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HomeMy WebLinkAboutSUS231' • • r QB 541.5 S7 W2E3 !.FF!CTS OF .RESERVOIR RELEASES ON TAILWATER ECOLOGY : A LITERATURE REVIEW C. H. Walburg, et al U.S. Fish 6 Wildlife Service BowLing Green, Kentucky September 1981 AD/A-105 058 • t .· . •· . , ..... , ... 4 ' ... ..... Vnclassifiell REPORT DOCUMENTATION PAGE Q~ . 5l.J t,S Sf] tJJJcb sus ~31 ... Jlt&AD DCITitUCTIOtfl · ... BUOU COIIPLETDJG FORM I'· III& .. OIIIT NUM.RIII E-81-12 12. GOVT ACCIIUION HOi ~:fC:1 .. 111NT'S C:ATA~OG NUIII.CIII Teclmic·al Report . AD/Al05 058 6. TITI..C (-,._,,,) . 'L TYPC OP' lllllf00111T a "111111100 COYCIIICD EFFECTS OF RESERVOIR RELEASES ON TAILWATER Final report .JCOLOCY: A LITERAl'URE REVIEW 1. AUTMOIII(OIJ I. ..CIU'OMIIINO OIIIG. IIIC .. OIIIT Nuti.Cit Charles H. Walburg, Jerry F. Novotny, Kenneth E. 'Jacobs, William D. Swink, Terry M. Campbell, L .:;ONTIIIMOT 01111 GIIIIAMT NU111••••• , . John Nestler, Gary E. Saul 1 ... CIIIP'01111111NO OIIIGANilATION NAIIII AfiD ADDIIIIIII u. S.; F.isll & Wildlife Service 110. "IIIOGIIIAM CLb&MT, PIIIOJ&C:T, TAll< East Centra~ Reservoir Investigations AIIICA a WO"K UNIT NWiellllll Bowling Green, Ky. 42101 EWQOS Task IIB u. s. Army Engineer Waterways Experiment Station Environmental Laboratory ... tz. ltllf'OitT OATil P. o. Box 631 September 1Q81 Vicksburg, Miss . 39180 IS. MU1118111t OP' "AGCS It, CONTftOI.I.INCI OP'P'ICE NAMII AND AOOitCSS 216 Of'fice, Chief of Engineers, u. s . Army IS. ICCUIIItYY CI.AU. (el IN• ,_., Washington, D. c. 20314 Unclassified ,._ -ITVtaNO AGCNCY NAME-a ltDOitU!(U ··-· -.C....IU•I Ollr-) Hie. ~.J~ASII P'ICATIOttl OOWIIICU.ADIItO KH DUI.C , ... DIITIIII8UTION ITATCMENT (el He Jt.-r) ,. ·~,,. ApproV'ed for public release; distribution unlimited. ' "· DIITIIII.UTIOR ITATbCNT (el llle -----M .,_, 20, It.,__.,_~) \ .. IL ~DI ... T·A'"' NOTES .•:~--. 5285 r.-Availabl e from National Technical Info~atio n SerVice, Port Ro)'ll] Road, Springfield, Va. .. 22151 . -· .. II. CE,. WOAOI (C..tl--,.,_ .. el._ II necee.-y -1-1)> fir .,..,. -) . ·'· I -.. ------·--.~ ..... t ..., ...... Biota Reservoirs •PIODutfD IT Discharge (Water) Tail water NATIONAL TECHNICAL . Impoundments Water qu ality IN FORM A liON SE.RVICE I U DlPAII.If Of C1)11Uict ..,. . SPIHilLD. YA. 22lfl .. . lA. ...... M:T~---·--·-,.·.,--) - ·This document presents a review of tile often contradictory literature de.cribing the effects of release waters on tBe tailwater environment and biota. The physical and chemi.cal conditions found in tailwaters downstream from varmwater and coldwater disd'J&rge impoundments are compared and con- trasted t o those found in natural streams. Reservoir disc barge. modify the pbysical, chemical, and biological (Continued) Unc l assified ·. . ... .,·. ' . , .... . ' -------·- Unclas sified 20. ABSTRACT (Continued) . cbaracteristiss or the stream ecosystem. Physical and ch~cal characteristics i.n tailvat.ers are prilllarily determ·ioed by the. depth; Yolume., and acbedul.e of vater releases. The m.agnitude of change is related to the tn>e of reservoir and to the design and operation of outlet s tructure~. The structure. of the biotic ca.nmity ret'lec.ta the pbysieal and chemical conditions existing io a pa:-ticular tailvater . 'l'be caamunity i.s composed or organics, including nonnative species, that are adapted to this enTiroDIDent . Tbe effects of the tailwater enviroDIDeot on the lire-history, physiology, and abundance of selected speci es are described. This information will aid in the development of reservoir discharge guideLines tha't will enhance the quality of the tailvater environment to in- crease project benefits. •':, .. -. II ttncJ ass1 f1 ed I .4 • . . PREFACE This report was prepared by the U. S. Department of the Interior, U. S. Fish and Wildlife Service, National Reservoir Research Program (NRRP), East Central Reservoir Investigations (ECRI), Bowling Gre.en, Ky ., with the assistance of the Environmental Laboratory (EL), U. S. Army Engineer Waterways Experimen.t Station ('WES), under Interagency Agreement WES 79-04 d.a:ted 1 April 1980. This study forms part of the Environmental and Wate·r Quality Operational Studies (EWQCS), Task IIB, Reservoir Releases . The EWQOS Program is sponsored by the Office, Chief of Enginee·rs, and is assigned to WES under the management of EL. This report was written by Messrs. Charles H. Walburg, Jerry F . llovotny, Kenneth E . Jacobs, William D. Svinlt, and Terry M. Campbell of ECRI and Drs. John M. Nestler and Gary E. Saul, EL, WES. Mr. Charles H. Walburg is the Chief of ECRI, and Mr. Robert M. Jenkins is the director of the NRRP. Preparation of this report vas under the direct supervision of . Drs. llestler and Saul snd the general supervision of Mr. Joseph L. Norton, Acting Chief, Water Quality Modeling Group, EL; Mr •. Donald L. Robey, Chief, Ecosystem Resear ch and Simulation Division, EL; Dr. J'erane L. Mahloch,. Program ·Manager, EWQOS, EL; and Dr. John Harrison, Chief, EL. · The Commander and Director of WES during this study was COL Nelson P. Conover, CE . The Technical Director was Mr. Fred R. Brovn. This report should be cited as follows: Walburg, C. H. , Novotny, J. F ., Jacobs, K. E., Swink, W. D., Campbell, T. M., Nestler, J., a .nd Saul, G. E . 1981. "Effects of Reserv:oir Rel.eases on Tailvater Ecology: A .Literature Review," Technical Report E-81-12, prepared by U. S. Department of~e Interior, Fish and Wildlife Service, National Rese i Researc h Program, East Central Reservoir Investigat ons, and Environmental Laboratory, U. S. Army Engineer Waterways Experiment Station, for the· U. S. Army Engineer Water- ways Experiment Station, CE, Vicksburg, Miss. 1 PREFACE PART I: INTRODUCTION • Problem. . • • • • Study Approach • • CONTENTS PART II: BASIC RESERVOIR LIMNOLOGY Hydraulic Residence Time and Settling within Reservoirs •• The,rmal Stratification Dissolved Oxygen • . . Nutrient Concentration • Reduced Compounds •••• PART III: RESERVOIR OUTLET STRUCTURES AND THEIR IHPACT ON THE TAILWATER ENVIRONMENT •• Design Considerations •• Operation Considera~ions • Summary. . • . • • • . . • PART IV: Ph"YSICAL AND CHE.\fiCAL DESCRIPI'ION OF TAILWATERS • Physical Characteristics Temperature . . .. . . . . -. . . .. ~ l 6 6 7 8 8 9 10 10 11 12 12 13 14 15 15 15 . Flow ••• . . 18 . . t ~~~ • ·Substrate • Turbidity •• Chemical Characteristics Dissolved gases •••••• •' . . . ' . Hydrogen ion concentration and alkalinity •. Particulate organic matter .•. Nutrients • • . . • . • . • • • Reduced compounds iD tailvaters PART V: META.BOLISt-1 AND TROP'rliC STRUCTUR.E • · · St'l'eams. • • Te.ilvaters . PART VI : AQUATIC INVERTEBRATES IN TAILWATERS • Inver~ebrate Ecology Streams • • • • Tai1vaters. • • Benthic Invertebrate Drift • Streams • • • • Tailvaters .•• 2 19 19 20 20 23 24 25 27 30 30 34 37 37 37 46 53 53 54 ~ •· .., I· .. ; ·· .. • ~-· . ·. ...... ~ ; .. Zooplankton. • . • • • • • • • • • • • • • • • . • Tai.lwater Effect.s on Specific Invertebrate Taxa •• Diptera . . • • . • • . Tricboptera. • Ephemeroptera P1ecoptera. • Mi s.cellaneous PART VII: FISHES IN TAILWATERS • Po1yodontida.e· (Paddlefishes) • Pa.ddlefish. • • • • • • • Paddlef'ish in tailwaters. Clupeidae (Herrings) •• Shad •••...•• Shad in tai1waters. Sa.lmonidae (Trouts) .•• · Rain bo·w trout . . ... Rainbo11 trout in tailwaters Esocidae (Pike~) • Pikes • • . . Pikes in tai1waters • Cyprinidae (Minnows) •. Carp. • .•.... Carp in tailwaters. Chubs . • • • • • • Chubs in ta.ilvaters True minnows •• True minnows in tailwaters. Shiners • • • • • • • Shiners in tailwaters • • • Stonerollers •.•••••• Stonerol1ers in tailwaters . Da c es in tailwaters . • • . Squawfishes and chiselmouths in Cyprinids in Russian tailwaters . Catostomidae (Suckers) • • . Buffaloes • • • • • • . Buffaloes in tailwaters Suckers • • . • • ·• • Suckers in tai1waters . • Redhorse ••••••••• Redhorse in tailwaters . 3 tailwaters. •. fY!. 55 57 58 59 60 60 61 63 63 64 65 67 67 68 69 70 13 · 85 85 86 88 ' 89 90 92 93 95 91 98 100 102 '103 103 104 104 106 106 108 109 113 116 ua .•· ·. ~ ..... '.~ ... - . ... --------------- Ictaluridae (Catfishes) ••. Bullheads • • • • • • • Bullheads i .n tailwater · Catfish .••••.•• Catfish in tailwaters Madtoms •••••.. Madtoms in tailwaters • • • • • • • • • Percichthyidae (Temperate Basses) •••••• White bass. . . • . • • • . • Temperate basses in tailwaters. • ••• Cent:rarchidae (Sunfishes). • . • • • • • • Black basses ..•..•.•• Black basses in tailwaters. • True sunfishes. • . • • . • True sunfishes in tailwaters. Crappie.s. • • • • • • . Crappies in tailwaters. Percidae (Perches) ••.•• Walleyes and saugers .. Walleyes and saugers in tailwaters. Darters • • . • • . • Darters in tailwaters Sciaenidae (Drums) ••.•• Freshwater drum Freshwater drum in tailwaters • Cottidae (Sculpins) .•••. Sculpins. • . . • • • . • Sculpins in tailwaters .• PART VIII: CONCWSIONS . • . • . Effects of Hypolimnetic Release on Downstream Biota. E!fec~s of Epilimnetic Release on Downstream Biota . Effects of Water Release Patterns on Downstream Biota. Past Tailwater Research and Suggestions for Future Study. REFEP.EN CES . APPENDIX A: ALPHABETICAL LIST OF THE 113 TAILWATERS MENTIONED IN THE TEXT WITH LOCATION BY APPENDIX B: RIVER AND STATE, PROVINCE, OR COUNTRY. LOCATION OF 105 RESERVOIR TAILWATERS IN Tli:E UNITED STATES MENTIONED IN THE TEXT • • • . • • • • • • • • • • 4 ·- ~ 119 119 121 122 124 125 126 126 127 128 130 130 133 135 138 141 143 145 145 147 150 151 152 152 153 154 154 155 157 157 160 162 165 168 A1 Bl .. ' " ~ . ., ·.~ ... ·. APPENDIX C: COMMON AND SCIENTI.FI·C NAMES OF FISHES MENTIONED IN THE TEXT, .ARRANGED BY F.AMILY • • • • • • • • • • • . • • • • .. Part I: Fishes From North American Tailvaters • Part II: Fishes from European Tailvaters. . . APPENDIX D: LIFE HISTORY INFORMATION FOR THE MJST COMMON FISH GROUPS MENTIONED IN THE TEXT APPENDIX E: GLOSSARY --. -. ··. 5 ---------------- Cl C2 C6 Dl El ...-;;.:"':~: :.t_ . ... ; •· . . ... r ------- EF?ECTS OF RESE.RVOIR RELEASES ON TAIL\lATER ECOLOGY : A LITERATURE REVImJ PART I : INTRODUCTION Problem 1. The Corps of Engineers (CE) normally operates reservoir projects to achieve downst.ream environmental quality objectives consistent with proje.ct purposes. Presently, there a·re no quantitative approaches or re- ~iable guidelines for determining water releases necessary to ensure the maintenance of a des.ired downstream aquatic environme.nt. Many en.viron- mental quality requirements fo·r downstream habitat and biota are not well understood or substantiated . 2. Reservoirs affect downstream aquatic habitats in a number of ways, depending on project design and operation and specific environ- mental requi:-em~nts of downstream biota. Large variations in flov associated wi~h power-peaking operations may adversely affe~~ ~~~­ stream fisheries during spawning periods, disrupt benthic communitie.s that serve as food for fish, and 1 imi t stream recreation. Change·s in temperature, dissolved gases, and other va.ter quality characteristics associated with reservoir releases greatly influence the species composition and abundance of the tailwater community . 3. The operation of reservoirs to achieve destred downstream objectives is often complicated by conflicting requirements to improve in-lake wa;:;er quality or demands of other proje.ct purpose.s. Periodi- cally, minimum releases required to maintain downstream aquatic habitat and associated stream recreation are greater than the releases required to meet o·ther authorized project purpose.s . During these periods, pro- blems associated with reservoir releases often become critical. Since the benefit:; of maintaining or enhancing downstream aquatic habitat and biota e.:-e difficult to quantify, justifying the allocatio·n of reservoir storage for do\mstream releases is complicated. Nevertheless., minimum releases are required to maintain adequate downstream habitat. 6 & • • r Study Approach 4. The first task of the U. S. Fish and WiJdlife Ser<ice in pre- paration of this report was to conduct an exhaustive literature search o u the e.ffects of reservoir water releases on T.ailwater biota. An ..mnota::ted b i bliography vas prepared from the literature th&;t most dir- ectly concerned tailwater problems (Walburg et al . 1980). 5. This report r e views available literature on the effects of reservoir releases on tailwaters that support populations of varmwater fish or trout. The extensive literature available on anadromous fish is not included. This "state-of-the-art" report documents the relation between changes in the quantity and quality of reservoir water releases and the quality of the downstream aquatic envirorment. 'l'he review also includes selected stream and river studies that have application to tailwater problems. Tailwaters cited in this report are listed alpha- .. betically in Appendix A together with name of river and l ocation (e.g., state of the United States or country). The geographic l ocations of the 105 tailwaters located in the United States are shown in Appendix B (Figure Bl). 6. To better understand the physical and chemical conditions found in tailwaters, this report begins with a brief description o f reservoir limnology, followed by a section on the design and operation of reservoir outlet structures a nd how this can impac t the tailwater environment. This is followed by a re'View of the physical, chemical, and trophic conditions found in tail waters. A general review at in- vertebrate ecology in both streams and tailwaters is then presented. Life history requirements of fishes found in tailwaters are r eviewed, together with a description of their response to the tailwater envi ron- ment. Tailwater environments created by various management schemes for reservoirs and tailwaters are discussed generally . Major physical and chemical alterations are indicated, together with descript i ons of how they affect organisms of the higher trophic levels. Finally, the nature and scope of studies necessary to comp l ete the develoiDent of conceptual models that can be used to predict the effect of changes in reservoir management on t he ta.ilwater environment are indicated. 7 -·· PART II: BASIC RESERVOIR LDmOLOGY 7. Knowledge of reservoir limnology is fundamental to under- standing water qual.ity characteristics i .n tailvaters. Wate.r quality in. reservoi r changes with season. The exten.t of cbaDge reflec.ted in the tailwater depends on the depth of wat.er withdrawal, project de- sign, morph~etry of the tailwater channel, and local atmospheric conditions. The following brief overview of reservoir limnology is intended to provide suf'ficient background information for a biologist or engineer to unders tand the relationship between reservoir biogeo- ~hemical processes and tailwater ecology; it is not inten~ed to be a detailed d.iscussion of limnology. For a canprehensive description, the reader should refer to the works of Hut.chin.so·n (1967), Wetzel (1975'), and Cole (1975). Hydraulic Residence Time and Settling within Reservoirs :. · 8.: In general, reservoirs with short hydraulic residence times · ~ •; have a reduc.ed impact on tailwaters because the water is discharged before the effects of impoundment becc:me established. This type of reservoir is often termed a flow-through or run-of-the-river pro·ject . The discharge is usua.lly similar to the inflow· in oxyg.en concentra- tion, temperature, turbidity, and nutrient concentration. Reservoirs vi th long hydraulic residence times undergo· processes sc:mewhat similar to those observed in lakes, although there are significant differences (Neel 1963; Baxter 1977). 9. Reservoirs with long hydra.ulic residence times act as set- tling basins in which suspended particles settle f'rc:m the water column. They are effective in removing suspended material frc:m inflowing streams or sediments washed into the reservoir during summer rains. Turbid water is usually discharged into the tailwaters only after long winter rains that are accc:mpanied by high runoff rates (Churchill 1967). 8 ,•. ..•... . ... , .. , -'a .. . .. ... :: .. .. . -. ... Thermal Stratification 10. The .PTOCess or thermal stratification is a major factor in altering the· wate.r quality of reservoirs. An understanding of thermal stratification a.nd its influence on wat.er as it flows through a rese.r- voir is essential to discu.ssion of water qualit·y in tailwaters. 11. Reservoirs strat,if'y thermally when solar radiation and in- flows from warmer tributaries during th.e spring heat surfac.e vaters more rapidly than the heat can be distributed throughout the water column. This produces temperature and density differences be:tween the surface water and the under~ying water, increasing t .he. resistance to _mixing. Shearing between the surface and deepe.r waters inhibits add.i- tlo.nal r;uxing and results in the formatio~ of an upper layer of warm water ( epilimnion) and a deep layer of cold wate-r (hypolimnion) , vi th a transitional layer be.tween the two (metalimnion). Following strati- fication, mixing effects caused by wind. and air tempe.rature changes are large·ly limited to the epilimnion. 12. Reservoir's destratify thermally when the loss of heat to t 'he cooler atmosphere in l .ate summer and early fall exceeds the ,heat input --~ , from solar radiation. Complete thermal mixing begins when the surface water cools and 'becomes. as dense. as the deeper water. Eventually tbe entire water column loses its resistance to mixing and_ the reservoir becomes thermally uniform (Wetzel 1975). Tributary inflows may accelerate destratifica.tion by prQviding add-itional cool water. 13. 'l'he vater column in warm temperate reservoirs is essentially of constant d-ensity during the· vinter, and little -thermal resistance to mixing occurs. Convection curre.nts and relatively little vi.nd action can thoroughly mix the entire water column (Churchill 1967). As a result, the. c·bemical and physical characteristics of the water r•~in un1form throughout the reservoir until spring. These reservoirs are termed monomictic, sinc·e they circulate freely only during the vinter months. 14. Reservoirs in cooler tempe.rate regions may beeome stratified during the vinter. The coldest water (0-3°C) remains at the surface 9 ---·--~-------- and the varmer, most dense vater (4•c) sinks to the bottom (Wetzel 1975). These reservoirs are termed dimictic because they circulate freely tvice during the year folloving spring and fall destratification. 15. Not all reservoirs thermally stratify during the summer. Shallov reservoirs vith relatively rapid rates of flov-through and exposure to extensive vinds usually remain vertically mixed. Temporary temperature "differences" vhich form after extended periods of calm veather and reduced discharge may be destroyed by vind action. These reservoirs may, hovever, undergo periods of vinter stratification after ice cover formation . Dissolved OXYgen 16. Concentrations of dissolved oxygen in reservoirs are closely associated ~lith the stratification process. In stratified reservoirs, tbe epilimnion is vell aerated due to vind action, mixing resul.ting from diurnal temperature fluctuation, and oxygen produced during photo- s.ynthesis. Dissolved oxygen in the hypolimnion, bovever,·is limited to that available at the time of stratification, and may be re·duced or eliminated by the oxidation of organic matter that settles into the hypolimnion from the epilimnion. These conditions persist until the reservoir mixes vertically. Lov dissolved oxygen concentrations are seldom a problem during the vinter, vhen lov 'vater temperatures sup- press metabolic activity, and oxidation rates of organic compounds are reduced. Hovever, lov dissolved oxygen can occur in ice-covered lakes during the vinter vith resultant fish kills. 17. In unstratified reservoirs, comple.te circul.ation and frequent aeration by vind action ensures that adequate oxygen is available throughout the vater column. Lov oxygen concentrations may occur in localized protected embayments and other areas not subject to frequent circul.ation. Also, nighttime algal respiration may lover oxygen levels. Nutrient Concentration 18. The import of nutrients from upstream or vatershed runoff is 10 ' I ' l ,. I .. I·· A. ·-. -.. :: : the main s.ource or reservoir enrichment. Me.xil:l.um inputs generally occur after heavy rains. Excessive amounts of nutrients (e.g. , f .roo municipal and agricultural sources) can cause· deterioration in overall water qual- ity. Nutrients can also be released from the sediments under anoxic conditions. These nutrients can be carried to the surface during lake ·curnover. In addition. wind-driven currents can incorporate sediments and associated nutrients into the water column in shallov, unstratifie.d reservoirs. 19. Phytoplankton growth is stimulated by an influx of nutrients. In stratified reservoirs, phytoplankters continuously settle out of the epili.mnion into the hypolimnion. Tbe loss of nutrients from the epilimnion resulting from settling may limit further phytoplankton pro- ·duction, whereas, the hypolimnion becomes increasingly enriched as the accumulation anc. decomposition of organic matter. proceeds. When the reservoir destratifies, the nutrients that were restricted to the hypolimnion are redistributed in the water column. The release of nutrients to surface waters, where light is sufficient to stimulate . •,: photosynthesis, results in increased phytoplankton production. In unstratified reservoirs, there is continuous circulation of the. Wa.ter column. Nutrients and organic matter are readily recyclable and remain ava:.lable in the productive areas for continuous assimilation. Reduced, Com-oounds 20. Anoxic conditions found in the hypolimnion of l&kes and reser- voirs result in the formation o·f reduced species of iron, manganese, .. sulfur, and nitrogen. These substances may become a nuisance to rec- reational and industrial water users, and may be <;}etrimental to aquatic li.f'e when they are released. These reduced compounds a.re c~nyerted to less noxious, more assimil.:s.ble compounds in the oxidizing en-rtronment of the epilimnion, and their effect. does not generally persist for more than 24 hours. Objectionable iron, manganese, nitrogen, and sulfur compounds are seldom a problem in well-mixed, unstratifie.d reservoirs. 11 PART III: RESERVOIR OUTLET STRUCTURES AND THEIR IMPACT ON THE TAILWATER ENVIRONMENT 21. Water quality conditions in reservoir tailvaters are deter-.• ... ' . ~ . mined by processes occurring in th~ reservoir (dis~ssed in the pre- ceding section) and the design and operation of the project outlet works. The following ·brief discussion is intended to provide general information concerning the design and operation of r e servoir.outlet structures. It should be emphasized that each project is unique in ·.design and-operation; therefore, many project features will not be specifically discussed in this section. Design Conside.rations 22. Most Corps of Engineers (CE) impoundments fulfill multiple purposes including navigation, flood control, hydropower, recreation, -·. Emphasis has been placed on effectively designing water supply, etc. BDd operating projects to meet all inte.nded purposes. For approxi-• •.•. o 0 ~' r •' .I • J 0 .. 1:~ mately the past 15 years, CE reservoir projects have been designed considering tr ~ water quality of project releases. 23 . Numerous design options are available to assure t h at project releases are compatible with tailwater habitat objectives. Most notable is_ the ~ncorporation of a selective withdrawal structure which .can j,,' release water from various strata within the reservoir to meet down- stream objectives. For selective withdrawal to be a viable alternative, density stratification must occur within the reservoir. This strati - fication may be due to vertical temperature differences within the impoundment, and/or the occurrence of strati fication due to the con- centration of dissolved constituents. Those reservoirs that are, vertically well mixed have little need for selective withdrawal. Typi- ·cally, such reservoirs are shallow, may have a short hydraulic residence time, and are often dominated by surface wind mixing . 24. Stratified reservoi rs provide an excellent opportunity for effective operation of selective withdrawal structures. R.eleases can 12 • ··'· .·. ~· ~.;I , .... ~ ~ ' . i ...... . ~ .... • ..... be made to meet downstream r equirements of tailvate~ ·fisher ies and also reduce the impact of flood control oper ations on the tailvater . 25. Most older flood control proje~ts were designed primarily to release vaters trca the bottom of t 'he r eservoir. Often, these water s are lev in 'di.ssolved oxygen, but because of aerat.ion that occurs in the outlet wor k.s it il .not. unc OJIIDon for dissolved oxygen in releas e waters to approach 95 percent saturation. Releases from hydropower projects receive little aeration due to the .r equirements to keep turbulence at a minimum. 