation from a recent feeding the JULY SEPT. NOV. JAN. MAR. MAY male began to court the female by Fig. 6—Monthly variations in ovarian nudging her with his snout along the weight relative to adjusted body weight of posterior regions of her body and con- Lepomis symmetricus. The vertical axis is on tinually nipping at her caudal fin. The a logarithmic scale. Ovaries from specimens collected June to April were from all age female did not respond to these actions classes, but ovaries from specimens collected but the male continued to nip at her in May were from 2-year-old (24 months) fins and nudged the female with his fish. snout between the pelvic fins while chasing her. The female remained un- Spawning responsive. After 3 days of this behav- In Wolf Lake breeding individuals ior the male began to charge the fe- were captured as early as 24 April and male at rapid speeds with his opercles as late as 30 May. Most spawning prob- flared out and with the irises of his ably occurred in May when water tern- eyes more intense in color than before. Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 453 Table 7.—Collections of breeding Lepomis symmetricus. Locality Collection Date Remarks Wolf Lake, Union Co., Ill. 24 April-30 May 1974 Males and females in extreme (INHS 75020, 75021) breeding condition. Pine Hills Swamp, Union Co., Ill. 27 May 1965 Female in breeding condition. (INHS 17583) Illinois River, Tazewell Co., Ill. 2 June 1880 Females in breeding condition. (INHS 75006) Swamp, Haywood Co., Tenn. 27 April 1974 Males and females in extreme (UT 90.140) breeding condition. Reelfoot Lake, Lake Co., Tenn. 8 April 1950 Males and females in breeding (FMNH 80532) condition. Roadside Ditch, Bradley Co., Ark. 23 May 1974 Male and female in breeding (UT 90.116) (HWR 74-8) condition. Ouachita River, Union Co., Ark. 25 April 1975 Females in extreme breeding (NLU 31455) condition. Big Hill Oil Field, Jefferson Co., Tex. 2 May 1970 Males and females in breeding (TCWC 3643) condition. Marsh, Orleans Parish, La. 15 April 1974 Males and females in breeding (NLU 29918) condition. Creek, Avoyelles Parish, La. 20 April 1975 Males in breeding condition. (NLU 31572)

