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University Microfilms International 300 North Zeeb Road Ann Arbor. Michigan 46106 USA St John's Road. Tylar's Groan High Wycombe. Bucks. England HP10 8HR 77-17,139 SMITH, Charles Gaylord, 1946- THE BIOLOGY OF THREE SPECIES OFJflXOSTOMA (PISCES:CATOSTOMIDAE) IN CLEAR CREEK, HOCKING AND FAIRFIELD COUNTIES, OHIO, WITH EMPHASIS ON THE GOLDEN REDHORSE, M. ERYTHRURUM (RAFINESQUE). The Ohio State University, Ph.D., 1977 Zoology

Xerox University Microfilmst Ann Arbor, Michigan 46106 THE BIOLOGY OP THREE SPECIES OP MOXOSTOMA (PISCES:CATOSTOMIDAE)

IN CLEAR CREEK, HOCKING AND FAIRPIELD COUNTIES, OHIO,

WITH EMPHASIS ON THE GOLDEN REDHORSE,

Mj. ERYTHRURUM (RAPINESQUE)

DISSERTATION

Presented in Partial Fulfillment of the Requirements

for the Degree Doctor of Philosophy in the

Graduate School of The Ohio State University

By

Charles Gaylord Smith, B.A. , M.S.

The Ohio State University

1977

Reading Committee: Approved By

Dr. Ted Cavender Dr. Tim Berra oy» a tyy* so / Dr. Bernard Griswold ///. Adviser Department of Zoology ACKNOWLEDGMENTS

I would like to thank my advisers Dr. Ted Cavender and

Dr. Walter Momot for their assistance during this study.

Appreciation is expressed to Dr. Tim Berra, Dr. Bernard

Griswold, and Dr. Roy Stein for their criticisms and sugges­ tions during the preparation of my dissertation, and to

Dr. Berra, Dr. Griswold, and Dr. Ronald Stuckey for serving on my final exam committee. I would also like to express my gratitude to Mr, Jack Nusbaum for his aid at Barnebey Center, to Mrs. Betty Porter for her excellent typing work, and to

Mr. Ken Covert for his help with field work. I am deeply grateful to my wife Carol and to my family and friends for their patience and support throughout this study.

ii VITA

6 June 1946 ...... Born - Syracuse, New York

1968...... B.A* in Zoology, The Ohio State University, Columbus, Ohio

1969-1972 ...... Secondary Biology Teacher, Cleveland Public Schools, Cleveland, Ohio

Summer 1971 ...... Ecology Instructor, Lake Erie Junior Nature and Science Center, Cleveland, Ohio

3972...... M.S. in Biology, The Cleveland State University, Cleveland, Ohio

1972-1976 ...... Graduate Teaching Associate, The Ohio State University, Columbus, Ohi o

Slimmer 1974 and 1975. . Mary H. 03burn Memorial Fund Fellowships, The Ohio State University, Columbus, Ohio

197 6 ...... Assistant Professor, Biology Department, Lake Erie College, Painesville, Ohio

197 7 ...... Ph.D. in Zoology, The Ohio State University, Columbus, Ohio

iii TABLE OF CONTENTS

Page ACKNOWLEDGMENTS ...... ii

VITA...... iii

LIST OF TABLES...... v

LIST OF FIGURES ...... viii

GENERAL INTRODUCTION...... 1

STUDY AREA...... 5

AGE AND GROWTH...... 15

MOVEMENT...... 58

FOOD HABITS ...... 88

SPAWNING BIOLOGY...... 96

SPECIES INTERACTIONS...... 120

SUMMARY ...... 130

APPENDIX

A ...... 137

B ...... 140

C ...... 141

D...... 147

LIST OF REFERENCES...... 149

It LIST OF TABLES

TABLE PAGE 1. Clear Creek data compared with published standard (3L), fork (FL) and total (TL) length conversion factors for golden and black redhorses. 23

2 . Backcalculated fork lengths at successive annuii for golden redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 27

3, Backcalculated fork lengths at successive annuii for black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 28

4. Successive annual length increments in millimeters for golden and black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 29

5. Population and mean individual instantaneous growth rates (by length) for untagged golden and black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 30 6. Yearly length increments in millimeters for golden and black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975 (averages do not include the first year1s growth). 32

Backcalculated fork lengths at successive annuii for immature male and female, mature male, and mature female golden redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 34

v TABLS PACK

8. Hmpirical (KM) and backcalculated (BC) average fork lengths (ram) for immature male and female, mature male, and mature female goLden redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1175. 56

9. Population and mean individual instantaneous growth rates (by length) for untagged immature male and female, mature male, and mature female golden redhorses captured in Clear Creek, Hock­ ing and Fairfield Counties, Ohio in April 1975. 57

10. Weight-Iength relationships for immature and mature Clear Creek golden redhorses electro­ shocked in the fall 1974, spring 1975, and fall 1975. 38

11. Weight-Iength relationships for immature and mature Clear Creek black redhorses electro­ shocked in the fall 1974, spring 1975, and fall 1975. 39 12. Weight-Iength relationships for immature and mature Clear Creek silver redhorses electro­ shocked in the fall 1974, spring 1975, and fall 1975. 40

15. Clear Creek data compared with published back- calculations of total length at successive annuii for golden redhorses. 49 14. Clear Creek data compared with published back- calculations of total length at successive annuii for black redhorses. 50

15. Clear Creek data compared with published weight-length relationships for golden red­ horses (determined by predictive linear regression analyses). 52

16. Clear Creek data compared with published weight-length relationships for black red­ horses (determined by predictive linear regression analyses). 55 17. Clear Creek data compared with published weight-length relationships for silver red­ horses (determined by predictive linear regression analyses). 54 vi iv.bbk 1AGR

-i 8 . Seasonal tagging and recapture records of golden, blacky and silver redhorses electro- shoctceu in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975. 66

19. Movement records (for each 10 mm size interval) of tagged golden redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975. 70

2 0 . Movement records (for each 10 mm size interval) of tagged black redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975. 71 21. Movement records (for each 10 mm size interval) of tagged silver redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975. 72 22. Movement records of tagged golden, black, and silver redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1975. 74

25. Upstream versus downstream movement records (as compared to the original capture site) for tagged golden, black, and silver redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975. 75 24. Clear Creek data compared with published infor­ mation concerning movement of golden, black and silver redhorses. 85

2b. Results of the analysis of intestinal contents (right loop) from golden, black, and silver redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1975. JI-92

?b. .Published data concerning food items consumed by golden, black, and silver redhorses. 95

27. Fecundity estimates for golden and black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in May 1975. 115

vii L 1 oT OF PIGLFHK3

FIGURF FAGF

1 . Map indicating the location of Clear Creek within the Hocking River drainage basin. 6 2 . Aerial view of the upstream, glaciated plateau region of Clear Creek, Fairfield County, Ohio (showing the covered bridge on Clearport Road). 7

5. Aerial view of the downstream, unglaciated valley region of Clear Creek, Hocking County, Ohio (3.5 km west of Route 33 on Clear Creek Road). 7

4. Map of the 12.4 km study area in Clear Creek, Hocking and Fairfield Counties, Ohio, indi­ cating the 53 sites at which redhorse were captured. 11-12

5. Fork length-scale radius functional regression lines as determined for 352 golden (L = 76.368 + 3-684S, r - .924) and 289 black redhorses (L - 76.650 + 4.6903, r = .893) caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 25

6. Fork length scale-radius functional regression lines for 100 immature male and female (L = 101.85 3 + 2.733S, r = .839), 115 mature male (L = 148.724 +■ 2.287 S, r » .674) and 137 mature female (L = 57.471 + 4.046 S, r = .711) golden redhorses caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 33

Weight-length functional regression lines for 117 immature male and female (LogW - -5.0849 + 3.0942 LogL, r = .9771), 240 mature male (LogW = -4.4989 + 2.8630 LogL, r = .8037), and 213 mature female (LogW = -5.4500 + 3.2520 LogL, r = .9294) golden redhorses caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 42

viii P J 6 ILL 8

Weighi-1 ctigth functional rt^reuuion lines for i Do immature male ana female (LogW = -6.2609 + 9.97ml Lo.'L, r = .0 999), 174 mature male (LogW = - 9. 22b9 + 9.1468 LogL, r = .9392), and 204 mature female (LogW = -9.9899 + 9.2867 LogL, r = .8895) black redhorses caught in Clear Creek, Hocking and Fairfield. Counties, Ohio in April 1979. 43

9. Weight-length functional regression lines for 19 immature male and female (LogW = -5.9202 + 3.2931 LogL, r = .9992), 16 mature male (LogW = -6.1128 +■ 3.5015 LogL, r = .9152), and 55 mature female (LogW = -5.5007 + 3.2711 LogL, r = .9808) silver redhorses caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 44

1 0. Weight-length functional regression lines for 639 golden (LogW = -4.9661 + 3.0495 LogL, r = .9813), 608 black (LogW = -5.3304 + 3.1728 LogL, r = .9832), and 60 silver (LogW = -5.4080 + 3.1990 LogL, r = .9955) redhorses caught in Clear Creek, Hocking and Fairfield Counties, Ohio in October 1975. 45 11. Mature male black redhorse (approximately 320 ramFL) captured in Clear Creek (site 48), Fairfield County, Ohio, tagged with a Floy T-bar anchor tag. 62

12. Prototype of the "artificial covers" placed at site 29 to determine the effect of stream cover on redhorse activity. 64

13. Redhorse capture site 29, located in Clear Creek, Hocking County, Ohio approximately six kilometers west of Route 33 adjacent Clear Creek Road. 78

14. The diel movement patterns of the redhorse aggregation at site 29 in Clear Creek, Hocking County, Ohio, approximately six kilometers west of Route 33 adjacent Clear Creek Road. 80

19. Tuberculate, male golden redhorse (approximately 280 mm FL) caught in Clear Creek, Fairfield County, Ohio in May 1975. 100

ix KUiUK'-: p a g j :

l.o. Tuberculate, male black redhorse (approximately jl70 mm FL) caught in Clear Creek, Fairfield County, Ohio in May 1975. 101

17. 'fubercuiate, male silver redhorse (approximately 400 mm FL) caught in Clear Creek, Fairfield County, Ohio in May 1975. 101

18. Dorsal view of the tubercles on a male golden redhorse (approximately 270 mm FL) caught in Clear Creek, Fairfield County, Ohio in May 1975. 1°2

19. Tuberculate, male golden redhorse (approximately 280 mm FL) upon removal from the spawning riffle at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. 104 20. Gravid, female golden redhorse (approximately 320 mm FL) upon removal from the spawning riffle at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. 104 21. Gravid, female golden redhorse (approximately 290 mm FL) caught in Clear Creek, Fairfield County, Ohio in May 1975, with a ventral orange blotch anterior to the left pelvic fin. 105 22. Spawning riffle at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. 106

23. Approximate locations of male defended terri­ tories on the spawning riffle at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. 108

24. Female golden redhorse flanked by two males just prior to the initiation of spawning, at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. 110

25. Female golden redhorse flanked by two males dur­ ing spawning, at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. 110

26. Fecundity-fork length functional regression lines as determined for 92 golden (LogF = -2.197 + 2.481 LogFL, r * .979) and 46 black redhorses (LogF * -7.536 + 4.501 LogFL, r =* .879) caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 114

x GENERAL INTRODUCTION

The research which I undertook involved the golden redhorse, Moxostoma erythrurum (Rafinesque), the black redhorse, Moxostoma duquesnei (LeSueur), and the silver redhorse, Moxostoma anisurum (Rafinesque), fish aptly cate­ gorized as non-game species. These redhorses belong to the ostariophysian family Catostomidae which consists of 11 genera with 57 species in the United States and Canada

(Bailey et al. 1970). The genus Moxostoma is distributed over most of North America east of the Rocky Mountains, with the golden, black, and silver redhorses restricted primarily to the mideastern United States. The latter boundaries, including the Mississippi and Mobile drainages and the south­ ern Great Lakes basins, are nearly identical for all three species (Jenkins 1970).

In Ohio the golden redhorse is present in nearly every permanent tributary (Trautman 1957) and often comprises the majority of fish biomass. Yet despite its widespread dis­

tribution and abundance, there is no definitive literature

on the golden redhorse in Ohio, and only one limited life history study outside of Ohio (i.e. Meyer 1962). Data con­

cerning the less abundant black and silver redhorses (which

1 are not as tolerant of siltation and pollution, Trautman

1957) are equally as sparse, with the sole major research contributions provided by Bowman (1959, 1970) in Missouri and Meyer (1962) in Iowa. This paucity of information has, from a literature standpoint, relegated these species to an insignificant position in fish communities. The references that are available are summarized in Carlander (1969) and

Jenkins (1970). These consist primarily of age and growth data, which are often insignificant due to very small sample sizes, and short, poorly substantiated statements of habitat preferences, food3, etc.

The biology of most members of the genus Moxostoma is very poorly understood (Robins and Raney 1956; Jenkins 1970; and Scott and Crossman 1975). According to Robins and Raney, the redhorses "have remained one of the most perplexing groups of fishes encountered by the American ichthyologist," This can be attributed to two major factors. First, redhorses are not game fish and therefore have received little attention from governmental agencies. Second, and most important, taxonomic problems (identification and nomenclature), involv­ ing the genus Moxostoma have been substantial, discouraging research and greatly complicating the acceptance of the limited literature available. Fortunately, Jenkins (1970) untangled much of the taxonomic disorder. His reexamination of specimens in the major American museums revealed numerous misidentifications. In addition he was able to correct many improper Moxostoma references in cases where specimens had been retained. The identification difficulties have also resulted in many references to "Moxostoma si£.111 this being especially true in survey publications (e.g. Brown I960).

The major complication in field work appears to be in differentiating the golden redhorse and the black redhorse, probably the two most confusing redhorses in Ohio. Although

a count of lateral line scales is usually sufficient for

identification, this procedure has often been considered

too time consuming for field work. Hubbs et al. (1943)

suggested that the identification difficulties associated

with the genus Moxostoma were due to: ”1. a tendency for

the species to be characterized by a combination of traits,

none of which by itself may be invariably distinctive; 2. a

common failure to recognize or appreciate all of these

characters."

With the aforementioned background and problems delin­

eated, in 1974 and 1975 I studied the golden, black, and

silver redhorse populations in Clear Creek, Hocking and

Fairfield Counties, Ohio. The intent was to examine in

detail certain aspects of the biology of these species, and

thus to fill some of the large and conspicuous voids in the

knowledge of the genus Moxostoma. It was a practical and

temporal impossibility to obtain a complete biological his­

tory, and therefore, special emphasis was accorded those

areas which would provide information beneficial to the understanding of the age and size composition, the growth, and the behavior of redhorse populations. In the majority of cases, "fishery biology” life history studies have been concerned with population parameters, while "ichthyological11 research has been more ethologically and ecologically directed.

Here I have attempted to combine important aspects of both.

The principle objectives of this work were as follows:

1. to establish the age structures of the three populations and to calculate the corresponding growth rates and weight- length relationships; 2. to ascertain the extent of seasonal movement, especially in regard to spawning migrations, and to examine diel activity; 3* to describe the feeding behavior and to determine the principle constituents of the redhorse diet; 4. to observe and photograph spawning behavior and to report associated factors, including environmental conditions, timing, sexual dimorphism, and fecundity; 5. to relate the interactions of the redhorse populations in the context of objectives 1-4.

Also of a practical necessity, research was conducted on three redhorse populations within a single section of a stream. This afforded an excellent opportunity for specific behavioral investigations, as I was able to become exceeding­ ly familiar with the location of aggregations of redhorses.

This was of greatest importance when attempting to observe spawning activities. STUDY AREA

Clear Creek is a major tributary of the Hocking River of the Ohio River Basin. With headwaters located in Amanda

Township, west-central Fairfield County, Clear Creek flows southeast through Clear Creek and Madison Townships (Fair­ field County) to its confluence with the Hocking River in

Good Hope Township, Hocking County (Figure l). The section upstream from the town of Revenge is located on the glaci­ ated Allegheny Plateau, which consists of rolling farmland

(Figure 2). The downstream portion flows through a steep, often gorge-like, unglaciated valley (Figure 3). The stream 2 is 37«1 km long and has a drainage area of 240.2 km . The average gradient is 2.83 m/km, with the greatest drop occurring in the first few kilometers. Headwater elevation is 334 m, dropping to 228 m at the mouth.

Historically, Clear Creek flows through one preglacial valley and part of a second, with the divide (now a very narrow gorge) located 1.4 km east of the Revenge Road bridge.

Before glaciation, a tributary of the Teays River ran west­ ward from the divide and a tributary of the Logan River flowed eastward from that point. The divide eroded through prior to the Wisconsin glacial period, thereby establishing

5 6

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!ioolville p U k f f“'| s\j Y 4 ’> S r T O , ,^wnpw?^^^ppf)r*7* - , m , i

Figure 1* Map indicating the location of Clear Creek within the Hocking River drainage basin, (Principal Streams and their Drainage Areas, Ohio Dept, Natural Resources, 1973) 7

Figure 2. Aerial view of the upstream, glaciated plateau region of Clear Creek, Fairfield County, Ohio (showing the covered bridge on Clearport Hoad).

Figure 3. Aerial view of the downstream, unglaciated valley region of Clear Creek, Hocking County, Ohio (3.5 km west of Route 33 on Clear Creek Road), the present drainage pattern. (Stout and Lamb 1958; Merrill

1950, 1953; Wolfe 1961; Wolfe et al. 1962)

The underlying rock strata for the entire drainage are

Mississippian sandstone. In the unglaciated valley, there

are numerous outcroppings of Black Hand sandstone of the

Cuyahoga Formation, which often forms the stream bed. Also

in the valley are deposits of Illinoian and Wisconsin out-

wash. Lacustrine deposits of Wisconsin origin occur west

of the former divide.

The geologic history of the Clear Creek watershed has

greatly influenced the water discharge pattern of the system.

The capture of land west of the preglacial divide greatly

increased the drainage area of the system. Considering this

increase in water supply and the gorge-like nature of the

lower portion of the drainage, Clear Creek discharge is that

of a flash flood stream. The United States Geological Survey

maintains a discharge monitoring station (03157000) 3-2 km

upstream from the mouth. Records from October 1939 indicate •5 a maximum discharge of 453 m /sec on 22 July 1945 and a mini-

mum discharge of .085 m /sec on 29 December 1947 (as a result

of a freezeup). Average discharge over 34 years has been

2.396 m /sec (Water Resources Data for Ohio). The stream

has a relatively constant discharge during dry weather due

to springs in the bedrock and to good ground water sources

in the glacial deposits of the headwater area. During the

period in which research was conducted, discharge ranged from .5 to 70.5 nrVsec with an average of 3.2 mVsec. The

Survey records indicate that the stream rises and recedes rapidly.

The vegetation of Clear Creek valley at the time of the

earliest land surveys was that of an Elm-Ash swamp forest.

Today the valley walls are heavily forested, with the valley

floor flora consisting of scattered, secondary deciduous growth. Large sycamore trees, PIatanus occidentalis, domi­ nate the streamside flora and horsetail, Equisetum fluviatile.

is the most common undergrowth.

During 1974 and 1975, 34 species of fish were captured

in Clear Creek (by seining and electrofishing), as compared

to 33 species (1939-1950) reported by Trautraan (1957).

Although the totals are similar, the species compositions

differ somewhat, especially in the Cyprinidae and Centrarch-

idae (Appendix A). The most unique feature of the Clear Creek

fish community is the presence of unusually large numbers of

catostomids, including golden, black, and silver redhorses,

white suckers, Catostomus commersoni (Lacepede), and hog

suckers, Hvpentelium nigricans (Lesueur). Large numbers of

quillbacks, Caroiodes cyprinu3 (Lesueur), were present only

during the spring, presumably due to a spawning migration.