26. Release ot minimlD flows to meet downstream ha.bitat objec- tives. is an important cons.ideration of project desi gn and operation. Projects vith large bottom sluice gates are generally unable to dis- charge low flows since the gates can not be operated with the neces- sary prec-ision or they vibrate violently when attempting to pass low nows und.er· high hydrostatic head . This problem can be over come with flow bypass gates that can regulate releases to Also, sluice gates can be designed to incorporate the addition of lev less than 1 m3/sec. lov nov piggyback gates t o re.l ea.se mini.m\iin lev nows. . .,: . ·: •. ··:::~'~·l·~.!l .. ~·!! ' . ' I , Operation Considerations 27. Flood control operation of a project is designed to attenuate and delay peaks in the inflow hydrograph, thereby r educ ing potential damage caused by i ncreased downstr eam va-..er levels. The attenuation of peak flood flows is obtained by storing water and releasing it throug'h time. Ther efore, the reduction in peak flow resu;J.ts in l.:Jnger periods. of high flov downstream. In many instances, reservoir disc harges are reduced as high flovs enter upstream to permit the downstream tribu- taries to discharge before r eservoir flood waters are released. 28. Flood releases from reservoirs are often hypoli mnetic because the bo ttom sluice gates generally have the largest. capacity . Thus, if epilimnetic withdrawals were occurring prior to the storm event, the downstream area may experience cold hypolimnetic release waters during the passage of the flood waters before returning to epilimnetic release 13 ~:··:;, .. • . .. . : I. schedules. If the project has a selective withdrawal structure, a portion of the releases can be made from near surface vaters to reduce. changes in ·downstream temperatures. 29. Hydropower is generated by tvo types of projects. Run-of-the- river projects generally provide baseload generation. The production rate. of power is determined primarily by the amount of vater flowing into the reservoir. Their relatively small capacity precludes their use in hydropower peaking operations. The hydraulic residence time of the water in run-of-the-river projects is usually quite short, and therefore, the water quality of reservoir releases is often quite similar to that of the reservoir inflow. Hydropower projects asso- ciated with large reservoirs are ideal peaking power plants because of their short response time. Thus, as demand peaks, the turbines can generate electricity almost immediately. In general, hydropower pro- ject releases reflect the demand for electricity, discharging only minimum .lov flows during the weekend and at night. These projects are . brought on line depend.ing on the demand for paver • ....... Summary 30. Efforts to manage a reservoir· tailvater to reflect conditions in an unregulated stream or river are impractical and often impossible because of constraints imposed by the design and operation of.the pro- ject. Thus, the quantity, quality, and t.iming of releases creates an environment vhicb differs from a natural str@'am. Understanding the .impacts of project releases on the tailvater environment, as vell as efforts to minimize detrimental effects and possibly improve down- stream conditions, are often determined by overall project design and operation. ' 14 ' ! ·. ..•. PA.~ IV: PHYSICAL AND CHEMICAL DESCRIPTION OF TAIL WATERS 31. Impoundments cause three major physical modifications in oatu=al stream conditions : (a) seasonal temperature changes are de- layed a.nd the amplitude of diurnal and seasonal temperat;ure· fluctua- tions may be reduced, (b) high natural streamf'lovs are reduced or eliminated and replaced by more moderate discharges o.,er an extended period of time; and (c) sediment transport is reduced (Neel 1963; Maddock 1976). In addition, discharges may degrade the streambed and banks, resulting in "armoring" of the streambed. Impoundments also affect the ch~ical characteristics of the discharge including concen- trations of dissolved oxygen, organic matter, nutrients, and reduced compounds. 32. The magnitude of these physical and chemical I!I.Odi ficatiorus is dependent on conditions vithin the reservoir (e.g., enrichment as related to age, duration, and degree of thermal stratification; hydrau- lic re.sidEmce time; density currents) and the de.pth and volume of dis- charge (Neel 1963). The water quality of reservoir releases can be further modified by conditions in the tailvater sueh as groundvater inflov, runoff, streamside vegetation, and atmospheri.c influences. Physical Characteristics Temperature 33. Water temperatures in the tailvater are determined primarily by ~limatic condjjJons and the ~th of release. Some tailvaters are subject to sudden, drastic temperature changes, vhe:r-eas. in others the, c·hanges are more subtle. Temperature alterations fre.quently result in the elimination of organisms from habitats vhere. they might otherwise survive. Many aquatic organisms present in a stream have distinct . temperature requirements, and changes of l°C can affect their existence (Britt 1962).-""' ... '"! ~l.e.s 34. Epilimnetic release dams on warmvater str~ams provide the tailvater vith vell-oxygene.ted vater near or at atmospheric 15 I I temperatures. Warmvater fish species found in these taiJvaters are vell adapted to release water temperatures. Pfitzer (1954) noted little difference in the varmvater fisheries in tailwaters below Tennessee Valley Authority dams built on varmvater streams vhen water releases vere from the epilimnion. Hovever, epilimnetic releases from. reservoirs built on coldwater streams can increase summer vater temper- atures in the tailwater and stress coldwater species . The temperatures of epilimnetic discharges from Ennis Reservoir, Montana, vere 4°C higher than those in the coldwater stream above the reservoir. This tempera- ture increase caused grovth retardation in trout more than 270 mm long, but did not affect smaller fish and invertebrates (Fraley 1978). 35. Water temperatures are levered belov hypolimnetic· release reservoirs built on historically warmwater streams. Faunal changes are generally :nore pronounced in rivers b e lov these reservoirs than in rivers belov epilimnetic release or nonstratifying impoundments. Sev- eral investigators have noted reductions in varmvater species caused by coldwater discharges belov hypolimnetic release reservoirs built on · ~water stre~s (Dendy and Stroud 1949; Edvard.s 1978) • 36. Coldwater discharges from d e ep-release dams on warmvater streams may result in tailwater temperatures as much as 20°C lover than the temperatures of Uriregulated streams of the region during the summer (Ward and Stanford 1979). A l0°C reduction in wate.r temperature below Norris Reservoir, Tennessee, changed the tailvater from a warmvater to a coldwater streac (Tarzvell 1938). Average water temperatures 11.3 km below Tenkiller Dam, Oklahoma, during June and July were reduced by 4. 3°C after impoundment (Finnell 1953) • Reductions in vat.er "bemperat,ure have permitted the establishment of put-and-take trout fisheries in some tailvaters that vere previously too warm to support trout. 37. During fall, hypolimnetic discharges from stratified reser- voirs may provide varmer than normal vater to the tailvater, effec- tively delaying the autumn decline in temper atures. River ice forma- tions nay be delayed .during winter by lags in temperature change, and some tailwe.ters may be kept completely ice-free by the release o·f 4°C bottom vater (Neel 1963). Delays of 20-50 days in the spring rise in 16 I 4 • -•JI'!· ·::. ' ·-. water temperature have also resulted from release of hypolimnetic water (Crisp 1977). 38. Seasonal temperature changes are also delayed in tailwaters below nonst·ratified reservoirs. Normal temperature changes are re- tarded because the time requir~d to cool or warm the reservoir is significantly longer than the time required to cool o.r warm an .mregu- la.ted stream. Additionally, diurnal and seasonal temperature fluctua- tions take place in unregulated streams, whereas temperature.s in tail- water areas are more nearly constant, especially near the reservo'ir outflow. Marked reductions (up to• 80 percent) in diurnal temperature fluctuation have been recorded (Fraley 1978). Before closure of Flaming Gorge Dam, seasonal temperatures on the Green River, Utah, ranged from 2°C in March to 22°C in July. After impoundment, water temperature fluctuations were reduced and ranged between 2 and 10°C (Vanicek and Krame.r 1969). 39. Severe vater temperature fluctuations may occur below dams . . during periods of low flow or no flow, because of atmospheric influence. Such periods are particularly characteristic of hYdropower projects where changes in water discharge depend on pover demand. Temperature fluctuations of 6-8°C may occur 2 to 3 times a day below these. dams (Pfitzer 1968). If 2 or 3 consecutive days of no flow occurs, water temperatures can approach mean air temperatures. The sudden release of ·· large volumes of cold bypolimnetic vater during the SUDIDer may cause thermal shock. Fish kills have occurred when cold, bypolimnetic waters, with reduced levels of dissolved oxygen, were suddenly released into a tailvater after several days of little or no flow (Krenke! et. a.l. 1979)·~· 4o. Thermal changes caused by bypolimnetic discharge can persist in a ta.ilvater for an extended distance downstream. The effects of an altered temperature regime below a hypolimnetic release reservoir in Canada were noted by the depletion of the benthic fauna 100 km down- stream (Lelmlkuhl 1979). Air t~erature, discharge volume~ groundwater and tributary additions, shade, and substrate type all play a role in modifying the tailwater temperature as the water !!loves downstream. At some point dovnstre~, where the influence of the reservoir lessens, 17 .· . . . .. ~ tbe interaction of these factors results in tbe return a ·f the stream to preimpoundrnent conditions . . Flow 41. flatural streams are subject to large fluctuations in flov as a result of variations in precipita.tion. Seasonally, flovs are highest in the sp::-ing and lovest in the late summer .or early autumn, although intermittent floods may occur as a result of periodic storms. 42. Impoundments can drastically alter the flow characteristics in stream systems. Tailwater flows may be relatively unifonn or may fluctuate frequently, depending on the method of dam operation and downstream water requirements. 43. Flood control and irrigation dams general1y reduce.the mangi- tude of flood flows, and release these flovs at reduced volumes over longer pe:riods of time. The reduction or elimination of floods reduces ban..'lt erosion and bed scour and decreases the amount o.f sediment washed. into the tailwater from the flooded bottom lands. The, resultant. bank and bed stability enhances the growth of aquatic and terrestrial vege-,....... . tat ion (Neel 1963). The encroachment of streamside vegetation, .. vt;i_ch is important in temperatw:-e regulation, shading, and in providing food. for invertebrates, can further increase bank and. floodplain st.abUity. However, such increased vegetative encroachment may resul·t ir+ the eventual loss of part of the wate.r-carrying capacity of the stream channel through a re.duction in channel size (Bovee 1975; Maddo.ck 1976). 44. Uniform flovs below ·nood control and irrigation dam.s often benefit'the invertebrate community through the establishment of dense mat.s of periphytic algae. These algal mats constitute both a habitat and a food supply for benthic organisms incapable of living in a more barren stream. Hovever, these mats may elim.inate species adapted to clean rock surfaces (Ward 1976c). 45. Flov fluctuations are more frequent an.d of greater magnit.ude below hydropove·r dams than in natural streams. Large daily flow fluc- tuations often preclude the establishment of permanent strei!JIIS'ide vegetation. The alternate inundation and exposure of the streambed., coupled with extreme variations in flov, may remove much of the aquatic 18 • • ,, • biota from the tai1wate~ (Neel 1963). A sudden increase in !"low :t.a.y remove algae, macrophytes, and sedimentary detritus, in addition to benthic invertebra~es. Sudden decreases in flow may strand attached or immobile species and result in their desiccation (Lowe 1979). Overall, the diversity and abundance ·of tailwater habitat and fish and inverte- brate food supply may be• s i gnificantly reduced by radically fluctuating flows (Neel 1963). Rec.ent studies (Matter et al. 1981) demonstrs.te that the surge of water from a peaking hydropower plant, and resultant bed scour may indirectly bene.fit tailwater fish by making benthic foods more available. Substrate 46. Because reservoirs act as sediment traps, there is usually little sediment in reservoir discharge. 'This loss of sediment in the discharge, coupled with the removal of fine particles by the current below the dam, results in a tailwater streambed composed primarily of coarse cobble and bedrock. Ultimately an equilibrium is reached be- "'·· tween the particle size of the remaining substrate and the stream's capacity to transport material. Upon reaching this equilibrium, further degradation of the tailwater streambed by scouring is halted (Komura and Simmons 1967). Increased flow rates below some hydropower facilities reduce bank and streambed stability, thereby causing in- creased bank erosion, streambed scour, and eventually armoring·. Scaller sediment particles are swept downstream and deposited in pools and slackwater areas . Turbidit Y 47. Turbidity can reduce or eliminate aquatic life in a stream . Decreased light penetration in turbid s~reams inhibits the establish- ment and maintenance of autotrophic plants, wi.ich may in turn effec- tively limit higher life forms (Ruttner 1963). Additionally, sedi- mentation resulting from turbid conditions eliminates invertebrate habitats by filling the interstices of gravel substrates. Sedimenta- tion may also cover fish spawning sites and interfere with oxygen transport to buried fish eggs (Fry 1960). 19 48. Tailwaters are usually clearer (less turbid) than the reser- voir inf1ow, particularly below deep-release reservoirs. Turb~dity below reservoirs is significantly affected by sedimentation within the reservoir-, density currents, discharge depth from the dam, and the in-oft-> flow from surface runoff and tributary additions. Turbidity was re-~~ \J' 0 .. duced up to sixtyfold in the tailwater below Yellowtail Dam, Montana, /. by the settling of suspended matter within the reservoir (Soltero et al. 1973). Density currents carrying fine suspended matter, however, may sometimes flov beneath or through the main 'body of wat-er in strati.fied reservo1rs and !be discharged directly into the tailvaters with little alteration within the reservoir (Churchill 1958). In these instances, mineral concentrations and turbidity may increase significantly in the tailvater. Turbid conditions may also result from the flushing of loose materials into tailvaters from unstable riverbeds and streambanks during periods of high discharge, and from tributary inflov. Chemical Characteristics ' ·. Chemical properties that may affect the tailwater biota are· . . the concentration of dissolved gases (i.e., oxygen, nitrogen), pH, particulate organic matter, available nutrients, and reduced compounds. Because of the variability of factors involved in altering water quality, rev general statements can be made that are applicable to all tailwaters. Chemical properties of the water immediately below a dam depend on vater quality within the reservoir at the level of release~ As the ve.ter moves downstream, local conditions influ~nce wat.er quality and tend to characterize each individual tailvater (Pfitzer 1954). Di .ssol ved gases 50. Dissolved oxygen. The concentration of dissolved oxygen in an aquatic system is dependent on water temperature, biological oxygen demand, atmosphe·ric exchange, and primary production. Water t~e.ra­ ture determines the solubility of oxygen, and thus the &mQUilt of avail- able oxygen in streams. This is an important factor in regulat.ing the metabolic rates of cold-blooded animals, since their rates of 20 I • . '· •. II!'. metabolism increase vith temperature. The solubility of oxygen .de- creases as vater temperature increases. At 100 percent satur.ation, 14.16 mg/1 of dissolved oxygen may be in solution at 0°C but only 7.53 mg/1 at 30°C (Boyd 1979). Decanposition rates of organic matter increase with increasing temperature, resulting in an additional deple- tion of oxygen content. The rate of decompcsition generally increases between 5 and 38°C. Temperature increases of 10°C o·f'ten double the rates of decomposition and oxygen consumption (Boyd 1979). 51. Mo.st st.reams are relatively vell oxygenated due to turbulent f'lovs and continual atmospheric exchange. In quiet pool areas with dense algal vegetation, diurnal va.ri·ations in the concentration of dissolved oxygen are directly linked to the amount of phot.osynthesis and respiration taking place in the system (Hoskin 195~). Oxygen con- centrations are highest during the day and lovest at night. 52. The concentration of O~Jgen in tailwaters depends on the ·type of reservoir, depth of v--ater releas~, vater mixing during re·lease fran t ·he. dam, and downstream flov conditions. Low dissolved oxygen concentrations normally do not occur belov surface-release reservoirs: - Water from the epilimnioh is usually well oxygenated as a result of photosynthesis :;.nd a.tmospheric gas exchange. 53. In deep-release reservoirs, biological decomposition of organic matter in the hypolimnion during the summer may result in the discharge of poorly oxygenated water into the tailwater. The low oxygen content of these waters may not satisfy the biological and chemical demand, especiall.y if there are additions of domestic and . industrial pollution downstream from the dam (Fish 1959). 54. Tailwater oxygen levels may also be reduced by the oxidation of iron and manganese present in hypolimnetic releases. This reduction may cause physiological stress to the aquatic community and further reduce the assimilation of organic wastes by stream organisms. Low- oxygen conditions may also intensify ~he potentially toxic effects of other chemical constituents-including ammonia and hydrogen sulfide, which are ott.en present in the anoxic hypolimnetic water. 21 ·' ·55.· Reaeration of deoxygenated water can be rapid, and serious oxygen depletions can be avoided if tailwater conditions are such that biologic·al ·and chemical oxygen demands are not excessive. Wirth et al. (1970) documented a consistent concentration of 7 ms/1 dissolved oxygen in discharges from a deep-release reservoir in which the hypolimnion was devoid of oxygen. Reaeration during discharge is credited with maintaining the high dissolved oxygen level. 56. The rate of reaeration below deep-release reservoirs depends on the turbulence of the flow in the tailwater, atmospheric influence, and extent of photosynthesis by aquatic vegetation below the dam. Low oxygen levels may persist farther downst.ream during peak flow periods, when riffle areas are inundated and more laminar now conditions exist. Be~ov an Oklahoma hydropower project, dissolved oxygen concentrations were low ( 1. 5 mg/1) for 3. 2 km downstream during peri.ods of moderate discharge, but extended 8 km downstream (2.0 mg/1) at peak flows (Summers 1954). 57. Oxygen concentrations greater than 5 mg/1 are generally pre- ferred ·by most stream fish (~·y 1 ~60). They appea~ to survive well in streams where dissolved oxygen content occasionally f~ls "below 5 mg/1 at night but !' ises abov·e this level during the day. Certain current- oriented aq·..tatic invertebrates can Yi thstand dissolved oxygen conc.en- trations less than 1 mg/1 if current velocities remain high (Bovee 1975). 58. When dissolved oxygen deficiencies occur in tailwaters, they adversely affect the health of fish through suffocation, growth retar- dation1 and decreased disease resistance. Macroinvertebrates may also be adversely affected,. but tolerant organisms usually replace those that have abandoned the a.reas because of lov oxygen levels. Oxygen reductions that are no·t great enough to retard fish grovth generally do not impair fish food resources (Doudoroff and Shumvay 1967). 59. Gas su·,ersaturat ion. Gas supersaturation can. occu.r in tail- waters when water is spilled over high dams, trapping air and plunging it to the stream below where hydrostatic pressure is sufficient to increase solubility of atmospheric gases (Weitkamp and Katz 1980). ~he 22 , • ·' : . ·~. . . ....... . ···.~ .• .. '·'. high levels of dissolved gases produce em.bolism.s in a variety of fishes and invertebrates. The ~ttpersaturated gases come out of solution within the fish's or invertebrate's body and for.n bubbles under the skin; severe cases can cause death. The condition is most caumon in tailwaters below hydropower reservoirs in the Pacific !lorth- west and has been attributed to both spill~ay and turbine releases ( Beiningen !!..."ld Ebel 1970; Ruggles and Watt 1975}. Crunkilton et a.l. (1980} reported on the' occurrence. of gas supersaturation in a '.tar::l- wa.ter tailwa.ter in Missouri. Spillway deflectors are reported to effectively reduce the level of gas supersaturatic!! in water pass:.ng over a spillway (Weitk.amp and Katz 1980}. Hydrogen ion concentration and alka.lini ty ·. ·. 60. The distribution of organisms in an aquatic system is dete!'- mined to a large extent by the hydrogen ion concentration (pH} of the water. Changes in the pH of surface waters a.re often brought about by addition or removal of co 2 during photosynthetic activity, decomposi- tion of organic matter • • and gas exchange. Fluctuations in pH, UDless . , ·, extreme, are not harmful in themselves, but variations may intensify or dec·rease the effect of toxic substances within the vater column (~ey 1960}. Waters with a pH from 6.5 to 8.6 are most productive, and fish populations are unaffected by slight deviations within this range (Fry 1960}. Additions of acid mine drainage or industrial wastes may result in extreme deviations fr-om these acceptable limits and stress some aquatic organisms (Edwards 1978}. Aquatic insects have been shown to have little tolerance for pH below 4 (Canton and Ward 1977}. Effects on !ish become lethal when the pH falls below 4 or.rises above 11 (Swingle 1961}. 61. Diurnal fluctuations in pH of unpolluted surface va~ers are· reduced through buffering by an alkalinity system of carbonates and bicarbO[tates, and tho! degree of buffering effectiveness depends on the concentration of these substances in the watershed. :otal alkalinity may range from less than 5 mg'/1 as Caco 3 (little buffer- ing capacity present} to several hundred milligrams per litre (Boyd 1979}. 23 Productivity in natural waters is related to their total alkalinity (Turner 1960; Bayes and Anthony 1964). 'Waters vitbin an alkalinity range of 20 to 400 mg/1 are generally considered biologically more productive than those vith higher or lower alkalinities (Moyle 1945). 62. There are no apparent trends in pH and alkalinity value~ in tailvaters, other than seasonal changes brought about b· thermal stra- tification (Vanicek 1967; Pearson and Franklin 1'968). Bacterial de- composition of organic matter in the hypolimnion increases the concen- tration of carbon dioxide, thereby reducing the pH in the tailvater. Turbulence in tailvaters increases gas exchange, which reduces the con- ·centration of carbon dioxide; consequently pH rises as the water flows downstream. Alkalinity values in tail~ters below bypolimnetic release reservoirs are reduced during spr'ing runoff but increase during the summer, when the reservoirs become thermally stratified (Vanicek 1967; Charles and McLemor• 1973; Ha~nan and Young 1974). Particulate organic matter 63. Particulate organic matter (POM), the ·organic component of the seston (Cole 1975); is the main food supply for de·tri ti vo;es in natural stream ecosystems. Concentrations of POM present in e st·ream are often positively correlated vith flow rates, which are, in turn, determi ned by the amount of rainfall in the watershed (Webster et al. 1979).. Concentrati,-,ns of POM are highest during lea.f fall in autumn and this source is gr<Wually depleted from the si.:-eam during the rest of the year. 64. The POM t'!!IS h! separated into coarse and fine fractions. The coarser materials tend ·.;o .move only a short distance downstream before they are trapped by obstructions and reduced in si.ze by mechanical Uld biologic.al processes. The finer particles are generally carried farther dovnstream until a reduction in the flow velocity allows the particles to settle· out. 65. Deep-relee.se reservoirs reduce the transport of most sus- pended POM to toe stream below. Settling and decomposition in the reservoir may .remove 70 tc· 90 per t!ent of tbe particulate organic matter introdut:e6. from the watershed (Lind 1971; Armitage 1977). Most 24 • .t ·'. allochthonous organic matter is washed into the reservoir during high winter or spring f l ows and is subsequently decomposed and transi~rmed into d.issolved nutrients during the period of SUliiDer stratification (Webster et al. 1979). The loss. or reduction of this food supply for bacteria, fungi, and certain macroinvertebrates can reduce the abun- dance of these organisms in the tailvaters. Density currents with high levels of POM may flov through reservoi.rs--und.erneath, through, o.r above the main body of vater--and be discharged into the tailvater, an.d when this occurs concentration of POM in the tailwater increases. 