A Life History Study of the Silver Chub, Hybopala atoreriana (Kirtland), in Western Lake Erie with Notes on Associated Species

DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University

by Edward Coyle Kinney, Jr. B.Sc. in Edu., B.Sc. in Agr., M.Sc. The Ohio State University

1951*

Approved by

XS VT ser ' s*--- TABLE OF CONTENTS Page Number A LIFE HISTORY STUDY OF THE SILVER CHUB, HYBOPSIS STORKRIANA (KIRTLAND), IN WESTERN LAKE ERIE...... 1-77 Introduction...... 1 Description...... k Synonyms...... 6

Collection Methods...... 11 The Otter Trawl as a Sampling Device...... 1$ Attempts to Locate the Young...... 21+ Age, Orowth and Abundance...... 28 Mortality...... Ul Distribution Records...... I4J4. Distribution Ecology...... ij.8

Food during the Period September, 195^ - August, 1953...... 55 Reactions to Decrease in Food Supply...... 61 Estimated Food Consumption...... 65 Reproduction Data...... 67 Economic Value...... 76 ASSOCIATED SPECIES...... 78-86 Introduction...... 76 Dorosoma cepedl ...... 78 Coregonus clupeaformis...... 79 Osmerus mordax...... 79 Catostomidae...... 60 i Cyprinus carplo...... 80 Caraaalua auratas...... 80

Notropla atherlnoldea...... 80 Notropl a hudaonlua...... 80 Notropla volacellua...... 8l Ictalurua punctatua...... 81 Amelurua ap...... 81 Noturua f lavua ...... Ql Schllbeodea mlurus...... 81 Lota lota lacuatrla...... 82 Percopala omlacomaycua...... 82 Morone chryaopa...... 82 Parca flaveacona...... 83 Stlzoatadlon vltreum...... 83 Stlzoattdlon v. Klaucum...... 8I4. Stlzoatedlon £. canadense...... 8I4. Hadropterua copalandl...... 8^ Perclna caprodea aemlfaaclata...... 8$ Gantrarchidae...... 85 Ambloplltea rupeatrla...... 85 Lepomla ap,...... 85 Aplodlnotus grunnlena...... ♦...... 85 Cottua rlcel...... *...... 85 Cottua Balrdl kumllonl...... 85 III. LITERATURE CITED...... 87 IV. AUTOBIOGRAPHY...... 99 11 ILLUSTRATIONS

Figure Pago Nunibor Number 1. Map of Wo stern Lake Erie...... 1

2. Variation in Scale Pattern...... 6

3. Measurements of Trawl Spread...... 13

4. Average Number of Silver Chubs per Haul...... 20

5. Number of Silver Chubs per Haul...... 22

6 . Scales from Representative Specimens from the September, 1952, Collection...... 29

7. Annulus formation...... 30 8 . Scales with Doubtful First Annull...... 31 9. Scales from Specimens Representing the Size Range of the One-Year Class...... 32 10, Total Longtn-Standard Length Correlation...... 35

11* Length-Welght Correlation...... 36

12. Standard Length-Anterior Scale Radius Correlation 37

13. Growth and Water Temperature...... 38 ll;. Distribution of the Silver Chub In North America. 1^8 15. Food of the Young of the Year...... 56

16. Food of the Adults...... 56

17. The 1953 Spawning Period...... 68

18. Variation in Weight of Mature Ovaries...... 69

19. Correlation of Body Weight - Ovary Weight During the 1953 Pre-spawning Period...... 70 20. Ovary Shape...... 71

21. Correlation of Ovary Weight and Number of Eggs... 72 ill TARLES

Table Page Number Number 1. Variation In Scale Pattern Between Pectoral Fins...... 7 2. Collection Data...... 21 3. Collection Data...... 23 4. Sex Ratios by Year Classes...... 39 5. Average Standard Length and Range...... 40 6 . Dead Collected on March 28, 1952...... 1+1 7. Dead Collected on May 23* 1992...... 42 8 . Food of the Young of the Year...... 59 9. Food of the Adults, September, 1992 - August, 1953...... 60 10. Food of the Adults, September, 1953 - March, 1951+*...... 64 11. Dorosoma cepedlanum Collections...... 79 12. Oamerus mordax Collections...... 79 13. Morone cnryaops Collections...... 02 14* Perea flavescens Collections...... 83 15. Stlzoatedion vltreum Collections...... 84

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Figure 1.— Map of western Lake Erie. A LIFE HISTORY STUDY OF THE SILVER CHUB, HYBOPSIS STORERIANA (KIRTLAND). mteSTm LAKE ERIE

The silver chub, Hybopsis storerlana (Kirtland), is a minnow common in Lake Erie, and in the large lakes, impound­ ments, and major streams of the Mississippi River drainage system. There are abundant distribution records of this species but little research has been conducted toward its life history. E. E. Good, while a student at Stone Institute during the summer of 1946, made a brief study of the silver chub. His work consisted of the examination of 35 specimens and a survey of the literature. Food habit studies were made on a few Lake Erie speci­ mens by Sibley (1929) and by Boesel (1937). Marie Poland Fish (1935) described the larval stages of the silver chub. A study of the parasites of Hybopsls atorerlana was made by Bangham and Hunter (1939). George A. Moore (1950) examined the cutaneous sense organs of this barbeled minnow.

The major portion of this study was conducted from the Autumn of 1952 through the Spring of 1954* A few collections were made during the period from 1946 to 1952. Most of the collections were made with a 30-foot otter trawl, within a 10-mlle radius of the Frans Theodore Stone Institute of Hydro-

-2- biology, Put-ln-Bay, Ohio (Pig. 1). The majority of the collections were made over mud bottom containing varying amounts of silt, clay, sand, and small gravel (Wood, 1953)* Most collections were made within two feet of the bottom at depths of 9 to 11 meters. The author wishes to acknowledge the assistance of the following persons: Dr. Thomas H. Langlols, who made this study possible and whose advice guided the study; Dr. E. E. Good, whose excellent preliminary work served as a starting point for this study; Dr. Jacob Verduin, who aided In statistical analyses; Dr. N. Wilson Britt, who rendered assistance In the identification of invertebrates found in the stomach contents; Dr. Milton B. Trautman, who helped with the fish taxonomy phases of this work; Mr. Paul V. Webster, who operated the Institute research boat, the BIO-LAB, and who helped Immensely with the collect­ ing activities.

-3- Descriptions of the Silver Chub, Hybopsla storsrlana (Kirtland)

During a meeting of the Boston Society of Natural History, on June 1$, 181+2, Dr. D. Humphrey Storer mentioned that he had received the descriptions of three species of fishes from Dr. J. P. Kirtland of Ohio (Kirtland, I8I4J4.K One of these species was Rutllua storerlanus# This original description of the silver chub, which Included a drawing, was published by the society in 18[4.7 . Kirtland had been submitting to the Boston Journal a series of articles, under the title, "Descrip­ tions of the Fishes of Lake Erie, the Ohio River, and Their Tributaries•" For some reason, possibly a printer's error, the words, "Lake Erie," were omitted from the title of the article containing the original description of the silver chub. I am convinced the omission was accidental rather than by the author's intention, because Kirtland gives the habitat for this minnow as Lake Erie. In a later publication (1850), in refer­ ring to the fishes of Ohio, Kirtland again gives the habitat as, "Lake Erie," and nowhere does he mention having collected the silver chub in the Ohio River* Kirtland's description of the adult silver chub does not Include a maxillary barbel, which is a key taxonomic character for this species* His description could be applied to Notropla hudsonlus were it not for the slse, given as eight inches* His description of the young, which is discussed later in this pa­ per. Is Incorrect. He gives the common name as, "Lake minnow." This name is currently used In the Lake Erie region when refer­ ence to Notropla a. atherlnoldea. the lake emerald shiner, is Intended. This minnow would fit Kirtland'a description of the young. Excellent descriptions of Hybopsla atorerlana may be found In Cox (1897)# Forbes and Richardson (1920), Jordan and Ever- mann (1896), and O s b u m (1901). There is some disagreement as to the location of the Inser­ tion of the dorsal fin with respect to the pelvic fins. Hubbs and Lagler (19L|.6) state that the dorsal fin is inserted dis­ tinctly In advance of the pelvic fins. Jordan and Evermann (op.clt.) state that the dorsal fin la Inserted over the ven- trals. In this study, the location of the dorsal fin was found to vary among Individual specimens. In some it was in advance of, in others It was over, and In a few it was slight­ ly posterior to the ventral fins.

No external difference between the sexes was found, other than the usual swollen condition of the females immediately prior to the spawning period. In the search for sexual differences, it was noted that the scale pattern on the ventral region, between the pectoral fins, varied from a complete lack of scales to a fully scaled pattern (Fig. 2). A record of this pattern was maintained because of the possible genetic significance. The soale pat- Figure 2.— Variation in the 3cale pattern in the ventral region between

the pectoral fins. tern on the Lake Erie specimens was compared with the pattern on the Ohio River specimens, from which they are geographi­ cally Isolated (Table !)•

TABLE 1

Variation In the Scale Pattern Between the Pectoral Pins Naked Partially Scaled Scaled Number Percent Sumber Percent Number Percent Lake Erie Males 191*. 2 8.0 205 29.5 296 14-2.5 Females 285 30.7 223 23.9 1*2 3 14-5. k Both Sexes 1*79 29.14. I4.28 26.3 719 54.2 Ohio River ko 71.5 15 26.7 1 1.8

It would be necessary to examine a larger number of Ohio River specimens before a significant analysis could be made. The approximate ratios, based on the above data, are, as follows: Lake Erie 1 naked 1 partially scaled 2 scaled Ohio River 3 naked 1 partially scaled 0 scaled The pectoral area Is richly populated with taste buds (Moore, 1950). The numbers of taste buds in this area are about the sane for scaled or naksd specimens. There is the possibility that the taste buds on the scaled specimens are less sensitive because they are underlain by scales.

Synonyms for Hybopsla atorerlana (Kirtland) There exists a great confusion of names applied to the silver chub, Hybopsla atorerlana (Kirtland). The following names are probably true synonyms for Hybopsls atorerlana; Rutllua storerlanua Kirtland, 18J+2 (I8I4J4.)* Lake Erie;

Leuclacus storerlanua Kirtland, I8I4.5 (I8I4.7 ), Lake Erie; Ooblo vernalla Girard, I8I4.5 (Jordan and Evermann, 1896); Ceratlchthya lucens Jordan (1879), Palls of the Ohio River; Hybopsls storerlanua Jordan and Meek (1885), Iowa; Erlnemus storerlanua Jordan (1921j.); Hybopsls storerlanua (Hubbs, 1929); Hybopsla atorerlana (Bailey, 1951).

The researcher la likely to encounter difficulty when working with this species because of the various generic names used, the various specific names used, and the con­ founding of Hybopsla atorerlana with Motropla hudaonlua (Clinton)•

In regard to the generic names used, Hubbs (1929) says, "The generic name Hybopsls has had a vacillating history since its proposal by Agassiz in 185(4.." Jordan (19214.) suggested

-8- that, "...the genus called Hybopsla by Jordan and Evermann (1896) was a composite group, and the name Hybopsls does not belong to any or Its members." He proposed the generic name Erlnemus for the silver chub. The name Erlnemus storerlanua was adopted by some authors and was used for several years. Hubbs (1929) suggested that provisionally, storerlanua should be retained in the genus Hybopsla. This latter suggestion was universally accepted until Bailey (1951) made what was prob­ ably the final change. For the period of some forty years (181*2-1885)» three different specific names were used. Since there are only three of these, they are easily kept in mind and do not cause too much confusion. Greater confusion resulted from the confounding of Hybopsls storerlana with Hotropls hudaonlus. Evermann and Bollman (1886) listed twenty-three synonyms for Notropls hudaonlus» among which were the following: Hybopsla storerlanua Cope"J Proc. Acad. Nat. Sci •, Phi la., 1861*, 299 (Michigan) Cope, Cypr. Penn. 1866, 386 (Delaware and Potomac Rivers) Jordan, Geol. Rept., Indiana, 1871+# Nelson, Bull. 111. Mus. Nat. Hist., I, 1*6, 1 8 7 6 . Leuclacus storerlanua Gunther, Cat. Fish., VII, 250, 1868 (Susquehanna River). Alburnops storerlanua Jordan, Cat. Fresh Water Fish. N.A., 1+19, 1 8 7 8 ; Hudaonlus storerlanua Jordan, Man. Vert., 290, 1880.