When he approached the female, he of time required for hatching, or the abruptly turned himself to a vertical morphology of hatchlings. position (with his snout pointing up- The smallest L. symmetricus individ- ward) and gently swam around her in ual from the study area was 12 mm, a close circle while fanning his tail. collected 21 June 1973 (Fig. 3) . At Similar courtship patterns were de- this size the nape, breast, and sides of scribed by Larimore (1957) for L. gu- the head were the only regions incom- losus. After 7 days of constant nipping, pletely scaled, but no definite pigment nudging, badgering, and displaying pattern was present. Small melano- other prenuptial behavior, the male phores outlined the scale borders on had succeeded in completely mutilat- the body and some of the fin rays but ing the uncooperative female's caudal were concentrated heavily on the top fin, and on the 8th day the female was of the head, on the lips, and around found dead. Even though an actual the eye. The soft dorsal fin ocellus was egg-laying session did not take place, it just beginning to develop (Fig. 3) . is evident that the nest building and A series of 43 young L. symmetricus prespawning behavior of L. symmetri- from 14.0 mm to 25.0 mm was collected cus does not vary greatly from that de- in the study area on 25 July 1973. At scribed for other species of Lepomis 14 mm squamation patterns were like summarized by Breder 8c Rosen (1966) . that at 12 mm, but many more melano- phores were present in the fins and they began to form patterns on the DEVELOPMENT AND GROWTH body. The ocellus was dark at this size. Mature ova ranged in size from 0.6 At 19 mm vague vertical bars had to 0.9 mm in diameter, were translu- formed, and squamation was nearly cent orange, and contained a single oil complete. At 25 mm the lateralis sys- droplet. No data are available on incu- tem was developed, and the overall pig- bation temperatures of eggs, the length ment pattern was similar to that of 454 ILLINOIS NATURAL HISTORY SURVEY BULLETIN Vol. 31, Art. 10 adults. Squamation was complete at DEMOGRAPHY this stage. At a slightly larger size ju- Density veniles began to take on the form, The nature of the habitat of L. sym- pattern, and coloration illustrated in metricus made population density mea- Fig. 3. surements difficult, since submerged L. symmetricus from Wolf Lake grew logs, brush, and vegetation prevented at a decreasing rate (Fig. 7) and reached thorough sampling of a given area. However, on two occasions approx- • •• • i Y.5.91+32.97 LOG X mately 5 months apart quantitative samples of L. symmetricus were taken in Wolf Lake by repeatedly seining a measured shallow margin of the lake until no more individuals could be collected. The number collected was translated into the number per square meter. The greatest density found for L. symmetricus in Wolf Lake was 0.69 sunfish per square meter (Table 8) . 2 14 16 18 20 22 24 34 36 38 In the nearby LaRue-Pine Hills MONTHS OF AGE swamp, the density of L. symmetricus Fig. 7—Size distribution by age of Lepomis may approach 0.72 sunfish per square symmetricus collected in Wolf Lake between 21 June 1973 and 30 May 1974. Data from meter (Table 8) or, at best, 1 individ- 27 May 1975 and 12 December 1974 are Table 8.—Number of Lepomis symmetricus included. Black dots represent sample means per square meter collected in vegetated mar- for both sexes combined. In total, 233 speci- gins of Wolf Lake and LaRue-Pine Hills swamp. mens are represented. Number one-half of the first year's mean growth of L. symmetricus in approximately 10 weeks. The rela- Number per square Date tionship between standard length (Y) Collected meter in and age in months (X) expressed for Wolf Lake the sexes combined is Y = 5.91 ± 32.97 and Pine Hills log X, with r = 0.943. Males grew at a slightly more rapid rate than females 25 October 1973, but were not significantly larger than Wolf Lake 9 0.313 females. At 13-18 months males aver- 28 March 1974, Wolf Lake 20 0.694 aged 45.9 mm and females averaged Mean 0.504 42.7 mm (t = 1.39, df = 11) . At 19-24 months males averaged 49.3 mm and 24 October 1973, Pine Hills 6 0.723 females averaged 47.5 mm (t = 1.00, 28 November 1973, df = 14) . The largest specimen exam- Pine Hills 4 0.542 ined from Wolf Lake was a 63.0-mm 27 March 1974, female collected 25 July 1973. In other Pine Hills 1 0.114 parts of its range L. symmetricus is Mean 0.460 known to attain a greater length, and specimens as long as 75.5 mm have ual per 3 square meters in optimal been collected (TU 148—St. Tam- habitat (Boyd et al. 1975) . Of 31 in- many Parish, Louisiana) . Based on dividuals collected during 9 months at the collections examined, such large several collecting sites in Pine Hills, L. size is unusual, with most adults rang- symmetricus made up 2.3 percent of ing between 55 and 60 mm. the total sample of fishes captured. Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 455 However, more than 80 percent of the Table 10.—Relative survival of year classes individuals were captured at one site of Lepomis symmetricus in Wolf Lake ex- where the habitat was judged to be op- pressed as proportions of the —1 year class (1 x') and the 1+ year class (1 X2). timal (Boyd et al. 1975) . Gunning & Lewis (1955) found that L. symmetri- Number Survival cus made up 5 percent of their total Sample Year of Class Speci- sample of fishes at Pine Hills. mens Ix' 1 x2 Composition Males —1 81 1.000 1+ 16 0.198 1.000 Of the 233 L. symmetricus collected 2+ 1 0.012 0.063 in Wolf Lake, 85.4 percent were up to 3+ •• • 1 year of age, 12.4 percent were over 1 Females 1.000 and up to 2 years of age, 0.8 percent 0.110 1.000 0.008 0.077 were over 2 and up to 3 years of age, 0.025 0.231 and 1.2 percent were over 3 years of Total age (Table 9) . sample 199 1.000 1+ 29 0.146 1.000 Table 9.—Distribution of sexes and year 2+ 2 0.010 0.069 classes in samples of Lepomis symmetricus 3+ 3 0.015 0.103 collected in Wolf Lake between 21 June 1973 and 30 May 1974, and on 27 May 1975 and 12 December 1974. Because of the difficulty in collecting in Wolf Lake, the numbers of 1+ and Number by Year Class older individuals in Tables 9 and 10 Sex Total —1 1+ 2+ 3+ are probably lower than their actual proportion in the population. Males 81 16 1 98 Females 118 13 1 3 135 The shapes of the survival curves for Total 199 29 2 3 233 males, females, and total sample were quite similar. All showed a very low Females predominated in the young- survival rate after the 1st year of life. of-the-year (-1) age class [1.5 females Only three individuals 3 years or older 2 to 1 male (x = 6.87; P < 0.01) ], and were found. The oldest L. symmetricus in the total sample (N = 233) the ratio from Wolf Lake examined was a fe- 2 was 1.4 females to 1 male (x = 5.97; male 3 years and 2 months old (as- P < 0.025) . Although predominating suming May hatching) collected 25 significantly in the —1 age class and in July 1973. the total, females were slightly less Specimens from throughout the common than males in the 1+ age range further confirm a 3+ -year life class. span for the species: INHS 17547— from LaRue-Pine Hills Ecological Area Survival collected 16 September 1959, contain- Relative survival values (Table 10) ing three individuals all 3 years for each year of life were calculated for and 4 months of age (assuming May males, females, and the total sample of hatching) ; NLU 31572—collected 20 L. symmetricus, using the data in Ta- April 1975 from a creek, Avoyelles Par- ble 9. It was assumed that each age ish, Louisiana, containing three indi- class was collected in proportion to its viduals 3 years of age. Most other spe- relative number in the population, that cies of Lepomis are much longer lived. the population was neither increasing nor decreasing, and that the number of DIET fry entering the population each year Stomach contents of 176 L. symmet- was constant. ricus from Wolf Lake were examined. 456 ILLINOIS NATURAL HISTORY S URVEY BULLETIN Vol. 31, Art. 10

Twenty-nine of these contained no food ation in diet (Tables 13 and 14) was items and eight contained green algal evident. Gastropods were eaten in the material. A large variety of food orga- winter and spring months. The largest nisms was found (Tables 11-14) . The percentages of most food items, includ- predominant food items of the Wolf ing gastropods, stratiomyids, chirono- Lake population were gastropods, cla- mids, and some microcrustacea, were docerans, ostracods, amphipods, dragon- eaten in the months prior to and dur- fly naiads, chironomids, and ceratopo- ing the spawning season, presumably gonids. reflecting an increase in consumption Small L. symmetricus (less than 21 associated with spawning preparedness mm) fed predominantly on microcrus- (Page 1974:17) . Aquatic Hemiptera tacea, dragonfly naiads, and chirono- were eaten exclusively in the summer mids; large individuals (more than 40 months, when they were most abun- mm) fed primarily on gastropods, dant. The presence in the diet of the dragonfly naiads, and amphipods (Ta- exclusively terrestrial hemi teran family bles 11 and 12) . Some seasonal vari- Fulgoridae reflects surface feeding by