66. In contrast, surface-release reservoirs tend to incr~ase. the concentration of POM in the tailvater through the discharge of large populations o·f sestonic plankton produced in the epilimnio·n of the reservoir. The amount of plankton received fran the reservoir fluctua- tes seasonally; it is greatest during peak production periods (turnover) in spring and fall. Microseston in the effluent of Lake Laurel, California, vas 85 percent more abundant than that in the infloving stream (Maciolek and Tunzi. 1968). Filter feeding invertebrates may floU!"ish below these epilimnetic release reservoirs due to the in- creased availability of food particles. 67. Concentrations o·f PO.M in tai.lvate.rs below reservoirs vith short hydraulic residence times may not differ substantially from those in reservoir inflows. Sho•rt hydraulic residence time does not allow sufficient time for production of enough phytoplankton or settlement and decomposition of enough suspended organic matter to significantly alter the POM concentration received by the tailvater. Nutrients 68. Minerals and nutrients characteristic of the surr0unding vaters.hed are carried into streams by surface runoff and tributary inflow. The amount of nutrients made available t:> a reservoir depe·nds on the nature of the soil, amount of rainfall, l o cal agricultural practices, and domestic and industrial sevage inputs . Significant short-term increases in minerals ~~d nutrients during periods of intense runoff are cl~acteristic of unaltered streams; whereas waters relee.3ed from reservoirs are more uniform in mine·ral and nutrient 25 content (Wirth et al. 1970). Nutrient enrichment of a tailwater is a funct.ion of the enrichment of the reservoir above, reservoir stratifi- cation, depth of release, and hydraulic residence time. 69. Stratified reservoirs that have surface outflows trap nutri-· ents in the deep hypoHmnetic waters (Wright 1968). Total dissolved solids, nitrogen, and phosphorus concentrations decrease in the epilim- nion during the summer through the assimilation of these nutrients by seasonally increased phytoplankton populations. In addition, adsorp- tion to clay particles and subse~uent settling may reduce phosphorous concentrations in the epilimnion. Moribund plankton from the epilim- nion and organic material c:il.rried by the inflowing water continually set,tle out, enriching the hypolimnion as decomposition and nutrient transformation takes place. 70. The retention of nutrient rich hypolimnetic water may in- crease the potential productivity of the reservoir ·(Murphy 1962) ; however, continual inflows. of dissolved and suspended nutrients may result in a noticeable deterioration in water quality (Johnson and 'Berst 1965). The . biological productivity of the . tailvater below a stratified surface-release reservoir is reduced during .tbe summer because of the decrease of dissolved nutrients in the discharge. In tbe fe.ll, however, after the reservoir becom.es vertically mixed and nutrients are uniformly distributed ?.n the water column, the avail- ability of nutrients in the tailvater is increased and productivity improved. 71. Deep-release reservoirs discharge the nutrients which accumu- late in the hypolimnion during stratification. The reduced oxygen concentration, resulting from decomposition, enhances the accumulation of diEsolved nutrients (Hannan and Young 1974). As a result, more nutr~ents are discharged from the hypolimnion in the form of readily usable ammonia and dissolved phosphate. The release of clear, nutrient- rich water, more fertile than that released from the surface, often results in increased productivity in the t .ailvater. Objectionable taste, odor, and excessive algal production are often associated with ' these releases. Dense algal mats, sometimes established below these 26 J ... ~ ... ...... · .. •• 't .~ . ' ; . ·. reservoirs as a result of the increased nutrienta, are usually asso- ciated vi th incre&sed numbers of invertebrates, providing a food base for tailwater fish. Increased algal growth and the subsequent increases in mac.roinvertebrates can aid in the establlshment of a trout fishery (Pfitzer 1954). Reduced compounds in tallwaters 72. Hypolimnetic discharges may contain high concentrations of reduced iron, mangane.se, sulfur, and ammon.ia produced by natural ly occurring anaerobic processes. High concentrations of these materials may be toxic to tailvater biota, or affect certain life stages at sublethal levels (Lehmkuhl 1979). Dissolution of these substances fraa the soil and decomposed organic matter occurs in the reservoir vhen the hypolimnion becomes anaerobic and the redox potential is !overed. A combination of these elements may have a synergistic effect, so that no one element or compound is solely responsible for toxicity to the biota (J. M. G~izzle, Auburn University, unpublished manuscript). There- lease of anoxic hypolimne~ic water may produce combinations of toxic elements that normally vould not be present in una.l tered streams c;;,r in.-~ .. .. . . • ~ .• . ~~;·~ .. ,. •....._ epilimnetic release tailvaters. ~-:-"'"" 73. Iron and manganese. Soluble reduced forms of iron and man- ganese begin to oxidize upon release from the reservoir and precipitate in the form of ferric a·nd manganic hydroxides that may stain concrete and rock surface·s in the tailvaters. These poorly solubLe hydrous metal oxides are a nuisance to dovnstream munic ipal water treatment plants . Deposits of these hydro·xides on the substrate below hypolim- netic release reservoirs may also af.fect the numbers and: types of organisms present. However, the effects of these deposits have not been sufficiently documented (Krenkel et al. 1979). 74 . Iron concentrations in neutral or alkaline waters usually range from 0.05 to 0.20 mg/1 (StW!DD and Lee 1960). The highest accept- able concentrations of iron are 0.30 mg/1 in domestic water supplie• and 1.00 mg/1 for treshvater a.quatic life (U. S. Enviromnental Protection Agency 1976). 27 ··~ -·-· -···------···--·-----, ....... 1 75. The chemical characteristics of manganese are similar to those of iron; however. manganese has a slower ox1dat.ion rate and forms more soluble salts than iron. Manganese concentrations are not as effectively eliminated from the water column by precipitation (Wetzel 1975). Soluble forms of manganese are therefore more persistent in tailwaters. Below stratified deep-release reservoirs, manganese con- , centrations may exceed l mg/1, which is greater t 'han concentrations found in most freshwater environments (Churchill 1958; Martin and Stroud 1973). Concentrations less than 50 ug/1 have been found to inhibit green and blue-green algae in streams and to favor diatoms (Wetzel 1975). 76. Hydrogen sulfide . Occurrence of hydrogen sulfide in bypolim- netic discharges is the result of the anaerobic bacterial decomposition of organic sulfur compounds and th.e reduction of sulfates to sulfides vi thin the reservoir. Hydrogen sulfid·e concentrations are highest and most toxic under acidic conditions (low pH). In neutral or alkaline waters • the sulfides combine with iron, forming insoluble ferrou.s sul~ide 9 which precipitates from the water. Thus, hydrogen sulfid~ · appears ·only when pH is low and oXygen content. is near zero. or when all available iron has been precipitated from neutral or alkaline water (Syoons et al. 1964). 77 . Discharges with hydrogen sulfide concentrations above 0.002 mg/1 have a!'l. objectionable odor and may re,sult in fish kills and in a reduction in diversity of benthos and algae. Smith et al. (194~) reported a 72-hour Lc 50 of 0.019 D.g/1 for bluegill eggs at 2l.9°C and a 96-bour tc 50 0.045 mg/1 for adult bluegills a:t l9.6-20.3°C. Fish die-offs a n d adverse benthic responses have been attributed to high levels of hydrogen sulfide (Wright 1968). 78. A.'!l!!lonia . Ammonia nitrogen (NH 3-N) is a by-produc·':. of organic decomposition (Cole 1975). In some situations, depending on the pH of the water, NH 3 may be toxic to living organisms. Concentration of NH 3 increases as pH increases, and NH 3 is most toxic when 'botb dissolved oxyge.n and carbon dioxide levels are low (Boyd 1979). Toxicity is also affected by temperature and alkalinit.y (Lloyd and Herbert 1960). 28 , .,· .. ' ·' ··- ·' .· Concentrations betveen 0.6 and 2.0 mg/1 are lethal to fish after short- term exposure (European Inland Fishery Commission 1973, in Boyd 1979). Pathological changes in fish organs and tissues have been noted after continuous exposure to sublethal concentrations between 0.006 and 0.34 mg/1 (Smith and Piper 1975, in Boyd 1979). 79. In late summer OF early fall, the concentration of 1m 3 in- creases in the anoxic hypolimnion of stratified reservoirs as the result of extensive anaerobic decompgsition of organic matter. Addi- tional JR3 may be released from the bottan sediments or may be. carried into the hypolimnion by density currents {Hannan 1979). Consequently, 81:11110nia concentrations in tailwaters below hypolimnetic release dams tend to increase during late summer or early fall. Ammonia concentra- tions also increase in tailwaters below epilimnetic release dams during fall, after the fa.ll overturn. . ' 29 . .. .. • ... PART V: METABOLISM AND TROPHIC STRUCTURE 80. The assemblage of organisms living in a particular reach of a tail~~ter may be considered a biotic community. In this community, the interactions between the various types of organisms (i.e., primary producers; primary, secondary, and tertiary consumers, etc.) are based primarily on their nutritional requirements and feeding habits . The canplex nutritional and energy cycle that results from this interaction is often referred. to as the trophic system, within which each type of organism occupies a particular level. 81. Energy enters the tailvater trophic system in the form of light energy and nutrients and detritus. These materials are used by the primary producers and consumers that make up the lover trophic levels of the system. Energy is progressively transferred upward through the system· when the organisms in these love.r trophic levels are eaten by the secondary and tertiary consumers that mak.e up the higher trophic· lc·;els • • !, .. .'.;; ·~ .. , . 82. As the energy is transferred from one level to another, • 4 ·~ • - lo~ses result from the partial use of the available energy requir~d for ' maintenance and reproduction. Generally, each successive trophic level contains only about 10 percent of t he energy available to the preceding one (Russell-Hunter 1970). The net result is that only a small .per- centage of the original energy is available at the highest trophic ·. level. Greater biomass production at the primary level, enabled -by · increased input (e.g., primary production, detritus), usually results in greater biomass production at the highest trophic level (Bovee 1975). Streams 83. In streams, metabolic essentials are not recycled and must be constantly supplied from upstream sources or streamside vegetation. Metabolic activity and productivity are addi.tionally governed by the composition and sources of the nutrients entering the stream and how efficiently thP.y are utilized. 30 I ... ·'· ·. .-'·· .. , 84. Stream organisms take up, transfonn, use and release organic materials, thus acting as processors of the organic material passing through the• system (Fisher 1977). Fish, the largest organisms in the stream community, us.ually represent the top level of the food chain. Their abundance reflects the quantity of primary and secondary produc- tion that takes place in lover levels of the trophic system. 85. The organic content of a natural stream system comes from allochthonous and autochthonous sources.. The metaboli sm for both sources is based on the supply of detritus vhich is used as food by de·tritivores and omnivores. and furnishes dissolved nutrients to the primary producers. An allochthonous system (heterotrophy) is based primarily on organic material that is carried into the system from the vatershed. Autochthonous systems (autotrophy) derive their energy ~ram photosynthesis that takes place in the streambed. These systems are most representative of slaver flovs, vhich enable a buildup of peri- phytic vegetation and, occasionally, vascular plants. 86. In contrast to the source of organic materials in most ecosystems. tbat in streams is d~rived primarily from allochthonous sources (H~es. 1970; Cummins 1974). A headvater stream, for example, may derive 99 percent of its energy inflov from allochthonous origins and the remaining 1 percent from photosynthesis (Fisher and Likens 1973; Cummins 1974). Higher order streams may also depend on alloch- thonous sources of energy . 87. As much as 60 percent of the total organic matter taken into the• stream from allochthonous sources may be in the form or leaf litter (Cummins 1974). Additional allochthonous materials may be in the form ., of twig~ and shoreline debris or dissolved nutrients from·vatersbed runoff. Leaf material may be found in suspension in the stream or deposited on the stre9JIIbed (Minshall 1967). The leaves are rapidly colonized by fungi and bacteria, vhich aid in p rocessing the material .into fine particles and dissolved organic matter . The dissolved nutrients released duri.ng decomposition are then available for u_se 'by primary producers . The colonizing fungi a nd bacteria are in turn used as food by the shredding and scraping invertebrates that are involved 31 ··- ... in 'the mechanical aspects of leaf decomposition. The fungi and bacte- ria may be the primary source of nourishment for these invertebrates, since some leaves have been shown to be of little food value. These invertebrates, in turn, make up the primary prey of predators in the higher levels of the trophic system ( CUIIIIlins 1974). 88. The structure and complexity of the benthic community may .. change vith the amount and variety of plant detritw~ present in the system (Egglishav 1969; Mackay and Kalff 1969). As the organic material breaks down, the variety of food becomes more diversified, producing a similar response in the invertebrate community. Seasonal shifts in the diversity of the stream invertebrates are related to changes in food supply and othe.r natural changes during the life cycle of the organisms (Mackay and Kalff 1969). 89. Since many aquatic invertebrates are able to process organic matter at lov temperatures, much of the organic material in streams is used during fall and winter . Hovever, not all of the organic matter entering the systec during fall is used. Sane of the material is stored in the slover depositional zones o~ the s 1tream, vhere it remains ·"·'"'· ... .. .. · · until used by the stream biota. These zones act as energy reservoirs and help maintain the biota annually. Material in depositional .. ~one15 .· · may be re.distributed during flooding. This redistribution may be important in slov-vate·r zones, in r ·educing the occurrence of oxygen deficienc.ies that could develop during extended periods of reduced flov. 90. Autotrophic production varies as a result of differing envi- ronmental situations (e.g., changes in shading, turbidity, water velocity, and water chemistry). Autotrophic production is greatest in streams with little shading from streamside vege·tation, and in small streams in forested areas before and after formation of a lellf' c.anopy (spring e.nd fall). Photosynthetic acti.vity is greatly reduced in heavily shaded streams, even vhen adequate amounts or nutrients are available. 91. Autotrophy may be the major contributor to the energy budget or large rivers or sm.all uncanopied streams (Minshall 1978). Removal 32 t ·• •• ... ·' ·. of a canopy results in a shi.ft from a heterotrophic to an autotrophic system~ as stream temperature and photosynthetic production both increase {Gelroth and Marzolf 1978). Photosynthetic activity by periphytic ·algae takes place primarily in shallov~ well-oxygenated, and vell-lit stream bottoms. The periphyton is usually the first auto- trophic component to become established in a stream, and is most preva- lent where flows are least variable. Phytoplankton become dominant in rivers and large streams where depth and turbidity inhibit benthic production by the attached algae {Fisher and Carpenter 1976). 92. Faster current velocity in riffle areas, as opposed to pools, results in increased rates of net primary productivity (Kevern and Ball 1965). Steep diffusion gradients between plants and available nu,trieata are formed in the fast-flowing water, allowing for more rapid assimila- tion of vi tal substances by the· attached algae or macrophytes. Because of these steep diffusion gradients, many algal species grow best in swift currents {>15 em/sec) (Whiteford 1960). Phosphorus uptake may be over 10 times greater and respiration over 70 percent greater in swift water {18 em/sec) than in still water (Whitford and Schumacher 1961). • ·\ .. -.:"":·!: 93. Dense, fel tlike, dark green or brownish communities, con..:·· .~. sisting mostly of diatoms, may be established in streams vi th svi.ft curreats (Mcintire 1966). Diatoms such as Ganchonema, Diatoma, or Navicula, which require high light intensities and solid substrates, are co~~~nonly found in swift streams in association with filamentous algae such as Cladochora . The accumulation of attached algae on gravel and rubble is more rapid in fast currents, but the growth stabilizes after a time and the total biomass per unit area is similar to that ia slower c.urrents. However, higher productivity is maintained in the faster current, allowing a greater export of biomass (Mcintire 1966). 94. Slow currents {<15 em/sec) may allow associations of green filamentous algae (including Stigeoclonium, Oedogonium, and Tr i bonema) to develop {Mcintire 1966). These associations appear as bright .green aggregations similar to those found in ponds. Concentrations of organic matter are usually higher in streams dominated by green algae than in those dominated by diatoms. Conversion from a diatom-moss 33 .. · . j assoc.iation to a c011111unity of filamentous green a.l.gae (possibly associated vi th rooted aquatic macrophytes or diat.oms) may indicate a shift to a more autotrophic stream syst~ (Cummins 1974). 95. Macrophytic vegetation develops in streams where flovs are relatively stable. This vegetation is seldom consumed and therefore does not directly contribute to higher trophic levels while alive (Cm:!mins et al. 1973~ r."ishe.r and Carpenter 1976), although they may serve as a surface for periphyton. However, when the vegetation dies 'Uld decomposes, it contributes organic matter to the syste;m. St.re.ams subject to severe changes in flows physically limit the development of significant plant growth. Additionally, these plants are not found 'where insolation to the stream is low or where the vater is relatively deep and turbid. Tail waters 96. A variety of types of organic matter, leaves, POM, algae, ·'etc :~ must be present in the stream to maintain ecosystem diversity :· . -:(CUIIIIlins 1972). However, reservoirs · act as particle traps, interfering vith the passage of detritus into the tailvater. As a result, detrital material from the watershed, important in energy transformation in natural streams, is largely unavailable to a tail water system. 97. Nut .rient material present in the clear disc.harges of bypolim- netic release reservoirs is primarily in a d.issolved state as a result of decompositi.on by-products that accunulate in the bypolimnion. (Odum 1971). Water d.ischarged from the .hypolimnion may be more fertile than that from the epilimnion because of the concentration of these dis- solv.ed nutrients in the deep vater. 98. Tailwaters immediately below hypoltmnetic release dams are autotrophic because concentrations of dissolved nutrients are increased and turbidity is decreased. The clear, nutrient-rich discharges are particularly important in tailvaters with stabilized f1ovs because they facilitate the production of dense algal growths (Stober 1963; Ward 1976b). However, as the water moves farther downstream, conditions 34 - • •• .. -' ••• (i.e. • increased turbid~-ty, reduced nutrient availability) become -less conducive to algal production,. The gradual increase in the detritus load due to allochthonous input results in a rev~rsion of th~ tailvater to a heterotrophic condition in vhich the algal community plays only a relatively l~ited role. It may. therefore. be possible for a section of stream belov a reservoir to shift from an autotrophic system in the i.Dmediate tailvater area to a heterotrophic system downstream. vi th ·'-0 area of transition in betveen. 99. Tvo genera of filamentous green algae, Clado~hora and Ulothrix, are commonly found belov deep-release reservoirs (Stober 1963; Ward 1976b). Mats of Clado~hora vere located in riffles in the first 9.6 km below Tiber Reservoir. Montana (Stober 1963). The for.u.tion of these algal mats may physically inhibit the production of the oost desirable fish food organisms but may attract othe-r taxa (Wel~h i96~; ·. · Ward 1976.b). The algal mats may act as barriers to organisms thFt.t · require deeper substrates for completion of their life cyclP.s (Armitaae 1976). 100. Water discharged from surface-release reservoirs, vbet~ stratified or not, may contain s,ignificant amounts of plant debris an4 .. detritus, but the primary source of organic matter in the tailvater is plankton produced in the reservoir. Plankton can be used directly by secondary consumers in the tailvater or, after de3.th and decomposition, may add to the particulate organic matter that is available to both primary and secondary consumers. Thus, most surface disc barges from reservoirs supply particulate organic matter to tailvaters, and deep discharges supply dissolved organic matter. 101. Phytoplankton uu.y occur in zones of the tailvater vhere the vater velocity is reduced and the str~ambed vid~ned, and adequate nutrients are available. Hovever, phytoplankton numbers decrease as the vater flovs farther downstream because of depletion of nutrients, increased turbidity, and simple mechanical dest-ruction (H~rt~ e.nd Himes 1961). 102. Storage of infloving vater for extended periods (e.g., as in flood control reservoirs), accompanied by reduced downstream releases, 35 . _,. -........ ~ ... ·· I 'I .. I J may result in bank stabili.z.ation and ~be eventual establisl:Dent of strea::tside vegetation in tailwater reacbe.s. Streamside vegets.tion in these e.reas increases both the bJilount of shade and the quantity of allochthonous materials in +he tail water. The establishment of macro- phytic vegetation BnJ periphytic algal growths are also enhanced vben stabilized flovs are accompanied by reduced tur'bidities (Wa...•d l976b). 103. M1.lllagement schemes for some reservoirs require that dis- charges be released at irregular intervals. For example, releases frOID flood control reservoirs may be erratic, since they usually depend o.n duration and amount of rainfall in the waterst.ed. These flov irregu- ·larities result in a brief buildup of detrital. material in the str~ambed of the tailvater during minimum re1eases. The subsequent relee.se of runoff vaters from the res~1w~ir may scour the tai.vater, resulting in a physical disruption and alteration of the stream simi1ar t~ that in a natural stream after a heavy rainfall (~otzky 1971). The periodic releases may sveep the streambed free of accumulations of leaf li~ter and detritus deposited by adjacent s t reamside vegetation (Ward 1976b): The tailvater is thus denied an important source of ·.energy -1·-# • i .. ~Radford and Hartlar.1-Rove 1971). 104. In tailvaters subject to daily variations in vater release (e.g., peak pover hydropower facilities), detritus and sediment are cons't.antly flushed avay (Ward l976a). Enrichment vitb perticulate organic matter is !llargi.nal at best, part.fcularly near the dam, and the lack o~ detritus places further stress on the trophic structure of these waters. Fluctuations in vater level increase streambed and bank erosion thereby increasing turbidity and discouraging the estab- lishment of s~reambanY. v~getation and streAmbed algal growth. These cor:-i ... t.i ons may critically lit:~it the tailvater biota, 'which might other- vise c.hrive. 