Jordan (1879) aptly described the silver chub as Ceratlchthya lucena. But three years later (1882), he described a apeciea which he called Hudaonlus storerlanua (Kirtland) Jordan, which waa suppoaed to have been the same species as Kirtland'a Leuclacus atorerlanua. This apeciea did not have maxillary barbela and so was probably not the silver chub* The lateral line count was correct for N. hudsonlus* but no mention was made of a caudal spot, and the size was given as eight Inches or longer, which la too long for Jl. hudaonlus * Since the larger specimens of !i. hud sonlua are closely associated with H. storerlanua. It Is possible that he had both species and took parts of his description from each of them*

-10 Collection Methods

l’he most successful collections were made with a 30-foot otter trawl* The method of operation of the 30-foot trawl, as described by Daiber (19$0), was changed In some respects. 1. The towline was shortened from 100 feet to 60 feet. This decreased the time consumed in lowering and raising the

trawl. Although the boards still dug into the mud bottom, less mud was thrown into the net. Approximately a 2 to 1 ratio between the towline and the depth was maintained. 2. The bridle line was shortened from 30 feet to about 22 feet, so the boards could be towed aboard with the power winch. 3* A pipe was used to connect the stern davits, then the towrope and bridle were pulled over this pipe, raising the otter boards to deck level. This eliminated the former, dangerous practise of reaching over the stern into the water to grasp the heavy otter boards. Formerly, two persons were required for operation of the trawl, one man being needed to raise each of the two otter boards. After the above changes were made, only one man was required to guide the boards as they were lowered to the deck. il* The towing speed of the workboat was increased from 800 r.p .m. to 1000 r.p.m. This increased the distance covered, compensating for the loss of width opening caused by the

-11- shortening of the bridle. It is reasonable to assume it also served to catch some of the fish that would have escaped at the slower speed. 5. The trawl itself was modified by the insertion of an 8-foot trawl net of quarter-inch mesh. This small mesh net was patterned after that described by Bullis (1951) «s a no-overhang flat trawl. The opening was made rigid by the use of a frame of half-inch water pipe. A short tunnel with a 6-inch iron ring at the end of it was enclosed in the mouth. This small trawl was detachable; it could be used separately with small otter boards to which 8-foot wings were attached. it was used for trawling over rock bottom and in shallow areas. In operation, the smaller net had much less tension than did the larger one, and the operator, by holding the towline, could tell whether it was dragging too much on the bottom. If this was the case, the towline was shortened until the trawl barely skimmed over the bottom, rfith tills net we were able to capture fry as small as 15 millimeters in total length. By trigonometric methods, the truwl opening was com­ puted to be about 13i feet. The angle at the apex of the bridle was computed by towing the empty trawl at 1000 r.p.ra. with the tow end of the bridle barely in the water, and then measuring across the bridle at a marxed distance from the apex (Pig. 3)*

-12- 11.5 ft

Figure 3*— Measurements of the trawl spread.

The actual trawl opening would tend to decrease aa the load increased. This decrease might be offset or even more than offset by additional friction between the board runners and the lake bottom.

The trawl speed was determined by timing several runs over a known-distance course. A speed of about 3*1 statute miles per hour was determined. Since we trawled with the waves and, therefore, usually with the wind, on rough days the speed was increased slightly. By trawling for 12 minutes we covered a rectangle 13i feet oy 3,2i>0 feet, or an area of i4-3*875> square feet, which is about one acre (^3,560 sq. ft.). In the roughest weather

13 wo experienced, wo covered an area of approximately *4.7,925 square feet, or about 1.1 acres per haul.

Other Collecting Devices Used 1. Four-foot by six-foot common sense seines and 4 -foot by 15-foot knotted seines were used to seine the shore areas along South Bass Island. No silver chubs were captured by this method. 2. A double funnel trap was constructed from a 55- gallon oil drum. This was effective in capturing rock bass, young smallraouth bass, spottail minnows, bullheads, crayfish (Orconectes proplnquus), and mud puppies (Necturus maculosus)• 3. The push-selne, which was devised and operated by Anthony Bodola (1952), was tried with success in capturing shallow-water species, but no silver chubs were taken. We also operated it as a tow seine In deeper water and caught small white bass and sheepshead.

if. A hoop minnow net was fished through the ice, but was effective only to depths of 15 feet; no silver chubs were taken. The Otter Trawl as a Fish Population Sampling Device

A total of 33 collections consisting of 90.5 Individual hauls of 12 minutes each were made with a 30-foot otter trawl in the vicinity of the Island region of western Lake Erie (Fig. 1, Table 2). Seven of these collections were made between 191J.8 and 1951* The remaining collections were made during the September, 1952 - September, 1953 period. The average number of adult Individuals per haul was 21.0, with a standard deviation of 15.9. After the exclusion of two collections which deviated greatly from the mean, an average of 18.7 adults was obtained, with a standard devia­ tion of 5*7 (Fig. I*). Of the two excluded collections, one, on May 27, 1953# about two and a half miles north of the mouth of Sandusky Bay, netted 293 silver chubs in three hauls. Evidently, this collection was made in the center of a large school returning to the open lake from a spring shoreward migration. A San­ dusky Bay haul the same day netted 18 chubs. The other collection was made near Green Island on April 7» 1953t and netted only six chubs in three hauls. During March and early April, the silver chubs leave the open lake and are present In large numbers along the south shore and in the lower reaches of the tributary streams. Evidence

-15- of this migration was noted by the author on March 19, 1951, at Catawba Dock, located at the northern end of Catawba Island* Sports fishermen were catching large numbers of silver chubs on minnows and worms* The dockman, 6111 Good, told me they catch them only during March. Harold Wasco, of the Toledo Aquarium, said silver chubs were taken In fyke nets In East Harbor and Rush Creek, from March 15 through early April in 1953* They were also noted by Frank Llgas, a Stone Institute student, in Turtle Creek, during the north­ ern pike spawning run in late March of 1952* Evidence that they are not present In these areas during the summer la sup­ ported by the Stone Institute seining records* Fyke net operations by Carter Gilbert of the Ohio Division of Wildlife, in East Harbor, during the period June 28 - July 3, 1953, failed to show any silver chubs* The area of the large collection was trawled again 15 days later, on June 11, 1953, and an average of 16*8 chubs per haul were taken* On the basis of the above data, the author feels justi­ fied in excluding these two collections from the standard deviation computation* Records of the numbers of Hybopsls atorerlana per haul were kept for 56*5 hauls made during 1952 and 1953 (Table 3)* A mean of 19*6 Individuals per haul was obtained, with a stan­ dard deviation of 6*7* When the four larger hauls were ex-

-16- eluded, a standard deviation of was obtained (Fig, 5). If all hauls are considered, the departure from Polsson distribution Is highly significant. If the hauls from the \ two largest and three amai^est collections ere excluded, a v chl>square of 12, 21* with eight degrees of freedom Is obtained Although this chi-square Is not significant, It Is rather high, and Indicates that the sliver chubs are not distributed entirely at random. It is probable that the silver chubs, \ except for the spring months mentioned, are scattered In loose schools throughout the open lake In the island region of the western basin of Lake Krie, The data gathered during the 1952-1953 period are sufficiently consistent to enable one to use them as a basis for future population comparisons. The possibility of using the otter trawl to make com­ parative population estimates of other species shows promise. On the basis of the work accomplished during the past five years It Is believed that with our present equipment, compare tlve estimates could be determined for the following species: Adults Young Hybopsls atorerlana Aplodlnotus grunnlens Notropla hudaonlus Morone chrysops Percopsls omlscomaycua Perea flaveaoens Percopsla omlscomaycua These species are caught in considerable numbers during all seasons of tAe year. Recently, through ^he use of a small otter trawl. which can ’-^>e ^owed more safely over rooky bottom, many young Stlgostedlon vltreum have been taken. Per­ haps this species could be added to the above list. If one were able to obtain an estimate of the number of fish that elude the trawl, it would be possible to obtain a fair estimate of the total population of the above-mentioned species. The British are attempting to do this through the use of frogmen observers and underwater movie cameras

(Anonymous, 1951* 1952). Thus far, they have worked only In areas having so few fish that no Justifiable conclusions can be drawn from their observations. They have made some excellent pictures of the otter trawl in action. The species list previously mentioned could be enlarged through the use of the following additional equipment: 1. Any-Depth Trawls (Anonymous, 1949; Kobayashi et al.,

1951). Trawls that may be towed at any desired depth are now in use off the northern Atlantic and Pacific coasts.

2. Echo sounders (Anonymous, 1949; Hasler, 1953)* Echo sounders are now being used In conjunction with the any-depth trawls. The sounder is used to locate the schools and to check the depth of the trawl. Conclusions: 1. The otter trawl can be used as a sampling device for yearly comparisons of Hybopsla atorerlana populations in western Lake Erie.

-IB 2. If the number of sliver chubs that elude the travl could be determined, a fair estimate of the total population could be made. 3. The populations of other mud-bottom-dwelling species could be determined, I4.. With the aid of additional equipment and appropriate methods, it might be possible to adequately sample Lake Erie fish populations by use of the trawl.

-19- OTTO TltUL COLLBCTIOM mi j-| I *~| I'M -51 CoiiMti or Bybowli itororlopo

I on Avoraf* por 1952-1953

of

toot of South M m Island

10 h i t of South a h » I iliod

Figure U*— Frequency of the number of silver chubs taken per haul

based on the average per collection. T A B L E 2

COLLECTION DATA Hybopals storerlani (Adults)

Coll • No. No. Av.No . ]Date Location No. Hauls Chubs per Haul

None 1 24 24 17 Aug 49 Between 3 . Bass 5c Rattlesnake 8699 1 21 21 24 Sep 48 Between S.Bass 5c Kelleys 8669 2 43 21.5 13 Sep 1 ml* west of Rattlesnake n it if8 8653 2 42 21.0 Between S.Bass 5c Kelleys 8682 3 56 18.7 20 Sep n ti it n w 8692 3 66 22.7 n n ft® West of Rattlesnake 1 1 20 20 26 Apr 51 Between S.Bass 5c Kelleys 2 6 106 17.7 22 Sep 52 n n it it it 3 2 29 14.5 6 Oct 52 n n it it it 4 2 15 7.5 22 Oct 52 it ii n it it 5 3 6k 21.3 13 Nov 52 Between Green 5c Rattlesnake 6 3 59 19.7 Nov 52 ti n it it 7 3.3 56 17.0 8 Dec 52 Between S.Bass 5c Kelleys 8 3 48 16.0 22 Dec 52 it n n it it it n It 9 3 1*7 15.7 Between Green 5c Rattlesnake 10 2.7 98 38.5 20 Jan 53 n it it n 11 2 42 21 10 Feb 53 Between S.Bass 5c Kelleys 12 3 29 9.7 12 Mar it it n it »t 13 2 15 7.5 it it P NW of Kelleys 14 3 6 2 3 Apr 53 Between Green 5c Rattlesnake 15 k 100 25.0 29 Apr 53 it n n n 16 5 122 15 May 53 South of Niagara Reef 17b 1 18 27 May 53 Sandusky Bay 17 3 293 •t it n South of Kelleys 18 3.5 66 18.8 11 Jun 53 South of Kelleys 5c Sandusky Bay 19a 3 32 10.7 25 Jun 53 Between Green 5c Rattlesnake 19p 3 36 12.0 ft it it Between S.Bass 5c Kelleys 20 3 75 25 26 Jun 53 South of Niagara Reef 21 3 56 18.7 9 Jul 53 Between S.Bass 5c Kelleys 2k 3 ko 13.7 7 Aug 53 East of North Baas Island 25 2 38 19 12 Aug 53 Between S.Bass 5c Kelleys 27 3 55 18.3 9 Sep 53 Between Green 5c Rattlesnake 28 3 63 21 11 Sep 53 Between S.Bass 5c Kelleys 90.5 J.BH2

1882 20.8 TO-

-21- OTTER TRAIL COLLECT I OSS HESTER* LAKE ERIE

1952-1953

Eraquancy of Nunbar of Hybopals storarlana Takan par Haul

Basad on Racorda Avallabla on Individual Hauls

— - C -----H Iiabar h a — Lass Hauls Containing 37 * 41 of Haula k ~--- (r ---- *

10

8

6

it 2 nn n r i Z 3 L J □ 5 7 9 11 U 15 17 19 21 23 25 27 W 37 1)1

Ruabars par Haul

- m m----* - w •

Figure 5.— Frequency of the number of silver chubs taken per haul based on individual hauls. TABLE 3 COLLECTION DATA Hybopala atorerlana (Adults) Collection Number per Collection Number per Number Haul Number inrui 5a 18 17b 16 b 19 18a 12 (i haul) c 27 b 19 6a 19 c 17 b 19 d 19 c 21 19a 12 7a 17 b 12 b 19 c 8 c 11+ d 8a 12 e 18 b 17 r 1+ c 19 20a 20 9a 17 b 19 b 15 c 36 c lk 21a 19 10a 37 b 20 b 37 c- 17 c 21+ 21+a 11a 20 b 16 b 22 c 10 15a 20 25a 22 b 28 b 16 c 25 27a 16 e 27 b 27 16a 21 c 12 b 20 28a 25 c 17 b 22 d i+o c- 16 e 21+