Centrarchids were not abundant. This delimiting situation

provided an excellent opportunity to examine the biology of

redhorses in a stream ecosystem, although not necessarily under "typical” ecological conditions. 1 0

The major study area consisted of a 12.4 km section of stream in the unglaciated valley, from the mouth of Clear

Creek at Route 33 westward to the Clear Creek Road bridge west of Revenge. This section of Clear Creek can be classi­ fied as a fourth order stream (Horton 1945; Strahler 1957), with an average gradient of 1.47 m/km. A 1.7 km upstream section, from Landis Road to Strickler Road, was surveyed to determine if the redhorse species of the glaciated plateau differed from the primary study area.

Redhorses were captured at 53 sites within the major study area (figure 4). All sites were pools ranging in area from approximately 30 (site l) to 490 m (site 35), in depth from .5 to 2 m, and in flow from .18 to .3 m/sec (as measured with a flow meter at the pool center). The 53 pools comprised less than 20 percent of the study section, floods during the

study did not significantly alter the location or geomorphol­

ogy of the pools. The stream bed within the pools consisted nearly exclusively of sand and silt, with only occasional

small areas of rubble. Three pools in the study section

(labeled A, B, and C in figure 4) had gravel bottoms and no redhorses were taken from them. Pools inhabited by red­ horses contained no aquatic vegetation and were devoid of

cover, with the exception of infrequent submerged drift material (e.g. tree branches). The lack of natural cover

can be attributed to the flash flood nature of the stream.

Cladophora. restricted to riffle areas, was the only aquatic MB* HOCKING CO

\ \

11 10

16 _c.

271

\22

■^T

Figure 4. Map of the 12,4 km study area in Clear Creek, Hocking and Fairfield Counties, Ohio, indicating the 53 sites at which redhorses were captured (Rockbridge and Clearport Quadrangles, Ohio, 7*5 Minute Series, Topographic). h / V

I I Old Citric C»

f>tl fl Barnebey Center

*—; 3 6 '

29\

IMI Figure 4.(Continued) Map of.the 12.4 km study area in Clear Creek, Hocking and Fairfield Counties, Ohio, indicating the 53 sites at which redhorses were captured (Rockbridge and Clearport Quadrangles, Ohio, 7.5 Minute N> Series, Topographic). 1 3 plant present. Streambanks were unstable, except where sandstone outcrops occurred.

The excellent clarity of the water in Clear Creek enabled unrestricted observations of redhorses everywhere but in the bottom of the deepest pools. There were no indi­ cations of pollution problems immediately detrimental to fish populations. However, silt eroded from the glaciated plateau was often evident. Heavy fertilization of upstream farmland in the spring undoubtedly increased dissolved nutrients.

Liquid fertilizer was observed running from drainage tiles into the stream during the spring, 1975. Although no attempt wa3 made to sample periodically, a Hach Kit was used to analyze water samples from sites 11 and 59 and from near the Landis

Road bridge on 17 July and 13 October 1975 (Appendix B). ■5 Discharge rates for the two days were low, .96 and .99 m /sec respectively. Conductivity measurements at the three sites ranged from 380 to 440 microohms/cm, well within the accept­ able range for the available power supply for electrofishing

(Novotny and Priegel 1971)* Higher readings were obtained only during flood conditions, when the water became quite turbid. Water temperatures varied from 2° to 23° C.

The usability of Clear Creek as a research area was enhanced by good access from Clear Creek Road, by limited land ownership (primarily the Columbus Metropolitan Park

System and The Ohio State University), and by the adjacent location of Barnebey Environmental Research Center which provided equipment storage facilities. The necessity of an excellent field study area was accentuated by the futility of laboratory work, a3 it was impossible to simulate the proper environment on a large enough scale. Black and golden redhorses were maintained in an 1800 liter tank, but feeding was difficult and the tank was too small for the fish to exhibit normal diel behavior. AGE AND GROWTH

Introduction

Age and growth determinations are necessary requisites for an indepth examination of a fish population. This infor­ mation comprises a valuable segment of studies of population dynamics, allowing fishery biologists to analyze fish popu­ lations. Age and growth statistics reveal how successful a fish population is in reacting to the physical characteris­

tics and the biological influences of its aquatic environment.

In fishery work, the phenomenon of "stunting11 of centrarchids in farm ponds is a familiar example. Additionally, knowledge of the ages and the corresponding growth rates of fish within a population permits a more meaningful evaluation of other

data relevant to that population (e.g. movement and reproduc­

tion information). And, intra-population and inter-population

comparisons can be accomplished with more confidence. Specif­

ically, in this study age, weight, and length data were

obtained from samples of Clear Creek redhorses in order to

determine the age and size composition, the growth rates,

and the weight-1ength relationships of the three populations.

15 16

Methods

All of the redhorses captured for age and growth, weight-1ength, movement, and other studies (nearly 4000) were taken by direct current electrofishing. A 110 volt McCullough generator served as a power supply, with a Fisher Electro­ shocker converting the alternating current to direct current.

By varying the voltage regulator on the Electroshocker, the current was maintained at three amperes. This provided an effective circular to ovoid electrical field (dependent on waterflow) of approximately one meter in diameter. Fish out­ side of the effective range displayed a fluttering motion and usually escaped capture. Those within areas of sufficient current densities exhibited a galvanotropic reflex and swam involuntarily toward the anode.

The cathode consisted of a one meter square of hardware cloth and it was placed downstream from the area to be fished, usually in the tailwaters of a pool. A single, handheld elec­ trode (anode) was attached to 45 m of #16 electrical cord, thus allowing full coverage of even the largest pool. Each pool was fished upstream as those redhorses not effectively shocked tended to move to the head of a pool, and would sub­ sequently again be encountered. Equipment was transported between pools in a small plastic boat towed by a two to three member electroshocking crew. The mortality of electroshocked

(and tagged) fish was checked by placing fish in a laboratory stream tank and in wire cages in the stream (site 38). Stunned redhorses were scooped up with dipnets and placed in holding tubs. When electrofishing of a pool was completed, the redhorses were weighed, measured, tagged, and immediately released. Wet weights were obtained with a

2000 g pan balance. A fish measuring board was used to determine fork lengths (i.e. the distance from the anterior most section of the snout to the end of the median caudal rays). Field work necessitated this type of measurement.

The thick caudal peduncle prevented standard length measure­ ment due to the difficulty in locating the base of the hypural plate (Jenkins 1970). Total length measurement was also inappropriate as the upper and lower caudal lobes were frequently damaged. During the fall 1975, a size stratified sample consisting of 57 (150-540 mm FL.) golden and 57

(180-370 mm FL.) black redhorses were measured by all three methods to establish length conversion factors (GM functional regression analysis; i.e. the predictive regression coeffici­ ent divided by the correlation coefficient, Ricker 1975)#

Of the three basic methods of age determination, length frequency analysis, known age comparison, and growth zone counts (Tesch 1968), the latter proved most appropriate for redhorses. Employment of the former two was not practical

(and in the case of length frequency analysis not applicable), as they were dependent upon requirements which could not be fulfilled. The annuli (yearly growth marks) on the cycloid scales of redhorses were easily discernable and measurement 18 of the distances between annuli enabled backcalculation of lengths at younger ages. The scale method assumed that an­ nul i were formed at regular yearly intervals, that the distances between successive annuli and annual growth incre­ ments were directly related, and that the scales were present throughout the life of the fish (VanOosten 1929; Hile 1941).

Scales utilized for backcalculations were collected during April 1975* Age and growth information was ascer­ tained for both golden and black redhorses. Insufficient quantities of scales prevented a complete age and growth analysis of silver redhorses. However, the scales available were read, enabling a partial description of the population.

The golden and black redhorse samples were stratified as to size, with a minimum of six fish from each 10 mm interval of the respective size ranges. Each fish sampled had approx­ imately five scales removed from the region between the dorsal fin origin and the lateral line. It was not necessary to clean or to in any way process the scales. Regenerated scales or those of abnormal size or irregular shape were not considered. With the aid of a Ray-O-Scope scale projector, two scales from each fish were read three times. This in­ cluded counting the number of annuli present and measuring the distances between the scale focus and the various annuli in the mid-posterior field. Whereas scale formation occurred in late spring, the outer scale margin (posterior) was regarded as the final age mark (Erickson 1967). 19

Before backcalculations of length could he derived, the

fork length-scale radius relationship had to he established*

A plot of these two factors indicated a linear relationship.

Using Lee*s Method, L = a + cS (Lagler 1956; Tesch 1968),

GM functional linear regressions of body length on scale measurement (Whitney and Carlander 1956; Ricker 1973) were

calculated for golden and black redhorses.

The fork lengths and the distances to the successive annuli were averaged for each age class. These means were

substituted into the proper functional regression equation in

order to determine the backcalculated lengths. This procedure required much less time than does the determination of length

for each individual fish at each annulus with a graphic or numerical nomograph (Tesch 1968). And importantly, there is no loss of accuracy (VanOosten 1953).

Upon completion of the backcalculation procedure,

successive annual length increments were arranged in tabular

form by age and by year. The population or apparent instan­

taneous growth rates were computed by multiplying the differ­

ence between the natural logarithms of the mean fork lengths

of two successive age classes by the functional regression

coefficient from the appropriate weight-length relationship.

The mean individual or true instantaneous growth rates were

calculated by multiplying the difference between the natural

logarithms of the backcalculated lengths at the last two

annuli of each age class by the regression coefficient

(Ricker 1975). The golden redhorse information was subsequently reworked with a separation of immature male and female, mature male, and mature female data. The fork length-scale radius relationship was determined for each separately and in turn the backcalculated lengths were derived. Addition­ ally, population and mean individual instantaneous growth rates were calculated and results from the three subdivisions were compared.

Weight-length relationships were employed for intra­ population and inter-population comparisons of Clear Creek redhorses. The data used to establish weight-length rela­ tionships were obtained from individuals three years of age and older. The smallest golden, black, and silver redhorses captured were 150 mm, 180 mm, and 230 mm respectively. The spring 1975 data, collected prior to spawning,were parti­ tioned into immature males and females, mature males, and gravid females for each of the three species. The fall 1974 and 1975 data were not subdivided as field identification of males and females was not sufficiently accurate. Only in the spring when males possessed nuptial tubercles and females were gravid could sex be easily determined. Forbes and

Richardson (1908) and Reighard (1920) suggested that the anal fin and lower caudal lobe were longer in males than females. However, an attempt to employ this character was unsuccessful, as the difference was small and could not be readily ascertained in the field. 21

The formula V/ = a l P was utilized to calculate weight-

length relationships. The values of the constants "a" and

"b" were derived with a linear regression analysis of the

logarithm of weight on the logarithm of length. The equation was thus utilized in the form Log W = Log a + b LogL. When graphed on double logarithmic axes "Log a" became the

Y-intercept and "b" the slope ("b" was the regression

coefficient). (Tesch 1968; Weatherley 1972; Everhard et al.

1975; Ricker 1975)

For each of the three species of redhorses, both pre­ dictive (for comparison and reference purposes) and GM func­

tional linear regressions (Ricker 1973) were calculated for

the spring subdivisions, the pooled spring data, and the pooled fall data. In regard to size, the golden and black redhorse samples were stratified, with a minimum of five

individuals in each 10 mm interval of the size range. The small sample sizes of silver redhorses prevented this requirement from being statistified. The test for signifi­

cant difference between various GM functional regression

equations was accomplished by an analysis of covariance

(Snedecor and Cochran 1967).

Results

Electrofishing proved extremely successful on the aggre­ gations of redhorses found in the open pools of Clear Creek. 22

The excellent water clarity aided in increasing the efficiency of this technique. Mortality due to electrofishing was negli­ gible, as fishing was done during periods of relatively low water temperature (6°-13° C» except those taken during spawn­ ing in May 1975) and low discharge (.66-2.66 m^/sec). A total of 47 electroshocked (and tagged) redhorses were held in captivity 161 days, and subsequently released with no deaths occurring. Unlike the observations of rainbow trout by Bouck and Ball (1966), redhorses did not demonstrate prolonged fasting after being shocked. Redhorses retained in a stream- tank were feeding within 24 hours. In addition, during the many hours spent working in Clear Creek, no dead redhorses were found. In a few instances, fish appeared injured by electroshocking, and therefore were not tagged.

The length conversion factors for Clear Creek golden and black redhorses were very similar to those reported from

Illinois, Iowa, and Missouri (Table 1)* Within the 200-400 mm total length size range, variations in standard length were

8-16 mm for golden redhorses and 5-9 mm for black redhorses.

The total length-standard length relationships for Clear Creek and DesMoines River (Meyer 1962) golden redhorses were identi­ cal. In general, the similarity of length conversion factors suggested a degree of homogeneity of external body structure.

Scale reading was enhanced considerably by the occurrence of distinct "crossover" marks found in the dorso-lateral and ventro-lateral fields. Since annuli were formed in May and Table 1. Clear Creek data compared with published standard (SL), fork (FL) and total (TL) length conversion factors for golden and black redhorses.

To convert To convert To convert SL to FL TL to FL TL to SL SDecies location multiDlv bv: multiDlv bv: multiDlv bv: Reference golden redhorse Clear Creek, 0, 1.163 .910 .782 This Study golden redhorse Hutchins1 and .820 Lewis & Elder Clear Creek, 111* 1953 golden redhorse Des Moines River, 1.149 .898 .782 Meyer 1962 Iowa black redhorse Clear Creek, 0. 1.164 .922 .792 This Study black redhorse Niangua River, .810 Bowman 1959 Missouri black redhorse Big Piney River, .815 Bowman 1959 Missouri

ro 24 early June, spawning checks were not present. The problems

of "shouldered11 scales and irregular scale margins as

reported by Meyer (1962) were not encountered. Regenerated

scales appeared frequently, however this did not prove to be

an inhibitory consequence due to the large number of red­ horses sampled (approximately 900), Parsons (1953) reported

that thirty-five percent of the golden redhorse scales

examined were regenerated.

The validity of the annuli as age indicators was corrob­

orated by the removal of scales from ten golden and nine black redhorses in both the fall 1974 and the fall 1975, In

each case, the 1975 scales clearly showed an additional

annulus. Furthermore, the similarity of empirical and cal­ culated lengths and of calculated lengths of various age

classes at a specific annulus validated the use of the scale method of aging.

The analysis of scale annuli from the three Clear Creek redhorse populations showed that golden redhorses ranged in age from three to eight years (150-340 mm) and the black and

silver redhorses from three to seven years (180-390 mm and

230-450 mm). The fork length-scale radius functional regression lines as determined for 352 golden and 289 black redhorses were L = 76.368 + 3,684 S and L - 76,650 + 4,690 S respectively (Figure 5), The correlation coefficients (.924 and .893) denoted the low degree of variability. Backcalcu- lations derived from these equations revealed that golden 400 -

300

Fork Length (mm)

200

100 golden redhorse black redhorse

Scale rtadius - mm Figure 5. Fork length-scale radius functional regression lines as determined for 352 golden (L * 76.368 + 3.684S, r ■ .924) and 289 black redhorses (L ■ 76.650 + 4.690S, r * .893) caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. ro 26 redhorses averaged 97 mm after the first year (Table 2), while the black redhorse mean was 101 mm (Table 3). The golden redhorse size ranges for each age class did not over­ lap more than 15 mm. The black redhorse data overlapped a maximum of 30 mm between the 6 and 7 age classes. Results of a limited investigation of silver redhorses indicated that individuals in the 230-255 ram interval were three years old, those 265-305 mm were four years old, and those 385-450 ram were seven years old.

The successive length increments between age classes

(as determined from backcalculated lengths) demonstrated that for golden redhorses the largest average increase in length

(40 mm) occurred in the 2-3 and 3-4 age intervals (Table 4).

After the fourth year, annual length increments decreased.

The maximum average increase for black redhorses (55 mm) took place between age two and three, followed subsequently by a steady decline. The population and mean individual instan­ taneous growth rates coincided with these results (Table 5).

The largest population instantaneous growth rate (.719,

169-205 mm) and the greatest mean individual instantaneous growth rate (.995, 157-20 5 mm) for golden redhorses were calcu­ lated for the 3-4 age interval. Similarly, the maximum black redhorse instantaneous growth rates were in the 3-4 age interval (.222, 209-251 mm, population growth rate and .227,

204-251 min, mean individual growth rate). Table 2. Backcalculated fork lengths at successive annuli for golden redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975 (all lengths in mm).

Backcalculated FL. at Annuli AGE SIZE RANGE N AV.FL. . 1 . 2 3 4 5 6 7 8 3 150-205 54 169 94 128 169

4 190-215 35 202 95 125 157 205

5 220-260 77 244 96 130 178 217 241

6 260-300 127 269 97 128 162 201 232 267

7 290-320 43 297 99 130 175 211 236 267 294

8 330-340 16 337 98 116 157 192 235 261 286 312

Average length mm 97 126 166 205 236 265 290 312

ro Table 3. Backcalculated fork lengths at successive annuli for black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975 (all lengths in mm).

Backcalculated FL. at Annuli

AGE SIZE RANGE H AV.FL.nf 1 2 3 4 5 6 --- 7l---- ....— ------3 180-220 43 203 100 147 209

4 205-280 96 254 105 150 208 261

5 265-305 30 287 99 143 196 250 300

6 310-365 71 331 103 146 195 239 280 317

7 335-390 49 357 100 145 197 245 292 325 346

Average Length mm 101 146 201 249 291 321 346

ro 00 29

Table 4. Successive annual length increments in millimeters for golden and black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975.

Golden Redhorse

AGE______1 2 5 4 5 6 7 8 3 99 29 41 4 95 30 32 48 5 96 34 48 39 24 6 97 31 34 39 31 35 7 99 31 45 36 25 31 27 8 98 18 41 35 26 26 25 26 Average 97 29 40 40 27 31 26 26

Black Redhorse

3 100 47 62 4 105 45 58 53 5 99 44 53 54 50 6 103 43 49 44 41 37 7 100 45 52 48 47 33 21 Average 101 45 55 50 46 55 21 Table 5. Population and mean individual instantaneous growth rates (by length) for untagged golden and black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975.

Golden redhorse Population Growth______Mean Individual Growth Instantaneous Instantaneous Age length Growth Length Growth Interval Interval Rate Interval Rate (mm) (mm)

3-4 169-205 .719 157-205 .995

4-5 205-241 .604 217-241 .391

5-6 241-267 .380 232-267 .525

6-7 267-294 .358 267-294 .358

7-8 294-312 .220 286-312 .324

Black redhorse

3-4 209-261 .222 208-261 .227

4-5 261-300 .139 250-300 .182

5-6 300-317 .055 280-317 .124

6-7 317-346 .087 325-346 .063 31 The successive length increments were rearranged so as

to indicate the year in which the growth was attained

(Table 6). For both species the average yearly increment

for all age classes (excluding the first year of growth) was

distinctly consistent. From 1969 to 1974, the average yearly

increases in length ranged from 31-37 mm for the golden red­ horse population and from 45-48 mm for the black redhorse

population.

The analysis of the three subdivisions for the golden redhorse data resulted in fork length-scale radius relation­

ships of I s 101.853 + 2.733 S (r = .859) for immature males

and females, L = 148.724 + 2.287 S (r = .674) for mature males, and L = 57.471 + 4.046 5 (r - .711) for mature females

(Figure 6). The age data and the empirical and backcalculated

lengths are contained in Table 7. The immatures comprised

28 percent, the mature males 33 percent, and the mature females

59 percent of the total golden redhorse sample (352). The majority of immatures (54 percent) were three years old, with 35 percent four years old and 11 percent five years old.

Mature males (230-310 mm) were present in age classes 5 (30

percent), 6 (53 percent), and 7 (17 percent). The oldest

individuals were eight year old mature females (330-340 mm).

The youngest mature females were in age class 5 (230 mm).