105 . Te.ilvaters are highly modified environments that may be subject to extreme conditions not normally found in natural streams. Be~ause of these modifications, the tailvater biota may be disrupted vr enhanced. Either development vill result in the creation of a trophic structure ~uch different f r om that normally found in an unaltered stream . 36 • • • : ... .. . .. -. ·•'.'•;. •, '· PART VI : AQUATIC I!fVER'1'EBRATE IN TAILWATERS Invertebrate Ecology Streams lo6. Benthic stream Ca!IIIIUllities are extremely dy'namic and are com- posed of a large number of species (Patrick 1970). Many have intricate life cycles adapted for survival in the changing environment found in most streams (Brusven et al. 1976). Their life cycles are characterized by short generation times, high reproductive potentials, and reduced, body sizes (Patrick 1970). The adults of most important stream insects are terrestrial, short-lived, and con.cerned only vi th breeding and dispersal (Hynes 1970). 107. The bottom fauna is not randomly scattered and its distribu- tion results from interaction of the invertebrate's habitat requirement• with the varying environmental conditions that exist in different areas of the stream (All.en 1959). The mosaic pattern of distribution ex-. . .. ,..•.:, hibit~d b,y benthic 'organisms is pri~ily determined by cur'rent veloc-· · · ity, substrate•.type, and food availability (Ward 1976a; Minshall and Minshall 1977). In addition, temporal and spatial temperature differ- ences affect the presence of both individual species and life stages. Temporary colonization of microhabitats produced by changes in these variables increases the taxonomic diversity of the stream benthos 'and ensures a year-rour.d food supply for fish (Ward 1976b). Outside inter- ferences (e.g., pollution, ilt.pound:nent) in a stream system tend to red'Uce the benthic diversity as a result of the reduced diversification ·· of available microhabitats (Ward 1976a). 108. ~ Current velocity ma.y h-ave the most influence on the regulation of invertebrate distribution and abundance, especially at specific sites in the stream (Chutter 1969; Giger 19i3). Hovever, the influence of current velocity on invertebrate densities in different sections of the stream may be masked by the effects of other variables (Chutter 1969). Some stream organislllS have morphologic:::a.l respirator;y and feeding demands that require them to position themselves in floving 37 .... water (Ward 1976a). As long as the stream velocity remains relatively high~ and the supply of food and oxygen is adequate, these organisms can survive (Bovee 1975). However, life stages adapted to fast water d.ie vhen subjected to slov current and reduced oxygen because they are phys- iol.:>gically unable to adjust to the altered conditions. Relatively high stream velocities are required to ventilate the delicate gills of these organisms and they must be exposed to the current (Gig~r l973t Armitage 1976). In· addition, filter-feeding i ·nsects are deprived of food if the'" current fails to carry materials into their food-gathering nets . 109. Many stream organisms move about in a "boundary layer" or dead-water zone (up to 1 mm in thickness) near the interface of the substrate and vater vhere current velocities are substantially reduced ( AI!Iouhl 1959; Ward 1976a). The boundary layers and dead-vater zones formed in and around substrate components provide many types of habitat and locations for invertebrates. Rough substrates result in an increase in thickness of these boundary layers and a tendency tovard ~re rapid reduction in the current velocity (Bovee 1975). 110. Strong stream currents discourage t'ree-sYiJIIIling invertebrates. Insects living in the svift-vater areas 'are .;;phologically modified· to vi thstand the mechanical forces of the· current. Most adaptations enable insects to avoid many of the adverse effects of strong currents by kee~ing their bodies avay from the force of the flov (Hynes 1970). Fusiform, flattened, and streamlined bodies aid survival in swift currents by enabling insects to "crouch" in the boundary layer. Because morphological modifications force them to face upstream, most voluntary movei:~ent is upstream (Hynes 1970). Temporary attachment structures are common adaptations that enable insects to release and reattach them- selves after a period of uncontrolled movement in the current. Some organisms construct c.ases of small stones, detritus, or silk that are anchored firmly to the substrate. These organisms catch organic particles frO!:l the water by either m~ing morphological filtering mechanisms or constructing nets. 111. Reduced current velocities li:ni t the abundance and diversity of "svift-vater invertebrate connuni ties, either physically because of 38 • • •• . . --... ... ..... :: ..... , • .... siltation or physi.ologica.lly because of inadequate oxygen and nutrient exchange (Giger 1973). An extreme reduction in current eliminates species that are dependent on flov for respiration and food procurement (Sprules 1947) . 112. Freshets, flov cessation, riffle sedi.mentation, water level nuctuations, and unstable substrates .result in changes in the species composition and may catastrophically reduce the stream fauna (Sprules 1947; Peterson 1977). Severe floods and spates scour and flush insects from the streambed, leaving only the species able to vithstand the in- creased flovs (Hynes 1970). Scouring may reduce the numbers of insects in a particular section of·a stream by 50 percent (Sprules 1947). 113. Spates that scour the stream substrate result in temporary dislocation and dispersal of invertebrates, but the density and struc- ture of the COIIIIIIWli ty may recover to pre flood conditions vi thin 30 days if the insects are not dispersed over great distances. After seven floods in Big Buffalo Creek, Missouri, the structure of the in.vertebrate riffle community vas very similar to that found in the same location bet'o·re the floods (Ryck 1976). Apparently, enough organisms are left after spates or. other disturbances to repopulate the stream section. They escape the ef.fects of the disturbances by maintaining themselves in the protected areas of the stream vhere flov is reduced (Patrick 1970) • 114. Most stream organisms are protected from the effects of severe flooding and relatively short periods of devater~ng because they ·<:::· .: .. ::~· .. ··· .. are imbedded in the upper 15 to 22 c:n of the gravel substrate (Hynes .. , .. 1974). The movement o·f trichopterans into deeper strata of the strella . bottom may be indicative of their response to scouring or to inc;eased sediment loads that accompany flood waters (Poole and Stewart 1976}. .... .... In addition, extended hatching periods and firmly attached e.ggs also en.sure that certai.n species are not eliminated during: str~am distur- bances (Hynes 1970). The repopulation of a riffle sections is probably a result of a combination of several factors, including downstream drift, upstream larval movement, ar.u upstream <:gg-laying. flights of adults (Ryck 1976). 39 : ··•··· .• 115. Substrate tYDe. In most streams, pools and riffles form the most _distinctive types of habitat. Formation of pools and riffles is a combined process of dispersion and sorting of the bottan materia.ls. Pools are ·associated vi th stream bends, and riffles vi th crossings and inflections in the streambed. Fine particles are washed away from riffles and deposited in pools, leaving only the larger graYels and rubble in the riffle area (Yang 1971). Ri.ff.l:es provide optimal environ- mental conditions for many species and are diverse, productive inverte- brate habitats. Invertebrates produced in riffles may pott:ntially be swept into the pools, where they are likely to 'be eaten by fish (Peterson 1977). More insects are usually produced in riffles than in pools or on bedrock or submerged vegetation because current velocities are higher in riffles and rubble substrates incre.ase the availS:bili ty and number of microhabitats. Insects inhabiting the· pool areas are . similar to those found in ponds and lakes and feed on the ac.cumulation of organic matter that forms sediment (Krumholz and Neff 1970; CUDIIlins 1972). }-~·· 116. Intermediate zones, which may oc.cur between stream riffles · ----and pools, a~e called "runs" (Luedtke et al. 1976.). ·:These are ~s of. moderate flow over relatively shallow stretches of ·the stream and may be depositional or erosional, depending on. the current velocity. The identity and abundance of benthic fauna present are determined by whether the area' is in a depositional or erosional :.one. · . . ll7. The benthic fauna associated wit'h specific substrate types generally forms a well-defined community. Any change in the substrate results in an accompanying change in invertebrate species (Sprules 1947; DeMa.rch 1976). Lar,ge rubble (45-70 111111 in diameter) has a variety of microhabitats, which me.y be inhabited by all sizes of insects. This type o:f subs,tre.te is found in swiftly !'loving areas of strE!ams where smalle.r sized substrate materials are vashed away. The organisms found here are ada-pted to living in habit-ats that offer reduced contact with the current ( CUIIIIIlins 1966). Invertebrates able t .o survive in small-particle ( 5-25 JIID) substrates are, by necessity, small and 4o • .•. '. .. . ...... ~. . •·. . . · . ~-=·.. ... •. ........ . . ' ' resilient (Sprules 1947). Their microhabitats ar~ commonly des1.royed· or altered during high, irregular flows. 118. Manipulations of streamflow may alter the detrit~U "trap" capacity of the substrate and ultimately affect species composition and stream productivity. Small detrital particles and silt tend to accumu- late excessively in the interstices of small-particle aubst.rates, ·whereas in rubble habitats the interstices are swept cl.eao by the current. As a result, many a.quatic insects prefer substrates composed of moderately si.zed ( 25-45 mm) particles because they serve as a better benthic food trap without reducing habitat diversit~r (Rabeni and M:Lnsball 1977) . 119. Some species of insects are able to take advantage of in-. creaaed sedimentation, but usually suspended and settled sedimeotl adversely affect the invertebrate population. At lov flows and reduced current velocities, silt and sand seal the interstices in a rubble substrate. This sealing restricts access to the undersurfaces of the ston.es and generally reduces the !lumber of usable habitats. The sedi- .. • ! • .. ... .. ... ........ ..--• •. .-\ -· ·.. ··~"-ifi:·s.l:::. ment ·is ttasily displaced into suspension at higher . flows and settles · .,; out in deposi tiona! zones where flows are reduced. The problems :. caused by sediments are sometimes lessened by the development of car- .pets of algae over the streambed, which r:ur.y replace microhabitats filled in with sediment (Brusven and Prather 1974'}: · .. · ·· ...... 120. Macroinvertebrates apparently migrate out of areas exposed to heavy sediment loacs. Their response to sedimentation is rapid, and only a fev days are required for numbers to decrease signi.ficantly. Conversely, their response to a decreas.e in sedimentation results in a ..... ·.rapid recovery of the c011111uni ty. Sandy substrates are generally un- suitable for insect production because of the instability of the sand particles and the thinness of the protective "boundary layer." Up- st.ream movement of riffle insects in sandy areas is generally prE:cluded by the effects ot current on the fine, loosely compacted sediments. L:w current velocities and rubble substrate facilitate upstream movement of · invertebrates (Leudtke and Brusven 1976) • 41 · 121. ~ A mature stream. ecosystem is highly diverse, and includes complex food webs (Krl.DDholz and. Beff 1970). The increased. ccaplexi ty of links. in the food veb increases the chances of survival of the stream community. A fluctuating autochthonous food supply 'supplemented by allochthonous nutrients increa.ses survival because . - species are not restricted to a single food item (Russell-Hunter 1970). 122. Although benthic crustaceans, mollusks, and other small invertebrate life might be present, aquatic life stages of insects are the most abundant f'orms of primary cons.l.DDers ·in most stre~. Their diets vary greatly (e.g., the forms represented may include herbivores, ·detritivores, carnivores, and omnivores) and some forms sh:if't from one source· to another seasonally (Chapman and Demory 1963). A change fran herbivory in the early insta.rs to ca.rnivory in the later insta.rs is common (Anderson and C'u:Dmlins 1979) . 123. Food gathering and the a.ss-:>ciated morphological and. be- havioral adaptations are the most important functions of' animal con- sumer& in a lotic system (Cummins 1972). Food ha~its are more closely related to the size of the organism than ~o :~e species. Most·early .. 'lite stages ~d small-sized insect's'.f'eed primarily on ·detrital can-:·''· ponents (Cummins 1972). Aquatic in.:.;•!cts. are opportunistic and are able to adJust to differing availabilities of food; very fev are strictly he.rbivores or strictly carnivores. Minshall (1967) noted the presence in a small benthic community of 14 percent herbivores, 3 percen.t carnivores, and 83 percent omnivores. Food availability is gene·rally not a. limiting factor for mobile stream organisms because they are able to actively seek food (Krumholz and Nef'f 1970). 124. Macrophytic coi:IIIIunities often shelte.r larger a .nd more varied populations of invertebrates vben they are composed of plants with finely divided leaves, rather than plants. with simple leaves. Herbi- vores harvest peripbyton from. the surface of the leaves and use the m~o:.~ of' vegetation for s'hel ter and protection (Harrod 1964). 125. The riffles may provide a. numbe.r of diYerse foods for inve.r- tebra.tes, ranging from plant detritus lodged under stones to cl.umps of algae and moss associated vi th the stream bottom. The accompanying 42 . . ... ·' .. ------ invertebrate communities may differ greatly aaong the various types of food sources (Egglishaw 1969). 126. The food base of invertebrates living in slow-water areas or pools primarily depends on the depos.ition of detrital material washed in fran up·stream. Water in pools may be too turbid and the current too slov to allow the establishment of sufficient benthic algal growths, and too deep to allow the development of' signifi,cant quanti ties of macrophytes. 127. The ·aquatic insect component of the stream benthos has been classified into four feeding t;rpes: shredders, scrapers, collectors, and predators (Cummins 1973). The life cycles of shredder-type aquatic ·· · ,•'· ····· · ·-··insects are closely associated vi th leaf fall in autumn. Most of their .. _ .. . . ~ ..,. -·. ·'• . .• ';"·· "" growth and . d:evelopm.ent occurs during late fall and winter. These organisms are able to adjust their growth rates to the relative avail- ability and quality of t'ood. Observations on shredders have indicated a positive selection of leaf material that is heavily colonized by . microorganisms, both on the surface and throughout the matrix of the ~ leaf particle {Anderson and Cumains 1979) . . . · · _:.;;· ·:;..;,;~-· . . 128. Scrupers are dependent on the production of an autochthonous · -~ ,~ food source from vh.ich they 11 terally "scrape away" required food particles . The nutriti~nal content of this potential food source (mostly algae) is much higher than t .hat of a detrital food source (Coffman et al. 1971) • These organisms are most often. in the _.-, ..• .-.--mainstream channel, where the dominant foods are diatoms and fila- . ...... , ... - • .::..-.;.~.-~"··~.: .. mentous green algae ( Cum!ri.1s 1972) . 129. Collectors filter fine particulate org~~ic material contain- ing surface colonies of ba~teria and fungi from the stream flow. The assimilation of nutrients by filter feeders is ineff.icient and much of the material is passed dOVTlstream, unmodified, wh~re it may be rein- gested by other organisms • This process results in a type of unidirec- tiona! "spiralling" cycle in which many organisms, each feeding on different components, are able to utilize !'ilterable organic material. Without this filter-feeding commun i ty, mu ch of the suspended organic material would pass through the system unused (Wallace et al. 1977). . .... 130. Predatory insects have high assimilation efficiences, largely because the nutritional quality of their prey is very high. However, the quantity of prey may vary and i s usually limiting (Anderson and Cummins 1979}. 131. Feedi~6 habits partly determine the location and limit the presence of invertebrate types in a stream. Filter feeders 11 ve in moving water with a seston load, scrapers in areas where algal growth is most apt to become established, shredders in areas of leaf litter and detrital accumulation, and predators in any or al.l of the available feeding locations, depending on the abundance of the potential prey ·population. 132. Temperature. Diurnal temperature fluctuations are generally required for normal growth and de.ve·lopnent of stream insects, whereas static temperatures are usually disadvantageous (Ward 1976c}. Life histories of most stream organisms a:~e geared to annual temperature cycles which synchronize the particular life stages and stimulate growth (Hynes 1970; Gore 1977}. Triggering mechanisms for excystment, .· . encystment, and other vital dP.velopmental processes may not be stimu- l ated when temperatures remain unchanged; differences of a .few degrees affect developmental times and durations.. Higher temperatures are accompanied by reduced concentrations of dissolve.d oxygen and increased metabolic demands, and cool tempera.tures produce a metabol ic slovdown (Bovee 1975}. The absence of a certain species from a stream section may result from one of the specific temperature requirements not being met. For example, Epeorus sp., a heptageniid ephemeropteran, has been shovn to require near-freezing temperatures for a certain length of time for the development of diapausing eggs . Additionally, nevly hatched nymphs require an extended period (2.5-4 months} of relatively high temperatures (18-28°C} for normal development and maturity (Britt 1962}. 133. Numbers and biomass of invertebrates in the streambed may fluctuate .-:-aCJ cal ly because of di f .fering rates of develoJDent and emergence (F~trick 1962). Invertebrates emerge in the same sequence each year, but the dates of first emergence and the duration of the 44 ' ' .•. 1 ··, ! -'i .; l ·-I . . :; '• ... , •'1 ....... ~..... • ••••• , ••• I ··-····· ..... -.. hatch vary, depending on. water temperatures (Sprules 1947). Insects may emerge to mate and deposit eggs all year, but the· greatest emergence takes place during the eveni.ng hours in spring and early summer ( Cbapmao 1966). During emergence, the population of a :particular species in the stream may be re<iuced to near zero (Hooper 191'3; Brusven et al. 1976). Alterations in stream temperature during this time may cause an extended period of oviposition, resulting in seve.ral ages and sizes of nymphs (Minshall 1968). Competition among the progeny of a species, as well as between vari.ous insect :;pecies with similar life requirements, may be partly averted by the stagge.red deposition of eggs and subsequent d.if'ferences in developmental ratee (Minshall 1968). The early stages of many aquatic invertebr~tes are small and essentially 'unavailable as fish t'ood ( Chapman 1966). 134. Densities of insects may vary seasonallY, dependjng on species composition. In northern temperate ·streams, the benthos genera:lly reaches a numerical maximum in the fell and earlY winter · a:rter ,eggs batch ( CUJIIIIlins 1973). Periods oi maximlll.' stream inverte- brate b .iomass may not occur concurrently vi th the numerical maxima . The greatest invertebrate biomass genera lly occurs in the spring and ··fall during peaks of growth. The abundance of organisms vith short life cycles v~ies considerably wi t .h environmental conditions (Patrick 1962). If they reach maturity by fall, additio.nal eggs &re deposited which undergo development during the winter months (Cummins 1973). Some species, however, require a full year of development to reach . ·matut'ity. 135. ......... . .......... . Environmental stress. .. ... .. ... ~. Benthic stream macroinvertebrates ·. . ~· ·have traditionally been good indicators of past a.nd present environ- mental str ess because of their long life cycles and relatively sedentary behavior. Changes in c011111unity structure are sensitive indicators of environmental alterations (Cairns and Dickson 1971). The most sensitive species are eliminated from the stream, resulting in diminished compe- . titian among ·the surviving organisms. Only those species able to tolerate a vi.de range or environmental conditions are able to survive when l.ife requi.rements become limiting (Bradt 1977). 45 ..~ : 136. Short-term expo,sure of tbe stream COIIIIIlWli ty to intolerable conditions may result. in alteration of the diversity and density of the fauna. More tolerant species increase in number, because of the lack of competition, until they reac.h their l .imits of space and food (Cairns and Dickson 1971}. Additional alteration of an already stressed environ- ment. will eliminate one or more of the remaining species, resulting in a major reduction in the standing crop. Tai~vaters 137. Invertebrate living conditions in a tailvater are different from those in a natural stream and are dependent prim~rily on the characteristics of the reservoir discharge. To survive, benthic organisms must be able to adapt to the changes in primary production arid the altered physical-chemical characteristics of the tailvater system (Krumholz and Neff 1970; Jonassen 1975}. 138. The effects of reservoir releases on the downs·tream biota depend on the type of dam and the subsequent flov patterns, release depths, and resiliency of the natural st.ream benthos. Macroinverte- brate species composition and diversity may either be substantially ··enhanced or reduced, depending on the characteristics of the flow in the tailvater (Ward 1976a}. The benthos belov reservoirs generally respond to the unnatural conditions vi t .h reduced taxa and increased numbers· of certain species (Spence and Hynes 197la}. Tailvater in- sects e.:re ma.l.ler and are considered to be of marginal food value for fish (Powell 1958; Bauer 1976). The number of species increases progressively downstream in response to the greater availability and variety of microhabitats and the presence of increased quanti tie.s of detritus from streamside vegetation and runoff (Hooper 1973; McGary and Harp 1973}. However, total numbers of individuals may be highest near the dam, since systems vi th fev species often are mo.re productive ~~d support a greater biomass than those vith many species. 139. Fe.ctors vhich inhibit be.nthic populations in tailvaters in- clude alterations in natural yearly temperature changes, isothermal temperatures, siltation (in some flood-control sites), daily water-level fluctuations, streambed sc.ouring, reduced concentrat ·on of particulate 46 • ~. c.,.ganic matter, altered vater quality, and seasonally altered flovs {Vanicek 1967; Hoffman and Kilambi 1970; Is om 1971; Ward 1976b). Con-\ versely, stabilized novs, decreased turbidities, the introduction of' seston from the reservoir, increased nutrient availability, and the grovth of algae and moss increase benthic standin~ crops. of the inhi.biting and beneficial effects depends on the schedule of vater release, the vithdraval depth, and the length of time water has b•!en retained in the reservoir. 140. Fluctuating flovs. High releases fol:oving periods of little or no flov result in scouring and turbidity, and ~luctuating vater levels cause increased bed and bank instability {Ward 1976a). Reduced flovs result in decreases in vetted perimeter, depth, surface a rc ea, and current velocity. Water temperatures during these periods become increasingly subject to ambient atmospheric influences {Pfitzer 1962; War d 1976a). The extreme changes in flov often create conditions that are unsuitable for most stream benthos. Because of this, invertebrates are least abWldant in inmediate tailvater areas that are subject to extreme periodic f'lov fluctuations . Insect densities in the ta.ilwater may be as much as 30 times less than those of streams floving into the reservoir {Povell 1958; Trotzky 1971). Hovever, dovnstream moderation of the effects of flov fluctuatio:1 often results in the gradual in- crease of invertebrate densities {Radford and Hartland-Rove 1971; Trotzky and Gregory 1974; Ward 1976b). 