1107 19*6 per Haul 5575 Attempts to Locate and Collect the Young Silver Chubs

From examination of the first few trawl collections made in 1952, it was apparent that the O-year class, or young of the year, were not being collected. The literature concerning Lake Erie fish was examined for possible clues to their location. Kirtland (1850) reported that the young, after midsummer, congregate in considerable numbers about the mouths of rivers and bays. He described them, as follows, "The young, when two inches long, are sil­ very and almost translucent, and are so much attenuated in their forms that they would not at first be recognised as any relatives of the full-grown specimens." More than a hundred young were collected In this study. They varied in total length from 26 to 80 millimeters. They did not fit the latter part of Kirtland*s description, but appeared to be miniature adults. Other investigators (Greeley, 1929; Fish, 1931) also reported that they are common at creek mouths. Since neither of these Investigators collected them at creek mouths, these statements may have been taken from Kirtland. The smallest silver chubs collected by Greeley were of the 1-year class. The young which Marie Poland Fish collected, which have not been positively identified as Hybopsls atorerlana. came from bottom hauls at depths of 16 to 20 meters. The fiah collection data which are on file at Stone Institute and cover the period from 1939 to the present were examined. They revealed that moat of the Ohio tributary streams as well aa the shoreline and Island region of Lake Erie have been thoroughly seined. The following records of young silver chubs were found: Date No. Place Collector 25 March 19^7 1 Peach Pt., Put-ln-Bay M.B* Trautman 28 June 191*8 2 Estuary, Portage R. T.H. Langlols & Class 5 July 191*8 1 East shore, Pelee Is. T.H. Langlols it Class No large numbers of young chubs were encountered during this study. Usually, a few specimens were found in each col­ lection. Since the island region was well covered with the trawls, It seems unlikely that they are In great abundance on the lake bottom, the lower part of Sandusky Bay, or near the mouth of the Portage River. The trawls used covered only an area two feet above the bottom and If the young chubs lived at intermediate depths they would be missed with such trawls. Those which were captured could have been taken during the lifting operation. The intermediate depths were trawled several times but only Notropls fttherlnoldea was taken. The otter trawl is designed to operate on the bottom and, because of its shallow mouth, is an Inefficient device when used off the bottom. To cover this area it would probably be necessary to construct a beam

25- trawl from quarter-inch knotted twine with a mouth about 15 feet wide and 20 feet high. This type of trawl would be difficult to handle and would require at least four men for operation. During the winter of 1952-1953 some adult chubs were kept In the laboratory In aquaria and In hatchery troughs. They were seldom observed to be more than a foot off the bottom, and were usually on the bottom. It was not until December of 1953 that success was achieved in keeping any young In the laboratory, A single specimen was observed during the Winter and Spring of 19514.* It seemed to show no preference as to depth and spent as much time near the sur­ face as it did near the bottom. Half of the food consumed by the young examined consisted of Eucopepoda and Cladocera. Chandler (1914.0) recorded the vertical distribution of the abundance of these orders. Prom his data It was determined that the greatest abundance of these organisms usually occurred at about five meters. Sim­ ilar conclusions were drawn by Wright and Tidd (1933)* On the basis of the above, the author believes that, in the open lake, most of the young live at depths more than a meter above the bottom. They might live in the upper bay areas which are too deep for seining and too shallow for operation of the trawl boat. Also, in the island region, they might live on the reefs.

-26- where it Is too deep for seining and too rocky for trawling* Although three trawl nets were thoroughly torn in the process, some successful trawling was accomplished on the reefs, but only one young silver chub was taken.

-27- Age, Growth, and Abundance of the Sliver Chub, Hybopsls atorerlana (Kirtland), In Western Lake Erie

In western Lake Erie, the male silver chubs, Hybopsls atorerlana (Kirtland), seldom live more than three years, while the females attain an age of almost four years. The maximum total length Is slightly over 200 millimeters, and the weight about 90 grams. The scale method of age determination was found to be valid (Fig. 6). The annuli formed during the last week of

May (1953)* and were clearly evident on most specimens by the third week of June (Fig. 7)* Some difficulty was encountered in determining the first annulus on approximately five percent of the specimens (Fig. 8), A series of scales from the one-year class was photographed to obtain an idea of the variation in site of the first annuli (Fig. 9). It was possible to age most of these doubtful specimens by assuming the first annuli, be­ cause the second annuli were too large to be mistaken for first annuli.

-28- o. 1-Year class, female, b. 2-Year class, male, 13i+mm. 113mm. standard length. standard length.

c. 3-Year class, ferrule, 167mm. standard length. Figure 6.— Scales from representative specimens from the doptember, 1952 collection. About ten times normal size. -29- a. Male, 126 millimeters In standard length, June 11, 1953* Second annulus Just forming on the edge.

b. Female, 131 millimeters in standard length, June 26, 1953* Second annulus well formed*

Figure 7*— Annulus formation*

-30- n. Female, 109 millimeters b. Male, 135 millimeters standard length, September standard length, September

2 2 , 1952. 22, 1952.

c. Female, 139 millimeters standard length, March 12, 1953-

Figure 8.— Scales with doubtful first annuli. 78 on. Standard length 93 mas. Standard length 110 mm. Standard length i u> ro i

a

115 mm. Standard length 120 mm. Standard length

Figure 9.— Scales from specimens representing the size range of the 1-year class,

September 22, 1952. All lengths were measured In millimeters and weights were made In grams. The followlrg linear regressions were computed (Snedecor, 194-0):

Total length = 1.8 f standard length x 1.227 (Pig. 10), with a standard error of estimate of 0.9 millimeters.

Log weight z 4-.876 f log standard length x 3.062 (Fig. 11), with a standard error of estimate of 2.2 grams. Standard length - 12.063 4- anterior scale radius x 1.676 x 20 magnification (Fig. 12), with a standard error of estimate of 5.79 milli­ meters, and a correlation coefficient of 98.1. The 1-year and 2-year classes dominated the samples collected, each comprising about 4-5 percent of the total. The 0-year and 3-year classes each made up about 5 percent of the specimens collected (Table 4-)* The sex ratios for the 0-year and 1-year classes were about 1 to 1. The ratio for the 2-year class was about 1 male to 1.6 females, and for the 3-year class there was 1 male to approximately 5 females. Among the 0-year and 1-year classes the size and weight were about the same for both sexes. The mature females were slightly heavier because of the ovary weight. In the 2-year and 3-year classes the females were larger and heavier, probably because the larger males had died. The average standard lengths and the range of variation

-33- are given in Table 5* There was little or no growth during the winter. Maximum growth occurred during the June- September period (Pig. 13)* The 1-year class more than doubled in length during this period. They increased about 60 millimeters, as compared to 25 millimeters of growth for the 2-year class and 15 millimeters of growth for tha 3-year class• A relative index of abundance, based on averages per collection for the 1952-1953 period, was determined to be about 20 adult chubs per acre. The total population was not determined, because there was found no method of computing the numbers which escaped the trawl. Experiments conducted with a model otter trawl in a hatchery trough containing minnows indicate that a major fraction of the fish in the path of the trawl escape. Al­ though there is very little visible light near the bottom of the areas trawled (Chandler, 191*2), the turbulence created by the otter boards may send the fish scurrying (Anonymous, 1952

Ketchen, 1914.1). Adult fish with fusiform bodies, such as 3tlzostedlon and some of the Catostomidae, are seldom taken with the trawl although, during certain periods of the year, they are common in the trawled areas. The silver chub is also a rapid swimmer; therefore, I would estimate the adult population in the island region, during the 1952-1953 period, to be somewhere between 20 per acre and 200 per acre. Figure 10.— Total length-standard length correlation. Lengths are In millimeters. The deviations from the regression line were insufficient to warrant plotting.

-35- - 1 ^ 5 7 64>3.04 Uo« L

Figure 11.— Length-weight correlation. Baaed on average weight in grama and atandard length in ml11imetera•

-36 ue 100

90

70

SO

30 gy*9»+li imrrrl mi 20

10 Anterior Seolo SmSlay Tl~»* » 10 !• 30 l|0 SO 00 70 80 90 10O

Figure 12.— Standard length-anterior scale radius correlation. Measurements were made In millimeters.

37 3 HO*TH Ctnmts B A U D OH AVEltAQK STANDARD LANOTH. 1 9 5 2 - 195 4

60

50

30

Ui^th 3-Y**r C l a n 1 * 0 140

120 IOO

SO *0 0 « l M r C l M * 40

20 *»pt 0 0 1 Nov tea /Ml P»» ter Apr Hoy Jun Julj (apt oat Ho* Do« Jan Fab Mar

Figure 13*— Growth and water temperature. Standard length is in millimeters and water temperature in Fahrenheit•

38 TArLE ^ NUMBERS OP MALES AND FEMALES PER AOE CLASS Month Total Zero-Yoara One-Yaar Two-Yeara Three-Years Aged Male Fern Male Pem Male Pem Male -■ PTrrl 19 52 Sept. lOi* 0 0 32 21+ 17 29 0 2 Oc t. 44 0 0 10 9 8 16 0 1

Nov. 121 1 0 32 21+ 26 31 2 5 Dec . 11+9 7 5 19 18 36 59 0 5 195,* J an. 97 0 0 31 28 15 21 0 2 Feb. 1+2 0 0 16 11+ 6 . 6 0 0

M e r . 54 3 2 15 11 7 12 1 3 Apr. 99 3 1+ 1+2 30 7 10 0 3 May 15 122 1+ 1+ 28 30 19 31+ 0 3 May 27 327 8 8 103 97 33 73 0 5 June 209 0 0 30 69 21+ 80 0 6

July 70 2 2 6 1+ 11+ 23 6 13 Aug. 77 1 1 6 8 26 31 1 3 Sept. 118 4 3 11 13 39 34 3 11 Nov. 52 7 7 3 6 16 6 1 6 Dec. 21 1+ 5 3 2 1 5 0 1

Mar. 11+ 0 1 2 0 5 5 0 1 Totals 1720 P 1+2 w r 299. I+79 TI+ 75

39 TAPLE 5 AVERAGE STANDARD LENGTH AND RANGE PER AGE CLASS In Millimeters 0 >4 e u Month • Class 1-Year Class 2-Year Class 3-Year Class Ave # Sange Av® * Range Av 6 * Range Ave. Range

J9 p t • 103 80-125 133 115-155 153 145-160 Oct. 114 80-145 141 120-160 150 150

Nov. 50 50 112 80-135 141 120-160 155 145-165 Dec • 48 30-60 102 80-120 139 115-160 161 150-165 Jan. 115 95-145 142 125-160 158 155-160 Feb. 116 85-140 144 135-165

War. US 40-50 114 85-130 143 130-150 163 155-175 Apr. 46 30-60 117 90-150 141 130-150 168 165-170 May 15 45 40-65 121 95-140 146 125-165 170 165-180

May 27 45 35-60 111 85-145 140 120-165 164 155-170 June 124 95-150 147 130-170 164 155-170

July 29 25-30 67 55-80 125 110-145 147 140-155 Aug. 40 35-45 96 80-115 129 115-150 152 145-160 Sept. 49 40-60 103 90-115 135 115-155 155 135-170 Nov. 41 30-55 116 105-130 142 130-150 163 160-170 Dec . 38 30-56 113 97-125 145 135-150 160 160 Mar. 32 32 115 110-120 145 135-152 160 160 Mortality of the Silver Chub, Hybopsls atorerlana (Kirtland), in Lake brie

Data obtained during the period 1952-1954 indicate that most of the male silver chubs, Hybopsls atorerlana (Kirtland), in western Lake brie die before the end of their third year of life. Many of the females live a year longer. McCormick (1892) reported finding dead silver chubs on the beach near Beaver Creek on July 8, 1891. Osburn (1901) found them on the beach at Sandusky. Van Oosten (1938) found dead specimens at Vermilion, Ohio, on August 10, 1931* He also found them on Euclid Beach, Cleveland, in July of 1928. During the summers of 1950, 1951, and 1952, Ligas (1952) reported that dead silver chubs were common in the island region of Lake brie. On March 28, 1952, 22 dead specimens were collected on the west shore of South Bass Island (Table 6).

TABLE 6 Dead Silver Chubs Collected on March 28, 1952 Number Sex Age ^Standard Length

k Female Almost 1+ yrs. 161-168 12 Female Almost 3 yrs. 135-156 6 Male Almost 3 yrs. 137-140 <*in millimeters Twenty of these fish contained net marks similar to those found on specimens obtained from Charles Mahler, a Put-in-Bay gillnetter. None had spawned. On way 23, 1953, 16 dead specimens were collected near the city dock at Put-in-Hay, Ohio (Table 7)*

TABLE 7 Dead Silver Chubs Collected on May 23, 1953

Number Sex Age Standard Length of Fish I4. Female almost {4. yrs, 155-167 6 Female almost 3 yrs, 1U4-150 2 Female almost 2 yrs. 119-122 1 Male almost 3 yrs. II4.6 3 Male almost 2 yrs. 120-127 ttin millimeters

Only two of these specimens contained obvious net marks. None had spawned. Van Oosten (op, cit.) and Ligas (1952) believe that much of this mortality is caused by commercial fishing.