Of the mature females, 2 3 percent were five years old, 48

percent six years old, 17 percent seven years old, and 12

percent eight years old. Table 6. Yearly length increments in millimeters for golden and black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975 (averages do not include the first year's growth).

Golden Redhorse AGE _ 1974_ _ 1973 1972 1971 1970 1969 1968 !9_67 3 41 29 (99) 4 48 32 30 (95) 5 24 39 48 34 (96) 6 35 31 39 34 31 (97) 7 27 31 25 36 45 31 (99) 8 26 25 26 26 35 41 18 (98) Average 36 31 34 33 37 36 18

Black Redhorse

3 62 47 (100) 4 53 58 45 (105) 5 50 54 53 44 (99) 6 37 41 44 49 43 (103) 7 21 33 47 48 52 45 (100) Average 45 47 47 47 48 45

V J 4 W 400

300

Pork Length (mm)

200

Immature males and females 100 mature males

mature females

1 2 3 4 5 6 7 Scale Radius (mm) Figure 6, Pork length-scale radius functional regression lines for 100 immature male and female (L = 101.853 + 2.733S, r - .839), 115 mature male (L * 148.724 + 2.287 S, r = .674) and 137 mature female (L * 57.471 + 4.046 S, r « .711) ^ golden redhorses caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. Table 7• Backcalculated fork lengths at successive annuli for immature male and female mature male, and mature female golden redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975 (all lengths in mm).

Backcalculated FL, at Annuli AG? LIFE STAGE SIZE RANGE N % of N AV.FL. 3 4 5 6 7 8 3 Immatures 150-205 54 54 169 172 M and F 4 190-215 35 35 202 162 196 5 220-250 11 11 225 173 198 225 Average Length 169 197 225 Average Increment 29.5 27

5 Mature 230-255 34 30 248 248 Males 6 220-290 61 53 262 242 264 7 285-310 20 17 293 249 268 285 Average Length 246 266 285 Average Increment 20.5 17

5 Mature 230-260 32 23 246 244 Females 6 260-300 66 48 276 233 272 7 290-320 23 17 300 238 267 296 8 330-340 16 12 337 231 260 288 31€ Average Length 237 266 292 316 Average Increment 52.3 28,5 28 35

A comparison of the average lengths (both empirical and

calculated) of the subdivisions (Table 8) indicated that at age five where overlap was complete, the immatures were

smaller than the matures and the mature males were larger than the mature females. At age seven the females were larger than males. The only age three and four fish were immatures.

In general, males and females matured at age five, the males being larger and having a shorter potential life span (to age

seven) and the females being smaller and having a longer potential life span (to age eight). The true instantaneous growth rates were largest for immatures in the 3-4 age inter­ val and for mature males and for mature females in the 5-6 age interval (Table 9). The growth rates of mature females were greater than mature males, but less than immatures.

The GM functional weight-length regression coefficients for the three redhorse species, as determined for the sub­ divided and pooled data (Tables 10, 11, and 12), were larger

than three in all but one case. Thus, growth was allometric and the weight gain per successive length increment was con­ tinually increasing. The single exception was mature, golden redhorse males which were captured in the spring 1975.

Covariance analyses revealed no significant differences between the functional regression lines of immature and mature (male and female) golden redhorses, immature and mature (male and female) black redhorses, mature male and mature female black redhorses, and mature male and mature Table 8. Qnpirical (EM) and backcalculated (BC) average fork lengths (mm) for immature male and female, mature male, and mature female gplden redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975*

A OF 5 4 5 6 7 8 ______EM BC EM BC EM BC EM BC EM BC EM BC______

Pooled Bata 169 166 202 205 244 236 269 265 297 290 337 312

Immature Males and Females 169 169 202 197 225 225

Mature Males 248 246 262 266 293 285

Mature Females 246 237 276 266 300 292 337 316

cn T a b le 9 . Population and mean individual instantaneous growth rates (by length) for untagged immature male and female, mature male, and mature female golden redhorses caotured in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975.

Population Growth Mean Individual Growth

Instantaneous Instantaneous Age Length Growth Length Growth Interval life Stage Interval(mm) Rates Interval(mm) Rates____

3-4 Immature 172-196 .405 162-196 .591 Males A Females 4-5 196-225 .427 198-225 .396

5-6 Mature Males 248-264 .180 242-264 .249

6-7 264-285 .218 268-285 .178

5-6 Mature Females 244-272 .354 233-272 .504

6-7 272-296 .276 267-296 .335

7-8 296-316 .211 288-316 .302

V>J -0 Table 10. Weight-length relationships for immature and mature Clear Creek golden redhorses electroshocked in the fa ll 1974, spring 1975, and fa ll 1975. Fish ranged in size from 150-340 mm fork length and in age from 3 -8+ years.

Weight-length Fork Len. Tyne of Relationship Correl, Date Life Stage (mm) size Reg session (g and mm) Coef.

Oct.-Nov. immature 150-340 545 Predictive logV/ = -4.8462 + 3.0011 Log! .9854 1974 and mature Functional LogW - -4.9531 + 3.0456 LogL

April 1975 immature 150-250 117 Prelictive LogW = -4.9223 + 3.0234 LogL .9771 Functional logW = -4*9849 + 3.0942 LogL

April 1975 mature 230-310 240 Predictive LogW * -3.1349 + 2.3009 LogL .8037 male Functional LogW = -4.4989 + 2.8630 LogL

April 1975 gravid 230-340 213 Predictive LogW - -4.8886 + 3.0223 LogL .9294 female Functional LogW = -5.4500 + 3.2520 LogL

April 1975 immature 150-340 570 Predictive LogW = -6.2463 + 3.5820 LogL .9613 and mature Functional LogW = -6.5886 + 3.7264 LogL

Oct. 1975 immature 150-340 639 Predictive LogW * -4.8519 ♦ 2.9925 LogL .9813 and mature Functional LogW = -4.9661 + 3.0495 LogL

v >j oo Table 11* Weight-length relationships for immature and mature Clear Creek black redhorses electroshocked in the fa ll 1974, spring 1975, and fa ll 1975. Pish ranged in size from 180-590 mm fork length and in age from 3+-7 years.

Weight-Length Pork Len. Sample Type of Relationship Correl Bate Life Stage (mm) size Regression (g and mm) Coef.

Oct.-Hot. immature 180-390 450 Predictive LogW = -5.2102 3.1286 LogL .9872 1974 and mature Functional LogW - -5.4134 3.1692 LogL

April 1975 immature 180-275 105 Predictive LogW = -5.7454 3.3631 LogL .9399 Functional LogW = -6.2605 3.5781 LogL

April 1975 mature 265-375 174 Predictive LogW ** -4*7481 2.9554 LogL .9392 male Functional LogW « -5.2285 3.1468 LogL

April 1975 gravid 265-390 204 Predictive LogW = -4.6607 2.9227 LogL .8893 female Functional LogW * -5.5855 3.2867 LogL

April 1975 immature 180-390 483 Predictive LogW * -5.1756 3.1252 LogL .9868 and mature Functional LogW - -5.2797 3.1669 LogL

Oct. 1975 immature 180-390 608 Predictive LogW * -5.2010 3.1195 LogL *9832 and mature Functional LogW = -5.3304 3.1728 LogL Table12# Weight-length relationships for immature and mature Clear Creek silver redhorses electroshocked in the fa ll 1974, spring 1975, and fa ll 1975- Pish ranged in size from 250-450 mm fork length and in age from 3+-7+ years.

Pork Len, Sample Type of Weight-Length Relationship Correl. Date Life Stage (mm) size Regression______(g and mm) Coef.

Oct,-Nov, immature 230-450 45 Predictive LogW xt -512997 + 3.1796 LogL .9931 1974 and mature Functional LogW a -5.4101 + 3.2017 LogL

April 1975 immature 230-320 19 Predictive LogW 3 -5.1662 + 3.1456 LogL .9952 Functional LogW X -5.5202 + 3.2931 LogL

April 1975 mature 380-420 16 Predictive LogW a -5.3436 + 3.2047 LogL .9152 male Functional LogW S3 -6.1128 + 3.5015 LogL

April 1975 gravid 410-450 35 Predictive LogW S3 -5.3366 + 3.2081 LogL .9808 female Functional LogW 0 -5.5007 + 3.2711 LogL

April 1975 immature 230-450 70 Predictive LogW S3 -5.0683 + 3.1033 LogL .9929 and mature Functional LogW m -5,1244 + 3.1254 LogL

Oct. 1975 immature 230-450 60 Predictive LogW = -5*2831 + 3.1846 LogL .9955 and mature Functional LogW S3 -5.4080 + 3.1990 LogL female silver redhorses. Significant differences were noted between mature male and mature female golden redhorses and immature and mature (male and female) silver redhorses. Yet, the latter differences were small, with the significance resulting from the closeness of fit of the data (a3 evidenced by the correlation coefficients). And for this reason, the pooling of the golden and silver redhorse spring data was rationalized. These intra-population relationships are best visualized graphically. Figures 7, 8, and 9 show the simi­ larity of the subdivided spring regression lines, despite the advanced gonadal development of the mature fish.

Comparisons of the pooled data, spring 1975-fall 1974, spring 1975-fall 1975, and fall 1974-fall 1975, for each species demonstrated a significant difference only for the spring versus fall golden redhorse data. The effect of gonadal growth was not as great as anticipated for black and silver redhorses. Nevertheless, in order to minimize this effect, the fall 1975 data were used for an interpopulation comparison. A covariance analysis revealed no significant differences among the regression lines of the three species

(Figure 10), and thus the increases in weight per unit of length were statistically similar. Comparisons among imma- tures, mature males, and gravid females (spring 1975 data) of the three redhorse species indicated significance for immature male and female and mature male fish. 794

/ 501 /

317 7/

199

126 immature males and females

79 mature males

mature females

50 \

160 199 251 317 398 Pork Length (mm) W Lght-length functional regression lines for 117 immature male and female C agW * -5.0849 + 3.0942 LogL, r * .9771), 240 mature male (LogV * -4.4989 + 2 3630 LogL, r * .8037), and 213 mature female (LogW = -5.4500 + 3.2520 LogL, r * .9294) golden redhorses caught in Clear Creek, Hocking and Fairfield unties, Ohio in April 1975. 794

501

317 Weif t

199 immature males and females

126 mature males

mature females

160 251 317 398 501 Fork Length (mm) Figure 8• Weight-length functional regression lines for 105 immature male and female (LogW * -6.2605 + 3.5781 LogLt r = .9399), 174 mature male (LogW ■ -5.2285 + 3.1468 LogL, r = .9392), and 204 mature female (LogW = -5.5855 + 3.2867 LogL, r = .8893; black redhorses caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. 1585

1000

631

398 Immature males and females

251 mature males

mature females

160

199 251 317 398 501 Fork Length (mm) Figure 9* Weight-length functional regression lines for 19 immature male and female (LogW = -5.5202 + 3.2931 LogL, r = .9952), 16 mature male (LogW * -6.1128 + 3.5015 LogL, r = .9152), and 35 mature female (LogW = -5.5007 + 3.2711 LogL, r = .9808) silver redhorses caught in Clear Greek, Hocking and Fairfield Counties, Ohio in April 1975. 1259

794

501

317

199

126 golden redhorse black redhorse

silver redhorse

160 199251 317 398 501 Pork Length (mm) Figure 10. Weight-length functional regression lines for 639 golden (LogW =* -4.9661 + 3.0495 LogL, r * .9813), 608 black (LogW = -5.3304 + 3*1728 LogL, r = .9832), and 60 silver (LogW = -5*4080 + 3.1990 LogL, r = *9955) redhorses caught in Clear Creek, Hocking and Fairfield Counties, Ohio in October 1975. 46

Discussion

Larimore (1961) reported that electrofishing captured

81 percent of the redhorses present in a study section of

Jordan Creek, Illinois. Results were verified by applying rotenone to the areas electrofished. Larimore attributed the high degree of success to the fact that the redhorses were in open water over clean bottoms and that these fish were easily seen and stunned. This situation was quite comparable to the study area, although the percentage of

Clear Creek fish captured (nearly 60 percent) was not as high.

The usefulness of electrofishing techniques for sampling redhorse populations was substantiated in Clear Creek. Yet, it is important to note that size is a critical factor.

Direct current electrofishing is very size selective, with larger fish (i.e. those with more surface area exposed to the electrical field) being much more readily stunned.

Initially, this was considered the explanation for the absence of small redhorses (less than 150 mm) from the electroshocked catch. However, subsequent sampling with a 30 ft bag seine and a 4 x 8 ft common seine during May, July, August, and

October 1975 and April 1976 (averaging four days per month) produced only seven redhorses under 150 mm (four black approximately 120 mm and three golden approximately 70 mm).

Therefore, it was believed that the redhorse populations in

Clear Creek consisted primarily of individuals 150 mm and larger and that electrofishing constituted a method of 47 sampling whereby the entire size range of redhorses was vulnerable.

Backcalculated lengths for golden and black redhorses did not exhibit the "Rosa Lee phenomenon" (i.e. data from the scales of older fish resulting in smaller estimates of length for younger ages than is actually correct; Ricker 1975).

Therefore, differences in mortality rates for various sizes within each age class were not indicated. But, a comparison of population and mean individual instantaneous growth rates did substantiate the presence of intra-class selective mortal­ ity. Within four of the five golden redhorse age intervals, three of the four black redhorse age intervals (Table 5), and five of the seven subdivided golden redhorse age intervals

(Table 9), population growth rates were smaller than mean individual growth rates, suggesting that in these age classes the larger fish have a higher mortality than smaller indivi­ duals. Thus, the apparent growth rates, which were based upon mean sizes of consecutive age classes, were less than the true growth rates.

The masking of the "Rosa Lee phenomenon" in the back- calculated data cannot be attributed to size selective samp­ ling. All available evidence supported the hypothesis that electrofishing was adequately effective on the full range of redhorse sizes (150-450 mm). To be of significance, sampling would have had to be selective for a specific size within an age class (Ricker 1975). Therefore, the complete absence of 48 the first three redhorse age classes could not have been a contributory factor. In all probability, the principle reason that the lee phenomenon was not detected was the variations in growth and mortality between males and females, as was specifically noted in the golden redhorse population.

One should be cognizant of the fact that this and other noteworthy factors were completely overlooked when the pooled golden redhorse data were analyzed. Much fishery and survey work does not allow for intra-population subdivisions, due to the extra time that would be involved in field work or simply to the lack of precise sex distinguishing characteris­ tics (as was the case for those redhorses captured in the fall). This has undoubtedly led to wrong conclusions regard­ ing mortality, growth rates, recruitment, and production, factors of extreme importance in analyzing sport and commer­ cial fish stocks. Practically, it may be impossible to obtain sufficient data, but as the golden redhorse example has shown, the consequences should be fully explored.

The comparison of growth in length of the Clear Creek golden and black redhorses with those from other locales vividly demonstrated the complexities of environmental regu­ lation. Clear Creek golden redhorses were in general the oldest, yet also the shortest and slowest growing (Table 13).

Conversely, the black redhorse data (Table 14) indicated that

Clear Creek fish were usually longer and faster growing than those reported from other streams. Thus, two closely related Table 13. Clear Creek data compared with published backcalculations of total length at successive annuli for golden redhorses. Those studies using less than 10 specimens were not considered.

Backcalculated TL at each annulus LOCATION N 1 2 ? 4 5 6 7 8 Reference Clear Creek, Ohio 352 107 138 182 225 259 291 319 343 This Study Hutchins1ft Clear Creeks, Lewis and Elder, Illinois 106 56 119 208 262 315 1953 Big Creek, Illinois 298 51 104 165 224 254 318 406 Lewis, 1957 Des Moines River, Iowa 1547 84 175 241 279 325 450 488 Meyer, 1962 Watts Bar Lake, Term. 42 157 284 368 455 500 Hargis, 1966 Chicakamauga Lake, Eschmeyer, Stroud, Tennessee 41 160 216 and Jones, 1944 White Oak Lake, Tenn. 74 140 226 279 375 Krumholz, 1956 Missouri 2549 79 150 218 272 310 333 Purkett, 1958 Salt River, Missouri 237 91 173 246 290 343 374 Purkett, 1957 Clearwater Lake, Missouri (Black River) Patriarche and pr e impoundment 61 155 257 323 Campbell, 1957 impoundment - 1948 76 249 373 457 impoundment - 1953 109 203 241 Lake Lawtonka, Oklahoma 549 94 218 305 378 445 521 579 625 Jenkins, Leonard, and Hall, 1952 Lake Spavinaw, Oklahoma 162 122 297 361 427 551 Jackson, 1966 Verdigris River Jenkins ft Finnell, tributaries, Oklahoma 48 130 206 279 325 1957 Lake Eueha, Oklahoma 89 145 302 391 434 490 Jackson, 1966 -p — .. — . ----Ki Table 14« Clear Creek data compared with published backcalculations of total length at successive annuli for black redhorses. Those studies using less than 10 specimens were not considered.

Backcalculatedl TL at each annulus

LOCATION N 1 2 3 4 5 6 7 8 9 10 Reference

Clear Creek, Ohio 289 110 159 218 271 316 349 376 This Study Missouri streams (statewide averages)2549 79 150 218 272 310 333 Purkett, 1958 Niangua River, Missouri 1625 84 127 173 213 244 269 287 297 302 325 Bowman, 1970 Big Piney River, Missouri 1775 94 168 229 277 290 310 320 325 338 Bowman, 1970 Clearwater Lake, Patriarche anc Missouri (Black River) Campbell, 195' pr eimpoundmen t 71 160 246 307 impoundment - 1948 91 239 353 434 impoundment - 1953 99 157 198 Lake Eucha, Oklahoma 109 119 284 384 484 480 Jackson, 1966

U) o species coexisted in the same habitat, yet one grew older

but slower than expected and the other faster and longer.

Importantly, it must be noted that the references for both

golden and black redhorses included data from streams of

varying size. The references which presented accounts of

golden and black redhorses over 400 mm total length obtained their data from impoundments. As noted by Carlander (1969),

growth in impoundments was considerably greater than in

rivers or streams. In evaluating preimpoundment and impound­ ment data, Fatriarche and Campbell (1957) reported a large

initial increase in growth of impounded golden and black red­

horses followed by a subsequent marked decline. This sequence most likely coincided with the productivity fluctuations after

impoundment.

The available weight-length relationships for golden, black and silver redhorses summarized in Tables 15, 16, and

17, presented an indication of the weight differences between various streams. Although not as correct statistically

(Ricker 1973), predictive regressions were employed to compare

Clear Creek data with published weight-length relationships

for redhorses, as a similar analysis was employed by the

other authors. Whereas the equations from the various refer­

ences incorporated different methods of length measurement,

(negating the possibility of directly comparing regression lines) an example was prepared for each species whereby weight was determined for two given lengths by utilizing the proper conversion factors. t

Table 15. Clear Creek data compared with published weight-length relationships for golden redhorses (determined by predictive linear regression analyses). Weights were calculated for 200 mm and 550 mm total length fish in order to evaluate the various relationships.

Location Weight-length Relationship Calculated Weights(g)Reference fg and mm 1)______200mmTL 550mmTL Clear Creek, Ohio This Study Fall 1974 LogW = -4.8462 + 5.001 LogFL 86 463 Fall 1975 LogW = -4.8519 + 2.9925 LogFL 82 433

Hutchins* and Clear Creek, LogW = -4.85 + 5.07 LogSL 89 497 Lewis & Elder Illinois - August 1951 1953

Meramec River, Missouri LogW = -4.881 + 2.975 LogTL 92 487 Purkett 1958

DesMoine3 River, Iowa LogW = -2.5057 + 5.098 LogTL 97 496 Meyer 1962 1960-1961 (lb and in)

U 1 N> Table 16. Clear Creek data compared with published weight-length relationships for black redhorses (determined by predictive linear regression analyses). Weights were calculated for 200 mm and 350 mm total length fish in order to evaluate the various relationships.