141. Continually fluctuating flovs interfere vith the establish- ment of a stable benthic community because of the preference of various species for a narrover range of environmental conditions {Pearson et al. 1968). Daily vater level fluctuations generally reduce the production and standing crop of stream invertebrates by eliminating both the benthic food base and the benthos (Bovee 1975; Ward 1976a). This situation is particularly obvious belov hydropover projects vhere maximtun discharges occur during periods of peak pover demand and minimum discharges vhen pover demands are lessened. In these instances, tvo enti.rely different stream habitats are cre&ted. T.he tailvater may change from a typical pool-riffle association during mini.mum releases 4'j . ,. I to a deep, swift stream during maximum rel.eases. Most affected is the "zone of fluctuation," which is composed of s ,fde channels, backwater areas, and shallows. These areas alternately undergo the physical disruption of microhabitats during high flows and dewatering and sub- strate expoSU!"e during reduced flows. Insects in these areas beccme disl::"~ciged and physically destroyed during high flows, and they are also subject to stranding and desiccation during reduced flows (Powell 1958; Trotzky 1971). In tail waters vi th fluctuating flows, the lack of permanent, clearly defined pools and riffles precludes the survival of most stream insects . 142. Some benthic tailwater communities subject to regular, peri- odic water-level fluctuations may event.uall.y attain a resonable level of production ( Odum 1969). In these "mature" tail waters • a few s.pecies (probably two or three) that have adapted to the flow changes make up the vast majority of the benthic community (Pfitzer 1962). Members of these benthic communities apparently tolerate brief periods of sub- strate exposure if it is not severe (Fisher and LaVoy 1972; Ward 19'i6a). No significant difference was found between numbe·rs of insects li ring on nonexposed substrates and those living on oocasional_ly exposed substrates ( 13 percent exposure time) in a "zone of fluctuation" below a hydropower d.am on the Connecticut River. Massachusetts (Fisher and LaVoy 1972). 143. Vegetative ~ats act as a refuge for insects in some tail- waters during brief periods of exposure because of the retention of moisture in the vegetation. Insects living near the· "mat-rock" inter- face are more likely to survive than those· found on the surface of the vegetation (Brusven et al. 1974). 144. Low air temperatures also increase the chan.ces of survival for organisms stranded in a tailwater and enatle them to tolerate dewatering for longer periods than during high ~ir temperatures. Brusven et al. (1974) reported that larvae of chironomids, lepidop- terans, and trichopterans. could survive dewatering for 48 hours in cool weather, but that high air temperatur.es and longer exposure periods resulted in high mortalities . 48 I I ' I ' I • I ·~ ' 145. In spite of the ability of some insects to survive periodic exposure., most insect species that inhabit tail waters are found in the permanently submerged habitats not subject to daily exposure (Powell 1958; Brusven and Trihey 1978). 146. The lack of flow fluctuation in tailwaters below flood control reservoirs can disrupt the benthic community. In fact, stable releases for 3 to 4 week s followed by moderate or extreme discharges .may cause· mor e stre·ss and mortality to an inse·ct community than the frequent pulsed releases .found below hydropower dams (Brusven and Tribey 1978). The stable periods encourage the colonization of the ta:ilwater sub- strate·, but benthic organisms vhich are not able to adjust to subse- quent severe flow increases are catastrophically swept downstream. 147. The high seasonal discharges below flood control reservoirs may also destroy the pool-riffle areas nearest th~ dam and replace them with an extended "run" section. Farther downstream, however, pools a~d deep channels may be present which act as buffers to fluc- tuating water levels and as refuges for invertebratea during periods of minimal re·leases (Powell 1958). 148. Chironomids are the most resilient group of insects founC: in unstable areas and are the first to recolonize denuded zones of fluc- tuation (MacPhee and Brusven 1976). They may be found in association with oligochaetes, amphipods, and isopods in coldwater tailwaters subject to rapid water-level fluctuations (Brown et al. 1968). These invertebrate associations are able to adapt to temporary periods of exposure by either migrating out of the exposed area or surviving in the thin layer of water which remains after the stream recedes . 149. Insects most affected by fluctuations i 'n flow are the groups · which are generally regarded as quality fish food, including mayflies, stoneflies, and caddisflies (Powell 1958; MacPhee and Brusven 1976). Nymphs and larvae of these species are most subject to desiccation, ·.·· since their eggs can usually survive extended periods of exposure. The number of insects affected by desiccation, therefore, depends on when a flow reduction occurs and the li:fe stages of the insects present. Desiccation of the streambed would affect the population drastically during a. larval hatch, but unhatched eggs may remain relatively un- affected (Hynes 1958). 150. Stabilized flows. Reservoirs vith stable releases have relatively stable tailwa.ter substrates (Ward 1976a.). The associated invertebrate fauna. stabilizes as flows and substrates become more con- sistent, resulting in reduced niche ove,rla.p and a shif't toward community equili br:!. um ( Blanz et a.l. 1970; Ward 1976c). Stree.m:flow stability com- bined vith reduced turbidity may enhance algal and macrophytic growth and provide additional food niches and microhabitat diversification for chironooids, oligochaetes, and mollusks (Armitage 1976; Ward 1976a). The stable environment and predictable .resources tend to eliminate some species, resulting in a less diverse faunal assemblage with higher standing crops of the species present (Ward 1976b). 151. In tailwaters subjected to stable seasonal discharges, sedi- mentation may become limiting to the benthic community. Extended periods of minimal releases accompanied by moderate detrital and silt input from rur.off and streamside vegetation effectively .reduce the av.a.ilable microhabitats. The lack of spates, necessary to flush the detrital material from the substrate interstices, may event'UI.llly result in the complete elimination of these productive invertebrate micro- habitats (Giger 1973). 152. Deen-release reservoirs. Cold water temperatures and poor water quality (low dissolved oxygen, reduced compounds) often occur in tai·lwat.ers below deep-release dams. Stratified deep-release reservod:rs typically produce benthic tailwater communities which are low in diversity, but which may have high standing crops. This situation i.s typical of a stressed environment (Ward 1974, 1976c; Young et al. 1976; Pearson et al. 1968). 153. Modification of the normal temperature regime can affect the diversity and quantity of the· benthic fauna. several kilometres down- stream from a. coldwater release dam (Pearson et al. 1968; .Lehmkuhl 1972, 1979; Ward l976c). The overall change in temperatures, coupled with the delay of seasonal fluctuations, often .results in the elimina- tion of many invertebrate species from tbe tailwaters. Minimum winter 50 i ... ·. ~·· .. ; ! .... .. ·•. • •' and maximum summer temperatures that would normally provide the thermal stimulus essential for the initiation of various life history stages of many stream invertebrates are never reached. Reduced growth efficien- cies a.t the lowered temperature.s may eliminate species which are unable to adapt metabolically to abnormally cold summer temperat';.U'es (Hannan and Young 1974). Alternatively, warm winter temperatures may accelerate growth rates and result in premature emergence and exposure to air temperatures that may be lethal or tha.t may complicate the mating pro- cess (Ward 1976c). These conditions, along with delays in spring warming and autumn cooling, may prevent the natural hatching and grovth of insects that have stringent thermal requirements (Lehmkuhll972). ..- 154. The lack of daily fluctuations in vater temperature may prevent the initiation of egg or larval development (Ward 1976c). Diel temperat.ure fluctuations in a 24-hour period were as great as l0°C ·in······ an unregulated stream in England, but rarely exceeded 1°C in a regu- lated section (Armitage 1977) . . 155. Stresses imposed on aquatic invertebrates below deep-release reservoirs may also be due to low dissolved oxygen, ·the presence or · high concentrations of' H2S and other decomposition by-products, o.r the · interaction of these factors (Coutant 1962; Hicks 1964; Young et al. 1976). Reaeration of the vater as it passes through a dam may increase the concentra.tion of dissolved oxygen to a level tolerable by the tail- water biota, but the by-products (e.g., iron, manganese, hydrogen sulfide) of hypolimnetic decomposition may be more persi stent and ha.•re a detrimental effect on t~ilvater organisms (Coutant 1962). If re- leases from the r-eservoir are increased, the harmful effects of the hypolimnetic discharges can be exte.nded downstream. r .. -.: •• 156. Deep-release reservoirs do not necessarily release water of poor quality (low or no dissolved oxygen, reduced compounds) but assuredly produce a new envi ronment which favors coldwater rather than warmwater organisms (Hoft'ma.n and Kilambi 1970). In st1•atified reser- voirs where anoxic conditions and pnor water quality do not develop in the hypolimnion, increas~d concentrations of dissolved nutrients, ) ·co2 , and lowered turbidities may favorably affect th~ living conditions 51 of tailvater invertebrates (Penaz et al . 1968). In these circumstances, the benthic community may have characteris.tics similar to those of communities in streams vith mild organic enrichment (Spence and Hyne~ 197la). • 157. Filterable food ma terial (plankton, some benthic organisms, and miscellaneous seston) produced in deep-release res~~oirs may be passed into the tailvaters during periods of' complete vertical mixing. However, this is not a reliable food source since the plankton and seston are congregated in the upper strata of most r~servoirs during stratification and are therefore unava.ilable to the discharge. As a result, filter-feeding invertebrates are often restricted to dovnstr.eam tailva~er locations, where discharge effects have been modified and s.ufficient amounts of filterable material have been int.roduced through runoff and tributary inflow (Ward 1976b). Plankton and benthos may be discharged from some deep-release reservoirs during stratification if the hypolimnion remains oxygenated. 158. Surface-release reservoirs. The downst.ream effects of a surface-release r;s~noir are usually similar to those produced by a .·~·~aturai surface-release lake (Ward and Stanfo~d 1979). Water quality is generally not a problem and tailvater temperatures may only be moderately influenced in comparision to deep-release sites. 159. Shallow, surface-release reser~oirs can produce abundant quantities of bottom fau."la and plankton. Invertebrates in the tailvater may subsequently receive much of their food. in the form of seston and i~sect.s from the ·reservoir discharge (Walburg et al. 1971). Because of the rich suspension of food in the rese.rvoir release, filter-feeding insects, particularly trichopterans, may be extremely abundant in the tailwater. Generally, the increase jn benthic density is accompanied by a reduced diversity and the elimination of some species. In formerly coldwater streams, the release of warm surface water from the reservoir during summer may result in a reduction or elimination of certain spE"cies frol!l the tailwater (Fraley 1978; Ward and. Short 1978). 52 · ... • •• ,•·:- •• Benthic Invertebrate Drift Streams 160. Interspecific competition for food a.nd microhabitats may result in an active downstream movement or "drift" by many benthic organisms (Waters 1969; Hildebrand 1974). The benthic invertebrate fauna and the drift i ng fauna are not distinct communi ties. Dr'ift is merely a temporary e'Vent in the l.ife cycle of most benthic organisms ('Waters 1972). 161. Benthic organisms enter the drift vhen they leave their pro- tective retreats and are swept downstream by the current. This process generally occurs a:t night during periods of fee.ding activity (Waters · 1969). Th~ feeding activity of each species is governed by a diel pattern based. on photope.riod and is generally initiated at the. same time each day (Elliott 1967; Waters 1972). The actual movement · downstream probably occurs as a series of short hops vith the turbulent flov assuring frequent contact vi th the substrate. The distance of passage depends primarily on velocity of flov and the species of inver-. tebrate involved (Gore 1977). 162. When flovs are suddenly increased, the physical disturba.nce ·of the streambed stimulates a catastrophic-like .response from the benthos and their p-re ~ence in the drift increases markedly (Elliott 1971; Brusven et al. 1976). The actual nwnber of organisms in the drift per unit volume remains relatively stable during these periods, but the ntunbers passing a certain point in the stream ove.r . a period. of ., .. time may increase significantly (Waters 1969). Natural spates in streams may disperse organisms over an increased living area or may displace them into stream environments such as pools and runs, they find survival difficult and vnere they are more likely to S\.une.d (Waters 1969). Many benthic organisms die vhen they are downstream into physically or chemically unacceptable z o nes. (Russell-Hunter 1970). vhere be con- svept 163. The drift may also be used as a means of escape from desiccation during low flows (Elliott 1971). Increases in the drift 53 .... ~ •,. --~· .... result from a decline in available living space associated with drastic reductions in streamflow (Minshall and Winger 1968; Armitage 1977; Gore . 1977). Areas dovnstream, relatively unaffected by reduced flows, may actually exhibit increased invertebrate populations as ~ result of the accumulation or organisms from devatered riffle areas (Giger 1973). 164. Recovery of the bottom fauna a~ter streambed exposure or other catastrophic. events may be rapid ( 19-28 days) after natural con- ditions return (Herricks and Cairns 1974-76). Drifting organisms from upstream areas quickly recolonize the affected streambed. Rec.oloni za- tion of denuded areas may also be aided by a certain amount of upstream "crawling" movement, which may· replace about 6 percent (by number) of the dovnstream loss due to drift (Bishop and Hynes 1969) • The renewal ·of drift out of a disrupted area may be delayed until the conmunity has recovered to a point where the benthic population exceeds the carrying capacity of the habitat (Dimond 1967). The failure of a formerly abundant species to repopulate an area after a stream. distur- :~·" bance may allow other species to become established, ultimately result- ing in a shift in community composition and abundance (Waters 1964) • Ta1lvaters . _ .. ~~;.~--~ 165. Drift in tailwaters is primarily compose.d of organisms from the reservoir above, particularly below unstratified r ·eservoirs and reservoirs with surface-release systems. Microcrustaceans and certain i~sect larvae (e.g., Chaoborus) produced in the reservoir commonly dominate the teilvater drift (Gibson and Galbraith 1975; Armitage 1977, 1978). 166. Drift of' benthic organisms that live in the tailva.ter occurs primarily as a result of extreme fL.tctuations in reservoir discharge. Even species not normally found in the drift (e.g., Chironomidae) may enter the drift because of these .fluctuations (Brooker and Hemsworth 1978). Increased drift may result in considerable reductions in the abundance of the stream faun.'!L vi thout actual streambed desiccation (\Tard 1976a). Repopulation of denuded zones is obviously not possible from sources upstream, because of the presence of the reservoir. Re- colonization is therefore dependent on several fa.ctors, including egg • • .,. . .. . . . ·~-..._. .. ~ -:'. ..... ~ .. .· deposition by adult insects which fly upstream, random oviposition of remai·ning individuals in the denuded zones, and upstream crawling or svimming by nonaerial invertebrates. Eggs deposited by insect species which are tolerant to the particular tailvater environment vill develop. . . Other species, which are not adapted to the specialized conditions, will be eliminated. 167. Minimal daily fluctuations and seasonally stable flows may result in inc.reased benthic populations (Ward 1976a). However, the stable flows that enhance benthic invertebrate development may reduce drift. Reduced drift in this instance may be detrimental to fish pro- duction by limiting the availability of prey (Chapman 1966). . ~. . .. 168. Terrestrial insects may make a significant contri.buti~n _to · available fish food in the taUvater drift; especially in 8.reS:s where streamside vegetation is abundant (Waters 1964). They may be attracted to the cool, moist areas of exposed streambeds below reservoirs ~uricg periods of reduced flows, and vhen discharges are suddenly increased. they are swept dovnstre9m. vi th the drift (Pfitzer 1962). Since .\. •""'"":"Y1•: ~ • · terrestrial insects-are. most active during daylight, they are mor~ · .. : c.ommonl.y found in the drift during the day -than at night (McClain 1976) •· ZooPlankton 169. River a:nd stream environments ·are poorly suited for the pro- duction and maintenance of zooplankton, and they characteristically harbor small zooplankton populations (Hynes 1970). Their abundance in ,.,_flowing waters is inversely related to the rate of the current. flow. Thus, the zooplankton present must be supplied to the stream trom adjacent quiet-water areas or other suitable productive habitats in t 'he drainage (Hynes 1970). 170. Most zooplankton found in tailvaters are produced in the upstream reservoir. Some zooplankton may be produced in backwaters and other quiet-water zones adjacent to the tailvater, but the input from .. these areas is not significant when compared with that of the reservoir. Therefore, the species and abundance of zooplankton found in the 55 ,. tailwater are dependent on the species and abundance tn the reservoir population (Brook and Woodward 1956). 171. The reservoir zooplankton community may vary as a result of seasonal population cycles . The community is also influenced to a large degree by the hydraulic residence time of the reservoir. Zooplankton nwnbers are generally higher in reservoirs with longer hydraulic. resi- dence time.s·. Both of these factors eventually affect tailvater zoo- plankton abundance. 172. The amount of zooplankton passed into a tailw~ter depends on the depth of reservoir release. The zooplankton migrates vertically within the vater col;.ann in response to changes in light intensity. This vertical migration may keep the zooplankton away from the level of discharge during certain periods of the day. Zooplankton abundance will also be altered in tailwaters belo'lt' selective withdrawal d.ams, vhere changes in the level of -~thdrawal are made on a seasonal or daily basis. 173. Zooplankton transported into a tail water f .rom the ·reservoir provides a more readily available source of . energy a:nd protein than the ·'· detri't~s normally found in. unregulated st~eam's (AnDita.ge 1978). Most of the zooplankton discharged from hypolimnetic-release reservoirs is already dead end simply contributes to the stream's load of organic debris. Moribund zooplankters settle out and decompose, providing a nutrient-r;i.ch d~tritus in the tailwater area (Armitage and Capper 1976). Seasonal inconsistencies in the reservoir discharge, hovever, preclude ;o.oplQ.l)kton from being a .reliable source of either nutrient inputs or food for benthic ~omswnption in the teilva·.'~er (Ward 1975). 174. Le.rger bodied :;,;ooplankters (e.g., copepods and d·aphnid cladocerans) may be coiiiillon in tailvaters immediately belolt surface- .release reservoirs, but they rapidly disappear as the water flows downst.ream. The progressive reduc.tion in densities downstream is characteristic and has been docmnented by several invest igators (Chandler 1937; Stober 1963; Ward 1975; Armitage and Capper 1976). 175. The zooplankton decrease vhich occurs between reservoir outlets and areas downstream is due to a combination of factors, 56 ·~ .. including the abundance of zooplankton discharged , the filterir~ effects of periphytic vegetation in the tailwater, p hysical destruc t ion, preC::Ltion, and adherence to or ingestion of silt and debr i s . Chandler ( 1937) indicated that zooplankton discharged fr~m a resei"!Foir in July was ·reduced 99 percent 8 km downstream; whereas, during February, when population levels were higher, abundance vas reduced only 40 percent. Algae and mosses in the tailwater essentially act as filters, r emoving the zooplankton as it flows through the vegetation (Chandler 1937). Armitage and Capper (1976) noted that 99 percent o f the zc oplankton discharged had disappeared within the first few kilometres, the greatest losses occurring in the first 400 m below t he dam . Larger ·-. organisms become entangled more easily and are eliminated from the streamflow sooner than smaller organisms. For this reason, greates t zooplankton reductions in tailwaters may occur when aquat i c vegetation i s abundant and the tailvat er levels are l o west. When water levels in the tailwater are high aDd aquatic vegetation is absent, the v e get a ti ve fi .ltering phenomenon is eliminated, and zooplankton may be more per- •• ~ •• 0 . sistent downstream (C~dler 1937). :-;:,';:_'· ~'-"~· ~ 176. Zooplankton is also highly susceptible to physical abrasion and-fragmentation. Some zooplankters a r e ut ilized as prey when f lushed from the reservoir, benefiting populat i ons of bent hic macroinvert ebrat es and fish in the tailwater (Ward 1975). Ot her zooplankters eith er ingest or adhere to sand and s i lt in the turbulent tailwaters and become heavier, tending to sink and die. Tailvater Effects on Snecific Invertebrate Taxa 177. The productivity and occurr ence of stream organisms in tailvaters are determined by physical-chemical stresses imposed upon the community. These stresses are related to the quantity and quali.ty of the releases from the re:1ervoir. Species normally present in a stream may be enhanced or reduced in a tailwater, but t he structure of the tailwater comnunity is generally much different from that of a natural stream. ., 178. Immature life, stages of insects from four orders, including Diptera, Trichoptera , Ephemeroptera, and Plecoptera, are prevalent ·in natural streams. Other aquatic insects, mollusks, benthic crustaceans, oligochaetes, and planktonic invertebrates may also be abundant. Di 'Dte!"a 179. !n tailwaters, dipterans appear to be the group best adapted to the al te!"ed conditions of reservoir releases (Ward. 1976c). Dense communi ties of Simuliidae and Chironomidae often occur in the immediate tailwater bel0w deep-release reservoirs becau.se major invertebrate pre- dators are few and fish di~ersities are low (Ward 1976c). They may be the only invertebrates present because of their ability to adapt to conditions in most cold tailwaters. Farther downstream in the tail water elevated water temperatures, predation, and mic·cohabi tat competition may result in reduced densities of both famil.ies and the possible disappearance of Simuliidae . 180. Simuliid larvae and pupae prefer cold water and are appar- ently tolerant of the poor water quality which occasionally occurs in tail~ters below dee.p-release dams (Hilsenhoff 1971; Goodno 1975). .•-' · .. They are.· not common below surface-release dSllls or' in ·tailwaters vhere <: water temperatures may become too high for their survival. The more diverse .fauna which exists farther downs t ream from these dams addi- tionally limits t'he establishment of simuliid populations because of increase.d invertebrate competition and predation . 181. · The various species in the fam:i,ly Chironomi.dae are .adapte~ to a •'ide range of environmental conditions, making them ideal tailwater inhabitants. Some prefer cold water, while others prefer warm water. Other species have more generalized requirements and may be found in a variety of conditions. The seasonally altered tempera- tures found in tailwaters below deep-release dams (winter warm, summ.er cool, and delayed maximtml and minimum temperatures) may a l low some species o f chi·ronomid larvae to survive during periods of the year when · they would not be found in natural streams. However, these same alterations may modify emergence patterns, which are controlled by water temperature and light intensity (Oliver 1971). Adults in t hese 58 -----------------------------------------~ ·. .. -· .. . ' ..... .. ·; :.