Table 1 1.-Stomach contents of Lepomis symmetricus from Wolf Lake, by size class of sunfish. Figures in parentheses are numbers of stomachs examined. Percent of Stomachs in Which Food Organism Occurred Food Organism <21 21-30 31-40 41-50 51-60 >60 mm mm mm mm mm mm (25) (44) (56) (19) (5) (5) Gastropoda 12.5 31.6 4.0 4.0 Arachnida Araneae 2.2 1.8 Acarina 10.7 Crustacea Cladocera 4.0 40.9 66.1 5.2 Ostracoda 12.0 34.1 42.9 • . Copepoda 8.0 18.2 32.1 Amphipoda 56.0 20.0 17.9 15..8 20.0 Insecta Odonata 36.0 29.5 16.1 10.5 20.0 60.0 Coleoptera Helodidae 2.2 20.0 Noteridae 4.5 Haliplidae 10.7 Diptera Psychodidae 3.6 Chaoboridae 1.8 Tipulidae 1.8 Stratiomyidae 8.9 15.8 40.0 Ceratopogonidae 4.5 5.4 5.3 20.0 Culicidae • 1.8 Chironomidae 52..0 4.5 25.0 Ephemeroptera 6.8 7.1 20.0 Trichoptera 5.4 5.3 Hemiptera Corixidae 9.1 8.9 21.1 Naucoridae 6.8 1.8 Fulgoridae 1.8 20.0 Pleidae 8.0 2.2 40.0 Mesoveliidae 2.2 Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 457 Table 12.-Stomach contents of Lepomis symmetricus from Wolf Lake, by size class of sunfish. Figures in parentheses are numbers of stomachs examined. Mean Number of Food Organisms Per Stomach Food Organism <21 21-30 31-40 41-50 51-60 >60 mm mm mm mm mm mm (25) (44) (56) (19) (5) (5) Gastropoda 0.29 0.84 0.60 2.40 Arachnida Araneae 0.02 0.02 Acarina 0.32 Crustacea Cladocera 0.36 6.41 2.14 1.26 Ostracoda 0.48 2.57 0.07 Copepoda 0.12 0.91 1.25 Amphipoda 1.92 0.91 1.20 0.4 12.8 Insecta Odonata 0.52 0.45 0.32 0.11 0.20 1.60 Coleoptera Helodidae 0.05 0.20 Noteridae 0.07 • • Haliplidae 0.54 Diptera Psychodidae 0.02 Chaoboridae 0.52 Tipulidae 0.04 Stratiomyidae 0.25 0.21 4.00 Ceratopogonidae 0.02 0.07 0.05 3.00 Culicidae 0.03 Chironomidae 0.16 0.52 Ephemeroptera 0.07 0.07 0.20 Trichoptera 0.05 0.(; Hemiptera Corixidae 0.18 0.29 0.26 Naucoridae 0.06 0.02 Fulgoridae 0.02 0.20 Pleidae 0.0.8 0.02 0.60 Mesoveliidae 0.02

L. symmetricus when these insects L. symmetricus has been reported to alight on the water surface. eat "dragon-fly nymphs and midge lar- Aquarium-held L. symmetricus fed vae" near Greenwood, Mississippi (Hil- in the typical Lepomis manner. When debrand 8c Towers 1927:134; Cook food was dropped into the water near 1959:180). In 22 specimens from LaRue- them, they sucked it in or swam up Pine Hills, Illinois, the major food near the food item and gulped it down items were "aquatic snails, green algae, before the food item fell to the bottom amphipods, and miscellaneous insects of the aquarium. Occasionally they and insect larvae" (Gunning 8c Lewis fed off the bottom by sucking up food 1955:556) . items. Spawning males and other indi- viduals fed readily on dragonfly naiads, INTERACTION WITH OTHER chironomids, and live and frozen earth- ORGANISMS worms. Miller & Robison (1973:184) reported aquarium-held specimens from Competition Oklahoma feeding on "daphnia and L. symmetricus occurs syntopically small earthworms." with all other described species of Le- 458 ILLINOIS NATURAL HISTORY S URVEY BULLETIN Vol. 31, Art. 10 pomis (including the introduced L. (71.6-240.3 mm SL) , four Pomoxis auritus) except L. gibbosus, from which nigromaculatus (76.4-144.8 mm SL) , it is geographically separated. Because one P. annularis (141.1 mm SL) , five of its preferred habitat of heavily vege- Lepomis gulosus (19.7-127.4 mm SL) , tated, shallow, lentic or slow-moving four L. macrochirus (131.8-140.1 mm water and its relative abundance there, SL), one Centrarchus macro pterus (82.4 it is doubtful that the species is geo- mm SL), and one Ictalurus natalis graphically limited to a great degree by (124.8 mm SL) were preserved and its several congeners. later examined for ingested L. symmet- ricus. These predators were collected Predation from all months of the year except July There are no literature reports of and December. A number of large gar predation on L. symmetricus and no (Lepisosteus oculatus, L. platostomus) evidence of such predation was found were seen during the summer and fall in the Wolf Lake study. As potential months but were not collected. Per- predators five Micropterus salmoides haps these large, relatively common

Table 1 3.-Stomach contents of Lepomis symmetricus from Wolf Lake by month of col- lection. Figures in parentheses are numbers of stomachs examined.