Weight-Length Relationship Calculated Weights(g) Location (g and mm) 20QmmTL 350mmTL Reference Clear Creek, Ohio This Study Fall 1974 LogW =* -5.2102 + 3.1286 LogFL 75 435 Fall 1975 LogW = -5.2010 + 3.1195 LogFL 74 422

Niangua River, Missouri LogW = -4.58 + 2.94 LogSL 82 427 Bowman 1959 1951-1957

Big Piney River, Missouri LogW = -4.59 + 2.95 LogSL 86 449 Bowman 1959 1951-1957

Meramec River, Missouri LogW * -4.977 + 2.998 LogTL 83 447 Purkett 1958

u i \ j i T a b le 1 7 . Clear Creek data compared with published weight-length relationships for silver redhorses (determined by predictive linear regression analyses). Weights were calculated for 200 mm and 400 mm fork length fish in order to evaluate the various relationships.

Weight-Length Relationship Calculated Weights(g) Location ______(g and mm)______200minFL 400mroFL Reference Clear Creek, Ohio This Study Fall 1974 LogW = -5.2997 + 5.1796 LogFL 104 942 Fall 1975 LogW = -5.2831 + 3.1846 LogFL 111 1008

LesMoines River, Iowa LogW * -2.53 + 3.1243 LogTL 114 967 Meyer 1962 1960-1961 (lb and in) 55

Comparing the weights of 200 mm and 350 mm total length

fish, the Clear Creek golden redhorses weighed less than

those from the Hutchins' and Clear Creeks, Illinois (Lewis

and Elder 1955), the Meramec River, Missouri (Purkett 1958),

and the LesMoines River, Iowa (Meyer 1962), Reported weights

of redhorses from Oklahoma (Jenkins et al, 1952), Tennessee

(Ward I960), and Alabama (Swingle 1965) were larger than

from Clear Creek. At 350 mm total length, the Clear Creek

golden redhorses were at least two years older than individ­

uals from the above locales. Weights of black redhorses

from Clear Creek were similar to those from the Niangua, Big

Piney (Bowman 1959), and Meramec Rivers (Purkett 1958) in

Missouri (although fish from the latter two rivers were

slightly heavier). Yet, Clear Creek black redhorses attained

comparable weights at a younger age than did Missouri fish.

Ages and corresponding weights and lengths of Clear Creek

and LesMoines River, Iowa (Meyer 1962) silver redhorses were very much alike. Thus, in terms of growth in weight and

length, the Clear Creek black redhorses were more successful

than either the golden or, to a lesser extent, the silver

redhorses. Ideally, more comparative information is needed

from throughout the ranges of the three species.

The weight-length relationships for Clear Creek red­ horses indicated differences in weight gain per successive length increment. For all three species, the heaviest fish per specific length, due undoubtedly to increased gonad weight, were captured in the spring 1975* Golden and black redhorses were heavier in the fall 1974 than in the fall

1975. Silver redhorses weighed more in the fall 1975. How­ ever, statistical comparisons revealed that the differences were not as significant as might be expected, especially when considering such variables as sex, maturity, season, and gonadal development. And, the weight-length correlation coefficients demonstrated the high degree of homogeneity of relationships within the redhorse samples (both pooled and subgrouped data)•

Golden redhorses can adapt to large, slow rivers and impoundments, whereas the requirements for black redhorses include higher gradients and clearer water. In Clear Creek, only golden redhorses were found on the glaciated plateau, as the gradient was less and the effects of agricultural runoff greater than in the unglaciated valley where both golden and black redhorses were abundant. Thus, the golden redhorse habitat requirements appeared less restrictive.

On the basis of inter-locale size comparisons, the unglaciated portion of Clear Creek was ideal black redhorse habitat.

There were no appreciable differences in the size ranges of the upstream (150-320 mm) and downstream (150-340 mm) golden redhorse populations. (Recaptures suggested the distinct possibility that there were two separate populations.) Clear

Creek may not have provided the best growing conditions for golden redhorses, yet in terms of age composition and abundancy (i.e. number of goldens versus blacks captured) they have successfully competed with the black redhorse population. The analysis of yearly population growth

(Table 6) for both species demonstrated the precision of population regulating mechanisms. M0VU4ENT

Introduction

The seasonal movement of anadromous and catadromous

fishes has often entertained the interest of ichthyologists

and fishery biologists. Yet, intra-stream seasonal movement

has. not received as an intensive or comprehensive research

emphasis, although there have been attempts to investigate

the seasonal activity of game fishes and certain non-game

species. Knowledge of movement patterns provide important

insights to such areas as density regulation, population

dispersal, fish production, spawning migrations, and homing behavior. In this study, the major objective was to deter­ mine whether redhorses were permanent, sedentary residents

of Clear Creek or seasonal migrants, exhibiting mass popula­

tion movements (especially spawning migrations), or if both

sedentary and mobile individuals were present. Additionally,

diel movement was examined, including activity patterns

exhibited and areas traversed.

Results of movement studies of stream fishes have been

conflicting. Some researchers, including Bangham and Benning­

ton (1938), Scott (1949), Tate (1949), Allen (1951), Larimore

(1954), Gerking (1953, 1959)* and Gunning and Schoop (1961), have concluded that movements of stream fishes are quite

58 59 restricted. Conversely, other studies have reported the sig­ nificance of stream fish movements (Stefanich 1952; Bowman

1959; Brown 1961; Behmer 1964; Bjornn and Mallet 1964; Hunt

1964; Shetter 1968; Lewis 1969). Berra and Gunning (1972) discussed both seasonal movements and home range. Substan­ tial evidence exists verifying the movement of stream fish into and from lakes (Shetter 1938; Raney and Webster 1942;

Carbine and Shetter 1943; Dence 1948; Brown and Graham 1953;

Warner 1959; Hartman et al. 1962; Niemuth 1967).

The majority of movement studies have incorporated the marking or tagging of fish in a specific locale and the sub­ sequent relocation of these individuals. Information regard­ ing the location of marked or tagged fish has been obtained from fisherman returns or from the active participation of the researcher in the acquisition of previously captured fish.

The latter has usually proven far more beneficial.

Many difficulties have been encountered in attempting to determine movement patterns. Most often, and probably unavoidably, arbitrary boundaries have been established and thus, movement has been defined as recapture outside of the original demarcations. However, as stressed by Gerking (1953), the limits of an individuals home range (i.e. the area in which an animal moves in its normal diel activity) may over­ lap a boundary. This naturally creates a bias in the number of fish considered to have moved. Yet, if sampling does not extend beyond the home ranges, the bias would favor the "no movement" group. Moreover, fish leaving the study area entirely cause significant statistical problems. Many of the detailed studies of fish movement have been limited to rather short sections of stream, and thus this becomes an important consideration. Additionally, fin clipping, a commonly employed marking technique, is restricted in the number of identifiable variations. Both marking and tagging often complicate the analysis of fish activity by altering normal behaviors. Finally, many fish movement studies are suspect simply on the basis of small sample sizes.

The study of the seasonal movement of redhorses described here is somewhat unique in that many of these difficulties have been eliminated as follows: 1* Each pool that held an aggregation of redhorses (age 3 and older) was considered a separate site, and thus there was no problem of where to establish boundaries. The redhorses were observed in riffle and run regions of the stream only when transferring between pools or during spawning. 2. The investigation included 53 sites over a 12.4 kilometer section of the stream (Figure 4), a distance sufficient to reduce the possibility of fish stray­ ing from the study area (especially since home ranges for golden and black redhorses were determined to be less than

.6 kilometers). 3. The tags employed were consecutively numbered, therefore enabling the identification of individual fish. 4. Hundreds of hours of observation of tagged red­ horses did not reveal any behavioral differences from unmarked 61 fish, 5. Sample sizes (2408) and tag recoveries (991) exceeded the levels of most detailed movement studies of freshwater fishes.

Methods

The major study area (Figure 4) was completely electro­ fished from the mouth of Clear Creek to site 43 in October and November 1974 and to site 53 in April and May 1975 and in October 1975. Seven sites (11, 14, 38-42) were sampled on 22 February 1975. The upstream section (Landis Road to

Strickier Road) was surveyed in April and May 1975 and in

October 1975. Redhorses captured during the fall 1974 and the winter and spring 1975 were tagged with consecutively numbered Floy, T-bar anchor, plastic tags (Floy Manufacturing

Company, Seattle, Washington). With a tagging gun, a tag was anchored between the interneural spines on the left flank near the dorsal insertion (Figure 11).

Tag numbers of redhorses recapturea in the winter, spring, and fall 1975 were recorded in conjunction with the number of the recapture site. Comparison with records of original cap­ ture sites enabled the determination of distance traveled for each individual recapture. Results were analyzed in order to ascertain size, seasonal, and species relationships. 62

Figure 11. Mature male black redhorse (approximately 320 mmFL captured in Clear Creek (~itc 4S), Fairfield County, Ohio, tagged with a Floy T-bar anchor tag.

Whereas the data from the winter and spring 1975 recap­ tures had suggested little seasonal movement, 109 redhorses

(63 golden and 46 black) were transplanted in May in order to determine if there was a homing tendency. Of the 41 fish re­ moved from site 29, 10 were transferred downstream to site 23

(1.5 km) and 10 to site 27 (.4 km) and 10 were released up­ stream at site 30 (.6 km) and 11 at site 32 (1.2 km). At site

38, 48 redhorses were electroshocked and 12 were replaced downstream at site 30 (2.0 km) and site 37 (.2 km) and up­ stream at site 39 (.5 km) and site 53 (4.0 km). In addition,

20 redhorses were removed from site 30, with 10 being placed 63 midway between sites 29 and 30 and sites 30 and 31 (.3 km downstream and .25 km upstream from site 30).

Field observations of diel activity were executed pri­ marily at sites 29 and 38, where the terrain provided excellent observational vantages. The superior clarity of the water and the lack of physical obstructions in the pools contributed to the success of field studies. Redhorse acti­ vity was viewed at distances of two to five meters (on occasion with the aid of binoculars). During August and

October 1975, the upstream and downstream portions of the activity cycle were timed.

In order to more closely observe and to photograph activity cycles and feeding behavior, a dock was built in

October 1975 at site 29. Projecting from the south bank three meters, at approximately 100 mm above the water sur­ face, it covered a section of the pool normally traversed by redhorses. The day following construction it was readily apparent that the redhorse aggregation had altered its move­ ment pattern by swimming around the end of the dock instead of going under it. With this in mind, three wooden and three glass "artificial covers" (.7 m sections on three legs) were constructed. (The prototype is pictured in Figure 12.) Com­ binations of three "covers" were situated in the stream in the normal redhorse movement routes and were systematically changed to determine the effect of stream cover on redhorse activity. Figure 12. Prototype of the "artificial covers" placed at site 29 to determine the effect of stream cover on redhorse activity.

Results

The Floy gun and tag provided an effective system for the individual marking of redhorses. Tag losses were not significant. Three golden and nine black redhorses which had shed tags were easily identified by the tag scar mark

(as compared to 991 recaptured with tags). More frustrating however were partial tag losses. The Floy tags used con­ sisted of two pieces of plastic glued together, a T-shaped anchor section and a colored plastic trailer. In 18 cases

(10 golden and 8 black) fish were caught with only the 65 plastic anchor remaining. A further difficulty was that of reading tags in the fall 1975, as they had become coated with a luxurious growth of algae which had to be scraped off before the tag number could be read.

Appendix C contains a complete accounting by species of the number of redhorses captured at each site in each of the seasons sampling occurred. The data for the upstream study area are in a composite listed as site 54. The quantities of redhorses captured per site varied tremendously, from six at site 1 to 488 at site 58. Both golden and black redhorses were caught at all 53 sites. Silver redhorses were only taken in the larger pools. As mentioned previously, the golden redhorse was the sole Moxostoma represented in the stream section surveyed on the glaciated plateau.

A total 2408 redhorses were tagged in October and Novem­ ber 1974 and in February, April and May 1975, 66 of which were in the upstream study area (1231 golden, 1061 black,

116 silver). Of those tagged, 865 or 35.9 percent were re­ captured, including 35.5 percent (437) of the goldens, 37.1 percent (394) of the blacks, and 29.3 percent (34) of the silvers. Total recaptures numbered 991 (494 golden, 460 black, 37 silver), including the 98 fish captured twice and the 14 fish taken three times.

The seasonal records (Table 18) indicated that that the probability of recapture decreased the longer a fish had been tagged, due to the increased chance of mortality. In 66

Table 18. Seasonal tagging and recapture records of golden, black, and silver redhorses electroshocked in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975.

GOLDENBLACKSILVER TOTALS 1. tagged 1231 1061 116 2408 2. tagged fall 1974 545 450 45 1040 5. tagged winter 1975 50 55 0 105 4. tagged spring 1975 636 556 71 1263 5. recaptured fall 1974 20 26 3 49 6. recaptured winter 1975 24 24 1 49 7. recaptured spring 1975 141 133 7 281 8. recaptured fall 1975 309 277 26 612 9. recaptures (not including multiple recaptures) 437 394 34 865 10. recaptures (including multiple recaptures) 494 460 37 991 11. recaptured twice 45 50 3 98 12. recaptured three times 6 8 0 14 13. tagged in the fall 1974 and recaptured in fall 1975 120 89 15 224 14. tagged in the winter 1975 and recaptured in fall 1975 10 17 0 27 15. tagged in the spring 1975 and recaptured in fall 1975 179 171 11 361 16. untagged captured in the fall 1975 640 625 63 1328 the fall 1975, 28.5 percent of the spring, 25.7 percent of

the winter, and 21.5 percent of the fall (1974) tagged fish

were recovered after approximately five, seven, and 11

months respectively. The number of recaptures in the spring

1975 (281) was 2.18 times less than the fall 1975 (612).

During the spring 1975, a total of 1544 redhorses were

taken, 18.2 percent being recaptures. In the fall 1975,

31,5 percent of the 1940 fish captured had previously been

tagged. This discrepancy may be attributed to spring move­

ment out of the study area. However, I believe it can (at

least partially) be explained by better electrofishing con­

ditions (lower discharge rates) and increased electrofishing

efficiency (enhanced by past experience) in the fall. Impor­

tantly, in the spring 1975, 24.5 percent of the redhorses

which had been tagged in Clear Creek were recaptured (281 of

1145), comparing similarly with the 25.4 percent taken in

the fall 1975 (612 of 2408).

The winter totals (Table 18) are based on one day*s

sampling of a limited number of sites (as previously listed).

The freeze-thaw nature of the weather kept discharge rates high during the winter months, and prevented a complete sur­ vey. Attempts at electroshocking during the late spring and summer 1974, when water temperatures were high, resulted in unacceptable redhorse mortality. Therefore, summer surveys were not conducted. Yet, observations made during June,

July, and August in 1974 and 1975 clearly showed that no mass exodus of redhorses occurred. 68

The recaptured golden redhorses moved an average of .425

Ion (0-9.2), the black redhorses .522 km (0-9.2), and the silver redhorses .782 km (0-4.75). Overall the 991 redhorse recaptures traveled an average of .484 km (0-9.2). A com­ parison of distances moved by fall 1975 recaptures versus spring 1975 recaptures demonstrated no significant difference

(Chi-square test of statistical significance) for golden

(.405 km versus .536 km) or black (.529 km versus .514 km) redhorses. Of the golden redhorse fall 1975 recaptures, 42.1 percent (130) had strayed one or more sections as compared to

39.0 percent (55) for the spring. For black redhorses, the percentages were 46.9 (130) and 47.4 (63). The 25 tagged golden redhorses electroshocxed on the spawning riffles in

May 1975 had traveled only from the adjacent pools where they had been originally captured. Therefore, over half of the recaptured golden and black redhorses in Clear Creek did not exhibit a spring spawning migration or any spring or fall mass movement pattern.

The information obtained concerning silver redhorses, even though lacking the extensiveness of the other two species, indicated the possibility of a spawning migration.

Of the spring recaptures, 85.7 percent (6) had moved, where­ as only 50 percent (13) of the fall recaptures had strayed.

Additionally silver redhorses were electroshocked upstream from site 38 only in the spring 1975. During the last week of April and the first week of May 1975, large aggregations 69 of approximately 20 to 30 individuals (male and female) were

observed on four separate occasions moving upstream between

sites 29 and 3 5 , These were the only homogeneous redhorse

aggregations observed during the entire study, with the

exception of groups on spawning riffles.

Electrofishing revealed that quillbacks definitely

exhibited a spring spawning migration. The fall 1974 and

1975 surveys captured no quillbacks. Yet, during the spring

1975, hundreds were shocked along the entire downstream and upstream study areas. White suckers and hog suckers were

extremely abundant during all seasons. But, none were marked, and it was thus impossible to determine if spawning migrations of these species occurred.

For the three redhorse species, there was no linear or distinguishable curvilinear relationship between average distance moved (for each 10 mm size interval) and length, although the longest average distances moved by golden and black redhorses were in the lower half of their respective size ranges (Tables 19, 20, and 21). Eight golden redhorses in the 220 mm interval moved sin average of 1,181 km, while

25 black redhorses in the 270 mm interval traveled an average of 1.306 km (one 210 mm recapture had moved 2.6 km). In general, average distances moved by silver redhorses were greater than for either golden or black redhorses. An exam­ ination of the recapture records revealed no direct correla­ tion between movement and fish captured more than once. In T a b le 1 9 * Movement records (for each 10 mm size interval) of tagged golden redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975.

Fall 1974 Winter 1975 Spring 1975 Fall 1975 Totals

Aver. Aver. Aver. Aver. Dist. Dist. Dist. Dist, Len. no moved no moved no moved no moved Total Av.Dist. mm move. moved km move. moved km move. moved km move. moved km Recaps moved-km 150 1 0 1 0

180 1 1.200 2 0 3 .400 190 2 0 5 5 .790 12 .658 200 4 .500 1 3 .963 1 6 .336 15 .547 210 1 0 3 3 1.250 7 1.071 220 1 .200 2 1 .917 2 2 1.625 8 1.181 250 1 0 4 0 3 7 .683 15 .455 240 1 0 4 4 .338 11 5 .091 25 .230 250 3 1 .100 8 3 .641 17 11 .261 43 .343 260 2 3 .260 5 3 .075 21 8 .547 33 25 .481 100 .457 270 3 3 .517 3 0 11 17 .708 40 37 .452 114 .507 280 2 .550 3 1 .025 22 6 .243 36 11 .180 81 .203 290 6 7 .762 16 13 .307 42 .448 300 1 .600 3 2 .720 6 2 .325 14 .486 310 1 0 1 0 1 2 .433 2 .45 7 .314 320 1 0 1 0 2 0 330 1 0 2 0 3 0 340 0 1 .800 1 .8 2 .800 O LTV 7 13 .405 18 6 .054 86 55 .536 179 130 • 494 .425

-j o Table 20* Movement records (for each 10 mm size interval) of tagged black redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975*

Fall i?74 Winter 1975 Spring J 3 T 5 M l 1^75 Totals Aver. Aver. Aver. Aver. Dist. Dist. Dist. Dist. Len* no moved no moved no moved no moved Total Av.Dist. mm move. moved km move. moved km move. moved km move. moved km Recaps moved-km 190 1 0 2 1.400 3 .933 200 1 .9 5 .400 6 .483 210 1 2.600 1 2.600 220 230 1 0 1 1 .650 3 .433 240 1 .8 2 6 .388 9 .433 250 1 1 .200 2 2 1.175 3 6 .422 15 .593 260 1 0 1 .100 1 2 .233 11 16 .822 32 .719 270 1 0 1 0 1 3 1.225 10 9 1.461 25 1.306 280 1 0 1 0 2 0 290 2 .150 3 2 .200 7 .186 300 1 .6 3 0 9 4 1.006 5 3 .444 25 .689 310 3 2.075 5 7 .290 6 9 .938 30 .793 320 1 7 .928 13 6 .521 17 7 .504 51 .577 330 2 .650 6 1 .057 11 12 .647 23 20 .421 75 .462 340 1 2 2.408 3 1 .025 15 13 .360 35 24 .351 94 .405 350 3 2 .240 3 2 .040 5 5 .150 17 10 .213 47 .184 360 1 .1 6 5 .291 8 6 .343 26 .312 370 1 0 4 2 .050 7 .043 380 1 0 1 0 390 1 .600 1 .600

8 18 .926 18 6 .050 70 63 .514 147 130 .529 460 .522

.-j H Table 21. Movement records (for each 10 mm size interval) of tagged silver redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975.