During the 1952-1953 winter, adult silver chubs were kept in a hatchery trough at the Ohio State Fish Hatchery. A continuous flow of lake water was maintained through the trough. The water temperatures ranged between two and three degrees (Fahrenheit) higher than lake temperatures. During May, seven of the larger specimens died (II4.2 -I6I millimeters standard length). It was noted that none of them floated to the surface.

Although much of the observed mortality on the lake and beaches can doubtlessly be attributed to commercial fishing, this may be only a fraction of the mortality resulting from other, as yet unknown, causes.

Because of its tendency to remain In relatively deep water, the silver chub In Lake trie Is rarely taken by minnow seine, the most commonly used collecting device of the Ichthyologist. So, to record the finding of silver chubs and other deep-water species such as the trout perch (Percopala omlscomaycus), on the beaches Is to provide valuable evidence of the fishes* presence in a body of water. If only the minnow seine Is used, many of these species will go unnoticed. Lake St. Clair is fished coimnerclally by the Canadians and was formerly fished by the Americans. If the silver chub rere present in this lake, the evidence of dead specimens on the beaches would almost certainly have been noted by

Canadian and Michigan biologists. Distribution Records of Hybopsls atorerlana (Kirtland)

The distribution of the silver chub, Hybopsls atorerlana (Kirtland), is confined almost entirely to the Mississippi River system. The silver chub is also found in the Red River of the North and in Lake Erie (Fig. 11+) . During post-glacial tines, the Red River and what is now called Lake Erie drained through the Mississippi Valley.

DISTRIBUTION RECORDS Key: (N)-No Record (?)-Doubtful Record (E)-Record is In Error (P)-Probably not Present Now ARKANSAS Hlack, 19t+0 Evermann & Kendall, 1892 Hubbs Sc Ortenbur^er, 1929 Meek, 1895 COLORADO Heckman, no date (1982 or 19$3)» (N) Ellis, 1911+, (N) Jordan, 1891, {N ) ILLINOIS Koroes, 1910 Forbes & Richardson, 1919 Hankinson, 1913 Luce, 1935 Nelson, 1881+ (E) O’Donnell, 193$ Thompson & Hunt, 1930 INDIANA Blatchley, 1936 Eigenman & Beeson, 1893 'Jerking, 191+5, (N)-Lake Michigan Drainage Hay, 1891+ Kiruch, 1895

kk IOWA Coker, 1930 Evemann & Cox, 1896 Marian 9r Speaker, 1961 Jordan & Meek, 1886 Meek, 1892 A: 1896 Jtarrett, I960 KENTUCKY Clark, 19^.0 Evemann, 1918 rtoolraan, 1892 KANSAS Hreakelman, 1914-0 Evermann .V Cox, 1896 day, 1868, (P) Jordan, 1891 MANITOBA

El;»enriann, 1896 Hlnks, 194 3 "flCHIOAN Creaser, 1963, (N)-Lake St. Clair Hubba Cooper, 1936, (N)-oxcept Lake Erie Hubba & Lawler, 1947, (N)-eacept Lake Erie

MISSOUHI Evermann V Cox, 1896 MINNESOTA Cox, 189? Friedrich, 1933, (N)-central Minn. MONTANA Evermann, 1892, (N) Henahall, 1906, (N)

NEBRASKA Evermann Ac Cox, 1896 Johnson, 194-2 leek, 1896

- 4-6 - NEW YOhK Pean, 1903* ( ?)-Lake Ontario Evermann Kendall, 1902, (?)-Lake Ontario Ireeley, 191+0, Lake Erie Hadforth, 191+4 * Lake Erie Haney, 195 3, (?) of Evermann & Kendall, 1902 Sibley, 1928, Lake Erie N HTH DAKOTA lanklnaon, 1929 tfvoolman, 1095, Red River OHIO Fi3h, 1935, Lake Erie ilensiiall, 1008 Jordan, 1880, 1882 Kirtland, 101+7, Lake Erie Kirtland, 1050, Lake Erie VcOormlck, 1(392, Lake Erie Osburn, 1901 Vun Oosten, 1938, Lake Erie OKLAHOMA Hubbs & Ortenburger, 1929 Jenkins, Leonard, & Hall, 1952 ONTARIO Dymond, 1922, Lake Erie Dymond, 191+7* Lake Erie; (N)-Lake Ontario (Canada) Fish, 1935, Lake Erie riubba and Brown, 1929; (N )-Lake Ontario (Canada) Sprulea, 1953, Lake Erie; (N)-Lake St. Clair PENNSYLVANIA

Evermann cc Bollman, 1866 (P) Fowler, 1919, (P) TENNESSEE Evermann, 1910 Kirsch, 1093 Kuhne, 1939 TEXAS Bonn & Kemp, 1952 -1*6- WISCONSIN Treena, 1935 Warner*, 1908, Mississippi River*

'WYOMING livermann, 1891, (N) Simon, 1939, (N)

IEN e h a l . disthibtttion Eddy & Surbar, 199-3 Forbes, 1910 Greene, 1935 Hubbs fie Lagler, 199-7 RadTorth, 199^4-

-U-7- ts

Figure It*.• — Distribution of the silver chub in North America based on distribution records. The record for Lake Ontario, shown by a circle, is doubtful.

48- Distribution and Ecology of the Silver Chub, Hybopsls storerlana (Kirtland)

The silver chub, Hybopala atorerlana (Kirtland), is one of the largest native minnows in central United States. Its distribution is restricted to the Mississippi River system, the Red River of the North, and Lake Erie (Fig. ll|).

From a study of the distribution and ecology of the silver chub, the following conclusions were drawn: 1. The silver chub is an Inhabitant of broad rivers of low gradient which flow through old or mature valleys. It rarely enters small streams, but is sometimes found in or near the mouths of small streams (Blatchley, 1936; Gerking,

19i+9; Hanklnson, 1913; Jordan, 1882, 1891; et al.). 2. Depth does not seem to be an Important factor in the limiting of its distribution. It has been found in depths varying from one meter to twenty meters (Woolman, 1891?; Fish, 193S). This might be a factor which would limit it to areas near the shoreline in eastern Lake Erie. 3. It ranges over bottoms which vary from gravel to silt, but is found more commonly over silt or mud bottom. This is the type bottom characteristic of mature and old streams. ij.. The silver chub Is a southern species which seems to require a growth period of six to seven months at water temperatures above k5°-50° Fahrenheit, with at least three

49 months above 70° F. (Fig. 13). Among the Great Lakes, this factor would limit its distribution to Lake Erie and possibly Lake St. Clair. Records indicate that the silver chub is not in Lake St. Clair. The summer water temperatures of Lake St. Clair, near its source, are only between three and five degrees (F.) colder than western Lake Erie and near its entrance to Lake Erie the average difference for the summer months is only one or two degrees (F.). The water temperature data were taken from Daiber (19^0), a 39-year summary for the Great Lakes; Millar (1952), a 5-year to 7-year summary for the Great Lakes; and Ohio State Fish Hatchery (19U4“1^5i+) daily records for Put-in-Ray, Ohio, from 19U4 to the present. 5. There is evidence that some spawning occurs when the water temperature reaches 65° F., but most of the spawning occurs at temperatures above 70° F. (Fig. 17).

6. In Lake Erie there is a shoreward migration in the spring which Is probably a positive thermotropic response. If the physiological make-up of the silver chub was the same during the Pleistocene Period as It is today, the following conditions would be necessary to explain its presence in the

Red River and In Lfke Erie: a. A large stream connection between these Isolated populations and the Mississippi drainage. b. Suitable water temperature conditions for growth and spawning. According to Radforth (1940) the silver chub entered the Red River during the glacial Lake Agassis stage. Lake Agassiz was in existence for a long period after the ice had receded from the Cireat Lakes region. Lake Winnipeg was not formed until the glacier retreated to the Hudson Bay region and water could drain northward through the Nelson River. Evidence is cited by Coleman (1922) to indicate that the cli­ mate during this period was, "not much more rigorous than now.n Data indicate that the silver chub is not present in Lake Winnipeg. The southern part of Lake Winnipeg is similar to western Lake Erie with respect to depth, bottom types, and bottom and fish fauna (Rajkov, 1930). The water temperatures, however, are such that the growth period for the silver chub would be at least a month shorter than in Lake Erie. Even if the climate were about the same during the early post-glacial period as it is now, it does not appear likely that the silver chub could survive in, or would enter, the outlet of a glacier-fed lake. The silver chub probably entered Lake Erie by passing from the Wabash River into the Maumee River, which enters western Lake Erie. CJreene (1939) and (Jerking (1945) indicate that this migration occurred during the glacial Lake Maumee stage. This would have been through the Ft. Wayne or Wabash outlet of Lake Maumee. According to Leverett and Taylor (1915)# Lake Maumee was

-51- a rather small lake which occupied much the same area as western Lake Erie, It was almost surrounded by glacial ice. It also seems unlikely that this southern species would migrate Into glacial stream. Radforth (op. clt.) states that the Fort Wayne outlet was In existence during the much later Lake Lundy stage. This writer has been unable to locate in her references any evidence to support this statement. She evidently had some basis for her statement, for she shows such an outlet on a map of Lake Lundy taken from Leverett and Taylor (op. cit.), although the original map shows no such outlet. The Lundy stage was much earlier than the Agassiz stage. Lake Lundy was fed by the glacial front which was only a hun­ dred miles away. Certainly, the habitat conditions existing at that time would not have been a suitable environment for the silver chub of today. An abundance of evidence concerning the post-glacial climates of the Great Lakes region has been presented during the past twenty-five years, by E.N. Transeau and J.N. Aolfe, of Tbo Ohio btate University; P.B. 3ears, of Vale University; J.E. Potz :er, of Butler University; the late Ira W. Wilson, of Heidelberg College; and many others. Most of these find­ ings are based on bog pollen analyses and the study of exist­ ing relict populations.

It Is generally agreed that there were at least five m a j o r post-glacial climatic periods, as follows: 1. A cool period, with Abies and Picea dominant; 2. A warmer and drier period, with Pinua prominent; 3. A warm and moist period, with Tsuga dominant and Fagus present;

4* A Xeric period with prairie vegetation dominant; 5. The present period. It would seem that the warm, moist period would have been a more favorable time for the migration of the silver chub into these northern areas. If the existing glacial outlet channels were filled with spring flood waters, suit­ able routes to these northern areas would have been open.

The migration into Lake iJrie could have been quite recent, i'nere are several records In Slocum's (1905) "History of the Maumee River Basin," which show there were temporary water connections between the northern and southern watersheds. .Judging from tne discharge at Ft. Wayne, Indiana, during the flood of 1903 (Cross, 1946)* the broad glacial Wabash outlet must have been full to the brim. In order to determine whether similar conditions exist between the Red River and Mississippi River drainage systems, I wrote to Mr. A. w. Gesterling, postmaster at Browns Valley, Minnesota. Browns Valley Is a town located on the east bank of the old glacial channel, on the present north-south divide. Mr. Gesterling (1954) informed me that during flood periods Lake Traverse, which is at the head of the Red River, becomes connected to Big Stone Lake, which la at the head of the Minnesota Hiver. Thus, fish migrations from the Mississippi drainage into the Hed River would not necessarily had to have occurred during the existence of the glacial Lake Agassis outlet. Three canals connected the Ohio River waters with Lake Erie before the silver chub was discovered in Lake Erie. These served as possible migration routes but may have been too narrow to serve as a silver chub habitat. Conclusions: 1. A study of the distribution and ecology of the silver chub, Hybopsls atorerlana. indicates that a. It Inhabits only large bodies of water; b. It is quite specific In its maximal tempera­ ture requirements for adequate growth and spawning; c. It has a positive photothermic tropism. 2. The maximal temperatures of the waters of the glacial outlets were probably lower than those now required for this minnow. 3. The more likely period of migration into the Red River and into Lake Erie would have been the more recent warm, moist, post-glacial period. I*. The eastern and western limits of the silver chub are determined by the width of the streams which flow from the Appalachian and Rocky Mountain divides. 5. The southern distribution is limited by the salt water of the Gulf of Mexico. -5k F'ood of the Silver Chub, Hybopals atorerlana (Kirtland), in Western Lake Erie