Percent of Stomachs in Which Food Organism Occurred Food Organism Jan. Feb. Mar. April June July Aug. Sept. Oct. Nov. Dec. (17) (18) (15) (9) (7) (43) (17) (10) (9) (13) (18) Gastropoda 29.4 11.1 13.3 55.6 14.3 .. 11.1 Arachnida Araneae . 7.7 5.6 Acarina .. 26.7 22.2 Crustacea Cladocera 17.6 44.4 20.0 33.3 .. 76.5 90.0 77.7 61.5 16.7 Ostracoda .. 50.0 73.3 22.2 11.6 52.9 11.1 23.1 16.7 Copepoda 5.9 33.3 33.3 33.3 4.7 5.9 10.0 . . 23.1 33.3 Amphipoda .. 13.3 11.1 71.4 44.2 35.3 10.0 .. 7.7 11.1 Insecta Odonata 5.9 11.1 26.6 .. 32.6 64.7 .. 30.8 Coleoptera Helodidae 4.7 Noteridae 11.8 Haliplidae Diptera Psychodidae 5.5 Chaoboridae 5.5 Tipulidae 7.7 Stratiomyidae 29.4 11.1 • • 33.3 Ceratopogonidae .. 5.5 6.6 11.1 14.3 11.8 7.7 Culicidae • • 5.9 Chironomidae 5.9 22.2 55.6 7..0 5.9 20.0 .. 11.1 Ephemeroptera • 14.3. .. 38.8 Trichoptera 33.3 .. 5.6 Hemiptera Corixidae 57.1 7.0 70.0 Naucoridae 14.3 4.7 11.8 Fulgoridae 4.7 5.9 Pleidae 14:i 9.3 Mesoveliidae 4.7 Stomach contents were not examined for May-collected specimens. Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 459 predators take some toll on the Wolf hybridization are small (Hubbs 1955: Lake population of L. symmetricus. 2, 18) .

Hybridization Parasitism Schwartz (1972) did not report any The Wolf Lake study population was accounts of hybridization involving L. rather heavily parasitized by plerocer- symmetricus. No evidence of hybrid- coids of the cestode Haplobothrium ization was found in the Wolf Lake glob uliforme. These plerocercoids oc- study area or in specimens examined curred in a total of 44 of 176 stomachs from elsewhere. The small size of L. (25 percent) examined. From one to symmetricus, its preference for shallow, five plerocercoids were found in each vegetated water, and its distinct breed- stomach. Usually the highest numbers ing coloration probably preclude mis- occurred in stomachs of the -1 year mating of the parental species. Since class. Specimens were found during all there is ample habitat available in months of the year except May and Wolf Lake and the fishes are presum- June. The plerocercoid stage of H. ably not unduly crowded, chances of globuliforme normally encysts in the

Table 14.-Stomach contents of Lepomis symmetricus from Wolf Lake by month of col- lection. Figures in parentheses are numbers of stomachs examined. Mean Number of Food Organisms Per Stomach Food Organism Jan. Feb. Mar. April June July Aug. Sept. Oct. Nov. Dec. (17) (18) (15) (9) (7) (43) (17) (10) (9) (13) (18) Gastropoda 0.71 0.17 0.93 1.56 0.29 .. 0.11 Arachnida Araneae .. 0.08 0.06 Acarina .. 0.93 0.44 Crustacea Cladocera 0.06 3.61 0.93 1.67 .. .. 11.35 20.00 13.22 17.23 0.72 Ostracoda .. 7.11 10.40 0.56 .. 0.44 3.00 .. 0.11 0.38 0.78 Copepoda 0.06 1.00 3.40 1.00 .. 0.07 0.06 0.40 .. 0.69 2.11 Amphipoda .. 0.13 0.11 14.28 1.79 2.59 0.10 .. 0.08 0.11 Insecta Odonata 0.06 0.11 0.73 .. 0.42 1.47 .. 0.31 Coleoptera Helodidae .. 0.07 Noteridae 0.18 Haliplidae .. 3.33 Diptera Psychodidae .. 0.05 Chaoboridae .. 1.61 • . . Tipulidae .. 0.15 Stratiomyidae 1.41 0.11 1.22 Ceratopogonidae .. 0.06 0.06 0.11 2.14 0.12 .. 0.77 Culicidae 0.12 Chironomidae 0.41 0.39 1.11 0.09 0.06 0.40 .. 0.22 Ephemeroptera 0.14 .. 0.39 Trichoptera 0.33 .. 0.06 Hemiptera Corixidae 1.00 0.14 1.60 Naucoridae 0.14 0.02 0.12 Fulgoridae .. 0.02 0.14 Pleidae 0.14 0.02 Mesoveliidae .. 0.02 Stomach contents were not examined for May-collected specimens. 460 ILLINOIS NATURAL HISTORY SURVEY BULLETIN Vol. 31, Art. 10 liver of fishes and has been reported name (A. symmetricus) that of the host from a number of other fishes in both instead of the parasite and that "A. this and the adult stage (Hoffman 1967: symmetricus" does not exist. She also 233) . No adults were found in the noted that Cleidodiscus diversus was study population. described from Lepomis cyanellus and One adult specimen of the acantho- that its listing for L. symmetricus was cephalan Pomphorynchus bulbicollz an error in the 1964 Index-Catalogue, was found in the stomach of an L. sym- Trematoda and Trematode Diseases, metricus collected 24 April 1974 at Part 2, that was perpetuated by Hoff- Wolf Lake. Neither the cestode nor man (1967) and the 1969 Index- the acanthocephalan had been known Catalogue. to parasitize L. symmetricus. One collection examined during this Dolley (1933) reported cestodes and study from Texas (TCWC 3643) col- trematodes from "Lepomis symmetricus lected 2 May 1970 was heavily infested in the St. Joseph River of Michigan, (all 32 specimens in the lot) with a but the misidentification of the host monogenetic trematode, presumably species is obvious, since L. symmetricus Anchoradiscus triangularis. No exter- has never occurred in Michigan. Hoff- nal parasites were observed during the man (1967) , who compiled a list of present study. fish parasites, cited for L. symmetricus the trematodes Actinocleidus symmetri- SUMMARY cus, Cleidodiscus diversus, and Ancho- The life-history information on L. radiscus triangularis. Dr. Mary H. symmetricus collected in Wolf Lake be- Pritchard informed me that Hoffman tween 2 June 1973 and 27 May 1975 is (1967) evidently cited as the species summarized in Table 15.