Fall 1974 Winter 1975 Soring 1975 Fall 1975 Totals Aver. Aver. Aver. Aver. Dist. Dist. Dist. Dist. Len. no moved no moved no moved no moved Total Av.Dist. mm move . moved km move. moved km move. moved km move. moved km Recans moved-km 230 1 .450 1 3.600 2 2.025 240 2 0 2 0 250 1 1 .600 2 .600 260 1 .200 1 .200 270 1 0 1 0 280 1 1 .45 1 3 1.538 6 1.175 290 1 1.300 1 1.300

350 1 2.400 1 2.400 360 1 1.200 1 1.200 370 2 1.638 1 2 .475 5 .940 380 2 0 2 0 390 1 1.0 1 1.000 400 1 .6 1 .9 1 2 1.117 5 .970 410 1 0 1 0 0 3 1 .250 6 .558 420 430 1 0 1 0

1 2 .500 1 0 0 1 6 .975 13 13 .793 37 .782

-J N> 75 some cases fish were recaptured two or three times at the

original tagging point and in others fish were recaptured at

two or three different sites.

Whereas redhorse habitat was restricted to pools, the number of sites moved were analyzed (Table 22) in addition to the actual distances traveled. The distances between sites averaged .24 km, varying from .075-.8 km. Of the 991 recaptures, 549 (55.4 percent) exhibited no inter-pool move­ ment (290 golden, 243 black, 16 silver). The redhorses which had strayed zero, one, and two pools accounted for

83.8 percent, 78.7 percent, and 54.1 percent of the recap­ tured golden, black, silver redhorses respectively. The winter data that were obtained revealed that 75.7 percent

(37) of the redhorses recaptured had not moved, with the rest having traveled no more than two sites.

The upstream versus downstream movement data (Table 23* for those fish which did travel between sites) demonstrated that deviation from random movement (i.e. 50 percent upstream and 50 percent downstream) was not significant for golden or silver redhorses. Yet, black redhorses demonstrated a sig­ nificant tendency to move upstream. In both instances, no seasonal trends were evident. The longest movement recorded was 43 sites (9.2 km), in one instance a black redhorse down­ stream and in the other a golden redhorse upstream. For each species, a majority of those which strayed from the original capture pool traveled upstream. This could possibly be T a b le 2 2 , Movement records of tagged golden, black, and silver redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1975. Movement is indicated by the number of pools traveled from the original capture site, minus signs signifying downstream and plus upstream.

Golden Black Silver Totals Movement Number Percent Humber Percent Number Percent Number Percent 0 290 58.7 243 52.8 16 43.2 549 55.4 -1 31 6.3 37 8.0 1 2.7 69 7.0 +1 43 8.7 45 9.8 1 2.7 92 9.3 -2 23 4.7 13 2.8 1 2.7 37 3.7 +2 27 5.5 24 5.2 1 2.7 52 5.2 -3 11 2.2 15 3.3 1 2.7 27 2.7 +3 10 2.0 16 3.5 1 2.7 27 2.7 -4 8 1.6 8 1.7 2 5.4 18 1.8 +4 6 1.2 1 .2 4 10.8 11 1.1 -5 3 .6 1 .2 1 2.7 5 .5 +5 4 .8 5 1.1 0 0 9 .9 -6 0 0 3 .7 0 0 3 .3 +6 4 .8 4 .9 1 2.7 9 .9 -7 0 0 2 .4 0 0 2 .2 +7 3 .6 5 .7 0 0 6 .6 -8 2 .4 1 .2 1 2.7 4 .4 +8 3 .6 2 .4 1 2.7 6 • 6 -9 1 .2 2 .4 0 0 3 .3 +9 0 0 2 .2 1 2.7 2 .2 -10 1 .2 1 .2 0 0 2 .2 +1°+ 2 .4 5 1.1 0 0 7 .7 12 2.4 8 1.7 3 8.1 23 2.3 +10 10 2.0 17 3.7 1 2.7 28 2.8

Totals 494 460 37 991 75

T a b le 2 3 . Upstream versus downstream movement records (as compared to the original capture site) for tagged golden, black, and silver redhorses recaptured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975.

Downstream______Upstream______Total Moved

Golden 94 (46.1%) 110 (53.9%) 204 (41.3%)

Black 91 (41.9%) 126 (58.1%) 217 (47.2%)

Silver 10 (47.6%) 11 (52.4%) 21 (56.8%)

Total 195 (44.1%) 247 (55.9%) 442 (44.6%) explained by the fact that electrofishing tended to force

fish upstream, although usually only to the head of a pool.

In the fall 1975, 24 (27 percent) of the fish trans­ ferred to pools were recaptured (13 golden and 11 black), with 17 taken at the transplant sites. Of the seven which had moved, the maximum distance traveled was three pools

(.7 km). From the 20 between pool transfers, 6 (3 golden and 3 black) were recaptured in pools downstream from their release point. (These 6 recaptures were not included in the movement data.) Thus, this first attempt at investigating redhorse homing behavior resulted in no indication of such an ability.

Another manifestation of the sedentary nature of Clear

Creek redhorses occurred at site 32, where 76 redhorses (26 golden, 41 black, and 9 silver) were tagged in the fall 1974,

Site 32 runs adjacent to Clear Creek Road. A flood (70.51 m /sec.) on February 2 3 and 24, 1975, resulted in a portion of the road being washed away. Approximately five dump truck loads of gravel were deposited on the streambank, with a sizable amount sliding into the pool (the deepest of the 53 sites). On April 10, 1975, when the spring survey reached site 32, only 8 golden (1 recapture) and 6 black redhorses were taken. The fall 1975 catch consisted of 14 golden (3 recaptures) and 4 black (2 recaptures) redhorses. Of those initially tagged in the fall 1974, 7 strays were recaptured in the spring 1975 and 7 strays were taken in the fall 1975. 77 Assuming that the pool regained its suitability as redhorse habitat (as all outward appearances indicated), the lack of homing behavior is again apparent, as well as an expected

(if indeed the fish are sedentary) slowness in the repopu­ lation of a pool.

The diel activity of redhorse aggregations (composed of goldens, blacks, and silvers) followed a similar pattern at all pools where observations were made. Basically, the red­ horses swam upstream through the length of a pool to the headwaters, reversed direction and traveled downstream to the tailwaters. This movement was repeated continually both diurnally and, as far as could be ascertained, noctumally.

On occasion the redhorses would not complete a cycle and would thus either return downstream before reaching the head of the pool or only complete a partial downstream trip. The time necessary to execute a cycle varied, but the downstream portion was much more rapid than the upstream segment.

At site 29 (Figure 13), the range of movement was approximately 36 meters. The time required to transverse this distance upstream ranged from 662 to 2578 sec, with an average of 1452 sec for 134 diurnal trials (Appendix D).

However, it was readily discemable that during daybreak and twilight hours, when feeding activity intensified, the up­ stream travel time increased. Upstream movement consisted of a series of starts and stops, with intermittent periods of feeding. During the intervals of no movement, individuals 78

Figure 13. Redhorse eanture site 29, located in Clear Creek, Hocking County, Ohio approximately six kilometers west of Route 35 adjacent Clear Creek Road. settled next to or on the substrate. The downstream move­ ment was much less complex. This component of the activity cycle lasted a maximum of 241 sec at site 29, and averaged

66 sec for 112 diurnal trials.

The greatest difficulty encountered was that of attempt­ ing to determine nocturnal activity. Initially, from the analysis of intestinal contents, it was hypothesized that redhorses left the pools at night, moving into the riffles to feed. But spring, summer, and fall observations during moon­ lit nights and with the aid of spotlights tended to discount this idea. A second check consisted of blocking off pools with nets before daylight to prevent the ingression of 79 redhorses (at sites 29 and 38), However, subsequent exam­

inations of the pools found the redhorses still present, and

thus nocturnal migrations from the pools were not considered

commonplace. Night observations of redhorse aggregations

suggested the existence of activity cycles similar to but

slower than diurnal patterns.

Within 24 hours after three wooden artificial covers

had been placed in movement routes at site 29, the redhorse

aggregation deviated from the typical design and had refused

to swim under the covers (Figure 14), In ensuing days, the

wooden covers were replaced by glass structures. At such

times, the redhorses modified their pathways and swam unhes­

itatingly beneath the glass covers. Alternating wooden and

glass covers were responded to as expected (i.e. avoiding

only the wooden covers). Lastly, when all artificial covers

were removed, the redhorse aggregation reverted to its orig­

inal movement cycle. As there were no known adult redhorse

predators (with the possible exception of an infrequent

osprey), stream cover did not constitute an essential survi­ val factor and it appeared to be actively avoided.

Discussion

The majority of golden and black redhorses age three and older (150-340 mm and 180-390 mm) were believed to be permanent, sedentary residents of Clear Creek throughout W 80 n A

36 m

V A, E B.

71

c. D.

wooden cover glass cover E 0 F ig u r e 14, The diel movement patterns of the redhorse aggregation at site 29 in Clear Creek, Hocking County, Ohio, approximately 6 km west of Route 33 adjacent Clear Creek Rd. A. Normal diel pattern B, movement pattern after placement of wooden "covers" C. Movement pattern with glass "covers" D. Movement pattern with alternating "covers" 8 1

the year (55.9 percent of the recaptures were taken at orig­

inal capture sites). Silver redhorses exhibited a tendency

to migrate upstream in the spring, although the data were

insufficient to be conclusive. The average distances moved

indicated that the home ranges of golden and black redhorses may in some cases encompass more than one pool (in regions where the pools were less than .6 km apart). Yet, these distances were relatively short, especially when considering that much of the catostomid literature has reported pro­ nounced spawning migrations. Dr. Milton Trautman has stated that his observations lead him to believe that, with the exception of salmonids, redhorses are the most highly migra­ tory (spawning) freshwater fishes (personal communication),

Contrarily, over 50 percent of the Clear Creek golden and black redhorse recaptures did not demonstrate any mass spawning migrations.

The question must be proposed as to whether or not the

Clear Creek redhorses undertook major movements during the winter and summer months when no fish were recaptured. If mass movements were to occur, one would expect them in con­ nection with the spring spawning period or possibly during the fall, as some fishes tend to move to larger pools or descend tributary streams. Clear Creek vas sampled during these periods and data from recaptures tended to discount the existence of major migrations. Summer observations

(i.e. noting large numbers of tagged redhorses within the 82

study area) and the few (49) winter recaptures supported this

hypothesis.

As with any mark and recapture study of movement, one

can discuss with certainty only those individuals which were recaptured. The fact that not all of the 2408 tagged red­ horses were recaptured may be attributed to mortality, migra­ tion, or incomplete sampling. Without the data, only an opinion may be expressed. Accordingly, on the basis of observations made while electrofishing, I ascribed the ab­ sence of 64.1 percent of the tagged fish primarily to the latter. Complete or 100 percent sampling success is nearly impossible, even in the smallest of streams. In Clear Creek, electrofishing was an efficient capture method. The 35.9 percent recapture record exceeded the results of many studies. However, the scope of the effective electrical field did not cover a complete transverse section of stream, thus allowing some fish to escape.

The information available concerning movement of golden, black, and silver redhorses is sparse (Table 24). The only partially extensive research of redhorse movement was by

Meyer (1962), although the study only spanned the period of

June to October I960. He reported that of 340 recaptured golden redhorses 45.6 percent made no movement from 1.07 km sections and of those which did stray, a significant number

(114) moved downstream. His results for silver redhorses indicated random movement of strays (45.5 percent did not move)• Table 24. Clear Creek data compared with published information concerning movement of golden, black and silver redhorses.

Percent Percent Number Percent Percent Moving Moving Locale Species marked Recaptured Not Moving Upstream Downstream Reference

Clear Creek, Ohio golden 1231 40.1 (494) 58.7 22.3 19.0 This Study DesMoines River, CO cn K\ Iowa golden 875 . (340) 45.6 20.9 33.5 Meyer, 1962 Missouri streams golden (19) 47.4 15.8 31.6 Punk, 1957 Richland Creek, Indiana golden 49 22.4 (11) 36.4 45.5 18.2 Gerking, 1950 Richland Creek, Indiana golden 89 31.5 (28) 85.7 7.1 7.1 Gerking, 1953 Clear Creek, Ohio black 1061 43.4 (460) 52.8 27.4 19.8 This Study Missouri streams black (15) 60.0 20.0 20.0 Punk, 1957 Clear Creek, Ohio silver 116 31.9 (37) 43.2 29.7 27.0 This Study DesMoines River, Iowa silver 297 18.5 (55) 45.5 70.0 34.5 Meyer, 1962

CD recaptured fish remained in one locale, whi; e t demonstrated a tendency to move. Punk ( 1 ^ t phenomenon by suggesting that a warn—water* s ~ r ~ lation was composed of both, sedentary an: neb:'*

Other researchers, including Moody ( 1 7*bC ) and 1 have extended this explanation to their resalts the major question still has not been answerer, certain fish stay in one area while otners t species stray? Berra and Gunning (1^71) stater difference in mobility "reflects the expected, indeed, necessary, biological variation ct:-ir.t respect to all characteristics”. T h i r v:-r_ : : : a species flexibility in .*•:-> :. . - :

The Clear Creek data revealed no r: e : i i r to how one would distinguish sedentary :‘r:r. ir:

Possibly, those individuals recarturei may re 3 with those not retaken comprising t h e m o t i l e . r were true, the mobile fish must have undertaken long downstream movements into the Hocking recapture data did not reflect a trend toward : movement. The 98 redhorse recaptured more t.vij nearly equally divided into three categories: remained in the same pool, those that traveled and those that strayed after one recapture and after another (or the latter sequence reversed) there possibly are genetic variations caur 1 n r . ; 84 In regard to seasonal movement patterns, Bowman (1970)

"believed that "black redhorses traveled to "larger1* pools to

overwinter, while in summer displaying an attachment to a

home pool. However, Bowman*s results were dependent on

visual sightings of tagged fish. Trautman (1957) stated

that golden redhorses migrate to "larger" pools to over­

winter. Berra and Gunning (1972) reported that longear

sunfish deserted their home ranges in winter, with many

returning in summer. Again, the evidence from Clear Creek

demonstrated that redhorses did not follow such seasonal movement patterns.

Minckley (1963) noted that a pool depleted of golden

redhorses by intensive electrofishing subsequently rarely

yielded any individuals, which "may indicate relatively

sedentary habits". Conversely, Larimore, et al. (1959) reported that golden redhorses repopulated a stream after a drought, although more slowly than other species. Gunning and Berra (1968, 1969) demonstrated that sharpfin chub-

suckers, Erimvzon tenuis (Agassiz), successfully repopulated

experimentally decimated sections of a stream within one year. In Clear Creek, redhorses were very slow in repopu­ lating a pool (site 32) and they did not exhibit homing behavior as has been reported for stream centrarchids

(Larimore 1952; Gerking 1953; Gunning 1959).

Many of the published accounts of stream fish movement display a distinct similarity in that a majority of the recaptured fish remained in one locale, while the others demonstrated a tendency to move. Punk (1955) explained this phenomenon by suggesting that a warm-water stream fish popu­ lation was composed of both sedentary and mobile individuals.

Other researchers, including Moody (i960) and Deacon (1961), have extended this explanation to their results. However, the major question still has not been answered. Why do certain fish stay in one area while others of the same species stray? Berra and Gunning (1972) stated that the difference in mobility "reflects the expected, normal, and indeed, necessary, biological variation encountered with respect to all characteristics". This variation would give a species flexibility in exploiting new habitats.

The Clear Creek data revealed no specific insights as to how one would distinguish sedentary from mobile fish.

Possibly, those individuals recaptured may be sedentary, with those not retaken comprising the mobile group. If this were true, the mobile fish must have undertaken relatively long downstream movements into the Hocking River. Yet, recapture data did not reflect a trend toward downstream movement. The 98 redhorse recaptured more than once were nearly equally divided into three categories: fish that remained in the same pool, those that traveled each time, and those that strayed after one recapture and did not move after another (or the latter sequence reversed). Whether there possibly are genetic variations causing physiological 86 differences, disease, injury, or parasitism factors, or 30me degree of social unacceptability or competition remains an extremely provocative question.

Observations of short term activity patterns exhibited by Clear Creek redhorses provided an introductory under­ standing of diel movement. The fact that redhorse movements appeared to demonstrate a marked degree of regularity is in agreement with the pronouncements of many researchers regard­ ing daily movements of freshwater fishes. However, a method is needed whereby activity may be quantitatively described over complete 24 hour periods* Reliable measurements have been exceedingly difficult to obtain, even in the laboratory.

Perhaps, a continuously monitoring telemetry system could be employed. Attempts at filming complete activity cycles in

Clear Creek failed.

The avoidance behavior displayed by Clear Creek red­ horses in reaction to the artificial wooden covers at site

29 coincided with observations of activity made at the few sites where natural cover was available. The movement patterns of redhorse aggregations were not viewed a3 being associated with cover. The amount and type of stream cover has long been regarded as an indicator of fish production.

Yet, for the three Moxostoma species in Clear Creek, the general absence of cover was not considered a limiting factor.

Mention must also be made of the 0, 1, and 2 redhorse age classes, which were not included in the movement study. As noted previously, electrofishing was effective for individuals age 3 and above, with those younger being conspicuously absent. Therefore, two possibilities exist.

First, the smaller redhorses may inhabit Clear Creek, but have been completely missed due to inefficient sampling techniques. Second, and most credible, the majority of redhorse fry may leave Clear Creek, with some eventually returning (homing?) at age 3. When the Hocking River was electrofished approximately one kilometer upstream and two kilometers downstream from the mouth of Clear Creek in

April, August, and November 1975, no redhorses of any size were captured (although coverage of greater distances would have been more desirable). Thus, the question of movement of redhorse young remains entirely unanswered. POOD HABITS

Introduction.

The energy sources available to a fish population constitute a major intra-population and inter-population regulating factor. The ability to obtain sufficient energy supplies directly influences growth, movement, and repro­ duction. Thus, it becomes important to understand what, where, and when energy sources are utilized. For Clear Creek redhorses, the analysis of food habits focused upon deter­ mining the major dietary constituents, the location of food supplies, and the times and areas at which feeding occurred.

Methods

The nature of the alimentary tract and correspondingly of the substances consumed created considerable difficulties in analyzing the food habits of the Clear Creek redhorses.

The redhorse alimentary tract consists of a long, anteriorly coiled tube of relatively uniform diameter. The tract does not contain a stomach, as indicated by the entrance of the bile duct posterior to the esophagus. Fish retained for study had the intestinal right loop removed and the contents from a 50 to 40 mm segment were subsequently examined. This

88 89 section was located between the esophagus and the posterior

bend in the intestine.

None of the standard quantitative methods of food analy­

sis were appropriate. Volumetric or dry weight measurements

of segments would have yielded little useful information due to the uniformity of fullness (i.e. either completely full or completely empty) and to the complexity of the intestinal composition. Frequency of occurrence or various other analyses based on enumeration of individual components were impossible. Even within the anterior portion of the alimen­ tary tract, aquatic insect larvae were usually broken apart due to the suctorial feeding action and the pharyngeal teeth.