A year-round study of the food of the silver chub, ■ Iybopsls atorerlana (Kirtland), in the island region of western Lake Erie, was conducted during the period September, 19S2 - August, 1953- It was found that the O-year class of silver chub fed principally on Copepoda, Daphnla, and Tendlpedidae (Cniron- ornidae) larvae and pupae (Fig. 15, Table 8 ). Except during February and March, semi-monthly collec­ tions were made, by means of a 30-foot otter trawl. All the i productive collections were made in water between 9 and 12 meters in depth, over bottom which was in large part mud, with varying small amounts of sand and gravel. Most of the collec­ tions were made within ten miles of South Bass Island, Ohio. Previous investigators (Sibley, 1929, and Boesel, 1937) examined the intestinal contents of a few (I4. and 13 respect­ ively) Lake Erie silver chubs. Their qualitative results are similar to those reported in this paper, but their samples were too small for any quantitative comparison. Since only 6^ percent of some fifteen handred chubs collected were young of the year, it is possible that Table 8 presents a distorted picture of the food eaten by

-55 POOD OP T in Yotmo OP TUX T U X IfTBOFSIS 3T0WXPIASA

1 9 5 2 -1 9 5 3 a n d A u g u a t, 19<*9

Ttndlptdldi* L irru, pupa*, a « u i 34.7#

SphMFtldM it

Figure 15*— Food of the young of the year silver chubs

p o o d o p h y b o p s is s t o h c t ia w a a d u l t s

Saptaabar, 1952 through Auguat, 1953

Bpbaaaroptar*

N yapha

(Baaaaartla) 6 6 ,7% s

Figure 16.— Food of the adult silver chubs the young. The possibility that most of the young live in a zone above the effective height of the trawl is still under investigation. It is conceivable that those specimens which contained bottom fauna were collected on the bottom, and that those containing Daphnia and Copepoda were collected during the time the trawl was being lifted. The bulk of the food of the adults consisted of Ephemerop- tera nymphs* Hexagenla comprised more than 99 percent of the nymphs. The decline of Hexagenla consumed during July (Table 9) coincided with the period of maximal emergence of this genus. It may have coincided also with an abundance of Daphnia. Because of ice conditions on the lake, only monthly collections were made during February and March. The high percentages of Ephemeroptera nymphs shown for these months may have been the result of a small number of samples having been taken over a small area. However, it was observed that during this sampling the trawl, when pulled aboard the work- boat, was covered with hundreds of Hexagenla nymphs. During other periods of the year the number of nymphs observed varied from zero to twelve. The data (Table 9) on the Gastropoda and Pelecypoda may be slightly in error because of the difficulty of separating the small shell fragments. Although the pharyngeal teeth of the silver chub show no signs of wear, it is believed that they are functional In crushing soma of the food eaten. It was observed In the laboratory that when they are fed particlea of food evident­ ly too large to swallow, there is a continuous exaggerated movement of the opercle as the food is drawn into the mouth. This movement is apparently associated with the movement of the teeth. A blunt probe was Inserted through the mouth and pharynx of a decapitated specimen; as the opercles were worked outward and inward the vibration caused by the scraping of the teeth on the probe could be felt. With one exception, only the immature, small Mollusca were found in whole condition. One thin silver chub 11+2 millimeters In standard length weighed 20 percent less than the mean weight for this length. An adult Sphaerlum was found lodged in the U-curve between the mid-gut and the hind- gut. The fore-gut was noticeably swollen. In two other speci­ mens, broken particles of thick-shelled Unionidae were found. Cyprinidae made up the bulk of the fish eaten during December and January. Percidae was found in one stomach. The fish eggs found In April were probably those of Stizo- atedlon. This decision was based on comparison of size and appearance, as well as availability. The silver chub, Hybopsls atorerlana, is a predaceous minnow. The young feed principally in micro-Crustacea and small, immature Insecta. The adults feed principally on hexagenla nymphs, small Mollusca, Daphnla, and Gammarua.

-56- TABLE 8

POOD OF THE YOUNO-OF-THE-YEAR HYP0PSI3 STORERIANA IN WESTERN LAKE ERIE, 1952-1953

SO£. Nov. Dec. Mar. Apr. Max Aug.

Total 17 15 9 5 12 21* 17 Number empty 0 3 0 0 1 0 0

Food - Percent by Volume

Insects Ephemeroptera nymphs l+.l* Trlchoptera larvae 11.1 16.7 1*.6 Dlptera I vn Tendlpedidae larvae 11*. 7 1*7.1 13.3 31*. 3 21.1 58.2 nD Tendlpedldae pupae 17.7 31*.1* i Tendlpedidae eggs 2.1 Mollusca Pelecypoda Sphaerlidae 1.2 16.6 11.1 Anno11da Oligochaeta 9.2 Crustacea Cladocera (Daphnia) 2.9 1*2.3 13*3 12.9 Ephippia (Daphni a) i*,7 Copepoda 61*.7 1*.7 53.3 60.0 72.3 10.0 12,1 Ostracoda 20.0 Amphipoda (Oamnarus) 5*7 Unidentified (Coleoptera) 3.0 (larvae?) TABLE 9

FOOD HABITS OF ADULT HYBQPSIS STORERIANA IN tfEoTERN LAKE ERIE, 1952-1953

Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug

Total 102 Lk 170 91 98 42 44 102 429 229 56 77 Percent empty 10.8 18.2 3^.1 43.0 45.0 7.2 36.4 18.6 3.7 12.0 5.4 2.6

Food - Percent by Volume (X - leas than one percent)

Insecta Ephemeroptera 67.3 85.0 85.0 85.7 46.8 100 91.4 61.0 52.0 70.5 21.3 34.9 Trlchoptera larvae and pupae 1.5 10.4 X 2.5 2.8 Diptera Tendlpedidae larvae 1.4 X 1.9 X 6.0 Tendlpedidae pupae X 2.0 2.0 Tendlpedidae eggs 8.7 Hemiptera Corlxldae adults 3.0 Homoptera adults 1.3 Mollusca Gastropoda 23.6 13.7 6.7 1.7 X X X 9.8 36.5 Pelecypoda Sphaerlidae 1.4 x 6.9 2.1 26.0 26.6 23.5 25.3 X 2.3 Annelida Oligochaeta 10.4 2.0 X 4«6 Hirudinea - 2,3 Crustacea Cladocera (Daphnia) 6.2 X 53.6 2.5 Amphipoda (dammarus) 1.2 2.8 16.3 2.1 XX 11.5 3.8 Chordata (PiseesJ 7.6 10.6 Pisces eggs 3.1 Reactions of the Silver Chub, Hybopsla atorerlana (Kirtland), to a Decrease in Food Supply

During the period September, 1952 - August, 1953» It was found that Hexagenla (Ephemeroptera) nymphs comprised two- thlrda of the food consumed by adult silver chubs, Hybopsla atorerlana (Kirtland)* During the first week of September, 1953, Dr. N.W. Britt (1954) encountered a widespread temporary thermocline, with accompanying low oxygen conditions, in the Island region of western Lake Erie. Britt found that In the Rattlesnake-Green Island area, one of the principal locations for the collection of the silver chub, the Hexagenla population had been almost completely destroyed. In the South Bass-Kelleys Island area the Hexagenla population had suffered a drastic decline. The unusual conditions (Chandler, 191+2), as noted by Britt, could affect the silver chub population in several ways.

1* There could be a population decline because of a. Low oxygen conditions; b. A food shortage. 2. The population could migrate to a more suitable area. Conditions similar to those found by Britt were encoun­ tered on a trawling expedition, July 21, 1953* In an area about 10 miles north of Vermilion, Ohio, on the western edge of the central basin of Lake Erie. No oxygen tests were run, but the temperatures were 2i+.5° C. at the surface, 21.5° c* at

61- 15 meters depth, and 17-3° C. at the bottom (17 meters). Although excellent collections had been made in this area during previous years, only four dead white bass were taken In two hauls. It was also noted that the net became clogged with a slime composed primarily of fungus, with some filamen­ tous algae and diatoms. The Rattlesnake-Green Island and South Bass-Kelleys Island areas were trawled during the second week of September, and no apparent decline In the population was noted (Table 10). Some Idea of the minimal oxygen requirements of the silver chub can be obtained from the results of an accident which occurred on June 1, 19*33* Six silver chubs of the 1-year class were being kept in an aquarium having a running water supply. Sometime between 10:00 p.m. of May 31 and 7s00 the following morning the water supply failed. Five of the chubs were found deed. The oxygen in the aquarium was p.p.m. Fresh tap water had an oxygen content of 9.1 p.p.m. The oxygen deter­ minations recorded by Britt near the bottom were considerably lower than I4..I4. p.p.m., but there was sufficient oxygen a few meters above the bottom. The greatest amount of oxygen was found at the surface. Evidence that silver chubs migrated vertically during this period is indicated by the stomach contents of the Sep­ tember 9 collection. Adult Tendlpedidae were found to compose 22 percent of the contents. This was the only time during the study that any appreciable amount of surface food was found in the stomach contents (Table 10). Collections made In November and December Indicated a possible SO percent . ^pulatlon decline In the two areas. It has not been determined whether this apparent decline Is the result of mortality or movement from the area. It Is difficult to summarise the food habits data for the second period because of the variation. 1. The food of the 0-year class remained unchanged. 2. The mayfly decline was evident except In the last two collections, which were made In the area recommended by Dr. Britt as containing a moderate mayfly population. 3- Daphnia pulex was an important source of food from September through December. Many of the Daphnla contained ephlpplal eggs. l4>. On the last collection, Gammarus fasciatus made up 29 percent of the volume. This was the largest amount of Gammarus encountered; however, the collection was so small it would be unreasonable to form any conclusions from such evidence. S* Tendlpedidae adults, larvae, and pupae formed a much larger percentage of the food after the Hexagenla decline. This would indicate that the Tendlpedidae are better adapted to conditions of low oxygen than are the Hexagenla.

-63- tj9- Total Percent Empty Percent Number of Hauls of Number Annelida Mollusca Insect* Chordata Crustacea rcotr larvae Trlchoptera Gastropoda nymphs Ephemeroptera Diptera Pelecypoda Oligochaeta mhpd (Gaamarus) Amphipoda ldcr (Daphnia) Cladocera Pisces Hlrudinea edpdde adults Tendlpedidae edpdde eggs pupae & Tendlpedidae larvae Tendlpedidae Sphaerlldae and pupae and (Hexagenla) ODO DL YOSSSOEIN (ITAD I WSEN AEEI, 1953-1951; ERIE, LAKE WESTERN IN (KIRTLAND) STORERIANA HYBOPSIS ADULT OF FOOD

et 9 et 1 Nv 1 Dc 2 Mr 23 Mar. 20 Dec. 10 Nov. 11 Sept. 9 Sept, 55 29.0 15.5 22.2 - Percent by Volume by Percent - 11.7 10 11.7 3 4 0 2.1 6.3 1.3 4 0 6.7 3.3 TABLE10 . 1 2.9 + 22.0 63 6C .0 3 4.0 9.5 4 1 5.8 1.5 3.6 0.7 0.7 4 0 1.5 52 32.7 4 7 5 11.0 17.0 6.5 9.5 2.0 1.4 3730.7 43.7 16 5 27.9 27.9 25.6 29.0 10.6 13 67.3 4 3.7 The Estimated Food Consumption of the Silver Chub, Hvbopals atorerlana (Kirtland), in Western Lake Erie

Feeding experiments carried out during 1952 and 1953 Indicated that adult silver chubs, Hybopsla storerlana (Kirtland), are able to consume, per day, between four and five percent of their body weight in food. In 1951*., it was found that the young can consume about ten percent of their body weight per day. These experiments were carried on at room temperature which varied from 18° c. to 22° C.

The average wet weight of the intestinal contents of 100 specimens of the 1-year and 2-year classes, having full intes­ tines, was 1.12 grams, or about 3 *214. percent of the body weight. This was the equivalent of 8.6 Hexagenla per specimen. On the basis of the work done by Lagler and Latta (1954)* Pearse (1924), Prather (1951)* and Surber (1935)* it is be­ lieved that 8.6 Hexagenla per adult silver chub would represent a conservative estimate of its dally food at summer temperatures. The average temperature of western Lake Erie is about

10.5° C. (Chandler, 1940* Daiber, 1950). For the purpose of this estimate, 10.0° C. will be used. If a for the rate of digestion, hence, rate of feeding, Is used (Battle and Huntsman, 1936; Chesley, 1934* Kinney, 1951* Nicholla, 1931), then, on a yearly basis, the consumption per day per chub would be about 4*3 Hexagenla.

-65- Using a silver chub population estimate of 20 per acre, based on the average number per trawl haul, the annual con* sumption would be equivalent to 31,390 Hexagenla per acre. Since Hexagenla made up two-thirds of the food eaten, a total of 20,927 Hexagenla were consumed per acre. Using an average of I4.OO Hexagenla per square meter, based on the work of Wright and Tldd (1933), Britt, Chandler, and Y/ood (Britt, 19$Li.), it is estimated that the silver chub population consumed about 1.3 percent of the Hexagenla pop­ ulation during the year September, 1952 - August, 1953* U»3 Hexagenla x 20 chubs x 365 days x 2/3 < I4.OO Hexagenla x I4.05O Since the silver chub population estimate of 20 per acre is a minimum, and the population may be as high as ten times this amount, the total yearly Hexagenla consumption is somewhere between 1.3 percent and 13.0 percent of the Hexagenla population.