·· .. · . • . ' .. .' instances me,y undergo premature emergence ~esulting in disorientation and reduced survival. Tric.hoptera 182. Most. species of Trichoptera found in tailwaters are net spinners or filter feeders. Their abundance is directly relat ed to the availability of seston discharged fl'om the ,reservoir which, in turn, is dependent. on the location of reservoir outflow (Rhame and Stewart 1976; Ward l976a). Deep-release reservoirs provide an extremely unreliable food source because of the lack of seston during periods of stratifica- tion. Unstratified and surface-release reservoirs may, in contrast, provide a rich food source, including plankton and other suspended organic matter. 183. The success of a certain species may depend on the si~e of ·· the food particles that are available. Individual species crop parti.- cles of specific sizes, depending on the mesh size of their gathering apparatus and their location in the streambed (Wallace e~ al. 1977). 184. Variations in flow alsc affect trichopteran survival. Ade- quate current v~locities are necessary to supply rood and oxygen to stat.ionary larvae and influence the design and construction ot food-gath~ing nets (Haddock 1977; Wallace et al. 1977). Sufficient ·· flow is required to keep :food-gathering nets extended!, but higher flows will result in their destruction (Haddock 1977). Large flow varbtions below s0111e dams may inhibit the survival and feeding success o 'f .some trichopterans, since food-gathering nets are swept away at high tlo~s and collapse at low flows. .. .......... , •u '•" 185. Larvae of some species of Trichoptera a:re found in. - quiet-water areas of natural streams. They are unable to vi thsta.nd swift water and are not dependent on net-gathering mechanisms for their food supply. Quiet-water areas are uncommon below most reser- voirs and if pre.sent they a:re subject to destructive periodic flow increases. As a result, most trichopterans found in tailvate'rs are adapted to living in svitt water. , . 59 ·'·· . . ~ .... .. Eoheme::-optera 186. Ephemeropte:::-~ {mayflie.s) typicaJ.ly inhabit small streams vi tb large-rubble substrates. The number of species present depends pri- marily on the habitat diversity, since each species exhib'its a high degree of habitat selectivity. Generally, the presence of more diverse habitats is reflected by an increase in the number of species (Macan 1957) •. 187. Ephemeropterans do not usually occur in ta.ilwaters be.cause the stable thermal regime found here does not provide ~he temperature stimulus required for life-stage development. They are additionally affected by the reduced habitat. diversity and the. abse~~e of. ai:i aae.:.. quate food supply (detritus). Downstream, where the effects. of reser- voir releases subside, mayflies gradually become more prevalent as the tailwater becomes more "stream-like" (Lehmkuhl 1972). 188. Ephemeropterans are rarely found in streams where velocities are below 15 em/sec or above 91 em/sec (Delisle and Eliason 1961). Fev species are able to survive in areas subject to extreme flow .reduction's , ., .(MacPhee and Brusven 1973; Ward 1976a) •. Mayfly species equipped vito .... ., ... hold-fa.st organs are able to exis·t in tailwaters vi th :noderately high current velocities (Ward 1976a). 189. Most e~hemeropterans do not readily colonize areas in a constant •state of vater-level ·fluctuation. since they cannot tolerate both lov and high flov extremes. One genus (Paraleptophlebia), however, bas been found abundantly in t .ailvat.ers exhibiting t 'hese types of fluctuations (Trotzky and Gregory 1974). Pleco-otera 190. Plecopterans are usually not found in tailvaters because · the lo•ss of habitat heterogeneity, changes in flov regime, and re- latively stable temperatures make conditions unacceptable for their grovth and development. Downstream their abundance gene.rally increases as the influence of the reservoir discharge declines (Ward 1976a). Several species of plecopterans (e.g., chloroperlids) are not affected by rapidly fluctuating flows, and may become abundant in tailvaters 60 . . where other environmental condi tions are suitable {Ward anq Stanford 1979). 191. Plecopterans are common predators on trichopteran eggs, cbironomid larvae, and simuliid larvae in natural streams (V,aught and Stewart 1974). The absence o~ these predatory species. contributes to the increased abundance of Simuliidae and Chironomidae in most cold tailvaters. Misce~laneous 192. Amphipods ., oligochaetes, i sopc.ds, mollusks, and turbellarians a:re often abundant. in tail waters. One fE•ctor common to these groups that may be significant to their abundanee in tailvaters is the lack of an aerial adult stage in their life cyclt~s. Becau.se they do not have an aerial adult stage, they are not subj·~ct to the problem of :pTemature emergence that often occurs among aquati·~ insects in thermally e.ltered tailvaters (Ward and Stanford 1979). 193. Ward and Stanford (1979) indi '!ated that amphipods arE often abundant in tailvaters that have reduced .summer temperatures anc . stable . . ·flow regimes. High nutrient inputs and '!"educed flood flovs alsc• favor amphi'pods because st-reambed siltation an•i increased macrophytic growth vhi.ch occurs in the:;e situations are hig:lly beneficial to thei.r develop- ment (Hilsenhoff 1971). 194. Oligochaetes may be present b•!lov deep-release· reserYoirs in pools avay from the strongest currents. T'he cool, nutrient-rich water and lack of destructive spates favor estt~blishment of dense populations in these areas (McGary and Harp 1973; Armitage 1976). 195. Isopods may be the· dominant organism in the riffle areas of cold tailwaters (McGary and Harp 1973). They are also less a!f~!Cted by minor vater-level fluctuations than other invertebrates because· they can migrate out of areas that are periodi ·::ally exposed. 196. The distribution of mollusks in natural st-reams is d~!ter­ mined primarily by substrate patterns and type~. The chemical nnd physical alterations that occur in tailwater environments have both enhanced and disrupted mollusk populat.ions. 61 197. .Enriched growths of attached algae and increased organic sediment were found to encourage the establishment of pulmona:te snails (Physa} during the absence of scouring releases or during extended. periods of reduced flows (Williams and Winget 1979). However. Harman (1972} found that chemical and biological alterations of tbe tailwater environment negatively affected t~e mollusk population and reduced sp~cies diversity. The number of mussel species was also reduced in the Tennessee River. Alterations to and loss of riverine habitat after extensive reservoir construction reduced the number of species from 100 to approximately 50 (Isom 1971). Formerly abundant species were re- ¢uced or eliminated and natural replacement was limited. The change in fish species that also occurred as a result of reservoir construetion further reduced mollusk populations through the ·obstruction of f'i:sh-host associations that are a necessary part of the molluskan life cycle ( Isom 1971) . 198. Turbellarians appear to be a minor member of the tailwater invertebrate community and are not widely studied. Based on the few studies tha:t .have been conductedt turbellarians apparently increased :in.tailwaters with stable flow regimes and cool summer tempera~ures · and declined in tailwaters with fluctuating flows (Ward and Stanford 1979). 199. Crayfish are common in some tailw&ters; however, no mention of this group was made in the li ter.ature except as .food for some fish s .pecies. 62 • ':.., .. 'l.-; • • 'j • ••.• ··.· ... .. ·.· PART VII: FISHES Ili TAIL'WATERS 200. In this section, fishes that commonly occur in tailwaters are discussed. The presentation i.s by family, followed by individual s~cies or groups. Fish species which occur in tailvaters but 'are of only mi.nor importance because few are generally captured or they are little mentioned in tailvater literature are not discussed. These include species from the folloviog families.: Petromyzontidae (lam- preys); Acipenseridae (sturgeons); Lepisosteidae (gars); Anguillidae (eels); Hiodontidae (mooneyes); Aphredoderidae (pirate perches); Cyprinodontidae (killifishes); and Atherini.dae ( silve·rsides). 201. A brief description. of enviromnental conditions nece.ssary for the successful completion of the various life bistory phases of eac.h species or group is presented under the following topics: habitat, reproduction, .food, and age and grovth. This is followed by a synthesis of tailvater literature pertaining to a particular species o~ group of fish. Emphasis is placed on how the species responds to .. · ·· · environmental conditions found in tailvaters· •. The depth of discU.ssi'on ..• :;-:·=~:.~. ~ .. varies by species and is dependent on the amount Of literature avail-' : able. Source material for the descri.ption of f'ish life history require- ments vas obtained fro111 Carlander (1969, 1977), Scott and Crossman (1973), Pflieger (1975}, and others. 202. The common and scientific names of fishes mentioned in this -report are listed in Appendix C . Fishes from North American tailvaters a.re· included in Part I and nomenclature follows that or Bail~ et al. (1970). Fishes from European tailvaters are included in Part II. Life history information for the most common fi.sh groups discussed in this report is' given in Appendix D. ..... Polyodontid&e (PaC.dlefishes) .. · • ..... 203. The paddlefish is one of the largest freshwater fish in North America. It is found only in the Mis-sissippi, Missouri, and Ohio rivers ,and their larger tributaries. Numbers of paddlefish have •• 1 declined greatly since the advent of the twentieth century. Major causes are believed to be dams,. over fishing, and pollution. Fish con- centrate below dams vhere they are especially vulnerable to fishing. Dams built to create reservoirs or for other pur·poses have inundated many former s ,pavning grounds or have prevented fish from reaching upstream spawning areas. The dec~ine of paddlefish stocks in past years followed the increased rele~se of domestic and indu~trial pollu- tion into the waterways. Paddlefish populations have increased in some areas where pollution abatement has been ef'fective (Eddy and Underbill 1974). Paddlefish 204. Habitat. The paddlefi sh is primarily found in the. open vater of sluggish pools and backwaters of large rivers where it svims about continuously near the surface or in shallow water. For spawning, it requires access to a large, free-flowing river vitb gravel bars vhi.ch are inundated during spring floods. 205. Orig.inally the large. free-flowing rivers of the M:tssissippi .;~-.. Valley provided ideal habitat for paddlefish. Nov some of the largest populations are found in man-made impoundments where tributary riTers .meet the fish's exacting spawning requirements. These conditions are met in some reservoirs in the states of Alabama, Arkansas, Kentucky, Missouri, OklabCIIIB., and Tennessee, and in several mainstream M:is.souri River reservoirs in North and. South Dakota and Montana. · ·206:· Reproduction. The reproductive habtts of paddlefish ·were descrioed by Purkett (1961), whose studies were conducted on tbe Osage River in Missouri . Spawning takes place in midstream, over sul:merged gravel-bars vhen the river is high and muddy in early spring at temper- atures of about 15.5°C. Meyer and Stevenson (1962) reported that female paddlefisb in Arkansas do not mature until they are over 11.34 kg in weight and that they may not spawn every year. The adbesi ve eggs stick to the first object they touch, normally stones on the streambottom. Eggs hatch in 9 to 12 days vben water temperat.ures are about 16°C (Purkett 1963; Needham .1965). Afier hatching, the. fry svim upward vigorously, then settle toward the bottcn. Frequent repetition 64 ~ .... of this activity by the fry is significant in that it permits the strong currents to swee·p the fry downstream out of the shallows and into deep pools before the gravel bars are exposed by receding vater levels. 207. Food. Paddlefish feed primarily on ~ooplankton and insect .larvae filtered from the water. They svim slowLy with their mouths open through areas whel"'e food is concentrated. Water passed through is filtered by the long, closely set gill rakers. The function of the paddle-shaped snout in relation to feeding is not knovn for certain, but its elaborate system or sense organs may enable it to function primarily as a device for locating concentrations of food organisms. 208. Age and growth . Paddle.fish grow rapidly. According to .. Pflieger (1975), yoUng nearly 150 ·mm :'tong have been collected from overflow pools of the Missouri River in early July. Two specimens kept in a fertilized pond reached a length of about 0.9 m and a veight of 2. 7 kg when they vere 17 months old. Paddlef'i sh in Lake of the OzP..rks, Missouri, attain a length of 250 to 350 mm in their .first year ~:• • .,., "• ·.;, • • :::0 "" " •· ''•. .....( ·~~···· •1 I ~1d about 530 mm in their second year. Seventeen-year-old fish average ne~ly 1. 5 m in length ana 16.8 kg in we.ight. The ·largest paddiefish are usually females. The species is also long-lived; many individuals live more than 20 years. Paddlefish in tailvaters 209. Large concentrations of paddlefish are found in some tailvaters es,pecially during winter and spring. Most studies on · · paddlefish in tailwaters are concerned vith spawning and reproduction.' ········ Paddlefish reproduction, prevented primarily by the blockage ~f lloPStream migration by dams, has also been affected by the altered flow regimes found in tail waters. Main-stem hydropov--er facilities on the Missouri River have not only reduced the amount of natural spavning - are.a available but have. also rendered most of the remaining river areas unusable. Large diel water fluctuations mask or delay the normal spring r .ise in water flows and temperatures. 210. Young-of-the-year fish have been reported in only one tailvater of the Missouri River reservoir system (Friberg 1974). Juvenile·s found 1:-. Levis and Clark Lake were produced in the 65-km section of the Mi s souri River above the reservoir, while those in t 'he tailwater either passed through Gavins Point Dam or were produced in the 97-lan section of natural river downstream from the dam (Walburg 1971; Friberg 1974). Recruitment from the reservoir above is believed to be the primary source for paddlefish found below Lake of the Ozark's hydropover dam on the Osage River in Missl')uri (Hanson 1977). 211. ·The effects of navigation or flood control dams on paddlefish reproduction are not as wel.l defined. Successful spawning on. eroded ving dikes below Lock and Dam Number 12 on the Missl~sippi River in· Iowa has been regularly observed (Gengerke 1978). Eroded ving dikes effectively simulate natural gravel bars. The control of spring . floodwaters below a Kentucky flood control reservoir appears. to inhibit downstream spawning activity (Branson 1977). 212. An alteration of the behavioral characteristics of paddle fish caused by impoundments is evidenced by their concentration in the svift-floving waters immediately below many dams (Friberg 1972; Boehmer 1973; Gengerke 1978f~ Concentration of these :fish into a relatively confined area increases their vu.lnerability to both commercial and sport ~ishing. Friberg (1974) reported high catch rates of paddleti.sh in tailvaters for three to four years following dam.closure, folloved by declining catches due to reduced numbers. These concentr·ations dur- ing the reproductive season may be explained by the blockage of upstream migration. During other seasons these concentrations are mo.st likely due to the increased availability of food i .n the. reservoir discharge·. 213. In tailwaters, zooplankton is most abundant in the reservoir discharge; :its abundance decreases rapidly downstream. The large quantity of food available would tend to attract feeding paddlefish from slow-flowing areas into the faster floving discharge, where they would normally not be found. 214. Except for the immediate discharge, there is general.ly less zooplankton available in the tailwater than in the reservoir. This relative scarcity of food is reflected in the reduced growth rates of tailwater paddlefish. Growth studies by Friberg (1974) indicated that 66 ~ .. _ .. ·, . .... . . . ·· ... paddlefish apparently raised in a ·reservoir suffered a·marked reduction in grovth following passage through the d.am into the tailvaters. Clupeidae (Herrings) 215. The herring family is composed, of species which are ma~nly marine or anadromous. Several species live in fresh water and ~re occasionally' found in tailwaters. They are skipjack herring, gizzard shad, and threadi"in shad . None are used for human food, but the skip- jack is sometimes sought by sport. fishermen because it fights specta:- ularly when hooked. Gizzard shad, particularly young of the year, are important forage for other fishes•. Thread.fin shad are important forage at all ages because t ·heir maximum length rarely exceeds 180 mm. The skipjack is not discussed. further here because it is little ntentioned . in tailwater litera.ture, ex~ept. for an occasional occurrence in fish species lists. Shad 216. Habitat. The gizzard shad in genera.l..l.y distributed o.ver the . ' eastern halt of t.he United States where it .is most abund8Jlt in reser- voirs and large rivers. -It inhabits'quiet-water habitats ~n lakes, ponds,. reservoirs, and backwaters of streams where· fertility ~d pro- ductivity are high. Shad. usually avoid high-gradient streams and those which lack large, permanent pools. Shad congregate into loose aggrega- tions, and large numbers are often observed near the water surface • During tall, winter, and spring, large numbers may be found in tailwaters.- .217. The thread..fi·n shad, whose babi.tat is similar to that of the gizzard shad, is generally confined to the southeastern states where it has been stoc·ked extensively in reservoirs for forage. It has also been stocked in reservoirs of' the Southwest. It is sensitive to low tempera- tures, a .nd extensive die-oft's ha,ve been reported at temperatures below 7.2°C (Pflieger 1975). Because of overwinter die-off, annuaA stockings of a...dults are necessary to maintain populations in many reservoirs. Large .rv.Jmbers of this ·species may also occur in tailwaters during fall and winter. 67 '· L 218. Reoroauction. Gizzard shad are very prolific and generally spawn during April, May, and June at temperatures between 17 and 23°C in shallow areas of protected bays and inlete. The scat:tered eggs sink to the bottom where they adhere to the first object they contact. Eggs hatch in about_4 days. and the young begin feeding when 5 days old. Young at-tain typical adult form when abo'.lt 32 mm ll"lng. · 219. Thread!'in shad begin spawning i n the spring vben the water temperature reaches 21.1°C and may continue throughout much of the summer.· Threadfin spawn in schools near shore. The -adhesive eggs stick to any submerged object and 'hateli in about 3 days. Young begin 'feeding when 3 days old. Individuals hatched early in the year com- monly mature ana spawn late i n their first summer of life . 220. Food~ The gizzard shad is almost 'entirely herbivorous, feeding heavily on microscopic plants, phytoplankton, and algae. The species is essentially a filter feeder, removing particulate matter from the water by passing it through its closely set gill rakers. 221. Threadfin shad are also filter feeders. Their diet consists ot microscopic plants and animals found in the water ·column . . 222. · Age. and growth. Gizz ard shad average• about 127 mm long at the end of ·their firs-t summer, 185 mm at age I, 257 mm at age II, and 302 mm· at age III (Car lander 1969) • The average' life span is 4 to 6 years but s.om.e live 10 or more years. Maturity is reached in the second or third year. Adults are commonly 230 to 360 mm long and veigh about 0.45 kg. Maximum. length and weight is about 520 mm and 1.6 kg. 223. Adult threadfin shad are usually 102 to 127 mm long, and . . few live more than 2 or 3 years. In Bull Shoals Reservoir, Arkansas, threadfin shad average 53 mm in length at the end of the first growing season and 124 (males) and 135 (females) mm at the end of three gro'l.'ing seasons (Bryant and Houser 1969). Shad in tailwaters 224. Shad are important forage species in tail waters. Both gizzard shad a .nd thread fin shad are eaten by striped bass, trout, wu.lleyes, sauge.rs, and other species (Parsons 1957; Pfitzer 1962; Walburg et al. 1971; Deppert 1978; Combs 1979). Threadfin shad have 68 . ~ ., ;, .. .. ....... .been stocked in sane reservoirs specifically to provide food in the tai.lwaters for piscivorous fish (Parsons 1957). 225. The number of' gizzard and threadfin shad is generally higher in tailwaters than in natural streams due to their mo'V'ement from re.ser- voirs either over or through dams (Clark 1942; Parsons 1957; Louder 1958; Pfitzer 1962; J. P. Garter 1968a, 1968b, 1969; 'Walburg 1971). Occurrenc:e of shad in tailwate.rs appears to be 1:10re related to season than magnitude of outnow from the reservoirs (Clark 1942; Parsons 1957; Louder 1958). ·The presence of impoundments has also increased shad distribution. In Oklahoma, shad became established in a tailwater where they did not appear in the natural s t ream prior to illlpoundment (Cross 1950). 226. Dams also affect anadromous members of the clupeid family by acting as barriers to upstream spawning migration ( 'W. R. Carter 1968; Foye et al. 1969). Large numbers of American shad concentrated below a ~am in Maryland during spring. Fish kills sometimes occurred when the turbines were shut down during normal peaking o·perations and dis- solved. oxygen in the tailwaters was reduced to lethal levels ('W. R. Carter 1968). ·Alteration of operating procedures to provide mainte--.. nance flows of 141..6 m3 /s.ec through each of two turbines has been required to avoid further low-oxygen fish kills. Maintenance of minimum flows has been credited with allowing the survival of American shad in the Ru ·ssian River in California (Anderson 1972). Sal.monidae (Trou.ts) 227. The trouts are coldwater fish which were originally found in the Arctic and North Temperate regions. Over the years they have been introduced into suitable waters throughout the world. Many coldwater tailwaters in the United States have been stocked with hatchery trout on a put-and-take basis. Stocking is done at regular intervals throughout the fishing season to maintain a satisfactory sport fishery. Catchable-size trout. (150-200 mm long) are usually ... stocked because smaller fish oft en have poor su.""Vival to the creel (Vestal 1954). 228. The rainbow trout is the, most common sal.monid stocked below dams bec,ause it is less costly to 'raise and easier to catch. Other salmonids planted in tail waters are brown trout, brook trout, a.nd coho salmon. Rainbow trout are usually stocked in the coldva.ter tailwaters belov rese:-voirs built on varmvater streams. Rainbow trout or other trouts may be st.ocked or may occur naturally in coldwater tailwaters below reservoirs built on coldwater streams. Discussion here is limited to the rainbow trout because it is the most common trout in tailvaters. Rainbow trout . ·229. Habitat. The rainbow trout is native to the streams of the Pacific coast where many varieties or subspecies have developed. The seagoing form is known as the steelhead trout and is believed to be identical to the strictly fre,shva.ter rainbow trout. Because of the ease wit-h which eggs of the rainbow trout can be transported, different strains have been distributed throughout, the world,. 230. This species lives in a variety CJf habitats, inc~uding streams, lakes, and reservoirs. The rainbow trout tolerates somewhat, higher temperatures than·other trouts but does best in, waters that remain more or less continuously below 2l.l°C. According t9 McAfee (1966), the upper temperature limit for the species varies fr.om about 23.9 to 29.4°C, depending on the oxygen content of the water, size of fish, and deg:-ee of acclimation. Cold waters discharged from the lover levels of so.me reservoi.:-s provide adequate tailvater babi tat for this species, provided year-round oxygen levels remain above 5 mg/1. 231. Rainbow trout thrive in small to moderately large streams and shallow rivers, with modere,te flow and gravel bottoms of the pool-riffle type. They generally prefer riffles and fast-water areas. Depth criteria have not been defined for trout in general; however, the depth of pools and amount of cover appear to be very important in terms of' fish size. Gunderson (1968) found that stream sections lacking.deep pools and adequate cover contained only fingerling trout, while stream sections with deep pools separated by riffle are:as 70 '· ···•· ....... ... ............ . ' contained much .larger trout. Trout. req~re adequate stream depths for normal intrastream movement. Riffles are extensively used for feeding areas and tor movement between pools (Giger 1973). Minimum ~epth requirements over riffle areas vary with the size of the stream and the trout inbabi ting it; hovever, minimum water depth over ri.ffles should probably not be leu than 0.18 m. Giger (1973) listed the optimal pool depth for cutthroat trout as 0.4 to 1.1 m, depending on age and size. Hooper (1973) stated that for trout in general, areas with velocities betveen 9.1 and 31 em/sec are preferred for resting. 