Additionally, a large portion of the intestinal contents con­ sisted of detritus and algal filaments. Therefore, a quali­ tative analysis was performed on each intestinal segment, microscopically noting the dominant constituents. And, the percentages of empty versus full intestines were compared in order to evaluate changes in diel feeding patterns. Drift nets, a Surber sampler, and an Ekman dredge were employed to discover the location of redhorse food supplies.

Results

Feeding was a natural component of the upstream portion of the activity cycle, although, as previously mentioned, appearing to be more prevalent during the twilight and dawn hours. The ventral mouth of the redhorses enhanced substrate

contact. Films showed that feeding consisted of sucking in

bottom materials and expelling the undesirable constituents.

The efficiency of the filtering system was evidenced by the

near complete absence of sand grains in the intestines. The

primary intestinal constituents were detritus, algal fila­

ments, and aquatic insect larvae (Table 25). Very signifi­

cantly, there were no discernable differences among the food

items of the three redhorse species nor among the age classes

sampled, indicating a reliance on the same energy sources.

The composition of the redhorse intestinal samples was

dependent on the time (season) of capture. The intestines

of redhorses captured on 27 November 1974 contained detritus

and algal filaments (Cladophora). Redhorses electroshocked

on 22 February 1974 had consumed only detritus. During

April, May, and July 1975, in addition to detritus and algal

filaments, aquatic insect larvae were found in redhorse

intestines. Ten redhorses taken on 26 August 1975, had con­

sumed tremendous numbers of chironomid larvae (Chironomus)

to the near exclusion of all else. On 13 October 1975,

intestines were dissected from 60 redhorses. With the

exception of four fish, all the sample intestinal segments

were replete with Cladophora filaments. These seasonal

variations in the dietary constituents strongly suggested

a relationship to the availability of aquatic insect larvae

and algal filaments. T a b le 25. Results of the analysis of intestinal contents (right loop) from golden, black, and silver redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1975. Collections were obtained by electrofishing, except for July and August samples which were seined. Time Pate SpeciesN Site % Bnotv of Day Primary Identifiable Constituents 11-27-74 golden 15 43 13 LA detritus, algal filaments black 9 43 11 LA detritus, algal filaments 2-22-75 golden 6 14 33 EA detritus black 4 14 25 EA detritus 4-11-75 golden 5 38 0 LA detritus, algal filaments, chironomid, and mayfly larvae black 5 38 0 LA detritus, algal filaments, chironomid, and mayfly larvae silver 5 38 0 LA detritus, algal filaments, chironomid, and mayfly larvae 5- 3-75 golden 5 0 ^ 44-48 12 LM detritus, algal filaments, chironomid, mayfly, stonefly, and caddisfly larvae black 50U) 44-48 8 LM detritus, algal filaments, chironomid, mayfly, stonefly, and caddisfly larvae silver 4 48 25 LM detritus, algal filaments, chironomid, mayfly, stonefly, and caddisfly larvae 5-17-75 golden 18(2 ) 48 17 LM detritus, algal filaments, chironomid and mayfly larvae 2(3) 5-29-75 golden 29 0 LA detritus, algal filaments, chironomid, mayfly, stonefly, and caddisfly larvae ?(3) black 29 0 LA detritus, algal filaments, chironomid, mayfly, stonefly, and caddisfly larvae

(1)females U3ed also for fecundity studies Key: EM - early morning ' 'spawning males and females LM - late morning (^individuals which did not survive EA - early afternoon

LA - late afternoon \D (Continued) Table 25. Results of the analysis of intestinal contents (right loop) from golden, black, and silver redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in 1975. Collections were obtained by electrofishing, except for July end August samples which were seined.

Time Date Species N_____ Site % Empty of Day______Primary Identifiable Constituents 7-23-75 golden 5 29 0 EM detritus, algal filaments, chironomid, mayfly, stonefly, and caddisfly larvae black 5 29 0 EM detritus, algal filaments, chironomid, mayfly, stonefly, and caddisfly larvae 8-26-75 golden 5 42 0 EM chironomid larvae, detritus black 5 42 0 EM chironomid larvae, detritus 10- 5-75 golden 3 15 0 EA detritus, algal filaments, chironomid larvae black 4 15 0 EA detritus, algal filaments, chironomid larvae silver 2 15 0 EA detritus, algal filaments, chironomid larvae 10-13-75 golden 20 38 5 LA algal filaments black 20 38 0 LA algal filaments silver 20 38 15 LA algal filaments (sand grains) (white 12 38 0 LA algal filaments, detritus suckers) 281 9

Key: EM - early morning LM - late morning EA - early afternoon LA - late afternoon

\£> rvj Qualitative benthic sampling during the spring, summer, and fall 1975 demonstrated that, excepting small quantities of chironomid larvae found in pool substrates, the bulk of redhorse food items (algal filaments and aquatic insect larvae) originated from riffle areas. Therefore, if prior conclusions concerning diel movement patterns are correct

(i.e. restricted to pools), the redhorses were very dependent upon drift into the pools. Drift net sets verified that substantial quantities of aquatic organisms and organic debris were continually drifting downstream. Presumably, the reduction in flow rates within the pools enabled the heavier drift components to settle near or on the substrate, and thus become available to the bottom feeding redhorses.

Of the 281 anterior intestinal segments examined, only

9 percent were empty (Table 25). These included winter samples and intestines from spawning individuals. All fish captured in the early morning had full intestines, while 11 percent of the late morning, 16 percent of the early after­ noon, and five percent of the late afternoon samples were empty. Thus, feeding was discerned to be a continuous activity of redhorses.

Discussion

The catostomid morphological adaptations related to suctorial feeding have been detailed by Weisel (i960) and Jenkins (1970). The frequent feeding and the exclusively alkaline digestion (Al-Hussaini 1949) reflect the morphology of the redhorse alimentary tract. The necessity for recur­ rent feeding correlated with the low incidence of unfilled intestines. The intestinal length corresponded with the omnivorous to vegetarian diet. The available literature concerning adult golden, black, and silver redhorse food items (Table 26) differed most notably from the Clear Creek data in that algae was represented but once in the former and molluscs were completely absent from the latter (and from Clear Creek entirely).

In summary, Clear Creek redhorses were believed to feed continuously on detritus, algal filaments and aquatic insect larvae that drifted into the pools. Prom a more quantita­ tive standpoint, what will probably be necessary are calo­ rimetry studies and specific analyses of the nutritional values of different food items. Special emphasis should be accorded the determination of the nutrient significance of detritus. And, whereas the Clear Creek study indicated that drift provided a major portion of the food requirements for the large catostomid populations, drift should be quanti­ tatively analyzed and its periodicity compared with peaks of redhorse feeding activity. (Waters 1961, 1962, 1965 discussed the quantitative and productive importance of stream drift.) 95

Table 26. Published data concerning food items consumed by golden., black, and silver redhorses.

REFERENCE SPECIES FOOD ITEMS

Bowman 1959 black insect larvae, cladocera, copepoda, nemathelminthes

Carter 1969 golden detritus, insect larvae, molluscs

Harlan & Speaker 1951 golden insect larvae, molluscs

silver insect larvae, molluscs

Meyer 1962 golden insect larvae, molluscs, oligochaetes

silver insect larvae, molluscs

Minckley 1963 golden detritus, algae, insect larvae

Roach 1948 golden detritus, insect larvae, molluscs

Stegman 1959 golden detritus, insect larvae, crustaceans SPAWNING BIOLOGY

Introduction

Spawning is probably the least understood area of red­ horse biology and yet possibly the most critical in terms of the successful maintenance of redhorse populations in fresh­ water streams such as Clear Creek, Breder and Rosen (1966) have summarized catostomid reproduction, but there have been no definitive studies published concerning the spawning biology of golden or silver redhorses, and Bowman's work (1959, 1970) remains the single source of black redhorse data. Unfortu­ nately, this study cannot make a sizeable contribution to the knowledge of black or silver redhorse spawning biology, never­ theless certain related data were obtained. However, a great deal of information was acquired relative to the spawning of golden redhorses. This section will emphasize environmental conditions and timing of spawning, sexual dimorphism (includ­ ing tubercle formation), spawning behavior, hybridization, and fecundity.

Methods

In May 1975* spawning golden redhorses were observed and filmed (8mm) at ranges of one to three meters. Unlike the

96 97 aggregations in pools, the redhorses on spawning riffles were easily approached. The results presented are "based upon these observations, conditions noted during the springs of

1974 and 1976, and analysis of films of spawning sequences from 1975. Water temperatures were taken with a Yellow

Springs recording thermometer.

During May 1975, fecundity estimates were obtained for golden and black redhorses. Egg masses were removed from gravid females and preserved with Simpson's modification of

Gibson's Fluid* Estimations were determined volumetrically and results for individuals of the same length were averaged.

Whereas the best fecundity relationship exists with loga­ rithmic transformations of length (Bagenal and Braum 1968),

GM functional regression lines were calculated for golden and black redhorses.

Results

The Clear Creek members of the genus Moxostoma were spring spawners and survey records (noting the condition of tuberculate males and gravid females) clearly indicated that the silver redhorse spawned the earliest. Silver redhorse males (11) captured on 7 April 1975 possessed well-developed nuptial tubercles, while the first male black (2) and male golden (3) redhorses displaying rudimentary tubercles were electroshocked on 8 April and 10 April respectively. Records 9 8

also demonstrated that the silver redhorses had spawned prior

to the 3 May collection. The hest estimate would place the

1975 spawning season in the last week of April.

Confusion arose when attempting to decipher the spawning

sequence of golden and black redhorses. In the spring 1975, golden redhorses were observed spawning 15-19 May with water temperatures between 18° and 20° c. (The temperature had reached 18° C by 12 May.) Whereas no observations were made of black redhorses spawning, it was ascertained from capture records that black redhorses spawned in May, and in all like­ lihood after the golden redhorses. Whichever species spawned first, the separation was exceedingly short. Compounding the difficulty was the length of the spawning period, only five days for the golden redhorse (Clear Creek) and four days for the black redhorse (Bowman 1959).

In the spring 1976, abbreviated sampling (by seine) indicated that silver redhorses spawned in mid-April. Black redhorse males were seen on a spawning riffle at site 23 on

11 May, but no actual spawning was observed. On the two previous days gravid black females had been seined, along with numerous spent golden females and golden males with only tubercle scars remaining. Therefore the spawning sequence of Clear Creek redhorses was silver-golden-black.

There were additional spring spawning catostomids to consider: white suckers, quillback carpsuckers and hog suckers. White suckers were not observed spawning, but ripe females were electroshocked during mid-April 1975, and thus probably spawned before silver redhorses. On 8 May 1975, quillbacks were viewed spawning at dusk on a riffle in the upstream study area with a water temperature of 16° C, Hog suckers were infrequently viewed spawning over an extended period, from 2-12 May at water temperatures of 14° to 18° C.

Therefore, the catostomid spawning sequence was determined to be white sucker-silver redhorse-quillback-hog sucker- golden redhorse-black redhorse. Quillbacks, hogsuckers, and golden redhorses were observed spawning on the same riffle (at different times) and indirect evidence indicated that white suckers, silver redhorses and black redhorses also utilized the riffle. However, there were no indications that spawning periods temporally overlapped.

V/ater temperature appeared to have been the dominant factor controlling the timing of spawning periods. Two cir­ cumstantial observations tended to support that assumption.

First, the Clear Creek water temperatures in mid-November

1975 had declined to 7° C. Subsequently, unseasonably warm air temperatures increased water temperatures to as high as

11° C, During this time, 13 golden and 5 black redhorses with developing tubercles were caught. The growth of tuber­ cles is normally a spring occurrence associated with the spawning period. Electrofishing during November and early

December 1974 revealed no tuberculate individuals. Second, the early warm temperatures of the 1976 spring advanced the 100

I sequence of spawning periods approximately two to three weeks.

Sexual dimorphism, a characteristic trait of the family

Catostomidae (Breder and Rosen 1966), is displayed within the genus Moxostoma "by nuptial tubercles and dichromatism.

Tuberculate golden, black, and silver redhorse males are shown in Figures 15, 16, and 17 respectively. As they develop primarily on mature male fish, nuptial tubercles were first present on age six silver redhorses and on age five black and golden redhorses. Black redhorses from an adjacent drainage (Salt Creek) possessed tubercles at age three.

Figure 15. Tuberculate, male golden redhorse (approximately 280 mm FL) caught in Clear Creek, Fairfield County, Ohio in May 1975. 101

cl‘iaiafcM.4

W I k As

Figure 16. Tuberculate, male black redhorse (approximately 320 mm FL) caught in Clear Creek, Fairfield County, Ohio in Kay 1975.

Figure 17. Tuberculate, male silver redhorse (approximately 400 mm FL) caught in Clear Creek, Fairfield County, Ohio in May 1975. 102

The distribution of tubercles on silver and black red­ horses was identical, being confined to the rays of the anal and caudal fins. On occasion minute tubercles were seen on the dorsal surface of the paired fins. At the approach of the spawning period, the male golden redhorse was completely covered with tubercles (to the touch feeling like coarse sandpaper), with the largest occupying the anal and caudal fins and the head region. The prominent head tubercles exhibited no regular distribution pattern (Figure 18). The tubercles were evident five weeks before spawning and within two weeks afterward were sloughed off. In rare instances, vestigial tubercles were located on the snouts of female golden redhorses. A review of the films of golden redhorse

Figure 18. Dorsal view of the tubercles on a male golden redhorse (approximately 270 mm FL) caught in Clear Creek, Fairfield County, Ohio in May 1975. 103

spawning indicated that the major selective advantage of tubercles related to the maintenance of body contact during

spawning, to the defense of territories on the spawning riffle, and possibly to the stimulation of females and to the recognition of sex.

Dichromatism was evident during the spawning season.

Silver redhorses captured from homogeneous aggregations in the spring 1975 were bright, metallic silver, with the bases of the paired fins reddish in color. Black redhorse males observed on a spawning riffle had nearly black dorsal sur­ faces with wide, yellow-gold, mid-lateral bands that also went across the nape. Beneath this band ran another dark band that gradually lightened vcntrally.

The golden redhorses filmed spawning were distinctly dichromatic. Male color patterns were similar to black red­ horses, except generally being more intense and the dorsal surface being less dark. (The intensity difference may have been due to the fact that the black redhorses observed were not actually engaged in spawning.) Golden redhorse males and females electroshocked on the spawning riffle lost their markings immediately upon removal from the water. In photo­ graphs taken as rapidly as possible after capture, the mid­ lateral band was barely visible (Figures 19 and 20).

However, films of the fish in the water clearly showed the vivid coloration. Golden redhorse females also exhibited the horizontal marking, although the coloration was not 1 0 4

Figure 19. Tuberculate, male golden redhorse (approximately 280 mm FL) upon removal from the spawning riffle at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975.

Figure 20. Gravid, female golden redhorse (approximately 520 mm FL) upon removal from the spawning riffle at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. 1 0 5 usually as intense as the males. The paired fins of males

and females were orange to red-orange in color.

A puzzling color phenomenon noticed on the caudal peduncle

and ventral flanks of approximately five percent of the golden

redhorse females was the occurrence of bright orange blotches, o the largest being nearly 50 mm (Figure 21). The only func­

tion (if any) that could be attributed to this coloration was

sexual recognition. It was also noted that occasionally dur­

ing spawning a female with no mid-lateral band (probably

spent) would appear on the riffle and would be ignored by

the males, in striking contrast to their reaction to gravid

Figure 21. Gravid female golden redhorse (approximately 290 mm FL) caught in Clear Creek, Fairfield County, Ohio in Kay 1975, with a ventral orange blotch anterior to the left pelvic fin. females. Thus, the dichromatic coloration appeared to play a

role in sexual recognition.

Golden redhorses were observed and filmed spawning on

15-19 May 1975, at site 48 (Figure 22) and at a riffle near

the middle of the upstream study area. (Approximately 550 ft

of super 8 mm film was shot.) The substrate of the spawning

riffles was composed of coarse gravel and occasional rubble

(Wentworth scale) and the water flow rates ranged from .9 to

1.2 m/sec. The riffles were watched from before sunrise to

after sunset, but spawning only occurred between 1000 and

1400. Over the initial hour period, the males arrived on the

riffle and immediately aggressive behavior ensued in the

establishment of territories (i.e. defended areas). The

Figure 22. Spawning riffle at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975* 1 0 7

number of males varied from a maximum of approximately 20 to

six or less on the last day of the season. The exact size

of territories was dependent on the number of males present,

but in general they were circular, about .5 m in diameter.

Aggressive behavior took the form of actual contact, with males making rapid dashes and butting (with their tuberculate

heads) against competing males and various egg preditors.

Prom the initial observation of males on the spawning riffle it was evident that the larger males dominated the

center-most territories where successful spawning occurred.

By standing downstream from the riffle, 52 males were indi­ vidually electroshocked. The majority (34), age six and

seven and ranging from 270 to 510 mm, were taken from the

central positions. The remaining 18 were five years old

(230 to 260 ram) and had been relegated to peripheral terri­

tories. (The approximate locations of male defended terri­

tories at site 48 on 17 May 1975 are diagrammed in Figure 23-)

Bowman (1959) reported the exclusion of small, black redhorse males from the spawning area. At the conclusion of the day's

spawning activity when the males return to the pools, there was no indication of aggressive behavior.

In Clear Creek, the females grew larger and lived longer than did males. In the spring 1975, the sex ratio was .62 males to .38 females, a statistic related to the spawning behavioral patterns. Spawning females were 230 to 340 mm in length and were five to eight years old. Periodically, a 1 0 8

POOL (Site 48)

Males Age 6 and 7 (270-310 mm) w Males Age 5 (230-260 mm)

RIFFLE

Figure 23. Approximate locations of male defended territories on the spawning riffle at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. The dashed line indicates a typical movement pattern of a female. 1 0 9

female from the adjacent, upstream pool drifted down through

the riffle to a position below the males and then swam back upstream to assume a stationary position alongside of, but not in contact with, the left flank of a male (Figure 23).

Instantly, a second male occupied the open female flank and the three fish moved together. On occasion at this time, a

spawning attempt would be aborted. If the spawn was to be

successful, the males initiated violent, tetanic, caudal vibrations.

At this point, the three fish, flanking each other, were at approximately a 30 degree angle with the stream bottom, their caudal fins touching the substrate (Figure 24). As their bodies arched, the heads of the two males were slightly above and bent toward the female. Maintenance of contact was undoubtedly aided by the tubercles. The vibrations lasted two to four seconds, with milt and eggs being expelled during the last one or two seconds. During the vibrations, the sedi­ ment was agitated, resulting in a plume of sand and silt

(Figure 25). (Spawning riffles were easily located by the

"clear” bottom created by the vibrations loosening silt deposits.) The heavy eggs settled to the bottom and were at least partially buried. At the climax of spawning, the pro­ tractile mouth of the female was open and fully extended.

When the vibrations ceased, the three fish rose, often with their heads breaking the surface. One male and the female usually veered to one side, and the second male to the Figure 24. Female golden rednorse flanked by two males just prior to the initiation of spawning, at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975

Figure 25. Female golden redhorse flanked by two males during spawning, at site 48 in Clear Creek, Fairfield County, Ohio on 17 May 1975. 1X1

other side. The female subsequently returned to the upstream

pool. It was difficult to trace the movements of females after they had left the riffle, but in one instance a tagged female was observed to have spawned six times within an hour.

Undoubtedly, females engaged in repeated spawning. At the height of activity, as many as five or six spawnings were occurring simultaneously. ^Frequently, a third, fourth, or even a fifth male unsuccessfully attempted to join a spawn­ ing trio. Additionally, on occasion two or three males would momentarily flank each other and then aggressively split apart.