66- Data on the Reproduction of the Sliver Chub, Hybopsla atorerlana (Kirtland), In Western Lake Erie

In western Lake Erie, In 1953* the silver chub, Hybopsla atorerlana (Kirtland), began to spawn during the second week of June, with the water temperature at 66° P. Spawning reached its peak during the last week of June and the first week of July, with the water temperature at 73° P. Spawning

was completed by the middle of August (Pig. 17). An examin­ ation of the 194-6 collections revealed that half the specimens had spawned by June 26 (temperature 70° P.), and that most

specimens had completed spawning by August Z (temperature

74-° P.). Greoley (1928) states that the sliver chub in eastern Lake Erie spawns late. On June 21 he found specimens which he considered about ready to spawn. Fish (1935) captured early larval sliver chubs at the eastern end of Lake Erie during the latter half of June and the first week of July in 1928. In the western end of the lake, early larval stages were

taken on June 7 and July 2, 1929. In Iowa streams, which warm earlier than Lake Erie, the silver chub spawns during late May and early June (Harlan and Speaker, 1951). Luce (1935) recorded that In the Kaskaskia River they spawn during June and July. Prom the above data, it Is concluded that the silver chub spawns at relatively high temperatures. This is a factor which would limit its northern distribution. -67- irMniN rwoo1*53 I t f l t f l i w Mttr T M M H t l l l

0 1 0 0

w • V ' • • 0 ,-.s V m 0 0 I 0 0 ■ A * . ?

2 0

0 0 rt amj i Mr 7 * •« » krt

PMt-

Figure 17,— A bar graph showing percentage of pre-spawning, spawning* and post-spawning individuals in six collections made between May 27 and

September 9, 1953*

-68- as. (lit) u 17

r * ■ I --- 4 . IS - t - I T * * - i - ■"-f- I 13 ■4— U

11 - 10 : Tr 41 f [- *

--

%0 00 TO

Figure 18.— -Variation in the weight of mature ovariea. All weights are in grama.

-69- COMBAT IM or MOT ■BIOBT-OYAJT H U B

11

12

11

1 0

*

Figure 19*— - Increase in weight of the ovariea during the pre-spawning period*

-70- >< Mttun Ovary UUMl Tlav

b. Firtldl; Spumd Ovtr;

a. Ovary with Spawning Saar Coaplation

Figure 20.— Diagrams showing change in ovary shape as spawning progresses*

-71- Ovmrj Might Ihi)

■o. Igg* M 3*5 4- 74*.** X Ofifj Might

12

Figure 21,— Correlation of the ovary weight in grama and the number of mature egga, A atandard error of eatlmate of egga waa obtained*

-72 A considerable variation was found in the weights of mature ovaries from specimens of similar weights (Fig. 18).

Because of this variation# the ovary weights of specimens

collected after March 12 and before completion of spawning were not included in the total weights used to compute the

length-weight correlation.

(Log weight = -4.876 f Log S.L. x 3.062)

To obtain the approximate weights of the females for this period, we must add the ovary weights obtained from Figure 19.

The ovary weights during June and July varied considerably because some specimens had spawned, some were spawning, and others were reaching maturity. For example, a group of speci­ mens weighing 60 grams would have ovary weights varying from

3 grama to 15 grams.

The stage of maturity of the ovaries was determined by the size and color of the eggs, and the shape of the ovary.

Most 1-year fish were sexually mature. In ovaries of fish of the 1-year class there were 10 eggs per centimeter, while in those of the older year-classes there were 9 eggs to the cen­ timeter. The immature eggs are light yellow; the mature eggs are light orange. The shape of the mature and the partially- spawned ovaries are shown in Figure 20.

The testes, which are cylindrical in shape and milky white in color, never exceeded one gram in weight. On the smaller specimens they appeared as white threads. The ovaries of the immature specimens were several times larger than the testes. By microscopic examination the eggs in the ovaries of the smallest specimens could be seen clearly. Karly in the study, some difficulty was encountered in distinguishing the eggs from oil globules located in small fat bodies near the ovaries. Puncturing the fat bodies would release the globules and they would float to the surface, thus distinguishing them from eggs. Moat of the egg counts were obtained by the sample method. Samples weighing between one and two grama were used. The total number of eggs in 11 ovaries were counted. The average variation between actual counts and counts based on samples was I4..3 percent. The approximate number of mature eggs per specimen may be determined through the use of the graphs on Figures 10, 11, Id, and 21. The place of spawning was not determined, but ripe females with free-runninr eggs were taken in the open lake on

June 26 and June 27# 1953. Marie Poland Fish (op. clt.) took larval forms in the open lake at depths between 10 meters and 20 meters. Because of the variation in size of the young, it is believed that some spawn early, in the warmer bay areas, and others spawn later, in the open lake. Attempts at artificial fertilization were unsuccessful• More than a half-pint of spawn was taken on the two above- mentioned days and was placed in hatcnery Jars at the Ohio State Fiah Hatchery* Evidently, during the third ni^ht, the water pressure Increased sufficiently to force the eggs away from the bottom of the jars and over the top of the fine screen at the outlets.

-75- Economic Value of the Silver Chub, Hybopsi a atorerlana (Kirtland)

The silver chub, Hybopsla atorerlana (Kirtland), is a forage fish. It la of little direct food value to man be­ cause of the numerous upper and lower Intermuscular bonea. Trautman (1939) examined the stomach contents of 198 Lota lota lacuatrla and found 25 silver chuba In 17 stomachs. The atomach contents of 5 Lota taken through the ice In January, 195U» were examined by the author. One silver chub was found. Doan (19i|-3) found 1+. silver chubs In 268 aaugers, (Stlso- stedion canadense), examined. Most of the saugera he examined were probably too small to have eaten these large minnows. In 57 pickerel, (Stlsostedlon v. vltreum). examined during this study, only one silver chub was found. Coker (1930) reported that a 108-millimeter silver chub was found in the stomach of a yellow catfish (Laptops ollv- arls), taken from the Mississippi River. Harlan and Speaker (1951) reported that in Iowa the silver chub is a prise bait minnow for large flah angling. Jordan and Evermann (1937) regard this chub as superior bait for largemouth and smallmouth black bass. Llgas (1952) reported that dead and dying silver chuba served aa food for piscivorous birds in the island region of western Lake Erie. In order to determine the value of the silver chub as a forage fish In Lake -trie, it would be necessary to make a comprehensive study of the food habits of some of the larger piscivorous species.

-77- Notes on Eish Species Associated with tne Silver Chub, Hybopsls atorerlana (Kirtland), in Western Lake Erie

During the period September, 1952 - March, 1954-, fi«h collections were made in western Lake Erie, with a 30-foot otter trawl, at intervals of two or three weeks. Most of the collections were -nade at depths of 9 to 11 meters. The Sandusky Bay collection was made at approximately 3 meters, and the Stone's Cove collection at 8-10 meters. Lack of sufficient time and space prevented a close examination of all the fish captured. Samples of the smaller specimens were preserved and placed in the Stone Institute fish collection. The notes on these specimens are presented for the information or interest they may provide to students of ichthyology. Clupeidae. Dorosoma cepedlanum. This species was captured In the deeper water (1-11 meters), only during the winter, when there was evidently a mass migration through the Island region. The specimens ranged from about 100 millimeters to 200 millimeters in total length.

-78- TAPLE 11 Dorosoma cepedlanum Collections Number Date Location Water Temperature of Fish (Centigrade J 1 8 Dec 52 S .Bass-Kelleys Is. 3.2 750(approx.)22 Dec 52 Green-Rattlesnake Is. 1.0 1500 22 Dec 52 S.Bass-Kelleys Is. 2.1 n ti n « Q#6 6 20 Jan 53 4 ( dead) 10 Feb 53 Green-Rattlesnake Is. 0.35 1 28 Dec 53 e " " 0.12 1 11 Jun 53 Sandusky Bay 21.8

Coregonidae. Coregonus clupeaformls. Only one specimen was taken, and that on March 12, 1953 (water temperature

0.5° C.), about a half-mile north of the northwestern shore of Kelleys Island. Its standard length was 165 millimeters; total length, 197 millimeters; weight, 62.2 grams. The stomach contents were four Tendlpedidae larvae. Osmerldae. Osmerus mordax. A few young smelt were taken almost evory month. Only one adult was collected.

TABLE 12 Osmerus mordax Collections Number Date Location Water ^Average »Length of Pish Temperature Total~ Range (Centigrade)Length

1 12 Mar 53 Ballast la. 4.2 183 6 27 Jul 53 Stone1s Cove 23.9 35.6 31-41 5 9 Sep 53 Rattlesnake Is. 23.2 48.3 40*52 28 28 Dec 53 S.Basa-Kelleys Is. 0.12 72.8 55-86 *In millimeters E i rht of the young were examined* Daphnla comprised almost 75 percent of the food eaten, and Copepoda the remaining 25 percent. On May 5 and May 6 , 1953, a large school of adulta was observed near the Stone Institute dock, feeding on young Notropls atherinoides. One specimen, 175 millimeters in total length, was captured. Catostomidae. ho suckers were taken in the trawl, though they are commonly caught b> gill nets in the trawled areas. Either they enter the area at night or outswim the trawl• Cyprlnua carplo and Carasslus auratus. Only a few were caught in the trawl in the Island region. These were taken during the summer. Hexagenla was the only food found in the intestines of three specimens examined on July 9, 1953* Notropls a. atherinoides. Emerald shiners are common near the bottom, in the open lake. Some 50 specimens were collected every trip. They fed primarily on zooplankton. Notropls h. hudsonlus. Spottail minnows are about as common as silver chubs in the deeper parts of western Lake Erie Their food habits are also similar to those of the silver chub. The largest specimen taken was 138 millimeters in total length. Fry were taken near the surface, along the western shore of South Bass Island, on June 29 end June 30, 1953* They ranged in total length from 11 millimeters to 19 millimeters. Thirteen young, taken on July 27, 1953, had a total-length average of 36.7 ; nineteen young, taken on September 9, 1953* averaged $4.2 millimetera In total length. Notropls v. volucellua. The mimic ahiner waa about aa common as the spottail shiner in the areas trawled. Young specimens less than 23 millimeters in total length were commonly taken in September.

Ameiuridae. Ictalurua jd. punctatua. Channel catfish were commonly taken throughout the year. More were caught in winter than In summer. The size varied from ijl* milli­ meters to 63$ millimeters total length. Seven adult speci­ mens examined in July contained Hexagenla. Most of those collected were given to William H. Coil, a student at the Stone Institute, who was using this species in hla parasit­ ology research. Amelurus sp. Only two bullheads were taken in the trawl. These were captured in an area two miles northwest of Rattle­ snake Island, on September 9, 19$3. Noturus flavus. Only 21 stonecats were collected. These were taken during September and October of 1952 and

September of 19$3. Schllbeodes mlurus. An occasional brindled madtom was taken, when the trawl passed over an area of gravel. Forty- five specimens were taken on May 1$, 1953# on the edge of Niagara Reef. Thirty of these were small (32-1^9 millimeters), but appeared to be sexually mature. The larger madtoms varied in length from 69 millimeters to 109 millimeters In total length. Gadidae. Lota lota lacustris. Eight lingcod, having a total-length rang® of 192 to 381+ millimeters, were taken during the September 22, 1952 - January 20, 1953* period. Five were caught through the ice between South Bass and Hattlesnake Islands during January, 1951+. Three of these were ripe females. Percopsidae. Percopals omlscomaycus. The trout-perch was the most abundant species consistently taken with the trawl. On July 9, 1953, 555 young, under 30 millimeters in total length, were taken in one 12-minute haul between South Bass and Kelleys Islands. For further information, see Kinney (1950). Serranidae. Morone chrysops. Only three adult white bass were taken, but the young were common.

TABLE 13 Morone chrysops Collections Number Date Location Water ^Average a-Length of Fish Temperature Total Range (Centigrade) Length 20 9 Jul 53 S.Bass-Kelleys la. 22.5 30 21+-36 21+ 27 Jul 53 S . Bass (Stone ’ s Cove) 23*9 31+ 31-51 11+ 9 Sep 53 Rattlesnake Is. 23.2 52.3 38-73 ■»In millimeters

-82- Percidae. Perea flaveacena. All size-groups of yellow perch were represented In most of the hauls* The young were more numerous than were the adults.