Table 1 5.—Summary of life-history information on Wolf Lake Lepomis symmetricus.

Characteristics Life-History Data Principal habitat Shallow, heavily vegetated margins of standing water Age at reaching sexual maturity I year Size at reaching sexual maturity Females about 34 mm; males about 40 mm Sexual dimorphism Adult males are darker on the head and body have duskier pelvic fins and longer pelvic fins; females tend to have more distinct vertical bars Number of mature ova in preserved females 219-1,600 Description of egg About 0.8 mm in diameter, translucent orange Spawning period From mid-April to early June Spawning habitat Presumably in shallow water, over soft mud bot- tom, near plant material Spawning site Shallow nest depression, about 90-120 mm in di- ameter Influence of sex on growth rate Virtually none Density Up to 0.69 sunfish per square meter Sex ratio among young 1.5 females : 1 male Longevity 3+ years Maximum size 63.0 mm standard length Principal diet Aquatic gastropods, insect immatures, and micro- crustaceans LITERATURE CITED ACKERMAN, K. 1975. Rare and endangered BOUDREAUX, J., K. STRAWN, and G. CALLAS. vertebrates of Illinois. Illinois Department [1959.] Fire ants, heptachlor, and fish kill. of Transportation, Bureau of Environmen- Southwestern Naturalist 3:7-12. tal Science. 50 p. BOYD, J. A., B. M. BURR, L. M. PAGE, and AVISE, J. C., and M. H. SMITH. 1974. Bio- P. W. SMITH. [1975]. A study of threatened chemical genetics of sunfish. II. Genic sim- and/or unique fishes within the boundaries ilarity between hybridizing species. Amer- of the Shawnee National Forest of southern ican Naturalist 108:458-472. Illinois. Pages 1-29 in Those on the brink BAILEY, R. M. 1938. A systematic revision of of doom: a study of rare fishes in the the centrarchid fishes with a discussion of Shawnee National Forest. Illinois Natural their distribution, variations, and probable History Survey and U.S. Department of Ag- interrelationships. Ph.D. Thesis. University riculture Forest Service. of Michigan, Ann Arbor. BRANSON, B. A., and G. A. 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and J. T. DAVIS. [19671 Checklist HANKINSON, T. L. 1908. A biological survey of the freshwater fishes of Louisiana. Lou- of Walnut Lake, Michigan. Pages 157-288 isiana Wildlife and Fisheries Commission, in Michigan Geological Survey state board Baton Rouge. 29 p. report for 1907. . 1975. Checklist of the fresh- HAY, 0. P. 1894. The lampreys and fishes of water fishes of Louisiana. Louisiana Wild- Indiana. Indiana Department of Geology life and Fisheries Commission, Baton Rouge. and Natural Resources Annual Report 19: 29 p. 147-296. EDDY, S. 1957. How to know the freshwater HESTER, F. E. 1970. Phylogcnctic relationships fishes. Wm. C. Brown Publishers, Dubuque. of sunfishes as demonstrated by hybridiza- 253 p. tion. American Fisheries Society Transac- . 1969. How to know the freshwater tions 99:100-104. fishes. 2nd ed. Wm. C. Brown Publishers, II ILDEBRA ND, S. F., and I. L. TOWERS. 1927. Dubuque. x + 286 p. Annotated list of fishes collected in the vi- EVERMANN, B. W. 1899. Report On Investiga- cinity of Greenwood, Miss., with descrip- tions by the U.S. Fish Commission in Mis- tions of three new species. U.S. Bureau of sissippi, Louisiana, and Texas, in 1897. U.S. Fisheries Bulletin 43:105-136. Commission of Fish and Fisheries, Part 24. HOFFMAN, G. L. 1967. Parasites of North Report of the Commissioner for 1898:285- American freshwater fishes. University of 310. California Press, Berkeley and Los Angeles. and W. C. KENDALL. 1894. The fishes viii ± 486 p. of Texas and the Rio Grande basin, consid- Hour, F. T., J. F. KEEFE, W. H. LEWIS, W. L. ered chiefly with reference to their geo- PFLIEGER, and M. H. SULLIVAN. 1974. Rare graphic distribution. U.S. Fish Commission & endangered species of Missouri. Missouri Bulletin for 1892, 12:57-126. Department of Conservation and U.S. De- FORBES, S. A. 1884. A catalogue of the native partment of Agriculture Soil Conservation fishes of Illinois. Illinois State Fish Com- Service, n. p. mission Report for 1884:60-89. IluBBs, C. L. 1955. Hybridization between 1909. On the general and interior fish species in nature. Systematic Zoology distribution of Illniois fishes. Illinois State 4:1-20. Laboratory of Natural History Bulletin 8: , and K. F. LAGLER. 1964. Fishes of the Great Lakes region. 381-437 ± 103 maps. University of Michigan Press, Ann Arbor. 213 p. and R. E. RICHARDSON. [1908.] The HuBBs, C. 1957a. Distributional patterns of fishes of Illinois. Illinois State Laboratory Texas fresh-water fishes. Southwestern Nat- of Natural History [Urbana]. cxxxvi +357 p. uralist 2:89-104. FOWLER, H. W. 1945. A study of the fishes of 19576. A checklist of Texas fresh- the southern Piedmont and Coastal Plain. water fishes. Texas Game and Fish Com- Philadelphia Academy of Natural Sciences mission, Division of Inland Fisheries, IF Se- Monograph 7. 408 p. ries 3. 11 p. GERKING, S. D. 1945. The distribution of the • 1958. A checklist of Texas fresh-water fishes of Indiana. Investigations of Indiana fishes. Revised ed. Texas Game and Fish Lakes and Streams 3:1-137. Indiana Depart- Commission, Division of Inland Fisheries, ment of Conservation, Indianapolis, and In- IF Series 3. 14 p. diana University, Department of Zoology, . 1961. A checklist of Texas fresh- Bloomington. water fishes. Texas Game and Fish Com- GOWANLOCH, J. N. 1933. Fishes and fishing mission, Division of Inland Fisheries, IF Se- in Louisiana. Louisiana Department of ries 3. 14 p. Conservation Bulletin 23. 638 p. . 1972. A checklist of Texas freshwater GREENE, C. W. 1927. An ichthyological sur- fishes. Texas Parks and Wildlife Depart- vey of Wisconsin. Michigan Academy of Sci- ment Technical Series No. 11. a ± 11 p. ences, Arts and Letters 7:299-310. . 1976. A checklist of Texas freshwater . 1935. The distribution of Wisconsin fishes. Revised ed. Texas Parks and Wild- fishes. Wisconsin Conservation Commission, life Department Technical Series No. 11. Madison. 235 P. ii ± 12 p. GUNNING, G. E., and W. M. LEWIS. 1955. The , and J. PIGG. 1976. The effects of im- fish population of a spring-fed swamp in the poundments on threatened fishes of Okla- Mississippi bottoms of southern Illinois. homa. Oklahoma Academy of Science Pro- Ecology 36:552-558. ceedings. 5:113-117. and . 1956. Recent collections JENKINS, R. E., E. A. LACHNER, and F. J. of some less common fishes in southern Illi- SCHWARTZ. 1971. Fishes of the central Ap- nois. Illinois State Academy of Science palachian drainages: their distribution and Transactions 48:23-26. dispersal. Pages 43-117 in P. C. Holt, R. A. Sept., 1977 BURR: THE BANTAM SUNFISH, LEPOMIS SYMMETRICUS 463