232. Reproduction. Rainbow trout spawn between early winter and late spring, depending on the genetic st·rain and stream conditions. The eggs are deposited in a shallow depression dug by the female on a · clean gravel riffle. Females deposit between 200 and 9000 eggs in the nest (redd), depending on fish size. After all eggs are laid and fertilized, they are covered with gravel. The incubation period varies with temperature, averaging about 80 days at 4.4°C and 19 days at Young remain near the hatching site for a while, tending to ' . -school at first and then become solitary and more widely. distributed._ 233. According to Hooper (1973), the optimUm spawning temperature for spring-spawning rainbow trout is 11.1°C, but ranges from 7.2 to 13-~3°C. _Preferred g;avel si~e is 0.6 to 3.8 em in diameter, and pre- fe~rred velocities for spa~ing a~~-between 42 .6 'and 82.0 em/sec. 234. Food. Rainbov trout eat a wide variety of foods but depend primarily on drifting insects. A compilation of' the findings of many . -. . studies indicat.es that immature and adult aquatic insects . (principally .. caddisflies, mayflies, and dipterans), zooplankton, terrestrial ·insects, and fish are usually the most significant foods, though their relative importance varies greatly between waters, seasons, and size of fish. Large trout include more fish in t -heir diet. Oligochae.tes, mollusks, fish eggs, a.mphipods, and algae are foods eaten less ex- tensively, but they may be important locally. For example, in upper Lake Ta:neycomo, below Table Rock Reservoir in Missouri, amphi.pod crustaceans make up almost 90 percent of the trout diet (Pflieger 1975). 71 ·•''- L I 235. Age and growth. Fev rainbov trout live beyond 6 years, and life expectancy for most is 3 or 4 years. Longevity is influenced by . marcy-.interrelated factors. Poor food conditions may result ·in poor survival after first spawning. Heavy angling pressure may crop most ... fish before they are 4 years old. This is especial1y true for fish stocked in ta.ilvaters, where most may be captured during the same year. 236. Maximum size of rainbow trout varie·s greatly among different environments. In small streams adults rarely reach 205 mm; whereas in large lakes individuals occasionally exceed 13.6 kg. Maximum size depends primarily on growth befo'!'e attainment of sexual maturity, which in turn is dependent on quantity and quality of available food . Most growth occurs during the first tvo graving seasons. Growt.h is fair for fish on a plankton diet, and is best where forage fish are abun- dant. Growth usually dec.lines after maturity is reached; this decline. is especially evident in waters where large forage organisms are lacking. 237. The rate of growth varies seasonally and at different ages, ·depending on water temperature, strain of trout, feeding conditions, ·age at maturity, and other factors. For example, in waters vith re- latively high temperatures throughout the year, such as the tailvaters of some reservoirs in the southeastern United States, growt.h is· fairly constant in all months·; whereas, in vaters vi th seasonal temperature changes, growth is slaver or ceases in the 'Winter. · 238. Hatc.hery-reared trout reach a length of 205 to 254 DDD vhen about one year old, at which time they are considered large enough to s~ock. In Lake Taneycomo, Missouri, where conditions for rapid growth are favorable, stocked rainbow trout grov about 19 DDD per month (Pflieger 1975). Naturally reared rainbow trout in Minnesota reach 125 DDD the first year, 230 mm the second, and 521 DDD the fifth year (Eddy and Underhill 1974). 239. The weight of rainbow trout varies greatly at lengths over 380 mm because of differences in feeding conditions and s.tage of maturity. Fish are heaviest where food is abundant, particularly in 72 • ... • .. ":.'. .•· ... · , .. lakes. According to McAfee (1966), weight of fish in average condition by 127 -mm fork lengt-h groups is as follows : Len~h 1 mm Mean veisht 2 kg 127 0.028 254 0.227 381 0.568 508 1.589 635 3.178 762 4.767 Most fish spavn by age III; .males one year earlier than femal.es. Size at maturity is extremely variable but usually ranges between 254 and 381 .mm. In California, mature rainbow trout typically veigh less than 0.45 kg (McAfee 1966). Rainbow trout in tailwaters 240. Habitat . Trout are found in most tailwaters throughout the ·. ... United States vbere s ummer water tempe·ratures normally do not exceed · :. -.. . . ~· . 21.1 °C. They may occur naturally below dams built on coldwater streams or the.y maY be stocked in tailvateTs below dams on warm:water streams '!lbere hypolimnetic releases maintain coldvater ·conditions throughout the year . Tail waters below most deep-release re·servoirs ha.ve lov turbidity, cold temperature, and s tabilized seasonal flov, thus pro- viding satisfactory trout habi ta'.:. These t a ilvaters often provide coldwater habitat on streams in areas such as Texas and Arkansas, where trout could not live before dams vere constructed . 241. The quantity and quality of col.dvater habitat in a tailvate!" suitable for trout are a function of reservoir design and operation, as vell as tailvater physical characteristics. ResE'rvoi r features that determine habitat suitability include rese!"-.roir stratification , reservoir storage capacity, reservoir storage and release patterns, and intake locati.on. Tailvater f eatures that influence habitat suit- ability are water tempera.ture, stream channel configuration, st!"eam substrate, stream cover, water depth, water velocity, water quality, 73 •·. turbidity, and tributary inflow. These factors act in combination with stocking to determine trout abundance and distributi·on. 242. NUI:lerous fish species are ·found in association. with rainbow trout in many tailwaters. Most compete with the trout for food and habitat. Game fish inhabiting these waters include largemouth and smal.lmouth bass, bl.uegills, longear sunfish, and catfish. Brown and brook trout. may alsc-occur in the same location. Cyprinids, suckers, sculpins, &nd sticklebacks are major nongame competitors. 243. Low flow, sometimes aggravated by elevated water temperature, is a coi!IDon factor limiting trout distribution and abundance in many tail waters. Low flows can decrease cover, increase· overwintering mortality, allow sediment accumulation, and cause stl"a.nding. Reduction of flow in a Tennessee tailwater limited habitat to a long shallow pool with bedrock substrate and 11 ttle c.over (Parson! 1957). Kraft ( 1972) found a 62 percent decrease in brown trout habitat when flow was reduced. 90 percent in another site. Weber (1959) found that when annual flow below a reservoir was reduced to 11 percent of the long-term average, trout habitat was reduced 82 percent.· Flows that reduce water .~.depth over riffles to 7.6 em or less make .these areu unusable to' large trout (Corning 1970). Decreased f l ows and thus decreased water velocity favor small trout and rough fish over large trout. Lov flows a l so cause a redistribution of trout to less suitable habitat, incre.asing competition with other fish. Corning (1970) also foU:Jld that re~uced flows concentrate. fish in the remaining habi tat ·.and intensify predation • . -. . . .. ·~ . : . 244. In the vestern United States, irrigation-storage reservoirs colleet surface runoff during winter and spring to be used during summe r irrigation. Holding runoff decreases stream flows during the winter and spring vhich can i ncrease trout mortality. Adequate reser- voir discharge is critical during winter to sustai n a tailwater· trout population . The survival of 3-year and older trout was di'rectly re- lated to the magnitude of flow from a reservoir during the winterr storage period (Nelson 1977). Vincent (1969) stated that dewatering below a da:n in Montana resulted in excessive mortality of young (age I) trout . Studies in Wyoming shoved that seasonal minimum •flo.,.s were . :·· .. . . ··· ··I . essential for trout survival. Winter flows of 14.2 m3 /sec wer~ recom- mended to maintain adequate cover for overwintering trout. Spring and summer flows of 22.7 m3/sec were recamnended to select against rough fish and for large trout, and flows of 45.3 m3 /sec were recommended periodically to flush silt from the tailvater. A 30-day survival flow of 8.5 m 3 /s~c was recommended for times of extreme water shortage (Banks et nl. 1974). 245. Low flows in tailwaters allow sediment accumulat.ion in holes and over· gravel sub~trate, thus limiting the quantity of trout. habitat. Sediment input into tailva.ter·s is usually from tributary inflow or the erosion of alluvial banks vi thin the tailvater. Reduced spring flows below a dam in Montana increased sediment accumulation by tributary . 'inflow. Timed spring discharges from the dam are used to wash sediment downstream (May and Huston 1979). Seasonal flood .flows over· the spill~ way removed. sediment from a tailva.ter in Alaska (Schmidt and Robards 1976). Sediment accumulations behind a dam in California wer·e reduced . . by periodic flushing into the tailwater, which caused destruction of trout .habitat for 1.6 t o 3.2 km downstream (Anderson 1972). ~.· 246. ·High flows and flow fluctuations in ·tailvaters can be limiting faetors for certain size classes of trout. Banks et al. ( 1974) found that water velocity increased sharply with increased dam dis- charge. Trou.t . 30 em long or longer were favored over smaller trout and rough fish because of lack of resting cover and high water velocities . High water velocities and sparse cover results in low standing crop (26.1 and 4·2.6 kg/ba) and harvest (7.7 and 8.6 kg/ba) of trout .. The trout th.a.t. remain are usually large and weigh "Jetween 0. 7 and 5.5 kg (Mullan et al . 1976). •. 247. Rapid flow fluctuation below both hydropower and diversion dams has caused stranding of both trout and salmon (Anderson 1972; Kroger 1973; Fowler 1978). Kroger (1973) suggested that decreasing flows at a ma.ximum rate of 2.8 m3 /sec/day would reduce stranding . of f1sh below· a dam in Wyoming. However, the most serious effec.t of flow fluctuations appears to be reduced trout reproduction, rather than a 75 ... ~ ..... I ; I dire.ct increase in adult fish mortality .(Parsons 1957; Baker 1959; Axon 1975). 248. The cold temperature of tailwaters below 'hypolimnetic re- lease dams built on warmvater streams helps trout to compete effectively with or replace native fish species. Vanicek et al. (1970) found native fish replaced by rainbow trout in the 42 km of river below a dam in Utah and Colorado. Lover water temperatures below a d.am in Texas caused partial replacement of 20 native species by stocked rainbow trout (Butler 1973). 249. Dams with hypolimnetic releases built on coldwater streams have altered the'habitat by lowering water temperature~ and stabilizing fiovs. Belo·v a Colorado dam, reduced water temperature caused a re- distribution of trout species, with brook trout moving into the colder water near the dam (5.0-8.3°C) and brown and rainbow trout moving into the warmer areas d.ovnstream. After completion of two Colorado dams, summer water temperatures in the tailwater were reduced by 3.2 to 5:0°C. The temperature reduction is believed responsible for trout appear_ing in areas where they were pr·eviously :r;are or absent (Mullan et al. 1976). P~naz et al. ( 1968) stated that . the Vir tail water on the Svratka River, Czechoslovakia, bad. reduce.d water temperatures and stabilized flows d.uring the summer, and these conditions favored brown tro·ut over case, a warmwater species. .Nase. we·.re reduced from 63.4 to 12.7 percent o·f' the tailvater fish harvest and brovn. trout increased to'76.8 percent. 250. In addition to reduced maximum temperatures, the seasonal rate of temperature increase is slowed in many cold tallwaters. In a Montana te.ilwate.r, tl;le normal spring water temperature of 12. 8°C is achieved 6 to 8 weeks later than it vas before the dam was built. This change delays the spawning of suckers and. gives trout a competitive advantage in the tailvater (May and Huston 1979). 251. Reduced stream flows (usually less 'than 10 percent average daily flow) during period,s of warm air t .emperat.ure can c.ause vater temperatures to exceed lethal levels for trout. This situation. often occurs in natural streams, in tailv-aters below deepwater release 76 -: I reservoirs on formerly warmvater streams, and belov hydropower dams that discharge only during periods of high e'lectrical demand. Dendy and Stroud (1949), Parsons (1958), Baker (1959), Kent (1963), Geen (1974), Axon (1975), Aggus et al. (1979), and others have stated that high water temperatures may 11mi t trout populat.ions in tail waters. Many reservoirs have incorporated minimum water releases in their opera~ting schedule to maintain suitable flows and water temperatures in tailwaters. The U. S. Bureau of Sport Fisheries and Wildlife (1969) rec<Dmended a minimum t'lo\1 of 7.1 to 11.3 m3 /sec to maintain the trout fishery in a Wyoming tailwater. Suitable trout water temperatures are maintained for 9.7 km below Canyon Dam, Texas, by cold hypolimnetic discharge (Butler 1973). 252. Extremely cold water temperatures were associated with the loss of a trout fishery in several tai.lwaters. A New Mexico tailwater ·experienced an 8-month average water temperature decline from 10°C in · 1968 to 5°C in 1971. This temperature change was associated with a decrease in trout harvest from 78,656 fish in 1968 to 10,642 in 1971 ( Mull~f et ·-al. 1976). In a similar situation, an average tr~ut harvest. of 109.5 kglha was recorded when annual water temperatures ranged from 3.1 to 12.00C; however, when the average annual temperature range decreased to between 4. 2 and 9. 2°C, the trout harvest dec.reased to only 6. 2 kg/ha (Mullan et al. 1976) . 253. Low dissolved oxygen concentrations and gas supersaturation resulting in nitrogen embolism can cause mortal.ity or trout in other- vise suitable tailvater habitat. Low dissol7ed oxygen is a problem ·often encountered in the southeastern United States in tailvaters of deep-release dams built on varmwater streams. This oxygen deficiency occurs when a reservoir thermally stratifies and decomposition of organic matter within the hypolimnion consumes available oxygen. "Nitrogen embolism may occur when fish inhabit tailwaters that are super- saturated with atmospheric gases and the body fluids of the fish also become supersaturated. 254. Trout generally prefer water with 5 mg/1 or more dissolved oxygen. The species may not survive when hypolimnetic release water · 77 conta_ins little or no dissolved oxygen. The penstock design at some dams does not allow for aeration of hypolimnetic water before release ' into the tailwaters. Low dissolved oxygen vas. found to limit the fauna in some tailwaters (Hill 1978). A dissolved oxygen concentration of . - 0.8 mg/1 presumably caused the death of about 100 rainbow trout in an Oklahoma tailwater (Deppert 1978). However, a dissolved oxygen concen- trat.ion of 2 mg/1 in November 1951 in a Tenne~see tailwater caused no apparent signs of distress in trout (Pfitzer 1962): Baker (1959) found low dissolved oxygen and distressed fish directly below a dam in Arkansas in the fall. The oxygen concentration returned to normal and · ... fish recovered · downstream over the first riffle. Turbulent flows in- crease the aeration process a .nd thus improve the tailvater habitat -by increasing dissolved oxygen concentrations. 255. Gas supersaturation has been a major cause of steelhead and salmon mortality during high-water years in the Snake River, Idaho and Washington (Raymond 1979). Graves and Haines (1969} stated that'dead ' . fish vi t~ gas bubbles on their fins were found vi thin 1.6 km of a 'dam _. i~ N;w Mexico on several occ~sions. · Nitrogen sup~;saturation below a · ·· ~ntana d~· p~~sisted fo~ ~ome · 32.2 ~ do~stream 'because'· flo.;s in ·tht·· .: ·· tailwater were not turbulent enough to dissipate the nitrogen. Of .39 rainbow trout held in a cage below this dam, 21 died, presumably of nitrogen embolism .(U. S. Bureau of Reclamation 1973). · 256. Reproduction. Spawning of rainbow trout has be-en observed in several cold tailwaters, although survival of eggs and juveniles .is low. Trout populations in tailwate.rs rely on stocking or recruitment from the reservoir above to maintain the.ir populations (Ston~ 1972). Few self-s.ustaining tailwater trout populations that support a fishery have been reported in the literature. Spawning may occur in tribu- taries where local conditions a ·re favorable, but the magnitude of tributary spawning has not been adequate to maintain fishable trout populations in most tailwaters. 257. The harsh environmental conditions (flow fluctuations, de- watering) encountered in tailwaters reduces reproductive success. Corning (1970) found that high flows washed out trout redds. Parsons 7R ... : :· :.· . •. .. ,. (1957) found the 3-m water-level fluctuation in a Tennessee tailvater from Oc tober through December detrimental to trout reproduction . Redds ve.re consistently scoured out at high flows and stranded on dry shoals during low flows. Rainbow trout spawning in December and January vas common on shoals and riffles, in an Arkansas tailvater, but vater-le.vel tlu~t·uation caused rolling gravel and destruction of redds. Spawning in tributaries vas successful, but the contribution of naturally pro- duced f.ish to the trout fishery in tailvaters vas insignificant (Parsons 1957; .Baker 1959; Pfitzer 1 962). Lov trout populations in the Beaverhead River, Montana, were caused by poor reproductive success due to low .flows. Spring and winter dewatering (low flows) of the r i ver · caused by. irrigation storage reduced egg and larvae survival. by de- creasing the cross-sectional area and water velocity of the stream (Nelson 1977). Corning ( 1970) found that J.ov flows exposed trout redds to the air and increased siltation, thus reducing egg survival !"rom 85 percent to only 17 percent. Siltation filled the int erstices of the substrate and smothered the eggs . Schmidt and Robards ( 1916) suggested' ·. . -:-. -· . ._ . staged flows in an Alaska tailvater to clean the substrate and increase trout and salmon spawning and rearing habitat. 258. Spawning of trout is common in a W)roming tailvater, but the lack of cover reduces survival of fry. This tailvater relies o.n an . . . . . annual stocking of 348 rainbow trout fingerlings per hectare to main- tain the fishery (Mullan et al. 1976). Mot'fett (1942) recommended stocking of 30,000 fingerling rainbow trout (127 to 178 mm long) per . ..... · ~ ........ :~ .. _... year to compe-nsate for low repr od.uction caused by vater-level f l uctua- tions in a tailvater in Nevad.a a:nd Arizona . 259. An obvious effect of dam. construction is the bloc.kage of .fish migration upstream to spavn.ing grounds. Construction of a dam on a California river, impounded 50 percent of the salmon spawning a r eas (Moffett 1949). Attempts to mitigate loss of spawning areas by hatchery construction have been only partly successful, particularly for anadromous ·species (Raymond 1979). When a hatchery for Bal ti.c salmon vas proposed for Narvskaya tail water in the U.S.S.R. to replace inundated spawning gro·unds, Be.rannikova ( 1962) argued against the 79 hatchery because he beli.eved the high number or predators in the tailwater would devastate the yearling salmon. Striped bass and saugers are significant predators on small trout stocked in some tailvaters during certain seasons (Boles 1969; Ari zona Game and F i sh Department 1972; Deppert 1978). 260 . Food . Food studies have been completed on trout from a .number of tailwaters and it is apparent that thei r diet is very di verse (Table 1). Cladoohora beds are an important food source for trout in ta'ilvaters (Moffett 1942; Mullan et al. 1976). Trout graze on thes.e mats and ingest the algae and the isopods, simulii.ds, and mayflies harbored therein. In some tailvaters, food organi,sms hav e been intro- duced to provide food for trout. Snails were introduced below Glen Canyon Dam, Arizona, and amphipods were introduced into Taneycomo Lake, Missouri (Table Rock tailwater) (Mullan et al. 1976; Ralph Burress, U. S. Fish and Wildlife Service, personal communication). 261. Trout are no·r totally dependent on. food production vi thi.n the tailvater. In Dale Hollow tailwater, Tennessee, 81 percent -of the organisms in rai nbow trout stomachs were cladoc.erans which were ··pro- '· duced in the reservoir above·(Li ttle 1967) •. In this same tailvater, rainbow trout weighing from 1. 8 to 2. 3 k g ate crappi es 102 to 152 mm long vhich had been stunned when they pas.sed through the dam (Parsons 1957). R~inbow trout belov Center Hill Dam, Tennessee, .fed heavily on 51-to 76-mm threadfin shad which had passed through the turbines (Parsons 1957). In Wyman tai.lvater, Ma i ne, rai nbow trout over 4o6 DID long were more piscivorous than smaller trout. Smaller t .rout ate Chironanidae (dipterans) which were produced in the reservoir and carried into the tailwater (Trotzky 1971). 262 . Food organisms eaten by rai nbow trout collected below a r eservoir d i ffered from that of fish 48.3 km downstream (Welcn 1961). Algae vere the primary food below t_he dam, and mayflies and small s,ton.eflies farther downstr eam. The influence o.f tailvat.er flow and t .emperature on food avai1'3.bility decreases downstream as tributary inflow, meteorological cond itions, and other influences moderate the effects of the discharge . Brown trout that were collected immediately 8o Tailvater Boulde:r (Hoover) Nev., Ariz . Canyon, Tex. Canyon, Tex. Cente.r Hill, Tenn. Cow Green, U.K. Dale Hollow, Tenn. Dale· Hollow, Tenn. Dale Hollow, Tenn. Davis, Nev., Ariz. Flaming Gorge, Utah, Colo. Fontenelle, Wyo. Glen Canyon, Ariz. '• ·:· . • < .. -. ··Table 1 Food of Trout in Tailwaters Food item Cladophora, mayfly, midge larvae Ephemeroptera, Diptera, Trichoptera Ephemeroptera, Diptera, Trichoptera, terrestrial insects Threadfin shad Ephemeroptera, terrestrial insects, zooplankt on from reservo:f.r Cladocerans, Diptera, Mollusca, terrestrial insects, isopods Cladocerans Algae, gastropods, Chironomidae, terrestrial insects, sow bugs, crayfish· Detritus .Algae ·.~· Cladophora, fish, Daphnia, copepods Algae, cladocerans, i:ntroduce~ snails (Continued) ·. Reference Moffett 1942 Butler 19'13 White 1969 Parsons 1957 Crisp et al. 1978 Little 1967 Bauer 1976 Parsons 1957 Fast 1965 Mullan et al. 1976 Mullan et al . 1976 Mullan et al. 1976 ;. Co 1\) Tail water Glen Canyon, Ariz. Granby, Colo. Gunnison, Colo. (several on river) Narvskaya, U.S.S.R. Navajo, N. Mex. Navajo, N. Mex. Norfoik, Ark . Tiber, Mont . Wyman, Meine • • Table !;(Concluded) .; ~ ·, .... J'ood item ... ' . Snails l· ·'. Diptera, Trichoptera, Ephemeroptera Cladophora , emergent insects .· Gemmaridae, opossum shrimp, Chironomidae Midge, black fly, caddisfly, mayfly, grasshopper, beetles, stonefly, dragonfly~-, ~nails Diptera, gastropods, Plecoptera, Tendipedidae, Ephemeroptera, fish , algae . Isopods, amphipods, crayfish, aquatic insects -~.· · . .-';· Simuliidae, algae' (Cladophora) Trichoptera, mayfly, Diptera,. terrestrial insects, Plecoptera ~·· ... ·' Reference Stone 1972 Weber 1959 .' Mullan et al. 1976 Barannikova 1962 Mullan et al. 1976 Olson 1965 Baker 1959 Welch 1961 Trotzky 1971 J ·": ••• , ; ...... ••• t . ,. : .. ·····:·· -~ . . . • • below a reservoir in Colorado a.te more. dipterans (small-bodied insects) than those collected downstream, whic.h ate primarily trichopterans and ephemeropterans (W'e'ber 1959). 263. Food is rarely limiting on the trout populations in cold tailwaters because of natural food production within the tailwater and food exported into the tailvater from the reservoir above. Fbod orga- nisms exported from the reservoir display seasonal abundance patterns; quantities available· in the tailwater are greater during spring and autumn than during midsummer and winter. 264. Age and growth. Trout grovt.h in tail waters is variable, · ··-..... -•... generally being comparable to that in reservoirs and often exceeding that in natural streams (Welch 1961; Trotzky 1971) • The average length of rainbow trout captured below Blue Mesa Dam, Colorado, in- ·. creased from 246 mm be'fore impoundment to 285 mm after impoundment · (W'ilt~zius 1978). 