TABLE 11+ P®.rc.a flavescens Collections Number Date Location Water ^Average ■B-Length of" Fish ---- Temperature Tota Range- ((Sentlgrade) Length

260 9 Jul 53 S,Bass-Kelleys Is. 22.5 35.2 28-45 35 27 Jul 53 S.Bass( Stone *s Cove) 23*9 45*9 38-53 10 9 Sep 53 N. of Rattlesnake Is.23-2 77.8 71-90 34 23 Mar 54 S.Bass-Kelleys Is. 0.9 88.9 ■a-ln millimeters

In the March 23rd collection, the first annull were in the process of formation. The young fed mostly on zooplankton and Tendlpedldae larvae. Stlzoatedion v. vitreum. Adults of this species were comnon throughout the year. The numbers taken per trip varied from 2 to 27. The young were identified by their three distinct pyloric caeca.

-83- TABLE IS 3tlzostedlon v. vitreum Collections Number Date Location Water ^Average oLen^th of Fish Temperature Total Range (Centigrade) Length

46 25 Jun 53 Green-Hattlesnake Is. 22.0 30.7 25-40 54 26 Jun 53 Niagara Reef 23.0 30.6 20-47 27 9 Jul 53 S.Fass-Kelleys Is. 22.5 54.1 42-66 3 20 J an 53 Green-Rattlesnake Is. 0.6 130.7 127-137 3 15 May 53 Niagara Reef 11.7 127.7 120-137 3 23 Mar 54 S .Pass-Kelleys Is. 0.9 145.5 142-159 •a-ln millimeters The young begin feeding on fish quite early in life. One 40-ml1limeter specimen contained a 19-millimeter fish. The smaller individuals contained mostly Daphnla. Among specimens in the 30- to 45-millimeter size range, Tendlpedidae larvae were common in the stomach contents. Ten specimens of the July 9th collection were examined and all contained fish. Stizostedlon v. glaucum. This species was not taken in the western basin. A few were taken in the central basin, about 12 miles north of Vermilion, Ohio, during the summer of 1952. Stizostedlon canadense. If the pyloric caeca count is valid for young Stizostedlon vltreum, then no young saugers were taken. Usually, two or three adult saugers were taken per trip. Hadropterus copeland!. Channel darters were taken occa­ sionally when the trawling was done over gravel bottom. One specimen of 53 millimeters1 total length contained a small minnow in its stomach.

-84- PercIna caprodes semlfasclata. Log perch were taken In the same areas in which channel darters were caught. They were taken more frequently than channel darters, but there were never more than two or three in one haul. Centrarchidae• Very few Gentrarchidae were taken in deeper water. On February 10, 1953# one 20-millimeter Ambloplltea rupeatris was taken between South Easa and Kelleys Islands. On March 12, 1953# one 29-millimeter rock bass was taken near the same area. On September 9, 1953* two 33-millimeter Lepomia sp. were taken In an area about one mile northwest of Rattlesnake Island. Sciaenldae. Aploilnotu3 grunnlena. On the basis of weight and volume, aheepshead constituted the bulk of most catches. On June 25# 1953 (water temperature, 22.0° C.), males with running milt were taken, west of Green Island. The first young were collected on July 9# 1953* Their total- length range was 15 to 22 millimeters. For further information, see Daiber (1950). Cottidae. Cottus ricel. One was taken near Niagara Reef, on May 15# 1953# and one was collected about three miles north of Green Island, on June 25# 1953. Cottus balrdl kumllenl. This species was more common in the collections than was tne ricel. Seven were taken in one haul, on September 22, 1952, about one mile northeast

-85- of South Pass Island. Evidently their habitat In deep water is limited to small areas, as none were taken in five other hauls made the same day in the same area. Except for the above-mentioned collection, there were never more than one or two specimens collected In any one day.

86- | | LITERATURE CITED

Anonymous• 191+9 . Floating trawls developed for mid-water fishing. Commercial Fisheries Abstracts from Atlantic Fisherman, Vol. 39, No. 8 (Sept., 1949), pp. 20-37. Anonymous. 1951. Underwater research on behavior of the otter trawl Commercial Fisheries Review, Vol. 13* No. 10 (Oct., 1951), pp. 37-39.

Anonymous• 1952. Comparative studies of trawl behavior by under­ water observation. Commercial Fisheries Abstracts from World Fishing, Vol. 1, No. 1+ (July, 1952), pp. 116-120. Bailey, Reeve M. 1951. A check list of the fishes of Iowa with keys for Identification. Reprinted from Iowa Fish and Fishing, Iowa State Cons, Comm., Des Moines, pp. 107-238. Bajkov, Alexander 1930. Biological conditions of Manitoban lakes. Contributions to Canadian Biol., Canada, Biol. Bd., n.s., Vol. 5* No. 12, pp. 303-1+22. Dangham, Ralph V. and George W. Hunter, III 1939. Studies on fish parasites of Lake Erie. Distri­ bution studies. Zoologica, Vol. 21+, Part i+, pp. 305-1+1+8. Battle, Helen I., A.G. Huntsman, et al. 1936. Fatness, digestion, and food of Passaraaquoddy young herring. Jour. Biol. Bd. Canada, Vol. 2, No. 1+, pp. 1+01-1+29. Bean, l’arleton H. 1903. Catalogue of the fishes of New York. N.Y. State Mus. dull. No. 278* Univ. of the State of N.Y., Albany, 701+ pp. Beckman, W. C. Guide to the fishes of Colorado. Colorado Fish and Game Dept. (1952 or 1953?)* 110 pp.

-87- i Flack, J. D. 1914-0. The distribution of the fishes of Arkansas. Thesis, Univ. of Mich., 246 pp., 69 maps. Platchley, W. S. 193o. The fishes of Indiana. Nature Pub. Co., Indianapolis, 121 pp. Bodola, Anthony 19B2. A push-seine, Its advantages and disadvantages. Paper presented at llj-th Ann. Midwest Wildlife Conf., Das Moines, Iowa, 6 pp. (mimeographed) Foesel, M. W. 1938* The food of nine species of fish from the western end of Lake Erie. Trans. Am. Fish. Soc., Vol. 67 (1937), pp. 215-223. Bonn, E. W. and R. J. Kemp 19B2. Additional records of freshwater fishes from Texas. Copeia, No. 3, pp. 2 OI4.-2 0 5 . Breukelman, John 1911.0. A collection of Kansas fishes in the state univ­ ersity museum. Trans. Kans. Acad. Sci., Vol. I4.3 , pp. 377-38ll. 191+0. The fishes of northwestern Kansas. Trans. Kans. Acad. Sci., Vol. I4.3 * pp. 367-37*?. Britt, N. Wilson 1951+. Stratification in western Lake Erie in summer of 1953. Effects on Hexagenla (Ephemeroptera) population. Ecology (accepted for publication), U 4. pp., typewritten. Bullis, Harvey R. 1951. Gulf of Mexico shrimp trawl designs. U.S. Fish 6c Wildlife Serv., Fishery Leaflet No. 3914-, 16 pp. Chandler, David C. 1914-0 . Limnologlcal studies of western Lake Erie. I. Plankton and certain physical-chemical data of the Bass Islands region, from September, 1938, to November, 1939. Ohio Jour. Sci., Vol. ij.0. No. 6, pp. 291-336.

191m ?• Limnologlcal studies of western Lake Erie. II. Light penetration and its relation to turbidity. Ecology, Vol. 23, No. 1, pp. 14-1-52.

-88- Chandler, David C* 19i+2. Limnologlcal studies of western Lake Erie. III. Phytoplankton and physical-chemical data from November, 1939» to November, 19ij.O. Ohio Jour. Sci., Vol. 42, No. 1, pp. 2i+—I4J4.• Chealey, Leon C. 1934. The influence of temperature upon the amylases of cold- and warm-blooded animals. Biol. Bull., Vol. 6 6 , No. 3# pp. 330-338. Clark, M. K. A list of the fishes of northeastern Kentucky. Kentucky Div. of Pish & Dame, 12 pp. (mlmeo.) Coker, Robert E. 1930. Studies of common fishes of the Mississippi River at Keokuk. Bull. U.3. Bur. Fish., Vol. b,S (1929) , pp. 141-225. Coleman, A. P. 1922. Glacial and post-glacial lakes in Ontario. Univ. Toronto Studies, Pub. Ont. Pish. Hes. Lab., No. 10, pp. 1-76. Cox, Ulysses 0 . 1897. A preliminary report on the fishes of Minnesota. Minn. Geol. and Nat. Hist. Surv., Zool. Series, III, St. Paul, 93 pp. Creaser, C. W. 1953. Letter to Louis B. Carrick, Stone Institute, Put-ln-Bay, Ohio, Jan. 21, 1953. Cross, William P. 1946* Floods in Ohio, magnitude and frequency. Ohio Water Resources Bd., Bull. No. 7, 154 PP> Daiber, Franklin C. 1950. The life history and ecology of the sheepshead, Aplodlnotus granniens Rafinesque, in western Lake flrie. Dissertatlon, Ph.D., Ohio State Univ., 150 pp. Doan, Kenneth H. 1914.3. The food of Stlsostedlon canadenae canadenae in in Lake Erie at Put-Tfn^l^ay^ Unpublished data on file at Stone Institute, Put-in-Bay, Ohio. Dymond, J. R. 1922. A provisional list of fishes of Lake Erie. Univ. Toronto Studies, Biol. Ser. No. 20, Pub. Ont. Pish, Res. Lab. No. 4i£(jpP* 56-86. Dymond, J. H. 19U7. A liat of the freshwater fishea of Canada eaat of the Rocky Mountains. Hoyal Ont. Mus. Zool. Misc. Pub. No. lt Toronto, 36 pp. Eddy, Samuel and Thaddeus Surber 1943* Northern fishes. Univ. of Minnesota Press, 252 pp. Ellis, M. M. 1911+. Fishes of Colorado. Univ. Colorado Studies No. 9, pp. 5-136. Elgenmann, 0. H. 1895. Results of exploration in western Canada and northwestern United States. Bull. U.S. Fish Comm., Vol. 1 (1095), pp. 101-102. Eigenmann, C. H. and C. H. Beeson 1893. The fishes of Indiana. Contribution No, 9, Dept, of Zool., Indiana Univ., Proc. Indiana Acad. Sci., pp. 76-100. Evermann, B. W. 1091. A reconnaissance of the streams and lakes of west­ ern Montana and northwestern Wyoming. Bull. U.S. Fish Comm., Vol. 11 (1091), pp. 3-60. 1918. The fishes of Kentucky and Tennessee: a distri­ butional catalogue of the known species. Bull. U.S. Bur. Fish., Vol. 35 (1915-1916), pp. 293-368.

Evermann, B. W. and C. H. Boliman 1086. Notes on a collection of fishes from the Monongahela River. Annals N. Y. Acad. Sci., Vol. 3 (1003-1805), pp. 335-34°* Evermann, B. W. and U. 0. Cox 1096. A report of the fishes of the Missouri River basin. Part 20, Kept. U.S. Fish Comm, for 109i+» App. 5, PP. 325-1*49. Evermann, B. W. and W. C. Kendall 1892, The fishes of Texas and the Rio Grande basin, con­ sidered chiefly with reference to their geographic distribution. Bull. U.S. Fish Comm., Vol. 12 (1094), PP* 57-126. 1902. Notes on the fishes of Lake Ontario. Part 27, Rept. U.S. Comm. Fish, for 1901, App., pp. 209-216.

-90 - Fish, Mari© Poland 1935- Contributions to th© early life histories of sixty-two species of fishes from Lake Erie and its tributary waters. Bull. U.S. Bur. Fish., Vol. 1+7 (1931-1933), pp. 293-396. Forbes, S. A. 1910. Article III.-On the general and interior dis­ tribution of Illinois fishes. Bull. 111. State Lab. Nat. Hist., Vol. 8, pp. 381— . Forbes, S. A. and H. E. Richardson 1919. Maps showing distribution of Illinois fishes (to accompany a report on the fishes of Illinois). 111. Nat. Hist. Surv., 111. State Jour. Co., Springfield, Plate 90, 1920. The fishes of Illinois. State of 111., Div. Nat. Hist., 397 PP. Fowler, H. W. 1919. A list of the fishes of Pennsylvania. Proc. Biol. Soc. of Washington, Vol. 32, pp. 1+9-71+. Friedrich, G. W. 1933. A catalogue of the fishes of central Minnesota. Copeia, No. 1, pp. 27-30. Good, Eugene E. 1914-6. Notes on the life history of the silver chub Hybopsls storerlanus in Lake Erie. Rept. for Zoology 623, Stone Institute, Put-in-Bay, Ohio, 8 pp. (typewritten) Greeley, J. R. 1929. F'ishes of the Erie-Niagara watershed. A biol­ ogical survey of the Erie-Niagara system. Suppl., 16th Ann. Hept., N.Y. Cons. Dept. (1928), Section b, pp. 190-179. 1940. Fishes of the watershed with annotated list. A biological survey of Lake Ontario watershed. Suppl., 29th Ann. Rept., N.Y. Cons. Dept. (1939), pp. 1+2-81. Greene, C. Willard 1939* The distribution of Wisconsin fishes. State Cons. Comm. Wis., Madison, Wis., 239 pp.