Paterson, and J. P. Hubbard, editors, The MCREYNOLDS, H. E. 1975. Threatened species: distributional history of the biota of the a review of the Eastern National Forests' southern Appalachians. Part III: Verte- studies of these animals. Indiana Academy of brates. Virginia Polytechnic Institute and Science Proceedings 84:250-257. State University Resources Division Mono- MILLER, R. R. 1972. Threatened freshwater graph 4. fishes of the United States. American Fish- JORDAN, D. S. 1884. List of fishes collected in eries Society Transactions 101:239-252. the vicinity of New Orleans by Dr. R. W. MILLER, R. J., and H. 'W. ROBISON. 1973. Shufeldt, U.S.A. U.S. National Museum Pro- The fishes of Oklahoma. Oklahoma State ceedings 7:318-322. University Press, Stillwater. xiii ± 246 p. . 1888. A manual of the vertebrate Mims, H. B., W. C. STARREIT, and F. C. BELL- animals of the northern United States. 5th ROSE. 1966. Man's effect on the fish and ed. A. C. McClurg and Company, Chicago. wildlife of the Illinois River. Illinois Nat- iii ± 375 p. ural History Survey Biological Notes 57. 24 p. . 1929. Manual of the vertebrate ani- MIZFLLE, J. P. 1938. Studies on monogenetic mals of the northeastern United States in- trematodes. IV. Anchoradiscus, a new dact- clusive of marine species. 13th ed. World ylogyrid genus from the bluegill and the Book Company, Yonkers-on-Hudson, New stump-knocker sunfish. Journal of Parasitol- York. xxxi ± 446 p. ogy 27:159-163. , and C. H. GILBERT. 1883. Synopsis of and R. C. HUGHES. 1938. The North the fishes of North America. U.S. National American fresh-water Tetraochinae. Amer- Museum Bulletin 16. LVI + 1018 p. ican Midland Naturalist 20:341-353. , and B. W. EVERMANN. 1896. The fishes MOORE, G. A. 1952. A list of the fishes of of North and Middle America. U.S. National Oklahoma. Oklahoma State Game and Fish Museum Bulletin 47:955-1240. Department, Oklahoma City. n. p. , and H. W. CLARK. 1930. 1973. Discovery of fishes in Oklahoma Check list of the fishes and fishlike verte- (1852-1972) . Oklahoma Academy of Science brates of North and Middle America north Proceedings 53:1-26. of the northern bouncpry of Venezuela and , and F. B. Caoss. 1950. Additional Colombia. U.S. Commissioner of Fisheries Oklahoma fishes with validation of Poeci- Report for the Fiscal Year 1928, Appendix lichthys pawipinnis (Gilbert and Swain) . X. 670 p. Copeia 1950:139-148. JURGENS, K. C., and C. HUBBS. 1953. A check- O'DONNELL, D. J. 1935. Annotated list of the list of Texas fresh-water fishes. Texas Game fishes of Illinois. Illinois Natural History and Fish 11:12-15. Survey Bulletin 20:473-500. KNAPP, F. T. 1953. Fishes found in the fresh- PAGE, L. M. 1974. The life history of the spot- waters of Texas. Ragland Studio and Litho tail darter, Etheostorna squamiceps, in Big Printing Company, Brunswick, Georgia. viii Creek, Illinois, and Ferguson Creek, Ken- + 166 p. tucky. Illinois Natural History Survey Bio- KUHNE, E. R. 1939. A guide to the fishes of logical Notes 89. 20 p. Tennessee and the Mid-South. Tennessee PFLIEGER, W. L. 1966. A check-list of the fishes Department of Conservation, Nashville. 124 p. of Missouri, with keys for identification. Mis- LAMB, L. D. 1941. A preliminary fisheries sur- souri Conservation Department Division of vey of the San Jacinto watershed. Texas Fisheries. D-J Series 3. 63 p. Academy of Science Transactions 24:42-48. . 1968. Checklist of the fishes of Mis- LAMBOU, V. W. 1962. Fishes occurring in Lake souri with keys for identification. Missouri Bistinou, Louisiana. Louisiana Academy of Conservation Department Division of Fish- Science Proceedings 25:75-79. eries. D-J Series 3. 64 p. LARGE, T. [1903.] A list of the native fishes of . 1971. A distributional study of Mis- Illinois, with keys. Appendix to Report of souri fishes. University of Kansas Publica- State Board of Fish Commissioners from tions, Museum of Natural History 20:225- September 30, 1900 to October 1, 1902. 30 p. 570. LARIMORE, R. W. 1957. Ecological life history . 1975. The fishes of Missouri. Missouri of the warmouth (Centrarchidae) . Illinois Department of Conservation [Jefferson City]. Natural History Survey Bulletin 27:1-83. viii + 343 p. LOPINOT, A. C., and P. W. SMITH. 1973. Rare PRATT, H. S. 1923. A manual of land and and endangered fish of Illinois. Illinois De- fresh water vertebrate animals of the United partment of Conservation Division of Fish- States (exclusive of birds) . P. Blakiston's eries, Springfield. 53 p. Son & Company, Philadelphia. xv + 422 p. McKAY, C. L. 1882. A review of the genera REEVES, J. D., and G. A. MOORE. [1951.] Lepo- and species of the family Centrarchidae, with mis marginatus (Holbrook) in Oklahoma. a description of one new species. U.S. Na- Oklahoma Academy of Science Proceedings tional Museum Proceedings 4:87-93. 30:41-42. 464 ILLINOIS NATURAL HISTORY SURVEY BULLETIN Vol. 31, Art. 10