Spawning was last observed on 19 May. There were no more than six small males on the spawning riffle at one time.

Spawning rarely occurred as females appeared very infre­ quently. Successful spawning involving only one male and a female was noted (on the last day only). Also, male-male encounters were much more frequent. Most noticeably, the spawning riffle contained hundreds of egg preditors, includ­ ing: hog suckers; stoneroller minnows, Campostoma anomalum

(Rafinesque); striped shiners, Notropis chrysocephalus

(Rafinesque); and rosyface shiners, Notropis rubellus

(Agassiz). During spawning attempts, these fish would mass around the redhorses. On previous days, the more numerous males kept the riffle relatively free from invaders. Thus, the golden redhorse reproductive strategy dictated that the older, larger, more aggressive males had the greatest 112

probability of spawning successfully and the eggs which they

fertilized had initially the best chance of survival.

Fecundity estimates were determined for 92 golden and

46 black redhorses (Table 27). An analysis of covariance

demonstrated that the regression lines obtained for golden

and black redhorses were significantly different (Figure 26).

At comparable lengths, the fecundity of golden redhorses was

substantially greater than black redhorses.

Discussion

In regard to the timing of spawning, Meyers (1962) noted

silver redhorses spawning in the first week of May at 13° C, which corresponds to the Clear Creek temperature for the last week of April. Available information concerning white suckers and hog suckers coincided with Clear Creek data. Geen et al.

(1966) reported that white suckers were early spawners

(10° C). In New York, hog suckers spawned in May at 16° C

(Raney and Lachner 1946). Jenkins (1970) estimated a range of 14° to 22° C. There is virtually no comparative infor­ mation on quillbacks.

Conflicting data exist concerning the spawning sequence of golden and black redhorses. Hubbs (1930) and Jenkins (1970) stated that black redhorses spawn before golden redhorses at temperatures of 13° to 23° C. Bowman (1959) reported average spawning temperatures in Missouri for five years ranged from 113 Table 27. Fecundity estimates for golden and black redhorses captured in Clear Creek, Hocking and Fairfield Counties, Ohio in May 1975.

Golden Redhorses Age Fhnrm (N) Average Humber of Eggs per Female______

5 240 6 5041 5 250 5 5334 5 260 13 6159 6 270 13 7089 6 280 16 8115 6 290 15 8131 7 300 10 9438 7 310 6 9125 7 320 2 9766 8 330 2 12,400 8 340 4 11,310

Black Redhorses

5 300 1 3250 6 310 4 5600 6 320 9 5706 6 330 7 7939 6 340 8 6360 7 350 7 8061 7 360 10 8677 12589 ‘

10000

7945

6310 Fecundity (average number of eggs per female) 5012 golden redhorse

black redhorse

3982

282 317 355 398 Fork Length-mm Figure 26, Fecundity-fork length functional regression lines as determined for 92 golden (LogF » -2.197 + 2*481 LogFL, r ■ .979) and 46 black redhorses (LogF * -7.536 + 4.501 LogFL, r * .879) caught in Clear Creek, Hocking and Fairfield Counties, Ohio in April 1975. tTT 1 1 5 17° to 21° C. He suggested that black redhorse spawning

correlated with the full bloom of the dogwood and with the

first day of the April full moon. In Kansas black redhorses

spawned in April or early May at 16+° C and golden redhorses

spawned in May at 21+° C (Cross 1967). Reighard (1920) described the spawning behavior of golden redhorses (at the

time incorrectly identified as Moxostoma aureolum, Jenkins

1970), which he observed on 17 May 1904, and 4 May 1905, in southern Michigan. Also in Michigan, Hankinson (1951) noted golden redhorses attempting to spawn on 10-11 May 1930.

Jenkins (1970) stated that golden redhorses spawned on 24-25

May 1969 in Virginia. The disagreement in spawning dates is naturally expected due to climate and seasonal differences, yet the opposing references to spawning sequence strongly suggest some interesting variations among populations through­ out the ranges of similar species.

Although temperature appeared to be the critical factor in initiating spawning, the possibility that a hierarchy of cues was involved was quite plausible. Por example, photo­ period has often been regarded as an influence on reproductive timing, as laboratory studies of some fishes have demonstrated that it can regulate gonad development (Andreasen and Barnes,

1975). The problem was that in order to verify the importance of environmental factors one either must have detailed infor­ mation spanning many years or must be able to conduct labora­ tory experiments where conditions can be controlled. In the 1 1 6

case of redhorses, the former was complicated by the short

spawning periods, and of course temporal restrictions, while

the latter was prevented by the difficulties in maintaining

redhorses in aquaria. Interest in factors regulating

Moxostoma spawning was stimulated by the nonoccurrence of

hybridization. Naturally, other isolating mechanisms played

a role in the temporal separation of spawning periods and

they will be discussed later*

As reported in the movement section, records indicated

that silver redhorses may make spawning migrations. But

recaptures of golden and black redhorses did not reveal a

spawning movement. Por example, in May 1975, 25 tagged

golden redhorses (18 male and 7 female) were taken directly

from spawning riffles and all had been originally tagged in

adjacent pools. Gerking (1953) also noted that golden red­

horses did not migrate to spawn. This conclusion is in

opposition to reports by Hankinson (1920), Trautman (1957),

and Hanson and Campbell (1963) concerning golden redhorses

and by Bowman (1959) regarding black redhorses. Records of

spawning migrations by species in the genus Catostomus have been presented by Raney and Webster (1942), Dence (1948),

Brown and Graham (1953), and Geen et al. (1966). But in

Clear Creek, the data strongly substantiated the non­

existence of spawning migrations by golden and black redhorses.

In some instances, Clear Creek spawning observations

differed from accounts by Reighard (1920) and Bowman (1959, 117 1970). Reighard (1920) indicated that spawning golden red-

horses had bright salmon paired fins and a white lateral

stripe. Bowman (1959) observed black redhorses with a "light

pinkish mid-lateral band", and he noted no color changes for

females. The aggressive behavior of Clear Creek male golden

redhorses was in sharp contrast to Bowman*s (1959) description

of black redhorse behavior in which he stated that "I have never seen contact between two males in territorial defense."

Also, Bowman (1959) observed pre-spawning jumping activity by black redhorses. Hackney et al. (1968) reported that the river redhorse, Moxostoma carinaturn, (the species thought to be most closely related to the golden redhorse by Jenkins,

1970), built redds and exhibited courtship behavior. Such a distinctive behavioral difference from other redhorses may indicate a pattern of divergent evolution. In summary, all of these observations again indicate the possibility of signi­ ficant inter-population variability and stress the need for comparative studies.

Tubercles on the golden redhorse and most probably the other species of Moxostoma are limited to males, although

Reighard (1920) reported small tubercles on a very few females

(sis was the case in Clear Creek). The tubercles form from the hypertrophy and hyperplasia of epithelial cells which create mounds. The resulting tissue which lies above a plane parallel to the surface undergoes keratinization. This type of tubercle is less complex than those of Erimvzon and some 1 1 8

cyprinids which have solid keratinized cones supported by

vascularized hypertrophied epithelium (Wiley and Collette

1970). These complex tubercles are arranged in definite

patterns, a condition which Branson (1961) considered primi­

tive. Therefore, if Branson (1961) and Wiley and Collette

(1970) are correct, the golden redhorse possesses tubercles

which exhibit an advanced distribution pattern yet are morpho­

logically less complex than those species with primitive

spacial arrangements (a hypothesis which needs further veri­

fication). Tubercular vascularization may possibly be related

to the row-like distribution.

Thompson (1935) stated that redhorse identification pro­

blems are due to occasional hybrids. Yet, unlike the tribe

Catostomini, there are no verifiable records of hybridization

occurring in the Moxostomatini (Hubbs 1955; Jenkins 1970).

According to Hubbs (1955), hybridization in fishes is a

function of the intergradation of spawning habitat, that is

Increasing frequency of hybrids with increasing proximity

of spawning. In Clear Creek temporal and habitat isolating mechanisms did not appear to function with the necessary

degree of efficiency to prevent redhorse hybridization. Mayr

(1963) stated that "ethological barriers to random mating

contribute the largest and most important class of isolating mechanisms in animals". At least initially, variation in

spawning behavior and distribution of tubercles as related to

the territoriality of males would seem to constitute a formidable barrier to the hybridization of golden and black

redhorses. What is needed is further detailed information

on redhorse behavior, with attempts to artifically hybridize

redhorses. If successful, investigation of hybrid mortality,

sterility, and inviability should follow. Interestingly,

Nelson (1968), working with Catostomus commersoni (Lacepede) and C. macrocheilus (Girard), ascertained that despite the

occurrence of hybridization there was no evidence of the reinforcement of isolating mechanisms and thus ’’natural

selection against individuals producing hybrids is apparently

slow".

The only comparative fecundity data available are from

Meyer (1962) and Bowman (1959), regarding golden and black redhorses respectively. In the DesMoines River, Iowa, golden redhorses were larger and considerably more fecund than those in Clear Creek. Conversely, black redhorses from the Niangua and Big Piney Rivers in Missouri were smaller and less fecund them Clear Creek fish. Thus, the fecundity comparisons paralleled those for growth. SPECIES INTERACTIONS

The interaction of golden, black, and silver redhorse populations occurred in sand-silt bottomed pools. This habitat coincided with reports by Thompson and Hunt (1930),

Greene (1935), Allen and Clark (1943) and Minckley (1963) of golden redhorse preferences. However, Gerking (1945) stated that adult golden and silver redhorses did not demon­ strate an affinity for hard or soft substrates. Cross (1967) noted that golden and black redhorse preferred firm bottoms, while Forbes and Richardson (1908) related that golden red­ horses avoided mud bottoms. Trautman (1957) reported that in Ohio golden, black, and silver redhorses preferentially inhabited streams where silt did not accumulate. In the study area, pool substrates were partially composed of silt, but periodic water fluctuations (after each moderate rainfall) prevented silt from collecting. However, as compared to the study area, the glaciated plateau region of Clear Creek

(Figure 2) was characterized by increased accumulations of silt, a lower gradient, and natural cover at each pool. These conditions did not provide suitable black or silver redhorse habitat. Therefore, the redhorse inter-population relation­ ships In Clear Creek were believed restricted to the

120 121

unglaciated study area (i.e. the area where the habitat

requirements of the three species overlapped).

The most obvious manifestation of the interaction of the

three redhorse species was their gregarious nature. Within

the study area pools, golden, black, and, in the larger sites,

silver redhorses exhibited no tendency to divide into homo­

genous groups. Composite aggregations consisted of as many

as 250 redhorses (site 58). The gregariousness of the red­

horses was evident during all observations, except at spawning

time. Aggregations contained redhorses age 3 and older, male

and female, yet no aggressive behavior of any magnitude was

ever observed. The compactness of redhorse groups varied from

individuals being separated by less than 100 mm to individual

spacings of over 500 mm. The latter occurred regularly during

the most active feeding periods. Without knowledge of prior

movements, it would at these times appear as if the redhorses

lacked any degree of unity. The infrequent nonconformist

(i.e. a fish not moving with the aggregation) was always either

a large black or silver redhorse. No strong cohesive relation­

ship existed between individuals, as any disturbance would

disperse an aggregation in all directions.

Aggregations of single and mixed redhorse populations are

certainly not uncommon. Kuehne (1962) and Clay (1975) reported

that golden and black redhorses were usually collected together

in fourth order streams. However, at times the term "school" may have been inappropriately used to describe groups of redhorses (e.g. Raney and Lachner 1946). Redhorse aggregates

In Clear Creek did not display the behavioral characteristics which specifically distinguish schooling species. Explicitly, schools are comprised of individuals of the same species, often of the same life phase, that actively maintain contact with each other and that may exhibit organized movement at any instant. In an aggregation there is "no mutual endeavor of the fish to keep together" (Radakov 1973). Bowman (1959) reported that a school of black redhorses "moved as a group and maintained their positions relative to one another".

Such was definitely not the case with Clear Creek redhorses, however, the aggregations were not homogeneous. On the basis of laboratory studies, Shlaifer (1940) stated that with goldfish the homotypic group effect was lost when vari­ ous varieties of goldfish were mixed, but not to as great an extent as when different species were introduced. The one characteristic of schools demonstrated by redhorse aggregations was the lack of dominant individuals. The apparent absence of any hierarchy, however, does not elimin­ ate the possibility of some form of social interaction, whether it be simple or complex.

On three occasions, once in October 1974 and twice in

October 1975, the redhorse aggregation (golden, black, and silver) at site 29 was observed in a near perfect rank and file formation. This very distinct arrangement was quite surprising and no significant reason could be ascertained 123 as to the function of such a grouping. Since individuals

were viewed actively maintaining this formation, it was

not considered a random occurrence.

Within an aggregation "flanking" by individuals was a

commonly witnessed movement, whereby a redhorse would roll

over horizontally brushing against the bottom and would then

push itself away with a strong caudal motion. In perform­

ing this activity, the soft substrate was agitated, first

suggesting a relationship with the location of food. How­

ever, further observations tended to discount this hypothesis.

Similar flanking motions have been reported for many species, in which cases it has been considered a reaction to external parasites or microbes. Visual examinations of Clear Creek redhorses revealed no external parasites or regions of infection, yet the possibility of microscopic irregular­ ities was not discounted.

The golden, black, and silver redhorse interactions in

Clear Creek tended to nullify the Eltonian "niche" and the

Gausian exclusion principles, as the three species comprised one aggregation and consumed the same food. Weatherley

(1963), in a discussion of niche and competition among freshwater fish, stated that extinction need not result from two species living in the same habitat. He maintained that instead of increasing the density dependent mortality, fish growth rates decrease. In the case of golden redhorses, the age and growth data support this supposition, as fish 1 2 4 were smaller and older than comparable populations. However, black redhorses were generally larger than other reported populations.

In terms of relative abundance (on the basis of 2408 fish tagged), golden redhorses were the most numerous (51.1 percent), followed closely by black redhorses (44.1 percent).

Silver redhorses comprised only 4.8 percent of those marked.

The mark and recapture study of redhorses in Clear Creek was structured primarily to examine movement patterns and was not designed specifically to provide data for population estimates. Nevertheless, considering this major limitation, the golden and black redhorse population sizes within the study area were estimated using the Bailey ’'point" census method (Ricker 1975). Whereas, the time between mark and recapture periods was relatively long, the number of recap­ tures was large, and the recaptured fish were individually identifiable, the "point*' census was deemed most appropri­ ate. Additionally, the Bailey method allowed for the determination of survival and mortality rates. Due to the small numbers of recaptures, the silver redhorse population size was not estimated. The three "points" used were the fall 1974, the spring 1975, and fall 1975 samples. Calcu­ lations indicated a golden redhorse population of 2391 and a black redhorse population of 1884 during the fall 1975.

This resulted in a 55.9:44.1 golden to black ratio as compared to a 53.7:46.3 ratio of tagged fish. Thus, the 1 2 5

size of the tagged golden and black redhorse samples appeared

to be representative of the respective populations*

Despite more favorable growth data, the black redhorse

population size during the study was smaller than the golden

(spring 1975 estimates were golden 2171 and black 1415).

The survival rate for golden redhorses (.78) was greater

than for black redhorses (.64) and the instantaneous total mortality rate was less for the golden redhorse population

(.00158/day) than for the black (.00246/day). In general,

the information obtained concerning the two populations revealed contrasting reactions to density dependent regu­

lation; the comparably larger golden redhorse population

composed of older, slower growing individuals and the smaller black redhorse population consisting of younger, faster growing fish.

Additionally, the interaction of the two populations did not result in character displacement (i.e. closely related species demonstrate more accentuated differences when they are sympatric, Brown and Wilson, 1956; Nurshall,

1974). And conversely, golden redhorses did not exhibit characteristics convergent with black redhorses as has been reported by Jenkins (1970).

The information obtained regarding growth and move­ ment demonstrated that the interactions of Clear Creek redhorses contributed to population stability. Gerking

(1957) reported that excluding barriers the three factors 1 2 6 contributing to the stability of stream fi3h populations were homing, social behavior, and recognition of the home range.

As discussed previously, the Clear Creek evidence negates the former for fish three years and older. Social behavior is exceedingly difficult to evaluate, although much obser­ vational data has been related. Experiments in group behavior of fishes led tfelty (1934) to conclude that the group con­ tributes an "inter-reassurance" which reduces inhibitions that have been environmentally induced. For example, he stated that grouped fish consume larger quantities of food.

Again, despite the belief that social interactions do con­ stitute an important stabilizing mechanism, the final verification will be dependent upon experimentation under controlled conditions.

An important factor demonstrating the stability of

Clear Creek redhorse populations was that home ranges were less than one kilometer (i.e. the average distance moved by recaptured, tagged fish). As is usually true for tagging experiments, the nagging question is what happened to the tagged individuals not recaptured. In Clear Creek, the conclusion, based on observations of tagged fish escaping electroshocking, was that they were still present in the stream, in most cases within the delineated home ranges.

An additional indication of stability was the similarity of average yearly growth increments. 127 It must be noted that due to pollution the Hocking River

may constitute a physical barrier to redhorse movement and

thus enhance population stability (by promoting the inter­

action of the redhorse species). Nevertheless, this remains

only conjecture as the extent of the redhorse populations in

the Hocking River is unknown. At least one catostomid popu­

lation, quillbacks, did migrate from the Hocking River into

Clear Creek and white and hog suckers did inhabit sections

of the Hocking River-Clear Creek confluence* A weir designed

to capture fish moving downstream was maintained across Clear

Creek (below site 1) during the last three weeks of September

1974. However, during that period, as the water temperatures

dropped from summer highs to below 10° C, no fish were

captured*

Spawning behavior was characterized by a lack of species

interactions, as no hybrids were found nor were heterogeneous

groups observed on spawning riffles* The reproductive strat­

egy of golden redhorses, as reflected in spawning behavior,

is a modification of the basic pattern in the animal king­

dom whereby the offspring of the larger, often older, and more fit individuals have the greater change of survival*

In order to clarify the redhorse strategy, quantitative data

on initial hatching success are necessary, although exceed­

ingly difficult to obtain in the field* In Clear Creek at

the end of the spawning season, drift nets were placed below

riffles to catch hatching fry. But, the drift nets 1 2 8

accumulated debris (not uncommon in spring runoff) too quickly

to be of any value, Artifically fertilized golden redhorse

eggs hatched in three days and the yolk sac was absorbed in

20 days; data that at least provide a time span within which

future efforts can be concentrated. A short hatching period

may have been necessitated by the short time intervals be­

tween the spawning periods of the various catostomids,

especially as they may utilize the same riffle as a spawn­

ing ground. Any overlap or simultaneous spawning would tend

to increase egg mortality.

The major component lacking from this study is in

formation regarding redhorse biology from the fry stage to

age three. The few young golden and black redhorses that

were caught in Clear Creek were seined from the shallower

areas of pools. These individuals were with heterogenous

groups of young white and hog suckers and small Notropis

species. No young silver redhorse were ever captured. The

reports regarding young golden redhorse indicated that they

inhabit slow water pools (Thompson and Hunt 1930; Larimore,

Pickering, and Durham 1952) or deep, fast water near riffles

(Martin and Campbell, 1953). In analyzing fish movements

(including Moxostoma robustum and M. collapsum). Hall (1972) noted that the distribution of a fish population was aided by larger fish moving upstream and smaller fish downstream.

The smaller fish hatched in upstream areas where the spring and early summer energy pulse was greater per volume than 129 downstream. This contributed to the rapid growth necessary

for early survival. Subsequent, downstream movements of

fry were to areas providing a more stable environment. Hall

stated that "migration and reproduction are coupled to

optimize the use of energy resources". This reasoning would

at least outwardly appear related to the redhorse species

and their interactions, since the drainage geology would

characterize Clear Creek as unproductive. Yet, seining has

exposed large quantities of young white suckers and hog

suckers, thus posing the question as to why the same scheme would not be beneficial to these two species. The funda­ mental necessity is to locate and track the redhorse fry.