-91 (Jerking, S. D. 1^45. Distribution of the fishes of Indiana. Vol. 3, Investigations of Indiana Lakes and Streams, Ind. Dept. Cons, and Dept, of Zool., Ind. Univ., Indianapolis, 137 pp. Hankinaon, T. L. 1913. Distribution of fish in the streams about Charles­ ton, Illinois. 111. Acad. Sci., Vol. 6, pp. 102-113. 1929. Fishes of North Dakota. Mich. Acud. Sci., Arts, and Lett., Vol. 10 (1928), pp. i4.39-4.6O, Harlan, James H. and Everett B. Speaker 1961. Iowa fish and fishing. State Cons. Comm., Iowa, 237 pp. Hasler, Arthur D. and James R. Villemonte 1993. Observations on daily movements of fishes. Science, Vol. 118, No. 3064# pp. 321-322. Hay, 0. P. 1888. A contribution to the knowledge of the fishes of Kansas. Proc. U.S. Nat. Mus., Vol. 10 (1807), pp. 242-263. 1894. The lampreys and fishes of Indiana. Indiana Dept. Geol. and Nat. Resources, 19th Ann. Hept., pp. 146-296. Henshall, J. A. 1888. Contributions to the Ichthyology of Ohio. Jour. Cincinnati Soc. Nat. Hist., Vol. 11, Nos. 2/3, pp. 76-00. 1906. A list of the fishes of Montana. Univ. Montana Bull. No. 3d# Biol. Ser. No. 11, 12 pp. Hlnks, David 1943. The fishes of Manitoba. Dept. Mines and Nat. Resources, Prov. of Manitoba, 102 pp. Hubbs, Carl L. 1929. Further additions and corrections to the list of the fishes of the Great Lakes and tributary waters. Mich. Acad. Sci., Arts, and Lett., Vol. 11, pp. 425-436. Hubbs, C. L. and D. K. ii. Brown 1929. Materials for a distributional study of Ontario fishes. Trans. Royal Canadian Inst., Vol. 17, No. 1, pp. 1-56. Hubbs, C. L. and G. P. Cooper 1936. Minnows of Michigan. Cranbrook Inst, o d . bull. No. 8, 99 pp. Hubbs, C . .• and K. P. Lagler 1947. Fishes of the Great Lakes region. Cranbrook Inst. Gel. Bull. No. 26, 186 pp. Hubbs, C. and A. I. Ortenburger 1929 • Fishes collected In Oklahoma and Arkansas In 1927. Univ. okla. Biol, iurv., Vol. 1 , No. 3, pp. 47-112. Jenkins, H M., E. M • Leonard, and G. E. Hall 1992. An investigation of the fisheries resources of the Illinois River and pre-impoundment study of Tenkiller Reservoir, Oklahoma. Okla. Fish. Res. Lab. Rept. No. 26, 136 pp. (mimeo.) Johnson, H E. 199-2. The distribution of Nebraska fishes. Thesis, Univ. of Mich., li|9 pp. , 91 maps. Jordan, D. S. 1880. Description of new species of North American fishes. Proc. U.^. Nat. Mus., Vol. 2 (1879), pp. 239-241. 1882. Report on the fishes of Ohio. Report on the Geol. Surv. of Ohio, Vol. 4# Zool. and Bot., Part 1, Zool., Nevins and Meyers, Columbus, pp. 739-1002.

1891. Report of explorations in Colorado and Utah during the summer of 1889, with an account of the fishes found in each of the river basins examined. Bull. U.S. Fish Comm., Vol. 9 (1889), pp. 1-40.

1924. Concerning the genus Hybopsl3 of Agassiz. Copeia, No. 130, pp. 51-92. Jordan, D. S. and B. W. Evermann 1896. The fishes of North and Middle America. Bull. U.S. Nat. Mus., No. 47, Part 1, pp. 314-322. 1937. American food and game fishes. Doubleday, Doran, and Co., N.Y., 574 PP» Jordan, D. •>. and . E. Meek 1886. List of fishes collected in Iowa and Missouri in August, 1884, with descriptions of three new species. Proc. U.S. Nat. Mus., Vol. 8 , No. 1 (1885), pp. 1-17. Ketchen, K. S. 1951. Prellm?nary experiments to determine the working gape ox* trawling gear. Pish. Res. Bd. of Canada, Prog. Repts. of Pacific Coast Stations, No. 88, p . 62. Kinney, Edwara C., Jr. 1950. The life history of the trout perch, Percopsls orals comay cue (Wallbaum), In western Lake £ri'e. Thesis, M• S7, Ohio State Univ., 75 pp. 1951. The effect of temperature on the rate of diges­ tion of Stizostedlon vltreum vltreum. Unpublished data, Stone Institute of rfyclrobiology, Put-in-Bay, Ohio. Kirsch, P. H. lo93. Notes on a collection of fishes from the southern tributaries of the Cumberland River In Kentucky and Tennessee. Bull. U.S. Pish Comm., Vol. 11 (1891), pp. 259-268. 1895. A report upon explorations made in Eel River basin of Indiana In the summer of 1892. Bull. U.S. Pish Comm., vol. 11+ (1891+) , pp. 31-1+1. Kirtland, Jared P. 181+1+. Proceedings of the Boston Society of Natural History (1842), p. 71* l8l+7. Description of the fishes of Lake Erie, the Ohio River and their tributaries. Boston Jour. Nat. Hist., Vol. 5, Art. 2 (1845-1+7), pp. 30-32. 18^0. Pishes of Ohio. The Family Visitor, Vol. 1, No. 32, p. 256. Kobayashl, K. and H. Takahashi 1951.Study of fishing trawl operated in any depth of sea-water. Bull. Faculty of Pish., Hokkaido Univ., Vol. 1, No. 3-1+, 139 pp. Kuhne, E • R . 1939. A guide to the fishes of Tennessee and the mid­ south. Tenn. Dept. Cons., Nashville, 124 pp. Lawler, Karl P. and William C. Latta 1951+. Food conversion of the northern rockbass, Ambloplltea rupestrls rupestrls, In Michigan. Cope la, No." 1 , pp. 59-7l.

-94- Laverett, Frank and Frank B. Taylor 191b*- The pleistocene of Indiana and Michigan, and the history of the Grent Lakes. Monograph, U.S. Geol. Sarv. , Vol. 53, 529 pp. Ligas, Frank J. 1952. Migration, nesting, and food studies of the piscivorous birds of the island region of Lake Erie. Thesis, M.S., Ohio State Univ., 107 pp. Luce, Wilbur M. 1935. A survey of the fishing of the Kaskaskla River. Bull. 111. Nat. Hist. Surv., Vol. 20, No. 2, pp. 71-123. McCormick, L. M. 1892. Descriptive list of the fishes of Lorain County, Ohio. Oberlin College, Oberlln, Ohio, 33 pp. Meek, S. 2. 1892. A report upon the fishes of Iowa, based on observations and collections made during 1889, 1890, and 1891. Bull. U.S. Fish Comm., Vol. 10 (1890), pp. 217-21+8. 1895. Notes on the fishes of western Iowa and eastern Nebraska. Bull. U.S. Fish Comm., Vol. 11+ (1891+), pp. 133-1U2. 1895. Report on investigations respecting the fishes of Arkansas conducted during 1891, 1892, and 1893, with a synopsis of previous explorations in the same state. Bull. U.S. Fish Comm., Vol. 11+ (1891+), pp. 67-91+. Millar, F. G. 1952. Surface temperatures of the Great Lakes. Jour. Fish. Res.. Bd. Canada, Vol. 9, No. 7» pp. 329-376. Moore, George A. I960. The cutaneous sense organs of barbeled minnows adapted to life in the muddy waters of the Great Plains region. Trans. Am. Micro. Soc., Vol. 69, pp. 69-95. Nelson, E. W. I08I+. A partial catalogue of the fishes of Illinois. Bull. 111. State Lab. Nat. Hist., Vol. 1, No. 2, pp. 33-52.

-95- ... Nicholls, John V. V. 1931. The Influence of temperature on digestion In Fundu) is heteroclitua. Contrib. Canadian biol., n . s ., Vol~ 7~» No. 5 , pp. U7-55. O'Donnell, D. J. 193!?. Annotated list of the fishes of Illinois. Bull. 111. Nat. Hist. Surv., Vol. 20, No. 5» pp. ^73-500. Ohio St&te Kish Hatchery 195i±. Water temperature records, 1 ^i(l|.-1954, Put-in-Bay, Ohio, 11 pp. (unpublished). Osburn, H. C. 1901. The fishes of Ohio. Ohio Acad. Sci. Spec. Papers, No. I4., Spahr & Glenn, Columbus, 105 pp. Fearse, A. S. 19^i+. Amount of food eaten by four species of fresh­ water fi3hes. Ecol., Vol. 5* No. 3» pp. 25i+-258. Prather, E. E. 1951. Efficiency of food conversion by young largemouth bass Micropterus salmoldes (Lacepede), Trans. Am. F*ish. Soc. , Vol. 60 CI950), pp. I54“l£7. Radforth, Isobel 19i+i4.. Some considerations on the distribution of fishes in Ontario. Contrib. Royal Ont. Mus. Zool., No. 25, 116 pp. Rane y, E . C. 1953. Letter from D. C. Chandler, Cornell Univ., Feb. k, 1953. Sibley, C. K. 1929. The food of certain fishes of the Lake Erie drainage basin. A biological survey of the Erle-Niagara system. Suppl., 18th Ann. Rent., N.Y. Cons. Dept. (1928), Part 7# pp. I8O-I0 8 . Simon, J. R. and Felix Simon 1939. Check list and keys of the fishes of Wyoming. Univ. Wvo. Pub. in Sci., Vol. b, No. I4., pp. ^7-62. Slocum, Charles E. 1905. History of the Maumee River Basin. Nitschke Bros., Columbus, Ohio, 638 pp.

96 Snedecor, George W. 191+0. Statistical methods. The Iowa otate College Press, Ames, Iowa, 1+22 pp. Sprules, tf. M. 1993. Letter, Jan. 23, 1993* from Dept, of Fish., Ottawa, Canada. Starrett, tf. C. 1990. Distribution of the fishes of Boone County, Iowa, with special reference to minnows and darters. Am. Mid. Nat., Vol. 1+3* No. 1, pp. 112-127. Surber, Eugene W. 1939. Trout feeding experiments with natural food (Qammarus faaclatus). Trans. Am. Fish. Soc», Vol. (193*0 , pp. 300-301+. Thompson, D. H. and F. D. Hunt 1930, The fishes of Champaign County. Bull. 111. Nat. Hist. 3urv., Vol. 19, pp. 1-101. Trautman, Milton B. 1939. Stomach contents from Lota maculosa. Rept., Stone Institute, Put-in-Bay, Ohio, 9 pp. (typewritten) Van Oosten, John 1938. Mortality of fish on Lake Erie. Combined Biennial Repts., Penna. Bd. of Fish Comm., pp. 92-99. /Vagner, George 1908. Notes on the fish fauna of Lake Pepin. Trans. Wis. Acad. jcI., Arts, & Lett., Vol. 16, No. 1, pp. 23-37. mood, Kenneth G. 1993. Distribution and ecology of certain bottom-living invertebrates of the western basin of Lake Erie. Dissertation, Ph.D., Ohio State Univ., li+9 pp. Woolman, A. J. 1892. Report of an examination of the rivers of Ken­ tucky, with lists of fishes obtained. Bull. U.S. Fish Comm., Vol. 10 (1890), pp. 21+9-292. 1899. A report upon lchthyological Investigations in weatern Minnesota and eastern North Dakota. Rept. U.S. Comm. Fish, for 1893* App. No. 3* pp. 31+3-373.

97- Wri Tht» Stillman and Wilbur M. Tidd lu33. Summary of limnological investigations in western Lake Erie in 1929 and 1930. Trans. Am. Fish. Soc., Vol. 6j (lv33)> pp. 271-28S.

-98- AUTOBIOGRAPHY

I, Edward Coyle Kinney, Jr., was born In Massillon, Ohio, September 27, 1917* I received my secondary edu­ cation In the public schools of the city of Massillon. My undergraduate training was obtained at The Ohio State University, from which I received the degree Bachelor of Science in Education in 1914-1. I entered the armed forces in 1914-1 and was released from active duty in 1914-6 , with the rank of major. I re-entered The Ohio State University In 19L|.6 and received the degree Bachelor of Science In Agriculture in I9I4.8 . My graduate work was begun at the Franz Theodore Stone Institute of Hydrobiology of The Ohio State University in 1914-8. In 1950, I received the degree Master of Science. During the period 1914-8-1952, I served as Assistant to the Director of Stone Institute. In 1952, I received an appointment as Senior Conservation Fellow at Stone Institute. I held this position for two years while completing the requirements for the degree Doctor of Philosophy.

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