RICHARDSON, R. E. 1904. A review of the sun- SMITH, P. L., and M. E. Stsx. 1969. The fishes fishes of the current genera Apomotis, Lepo- of west Kentucky. II. The fishes of Obion mis, and Eupomotis, with particular refer- Creek. Kentucky Academy of Science Trans- ence to the species found in Illinois. Illinois actions 30:60-68. State Laboratory of Natural History Bulletin SMITH, P. W. 1965. A preliminary annotated 7:27-35. list of the lampreys and fishes of Illinois. ROBERTS, F. L. 1964. A chromosome study of Illinois Natural History Survey Biological twenty species of Centrarchidae. Journal of Notes 54. 12 p. Morphology 115:401-418. 1971. Illinois streams: a classification ROBISON, H. W. 1974. Threatened fishes of based on their fishes and an analysis of fac- Arkansas. Arkansas Academy of Science Pro- tors responsible for disappearance of native ceedings 28:59-64. species. Illinois Natural History Survey Bio- . 1975. New distributional records of logical Notes 76. 14 p. fishes from the lower Ouachita River system [1973.] A key to the fishes of Illinois. in Arkansas. Arkansas Academy of Science Illinois Department of Conservation Fishery Proceedings 29:54-56. Bulletin 6. 43 p. , G. A. MooRE, and R. J MILLER. 1974. , and D. W. BRIDGES. 1960. Ichthyolog- Threatened fishes of Oklahoma. Oklahoma ical type specimens extant from the old Illi- Academy of Science Proceedings 54:139-146. nois State Laboratory of Natural History. ROZENBERG, E. R., R. K. STRAWN, and W. J. Copeia 1960:253-254. CLARK. 1972. The composition and distri- , A. C. LOPINOT, and W. L. PFLIEGER. bution of the fish fauna of the Navasota 1971. A distributional atlas of upper Missis- River. Texas Water Resources Institute sippi River fishes. Illinois Natural History Technical Report 32. iii ± 120 p. Survey Biological Notes 73. 20 p. SCHLAIKJER, E. M. New fishes from the 1937. SUMMERS, W. A. 1937. A new species of Tetra- continental Tertiary of Alaska. American onchinae from Lepomis symmetricus. Jour- Museum of Natural History Bulletin 74:1-23. nal of Parasitology 23:432-434. SCHWARTZ, F. J. 1972. World literature to fish , and H. J. BENNETT. 1938. A prelimi- hybrids with an analysis by family, species, nary survey of the trematodes from the gills and hybrid. Gulf Coast Research Laboratory, of Louisiana fishes. Louisiana Academy of Ocean Springs, Mississippi. 328 p. Science Proceedings 1:247-248. (Abstract) . SEAMSTER, A. 1948. Gill parasites from Louisi- WALKER, J. M. Fishes in north Louisi- ana fishes with a description of Urocleidus 1962. wadei n. sp. American Midland Naturalist ana. Louisiana Academy of Science Proceed- 39:165-168. ings 25:35-41. SCHRENKEISEN, R. 1938. Field book of fresh- 1963. Fishes in Choudrant Bayou. water fishes of North America. G. P. Put- Louisiana Academy of Science Proceedings nam's Sons, New York. 312 p. 26:45-48. SEEHORN, M. E. 1976. Fishes of Southeastern WEBB, D. H., and M. E. Sisx. 1975. The fishes National Forests. Southeastern Association of of west Kentucky. III. The fishes of Bayou Game and Fish Commissioners Twenty-ninth de Chien. Kentucky Academy of Science Annual Conference Proceedings for 1975: Transactions 36:63-70. 10-27. WHITAKER, J. 0., JR. 1968. Keys to the verte- SHARMA, M. S. 1964. The cephalic lateral-line brates of the eastern United States, excluding system in Notopterus chitala (Ham.) . Copeia birds. Burgess Publishing Company, Minne- 1964:530-533. apolis. iii ± 256 p. INDEX A juvenile, 444 Adjusted body weight, 438, 452 Age attained (see longevity) Age composition, 455 Aging method, 438 Associated species (see species associates) Largest specimen examined, 442, 445 Lectotype designation, 442 Life history in Wolf Lake, 449-460 Locations and dates of breeding, 452-453 Longevity, 455 Breeding dates (see locations and dates of breeding) Breeding male, 443, 450-451 Museum abbreviations, 437

Collecting methods, 438 0 Coloration (see description) Ovarian size and weight variation, 451-452 Competition, 457-458 Conservation status, 448-449

Parasitism, 459-460 Predation, 458-459 Density per square meter, 454, 455 Prejuvenile, 444 Demography, 454-455 Description coloration, 443-445 general, 443-445 Relationships, 446, 450 Development and growth, 453-454 Relative survival, 455 Diagnosis, 442-443 Relative survival calculation, 439 Diet Reproductive cycle general, 455-459 of female, 451 of young, 456-457 of male, 450-451 seasonal variation, 456, 458-459 Distribution, geographic, (iv) Drainages, 445-447 Sexual differences (see variation, sexual) Size distribution, 454 Eggs Spawning, 452-453 number of, 451-452 Species associates, 450 Specimens studied, 446 447 Study area, 449 Survival (see relative survival) Synonymies and synonyms, 439 440 Genital papillae, 445, 450-451 Syntopic species of sunfishes, 457-458 Geographic distribution (see distribution, geographic) Growth (see development and growth) Types, 440, 442

V Habitat, 449-450 Hatching (see development and growth) Variation Hybridization, 459 allometric, 445 geographic, 440-442, 445-446 meristic, 440-441, 445 morphometric, 442 Illinois Fish Code, 438 ontogenetic, 444-445 Interactions with other organisms, 457-460 sexual, 445 465 466 ILLINOIS NATURAL HISTORY SURVEY BULLETIN Vol. 31, Art. 10

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CONSULTANTS AND RESEARCH AFFILIATES: SYSTEMATIC ENTOMOLOGY, RODERICK R. Dtwnr, Chicago, Illinois; WILDLIFE RESEARCH, WILLARD D. KLIMSTRA, Ph.D., Professor of Zoology and Director of Cooperative Wildlife Research, Southern Illinois University; PARASITOLOGY, NORMAN D. LEvINE, Ph.D., Professor of Veterinary Parasitology, Veterinary Research and Zoology and Director of the Center for Human Ecology, University of Illinois; ENTOMOLOGY, ROBERT L. METCALF, Ph.D., Professor of Biology and Research Professor of Entomology, University of Illinois; and GILBERT P. WALDBAUER, Ph.D., Pro- fessor of Entomology, University of Illinois; STATISTICS, HORACE W. NORTON, Ph.D., Professor of Statis- tical Design and Analysis, University of Illinois.