As v/ith any research project, more questions were gen­

erated than were answers provided. Yet, the most essential consideration is that relevant information was obtained which contributed to the basic understanding of Moxostoma biology and of interactions between redhorse species. The specific areas examined provided data that will begin to fill the voids in the present knowledge of redhorses and will enable more precise definitions of the role of red­ horses in stream fish communities. SUMMARY

The golden, black, and silver redhorse populations in

Clear Creek were studied during 1974 and 1975- Despite

their widespread distribution, these fishes are poorly understood members of the Catostomidae family. The scarcity

of information can be attributed to the taxonomic problems

associated with the genus and to the non-game status of redhorses. In this research project, the principal areas

of investigation included age and growth, movement, food habits, spawning biology, and species interaction.

The major study area was a 12.4 km seotion of Clear

Creek in Hocking and Fairfield Counties, Ohio, running from

the mouth of Clear Creek at Route 35 westward through an unglaciated valley to the Clear Creek Road bridge west of the town of Revenge. Redhorses were found in 53 sand-ailt bottomed pools, ranging in depth from .5 to 2 m and in area 2 from 30 to 490 m . Direct current electrofishing proved an effective capture method. Redhorses electroshocked were weighed, measured, and tagged with consecutively numbered

Floy T-bar tags. Studies demonstrated that mortality due to tagging was low.

1 3 0 Prom scale samples collected during April 1975, red­

horse ages were determined by counting scale annuli. The

validity of the annuli as age indicators was indirectly

substantiated. Golden redhorses captured ranged in age from

three to eight years (150-340 mm), black redhorses from

three to seven years (180-390 mm), and silver redhorses

from three to seven years (230-450 mm). Backcalculations

for golden and black redhorses indicated average first year lengths of 97 mm and 100 mm respectively. Successive length increments between age classes and individual instan­ taneous growth rates revealed that after the first year the greatest increase in both golden and black redhorse size occurred in the 2-3 and 3-4 age intervals. Yearly compar­ isons from 1969 to 1974 of all age classes demonstrated that the average increase in length was distinctly con­ sistent, 31-37 mm for golden redhorses and 45-48 mm for black redhorses.

The backcalculated lengths did not exhibit the Rosa Lee phenomenon. However a comparison of population and mean individual growth rates did indicate intra-class selective mortality. The masking of the Lee phenomenon was attributed to variations in growth and mortality between males and females.

The subdivision of the golden redhorse scale data revealed that immatures were three to five years old (150-

250 mm), mature males five to seven years old (230-310 mm), 132 and mature females five to eight years old (230-340 mm).

The growth rate of mature females was greater than mature males, hut less than immatures.

Weight-length relationships for golden, black, and

silver redhorses showed few intra-species or inter-species differences. This was unexpected, especially when con­ sidering seasonal variations in gonadal growth.

In comparison to published data from other redhorse populations, Clear Creek golden redhorses weighed less and were older at comparable lengths. Black redhorses from

Clear Creek achieved comparable lengths and weights at younger ages than was reported from other locales. Clear

Creek silver redhorse data compared similarly to published accounts. Thus within the unglaciated valley, the black redhorse was the most successful Moxostoma species in terms of growth in length and weight.

Prom a total of 2408 redhorses tagged during the study

(1231 golden, 1061 black, and 116 silver), 991 recaptures were recorded (494 golden, 460 black, and 37 silver). The home ranges (i.e. the average distances moved as measured from the original capture site) were .425 km for golden redhorses, .522 km for black redhorses, and .782 km for silver redhorses. No significant differences were found between average distances moved in the spring as compared to the fall, nor could any relationship between fish length and distance moved be established. Recaptured golden and 133 black redhorses did not exhibit any mass seasonal movement

patterns and the majority were believed to be permanent,

sedentary residents of Clear Creek throughout the year.

In terms of inter-pool movement, less than 20 percent

of the recaptured redhorses traveled more than two pools

from the original capture site. Black redhorses which had

strayed from the point of capture demonstrated a significant

tendency to move upstream, while golden and silver redhorse

movement was random. Golden and black redhorses transplanted

in pools away from the original capture site did not exhibit

a homing ability.

The diel movement of a redhorse aggregation consisted

of a cyclic upstream-downstream pattern within a pool. The upstream portion of timed cycles averaged 1452 sec and the

downstream segment averaged 66 sec. Placement of artificial

covers over regularly transversed routes resulted in altered movement patterns.

The primary constituents of the redhorse alimentary tracts examined were detritus, algal filaments, and aquatic insect larvae. The specific food items found during a particular sampling period were similar for the three red­ horse species and for all age classes of each species.

Seasonal variations in food items were observed, especially as related to the availability of aquatic insect larvae.

Whereas only nine percent of the intestines examined were empty, feeding was discerned to be a nearly continuous diel 134 activity. Specifically, redhorses fed on drift from riffle

areas during the upstream segment of each movement cycle.

In Clear Creek the catostomid spawning sequence was

determined to he white sucker-silver redhorse-quillback-hog

sucker-golden redhorse-black redhorse. These six species were believed to utilize the same riffles as spawning grounds, however spawning periods probably did not overlap.

In the spring, sexual dimorphism was evident in the

Moxostoma species. Mature golden redhorse males had tubercles covering the entire body. Tubercles on mature black and silver redhorse males were restricted to the anal and caudal fins. The tubercles aided in maintaining body contact and in stimulation of females during spawning and, for golden redhorses, in the defense of territories.

Dichromatism was displayed by males and females of the three redhorse species. Both color and tubercles were believed to be involved in sexual recognition.

Golden redhorses were filmed spawning on 15-19 May

1975 in riffles below large pools. Mature males aggres­ sively defended territories on the spawning riffle. The larger and older males (six and seven years old, 270-310 mm) occupied the centermost territories, while the smaller, mature males (five years old, 230-260 mm) were relegated to peripheral positions. All 25 tagged golden redhorses removed from spawning riffles had been originally captured in adjacent pools. 135

During the spawning period, a female golden redhorse

would drift from the pool down through the spawning riffle

and then would swim back upstream to a position alongside

a male. Instantly a second male would occupy the open

female flank and spawning would occur. The majority of

spawnings observed involved the two male-one female trio, but on occasion four or five males would attempt to 3pawn with a single female.

During the last day of the spawning period, only a few small males were on the riffles, successful spawnings were infrequent, and, unlike the previous days, egg preditors

(hog suckers, stoneroller minnows, striped shiners, and rosyface shiners) were very abundant. Thus, the golden redhorse reproductive strategy dictated that the older, larger, and more aggressive males had the greatest proba­ bility of spawning successfully and the eggs which they fertilized had initially the best chance of survival.

Comparisons of Clear Creek spawning data with the few published sources suggested the possibility of inter­ population variations. Additionally, Clear Creek golden redhorses were less fecund and black redhorses more fecund than fish reported from Iowa and Missouri respectively.

The interaction of the three redhorse species in Clear

Creek was limited to the unglaciated study area, as only golden redhorses were found on the glaciated plateau. The most obvious manifestation of redhorse interaction was the 136

gregarious nature of the three species. However, schooling

was not observed. It was believed that the gregariousness

and the small home ranges contributed to the stability of

the three redhorse populations,

A Bailey point census indicated that within the study

area the golden redhorse population was slightly larger than

the black redhorse population (2391 to 1884 in the fall

1975), The silver redhorse population was considerably

smaller. Survival rates for golden redhorses were greater

than for black redhorses. Thus, corresponding with the

age and growth data evidenced earlier, the golden redhorse

population responded differently than the black redhorse

population to density dependent regulation. The larger golden redhorse population consisted of older, slower growing individuals, whereas younger, faster growing individuals comprised the smaller black redhorse population. Appendix A. 1939-1950 (Trautman 1957) and 1974-1975 (this study) surveys of fishes in Clear Creek, Hocking and Fairfield Counties, Ohio.

(Trautman 1957) (This Study) FAMILY GENUS-SPECIES COMMON NAME 1219-1950 1974-1975

PETROMYZONTIDAE Lampetra aepyptera least brook lamprey X SALMONIDAE Salmo gairdneri^ rainbow trout X ESOCIDAE Esox americanus vermiculatus grass pickerel X CYPHINIDAE Cyprinus carpio^ carp X X Notemifronus crvsoleucas golden shiner X Phoxinus ervthro/raster southern redbelly dace X X Rhinichthvs atratulus blacknose dace X X Camporstoma anomalum stoneroller X X Semotilus atroraaculatus creek chub X X Ericvmba buccata silverjaw minnow X X Pimephales notatus bluntnose minnow X X Notropis photogenls silver shiner X Notropis rubellus roayface shiner Notropis urabratilis redfin shiner X Notropls chrvsocephalus striped shiner X Notropis spilopterus spotfin shiner X Notropls stramineus said shiner X X Notropis volucellus mimic shiner X 137

introduced (Continued) Appendix A. 1939-1950 (Trautman 1957) and 1974-1975 (this study) surveys of fishes in Clear Creek, Hocking and Fairfield Counties, Ohio. J

(Trautman 1957) (This study) FAMILY GENUS-5PECIES______COMMON NAME______1939-1950 1974-1975 CATOSTOMIDAE Catostomus commersoni white sucker X X Carriodes cyprinus quillback X Moxostoma anisurum silver redhorse X Moxostoma duauesnei black redhorse X Moxostoma ervthrurum golden redhorse X X Hvpentelium nigricans northern hog sucker X X ICTA1URIDAE Ictalurus melas black bullhead X Ictalurus natalis yellow bullhead X Noturus gyrinus tadpole madtom X Noturus miurus brindled madtom X X PERCOPSIDAE Percopsis omiscomaycus trout-perch X CYPRINODONTIDAE Fundulus notatus blackstripe topminnow X CENTRARCHIDAE Micronterus dolomieui smallmouth bass X X Micronterus sa]moides largemouth bass X Ambloplites rupestris rockbass X X Leoomis cyanellus green sunfish X X Lepomis macrochirus bluegill X Lepomi3 megalotis longear sunfish X

Pomoxis annularis white crappie X 138 (Continued) Appendix A. 1939-1950 (Trautman 1957) and 1974-1975 (this study) surveys of fishes in Clear Creek, Hocking and Fairfield Counties, O hio.

(Trautman 1957) (This Study) FAMILY OENUS-SPECIES______COMMON NAME______l?39r1.952-- 3-974-1225. PERCIDAE Percina maculata black3ide darter X Etheostoma nirrum Johnny darter X X Etheostoma hlennioides greenside darter X X Etheostoraa caeruleum rainbow darter X X Etheostoma flabellare fantail darter X X Etheostoma zonale banded darter X X COTTIDAE Cottus bairdi mottled sculpin X X

H \D Appendix B. Chemical analysis (Hach Kit) of Clear Creek (Hocking and Fairfield Counties, Ohio) water samples taken on 17 July and 13 October 1975 at sites 11 and 39 and at the Landis Road bridge.

17 July 1975 13 October 1975 Test Lte 11 Site 39 Landis Site 11 Site 39 Landis Rd. Total Alkalinity (ppmCaCO^) 110 110 180 100 100 190 Alkalinity - phenophthalcin 0 0 0 0 0 0 Carbon dioxide (ppm) 8 10 15 10 8 20 Hardness - calcium 10 10 10 10 14 10 Hardness - magnesium 8 8 10 10 7 10 Hardness - total 18 18 20 20 21 20 Iron (ppm Fe) .22b .15 .12 .19 .15 .10 Nitrate (ppm) 35.2 48.4 44 Nitrite (ppm) 0 0 0 Dissolved oxygen (ppm) 10 11 8 8 13 7 PH 8.4° 8.0 8.2 8.2 7.8 7.9 Ortho-phosphate (ppm) 1.2 1.4 1.6 1.6 1.8 2.1 Poly-phosphate (ppm) .8 .8 1.4 1.2 1.2 1.4 Total phosphate (ppm) 2.0 2.2 3.0 2.8 3.0 3.5 Silica (ppm) 6 6 6 4 4 4 •Sulfate (ppm) 125 125 50 Turbidity (JTU) 0 0 0 0 0 0

aAn iron hydroxide precipitate was present in numerous backwater areas bWhite and Dilenschneider recorded a value of *242 in May, 1973. cWhite and Dilenschneider recorded 8.2 in May, 1973. Appendix C. Records of golden (G), b la c k (3), and silver (s) redhorsea tagged and recaptured from Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975 indicating the number of each species caught at each site.

POOL SITES 1 2 3 4 3 6 7 8 9 10 11 15 14 15 16 17 18 19 20 21 22 23 24 25

G 2 1 5 5 4 3 1 10 41 10 4 4 9 26 17 2 17 6 B 1 1 5 3 15 2 1 7 6 2 28 5 2 2 7 10 4 S 5 2 1

Tagged G 3 2 Winter 1975 B 6 10 3

Tagged G 2 3 1 9 4 13 8 7 14 3 3 13 43 8 8 2 5 4 42 6 Spring 1975 B 1 16 5 20 5 13 5 7 4 33 4 3 11 2 2 1 38 13 S 1 1 2 1 4 17 1 4 1 15

Recaptures Fall 1974

Recaptures Winter 1975

Recaptures G 4 2 1 6 7 2 4 3 Spring 1975 B 14 2 6 10 1 2 S 2 1 2 141 (Continued) Appendix C. Records of golden (G), black (B), and silver (s) redhorses tagged and recaptured from Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975 indicating the number of each species caught at each site.

POOL SITES (continued) 26 27 28 39 50 31 52 53 34. 35_36_37 38 39 40 41 42__43 44 45 46 47 4.8 49 50 Tagged Fall 1974 G 10 9 50 29 24 26 15 19 21 29 20 71 9 13 13 14 B 11 5 18 35 2 34 20 21 33 24 12 58 6 19 25 15 S 2 2 2 3 8 7 3 3 6 1

Tagged G 37 4 2 2 Winter 1975 B 15 4 10 10 S

Tagged G 9 12 12 40 37 8 6 3 18 2 28 10 2 9 13 2 15 6 12 14 31 6 25 Spring 1975 B 2 13 14 20 37 31 7 4 8 3 23 12 7 22 1 7 11 14 5 37 13 29 S 1 1 2 4 3 1 1 2 8 1

Recaptures G 4 3 1 3 1 3 2 Fall 1974 B 1 1 5 1 7 2 1 4 2 S 1 2 1

Recaptures G 14 1 3 1 Winter 1975 B 11 2 4 2 S 1

Recaptures G 2 10 6 8 1 1 6 12 6 3 3 6 2 21 2 Spring 1975 B 1 1 1 9 12 6 1 3 1 12 11 4 1 18 1 2 1 1 9 3 5 S 1 1 A ppendix C. Records of golden (G), b la c k (B)# and silver (s) redhorses tagged and recaptured from Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975 indicating the number of each species caught at eachsite.

FOOL SITES (continued) 51 52 55 54 TOTALS

Tagged G 545\ Fall 1974 B 450 > 1040 S 45x

Tagged 0 5 0 ^ Winter 1975 B 55 > 105 S 0 - ^

Tagged 12 9 31 66 G 6 3 6 ^ Spring 1975 14 4 35 B 5 5 6 ^ > 1236 S 7 1 ^

Recaptures G 20. Fall 1974 B 26\ 49 S 3 - ^

Recaptures 0 Winter 1975 B 24 > 49 S 1 ^

Recaptures 1 3 14 G 141%. Spring 1975 1 B 133 > 281 S (Continued) Appendix C. Records of golden (G), black (B)f and silver (s) redhorses tagged and recaptured from Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975 indicating the number of each species caught at each site.

POOL SITES 1 2 ? 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 Recaptures g 6 i 1 3 2 4 4 6 1 16 13 1 5 4 1 17 2 1 Fall 1975 B 3 4 6 1 5 2 2 6 5 8 1 2 4 8 2 2 s 1 3 3 3 1

Unmarked 6 1 3 15 12 1 30 9 5 3 18 16 2 7 9 6 2 5 2 3 1 2 Fall 1975 B 3 11 26 23 7 17 13 3 5 15 13 2 11 12 10 8 6 5 8 1 S 1 4 2 5 1 10 1 2 1 6 1 1 2 5 (Continued) Appendix C, Records of golden (G)f black (B), and silver (s) redhorses tagged and recaptured from Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975 indicating the number of each species caught at each site.

POOL SITES (continued) 26 27 28 29 50 51 52 53 34 55 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50

Recaptures G 7 3 4 26 19 5 2 3 1 11 11 2 25 6 5 1 9 4 6 3 1 4 24 3 8 Fall 1975 B 3 3 4 11 32 18 3 3 1 1 6 1 24 10 9 17 2 3 5 2 5 12 9 14 S 1 1 1 1 2 9

Unmarked G 8 2 3 29 38 40 2 6 6 13 14 64 3 6 8 29 17 3 7 3 5 18 8 25 Fall 1975 B 4 6 8 18 33 32 11 13 7 7 8 67 9 16 9 40 9 7 15 4 8 12 17 8 S 3 1 9 1 1 6 (Continued) Appendix C. Records of golden (G), b la c k (B), and silver (S) redhorses tagged and recaptured from Clear Creek, Hocking and Fairfield Counties, Ohio in 1974 and 1975 indicating the number of each species caught at each site.

51 52 5? 54 TOTALS Recaptures 2 4 8 14 G 309 Fall 1975 4 3 11 B 277 S 26

Unmarked 12 26 23 70 G 640 Fall 1975 13 17 28 B 625 1328 S 63 9*1 147

Appendix D. Times recorded for 134 upstream and 112 down­ stream segments of the redhorse diurnal activity cycle at site 29 (36m) during August and October 1975.

Upstream Tlmes-sec

8-1 2250 8-7 1945 8-26 1858 10-7 1445 2111 1581 1518 1726 1279 1124 1264 1617 1446 1818 1422 1131 1524 2115 1096 1557 1578 1743 1805 1762 1428 1222 2340 1322 1366 1971 10-8 1114 2015 1229 2578 1100 1539 1631 8-2 1699 8-27 2085 1711 1056 8-8 1796 1492 1364 1419 1332 1883 1590 1365 1812 1938 1483 1695 1252 1350 1599 1361 1417 10-14 1354 1760 1288 1025 1245 1518 1221 1128 1319 1851 1247 1136 8-3 1722 2023 1856 1583 1144 1875 1018 8-9 1792 8-28 1247 1447 1826 1058 10-15 1178 1145 1477 814 1001 1736 1103 992 799 1534 1569 1240 640 2002 1453 1175 984 1566 1405 1389 8-4 936 1215 930 1233 662 1303 1168 855 1246 1012 8-29 1265 1146 8-10 1583 737 1409 1286 835 1757 1799 1187 1982 1757 1424 1576 1075 1015 1359 1656 1806 1200 1636 1761 ] 244 1871 148 (Continued) Appendix D. Times recorded for 134 upstream and 112 down­ stream segments of the redhorse diurnal activity cycle at site 29 (36m) during August and October 1975.

Downstream Times-seo

8-1 92 8-9 52 8-29 78 75 34 62 61 38 82 40 gg 34 51 88 25 43 115 79 69 79 59 48 8-2 59 8-10 64 35 58 10-7 54 84 93 31 30 71 83 143 96 38 42 69 40 78 87 98 71 105 33 55 8-3 24 10-8 58 17 8-26 58 43 91 7 i 133 29 61 70 51 70 55 88 4 4 67 29 52 8-4 62 |1 70 241 10-14 35 69M 5 3 875 6 75 8-27 125 63 59 38 57 80 10-15 37 45 86 45 54 47 72 92 63 8-8 77 66 49 38 69 84 51 84 8-28 29 28 44 50 65 82 158 46 75 56 33 50 109 LIST OF REFERENCES

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