265 ~ Rapid growth of rainbow trout stocked in tailvaters has been reported in & number of studies. Mullan et al. (1976) reporte~ that ~ ~ ' . stocked fingerlings averaging 191 mm long grew 76 to 102 mm. per year in . Glen Canyon tailvater, Arizona; 76-mm fingerlings grev to 254 mm in length in one year in Flaming Gorge tailvater, Utah; and 76-to 127-mm fingerlin(iis grew to 254 to 381 mm in length in Fontenelle tail water, 'Wyoming. Rainbow trout fi~erliilgs. released at an average length of 83 mm· grew.l78 mm from July to December in Navajo tailvater, New Mexico (Ol~on· 1965). ·stevenson (1975) reported that the average length of rainbow and brovn· trout-below Yellowtail Dam, Montana, increased 152 mm between May and December • . _,., 266. The abundance of food organisms coming from a reservoir and of 'inv~rtebrates produced in the tailvater is associated vith rap~d growth of trout. Parsons (1957) stated that rainbow trout in Center Hill tailvater, Tennessee, fed on thre11.dfin shad· coming through the dam and grew 25 mm per month. The availability of the higher quality food ·(fish versus insects) in Center Hill. tailwater apparently produced the · rapid growth. Rainbow trout growth decreased in Dale Hollow tailvater, Tennessee, in 1953 and 1954 when the number of fingerlings stocked vas .· · .. . . . , .... .. inereased·from 20,000 to 30,000. This indicated high utilization of available food in the tailwater. Temperatures in Dale Hollov tailvater, vhieh range ~rom 7 . 2 to 13 . 3°C vi t.h a maximum monthly variation of 1.7°C, are considered excellent for trout growth (Parsons 1957). Rain- bov trout that fed on the large numbers of arthropods harbored in the vegetation in Norfork and Bull Shoals ts.ilwate:r"s, Arkansas, during 1957, grev 23 mm :per month. In 1958, vhen flood flows washed out .much .of this vegetation, trout growth decreased to 17 mm :per month (Baker 1959). · 267. Trout growth in tailwaters and other areas bas been discussed by several authors. Irving and Cu:plin (1956) re:ported only small dif- . ferenees in g~ovth of native and stocked rainbow trout ca:ptured in ... ,.. ..·· several Snake River tailwaters: .··· .. Total length, mm ~ Native fish Hs.tcher;r fish I 130 127 II 262 244 .-. III 351 .. 333 . . • ,,;•\' .. IV 467 445 v 488 Also, no differences in growth could be sh~ f pr rainbov trout take~ ·. . f:r.om ,"t;ai1vaters or im:polll\d.ments on the Snake River, Idaho . Trotzky , ... · ... (1971) found that rainbow trout in Kennebec River tailvaters, Maine, grew. f~.ster than rainbow trout in streams and lak.es ·from other areas of the United ·States. Large trout have been reported from several tailwaters . Rainbov and brovn trout veighing ·6.8 kg have been captured in the White River tailwaters, Arkansas (Baker 1959), and trout u:p to 5.5 kg are commo n in Fontenelle tailvater, Wyoming (Banks et s.l. 1974). 268. Reduced water temperature ereremes in vinter and summer and a more homogeneous temperature regime throughout the year appear to have resulted in year-round growth of ra.inbov trout in some tailwat.ers . This eo~clusion is supported by the inconsistent age :readings and lack of annulus formation on scales of trout from some tailwaters (Moffett 84 • • , 1942; Parsons 1957; Pfitzer 1962; Olson 1968). In Watauga tailvater, Tennessee, where winter water temperatures are moderate, year-round growth of rainbow trout averaged 15 mm per month. Little or no fish growth vill occur in tailwaters where vinte:r water temperatures are extremely cold. Esod.dae (Pikes) 269. The.re are four spedes of the pike family in North America: grass pickerel, chain pickerel, northern pike, a.nd muskellunge. The natural range of the_ chain and grass. pickerels is the eastern United States; whereas, that of tbe northern pike is north-central United States and Canada, and taat of the muskellunge is the Great Lakes stat.es south to Kentuclr.f. Northern pike and muskellunge reach a large size and. are highly regarded as game fishes. Both specie.s or their hybrids have been stoc ked extensively in some reservoirs. The chain and grass pickerel are smaller fi.sh and though chain pickerel provide · ·, f fishing in som.e streams, gn'.ss pi~ke.rel seldom .reach catchable ·size. •, .. The pickerels occur in SO!l'.e tailwaters, but they receive littl':! mention in the literature' and vill not be discussed further. ~ 270. Habitat. All pikes have similar requirements in that they prefer clean, quiet-water areas of lakes and streams where there is an abund..an.ce of aquatic vegetation. In streams, they prefer cove.r along the margins in patches of vegetation, beside submerged roots or branc·hes, or in patches of shade. Northern pike are commonly found in a variet.y of habitats in lakes, reservoirs, and large streams , and muskellunge in lakes and pools and backwaters of slow-moving streams. 271. Reoro duction . Pikes spawn in the spr:ing soon after ice-out. They move. into marshes or other shallow marginal waters vher·e vegeta- tion is abundant. No nest is built, and eggs are broadeast and aban- doned. The adhesive eggs sink and adhere to the bottom or to vezP.ta- tion. They hatc·h in 10 to 14 days, and the larvae remain inactive but attached to vegetation for 6 to 10 days or until the yolk sac is absorbed. Both eggs and young may be stranded if water levels drop in the shallov breeding areas. Studies suggest that spring water levels must remain high for at least a month afte.r pikes spavn to obtain good year-class survival. 272. Food. Pikes are carnivorous, feeding principally on other fishe.s. They remain motionless near cover and dart out t,o capture unwary· passing prey. Pike larvae eat zooplankton and the. larger young eat aquatic insects and small fish. 273. Age and growth. Grovth of both the northern pike and muskellunge is extremely rapid. Norther.n pike average 251 mm long at the end of their first_year and 777 mm at the end of their sixth year (Karvelis 1964). The maximum length of male.s probably does not exceed 760 mm (Threinen et al. 1966). Maximum length of females may exceed 1016 mm, bu.t fev survive beyond 12 years. Muskellunge. average 267 mm long at the end of their first year and 769 mm at the end of 5 years (Karvelis 1964). Maximwn age is about 20 yeal's, and maximum repol'ted length is in excess of 1270 mm. For all species or pike, females grow '·'·mOre rapidl.y and live longer than males. or 3 years old.' Pikes in tailvaters Most .fisb are mature vhen 2 274. None of the pikes are common tail waters. They may be found there for several years after construction of a reservoir on. rivers vhe.re the species occurs naturally or after stocking within the. reser- voir. They may move upstream into tailwaters from downstream locations at certain'ti.J;les of th~? year to feed or spavn • . 275. Most. north~rn pike and muskellunse found in tailwaters have passe~ over or through a dam from the reservoir above (Diuzhikov 1961; Hanson 191'7; Wilt:.ius 1978). During or immed.iatly after the filling of a nev reservoir, pike may become common in the tailvater for a short period!. Di'-!zhikov (1961) reported large year cla.sses of pike being produced in Kuibyshev Reservoir, Volga River, as it wa.s being filled. Many of these fish passed over the dam and congregated .in the tailwaters, feeding on the large. numbers of small fi.sh t 'hat passed over the da:n or vere blocked during upstream migrai.iuns. During the tvo 86 • . . -' ....•. ~ to three years following impoundment, up to 40 percent of the number and 90 percent of the biomass of fish caught in the tailvaters were pike. Th.ese high catches were followed by .a rapid decline in numbers within 5 years {Chikova 1968). .. 276. The loss of p.ike from. Kuibyshev tailvaters can be attributed to both reduced reproduction in the reservoir and la~-k of spawning success in the tailwater. Spawning in. the tail water vas delayed from early May to late May because of slower warming of water and high daily and. weekly vate.r-level fluctuations caused by hydropower production, which further inhibited spawning. Additionally, a seasonal drop in water levels of 4 to 5 m occurred during embryonic development, re- sulting in massive desiccation of the deposite'd eggs and larvae {Eliseev and. Chikova 1968). 277. A similar situation occurred in Narvskaya tailwaters on the Narova River, where regular, sharp water-level fluctuations due to hydropower peaking activity result.ed in the disruption or cessation of pike spawning. This vas evident .from the large number of ad?lt pike found to be resorbing sexual products (Ba.rannikova 1962). Dilappear- anc.e of northern pike from the tailwaters below four hydropower im- poundments {e.g., Cooke, Five Channels, Foote, and Mio) on the Au Sable River, Michigan, is believed to have been caused by the lack of spawning success in the tailvaters, coupled with a general decrease in produc- tivity of the upstream reservoirs (R_ichards 1976) •. 278. Dams have also effect.ively limited reproduction by blocking movement of fish to their upstream spawning grounds. This occurred on the Middle Fork of the Kentuck.y River where muskellunge numbers have steadily declined since the construction of Buckhorn Reservoir, Kentucky (Branson 1977). · 279. The ef:t"ects of flow regulation on pike are not limited to the immediate tailwaters of certain dams. Reduced river flows result- ing from reserv oir filling can affect pike distribution and reproduc- tion hundreds of kilometres downstream. Low vinter flows below Bennet:t Da:n on the Peace-Athabasca River in Canada during the filling of W'i.lliston Lake threatened the northern pike populations in Lake .... .. ·· Athaba.sca through severe freezing of shallow-water areas and increased oxygen de,pletion which caused a winter-kill {Townsen~ 1975). 280. Similarly, flood control below the Volg'>g:;.-ad hydropower facility on the Volga River has affected the northern pike population ·in--the delta Oil the Caspian Sea far downstream. The lov \o"ater resulting from spring flood control has shifted the pike's distribution vi thin the delta and. bas caused a. delay in the spal.~ing season from March-April to May ,and a corresponding decrease in spawning success. The delay in spavning shortened the foraging season, and pike grovth ·rates declined. '···. .. . ·Additionally, the reduction in numbe,rs of the pike's prime forage species, mai nly ·spring spawning vobla, pike-perch, and bream, resulted in a decrease of its annual food supply (Orlova and Popova 1976). Cyprinidae (Minnows) 281. The minnow family is the largest of all fish famil.ies. Most m•bers are small, but some (carp, squavf'ish, chubs) attain large size. Minnows are found in all natural wat .ers, but are more common in streams than in lakes or ponds. In s t reams, they are often more numerous than all other fishes combined . Minnows are efficient in transforming minute aquatic food into sizable food for larger game fishes . 282. As a group, the c.yprinids vary greatly in. food babi ts; some feed on insects, some on algae, and some on the organi.c mud of the bottam;··st-ill others are omnivorous: Habitats may include silty, clear, a~·. bog waters; quiet or rapidly fldwi.ng streams; sand, mud or gravel bottoms. 283 . Spavning migrations by minnows. are limited; no species moves more than a short distance upstream, or beyond the shoals of a lake. All spavn in spring or summer and the incubation period of the eggs is relatively short. Some species merely sca.tter thei.r · eggs in a sui table habitat; others deposit th!!lll 'in specially prepared nests; and still others guard the eggs until they hatch. 284 . Many different species of minnows often occur in the same waters. In such instances, the various species are found over 88 different types of bottoms: mud feeders over mud bottom, algae feeders over algae-covered rocks, and insect feeders over sand and gravel or other types of bottoms. 285. With the exception of carp and some chubs, minnows are not sought by sport fishermen. Most minnows are effective baits for the taking of sport fishes. Because of the large number of minnow species, this discussion of life histo-ry is limited to the following general cyprinid groups: carp, chubs, true minnows, • shiners, and stone roller's. Spec.ies considered. vi thin these groups are those most often mentioned in tail water literature. Discussion of cyprinid occur.:rence in t&i.lvate-rs includes the above five groups plus daces, squavfishes and chiselmouths, and cyprinids in Russian tailvaters. Daces, squavfishes, and chiselmouths are of local importance, and Russian cyprinids are included. to illustrate tailvater-fish problems simi.lar to those found in the ~nited States. ~ -286. Habitat. Carp are found throughout the United States. The • _. -,·r • •, species is very adaptable and occ.urs in most aquatic habitats b\it is ; . most COIIIIDOD in large streams, lakes, and man-made impoWidments that a.t:-e highly productive because of natural fertility or organic pollution. In streams, adult carp are usually found near submerged cover such as brush piles or logs and where the current is the slowest. In lakes and reservoirs, carp e.re usual.ly found near shore or in shallow embayme.nts; an occasional fish may be found in depths exceeding 10 m. 287. Carp do not school, but they ·do form lo.ose aggregations. The species is not considered migratory, but some individuals move for · long distances. Where carp be.come abundant there is usually a general deterioration of the habitat because of increased. turbidity and destruc- tion of aquatic vegetation caused largely by the fish's feeding habits. 288. Reuroduction. _Carp move into shallows to spawn between late March and late June. Spawning starts at va.ter temperatures of 14.5 to *Members of the family Cyprinidae whose common name includes the word "minnow" (Bailey et al. 1970). 89 . ~. .. ·· I~ 'I I 17°C but peaks at 18.5 to 20°C. The eggs are scattered over logs, rocks, or submerged vegetation . Eggs ha.tch in 4 to 8 days, depending on temperature; there is no parental care of eggs or fry. Carp o.ften spa'Wll in water so shallow that their backs are exposed and the noise created as they thrash abc•1t can be heard for considerable distances . 289. Food. Carp fee.i on a variety of an.imal and plant mate.rial. Aquatic insects are the most coDDDon diet item and plant material ranks second. Most active feeding occ~s in late evening or early morning and food is probably located more by taste tha:n by sight. Carp are mostly bottom feeders, but they have also been observed taking floating objects en the .water surface. 290. Age and grovt h. According to Pfli eg·e:r ( 1975) , carp in Missouri streams average 165 l!llll long at the end o·f their first year o f life and reach lengths of about 279, 361~ 424, and 457 mm in succeeding years. On the average they veigh 0.45 kg when 305 nun long, 2.3 kg when 546 mm long, and 5.4 kg when 698 mm long. Carp from cool, infertile vate~s grow more slowly and veigh less at comparable lengths than do those from warm, productive waters. Few live more than 12 years. Most •. males are mature at ages II-IV and most females at ages II!-V. · · CarP in tailvaters 291 . Carp are common to abundant .in many varmv~ter tailvaters. Tbey were 'abundant in the anglers catch in Lake of the Ozarks from 1965 to 1974 and were 6 percent of the catch ~ 1964 in Clearwater Dam tailvater, Missouri (Hanson 1965, 1977). Carp made up 57 percent of the 'fish captured by anglers in 1968 in Carlyle tailvater, Illinois (Fritz 1969), and. constituted the bulk of the weight of fish collected in Holyoke Dam tailvater, Massachusetts (Jefferies 1974). They were also numerous below Fern Ridge Reservoir, Oregon {Hutchison et al. 1966). 292. Carp have also adapted well to conditions in a number of cold tailvaters. Creel surveys in Kentucky on the Barren Reservoir and Nolin Reservoir tailvaters from 1965 through 1971 recorded catches of carp rangi.ng from 1,300 to 6,200 and 143. to 10,000 fish, respec- tively. These catches accounted for 3 to 30 percent of the total 90 ·• . • ... ' ; .... •• ~· •:. I .. sport fishing catch on these tailvaters during those years (J. P. Carter 1968a; Charles and McLemore 1973). In addition, carp composed 50 per- cent of the fish population (estimated from samples collected by electrot'ishing) in Nolin Reservoir tailvater in March 1966 (J. P. Carter l968a). Carp are abundant in Chilhowee and Norris tailwaters, Tennessee (Hill 1978). They are one of the dominant species in Dale Hollow tailwater, Tennessee (Bauer 1976). Though not numerous in the immediate tailvater, carp were found to make up 18 percent. of the fish population 48.4 km downstream below Cumberland Dam, Kentucky (Henley 1967). Carp are commonly found in angler catches in a series of tailwaters on the Snake River, r daho, including Lower Salmon Falls,. Bliss, and C. J. St.rike (Irving and Cuplin 1956}. Holden and St~lnaker ·(1975) also re•ported carp as common be-low Glen Canyon Dam, Colorado. Not • ..... ·all cold tailvaters continue to pro!id~ good carp ,habitat! Carp. h&ve.····, .... decreased in abundance in a Wyoming tailvater, apparently because of declining temperatures (Mullan et al. 1976). 293. · Dam construction has limited the distribution of carp. The upstream migration of carp out of some Tennessee reservoirs has been .:; ., blocked by mil.l dams. This helps maintain t .he populations of small.::; --. . . mouth bass and. other game fish in these streams by limiting competition (Ruhr 1957). In the Kennebec River, Maine, carp have been kept out of the productive upper river by the presence of Augus~a Dam (Foye et al . 1969). 294.· Successf'ul. carp reproduction may occur in tailwaters, but it has not been verified. Running ripe c.arp we·re caught in Lewis and ·· Clark tailwater, South Dakota a n d Nebraska, at temperatures of 22°C, · and spent females were found after temperatures of 25°C were .reached. No young-of-the-year carp were collected from this tailwater (Walburg et al. 1971} . 295. Most recruitment of carp to tailvaters appears to come from the reservoir above or the river downstream. Large numbers of 'young- of-the-year carp-as ms.ny as 310,000 in 24 .hours--were found in the· discharge at. Levis and Clark Lake, but these fish did not stay in the 'tailvater (Walburg 1971; Walburg et al. 1971). Adult. carp were the 91 second most numerous species lost over the spillway at Little Grassy Lake, lllinoi-3, amounting to 14 percent of the total fish transported into the tailvater (Louder 1958). 296 .. Food habits of carp in ta.ilvaters were examined only below .. Levis and Clark Lake. Zooplankton, algae, and bryozoans were all utilized. ZooplaDkton was the main food available during the period vhe_n carp were most abundant, from early spring through July (Walburg et al. 1971). · 297. Carp grow well in most varmwa.t er tail waters. The average ve.ight of carp taken by anglers in Car lyle tail water, Illinois, vas . 0.66 kg _in 1967 and 0. 50 kg in 1968 (Pritz 1969). The average weight or this species collec.ted in Canton Reservoir stilling basin, Oklahoma, vas 0.50 kg (Moser and Hicks 1970). Carp collected in Lewis and Clark tailvater averaged 8 percent larger than those from the reservoir (Walburg et al. 1971). ~ 298. Habitat. Most species of chubs inhabit clear. streams having penaanent flov and a predominanc.e of clean gravel or rubble -bottoms. _., _'. 'Adults -&re usually found near riffles or in Jiools but not in the swifter current. The young are usually found ill qui.et:-vater areas 8J:1.d most often in association with higher aquatic pl~ts. 299. Re.production. The following description of spawning _by the · hornyhead chub (Scott and Crossman 1973) is believed typical for stream chubs. Spawning takes place in the spl_"ing of _the yeS!, probably when water temperatures reach about 24°C. Nests of stones and pebbles are built by the males on a fine-gravel or pebble bottom, often below a riffle, and in relatively shallow water that covers the top o'f a com- pleted nest to a depth of 15.2-45.7 em. Nest building is usually begun after May 15. As construction progresses, females may approach the, nest and be enticed to it or d.ri ven over· it by the male. Spawning tak.es place in a few seconds and the female moves quickly downstream. The re.lee.sed eggs settle among the stones and the male cont~nues to add more stones to the nest, thus ensuring additional protection for the eggs. The nest-building male. carries stones in his mouth or rolls 92 .. : .... _,.-.. .. -.. · .. ' -~ .- ·~. .· and pushes them with lips and snout. The total egg complement is not deposited in one nest at one time, since at each spawning a female deposits only ripe eggs. As many as 10 females may spawn in one nest. The nest mou:nd incr~ases in size as more stones are added by t .he male after successive spawnings. The sizes of nests are irregular and vary from. 30.5 em to 91.4 em vide, from 61.0 em to 91.4 em long (with the current), and from 5.1 to 15.2 em deep. The nesting sites may be used as spawning grounds by other species of minnows, even while the chub is still using the nest. 300. ~ Young chubs feed on cladocerans, copepods, and chironcmids. Adults eat mainly immature and adult aquatic insects, terrestrial insects, crustaceans, and plant material. Crayfish, vorms, and mollusks are also consumed by som.e species. 301. A.ge and grovth. The ultimate size attained by chubs varies with the genus. Total lengths of adults are generally 65 to 75·mm in · some ge:nera and 100 to 200 mm in others. Maximum length ranges from 90 mm for some genera to 260 mm for others. Males groW' tnore rapidly · ... · than teauues and rea.c.h a larger .maximum · si"ze. • · ,. Chubs in ta.ilwaters 302. The common name "chub" is used for fish of several genera. This discussion is limited to chubs from the genera Hybopsis, Gila, ~loc heil us , Nocomi s ., and Semot il us . 303. The various chubs respond differently in tailvater environ- ments and their presence or absence depends primarily on the charac- teristics of the particular. tailvater. In Tennessee, the river chub (Nocomis microPOgon) is abundant in the coldwater Apalachia and Chilhowee tailvaters, vhere it is used as forage by other fish species (Ptitzer 1962; Hill 1978). This chub bas disappeared from the Grand River below Shand Dam, Ontario, but it remains common in the river above the reservoir.· Delay of the spawning season because of a 7°C decline in m&Ximum temperature, from 28°C above the reservoir to 21°C in the tailvater, is cited as the reason for the river chub's dis- appearance (Spence and Hynes l97lb). The river chub has also dis.- appeared from the warmvater tailvater·s below tour hydropower 93 ··-... .. ' ..... ·• .. j !: impoundments on the Au Sable River, Michigan (Richards 1976), but the reason for this disappearance is unknown. 304. The roundtail chub (~ robusta) is found below Blue Mesa Dam, Colorado, and has apparently suffered no ill effects from the reduced temperature and turbidity in this tailwater (Mullan et al . 1976; Wiltzius 1978). However, this species of chub appears to be disappear- ing from a tail vater in New Mex i co (Mullan et_ al. 1976) •. In & tailwater in Utah and Wyoming, t .he round tail chub did no~ reproduce in two of three years following impoundment, presumably because or the reduction ·of summer water temperat.ure from 8. 3 to 2. 8°C that resulted from in- creased summer hypolimnetic discharges in 1964 and 1966 •. -. The growth ~ •• ~ ''o I 'II o ... -·· • : • o I o o rate o f this speci es has also declined since impoundment (Vanicek and · Kramer 1969). ·· The similar bonyt.a.il chub (Gila elegans) , once con- sidered a_ subspecies of the •oundtail chub, has disappeared from~ cold tailwater in Wyoming because water temperatures and turbidities suitable for this spec·ies no longer occur (Mullan et al. 1976) • . ' . 305. The humpback chub (Gila CYJ>ha) :was occasionally found in. an Arizona tailwater in 1967, but .by 1973 numbers of this. specie~ in .this ~old tail water h~d further declin~d, most lik-;~Y beca~se of habitat'_::; i changes resulting from dam construction and operation (Holden and Stalnaker 1975). 306. The hornyhead chub (Nocomis biguttatus) has a large popula- lation in the cold tailwater below Hoover Dam, Ohio, where temperatures r~~ge from 14 .4 to 21.7°C. Overcrowding in the tailwater may be aiding expansion of this species in the drainage, where it was rarely found before impoundment of the reservoir ( Cave.nder and Cru:rik:H ton 1974). Spawn i ng delays caused by the 7°C t .emperature reduction below Shand Dam, Ontario, have eliminated the hornyhead chub from this tailwater . This species is one of the more common cyprinids above the reservoir (Spence and Hynes 197lb). 307 . The creek chub (Semotilus atr omaculatus) has thrived in cold tailwaters. It is common in Hoover tailvater, Ohio (Cavender and Crunkilton 1974), and in the tailwater below Granby Dam, Colorado, in spite of a reduction in flows to only 11 percent of the hist orical 94 : ~. .. ~· .• .. .. ~ . .......... ·:·· .-; ... ·.· average (Weber 1959). Cold wate.r temperatures appear to have aided the creek chub in Shand Dam tailvater, Ontario. The. species has continue.d to reproduce successful.ly.here, whereas the reproduction of other cyprinids has ceased. This has allowed the creek chub to become numer- ous in the tailvater, although i t i nfrequently occurs above the ~eser­ voir (Spence and Hynes 19Tlb). 3o8. A 6-to 8-week delay in spawning caused by reduced tempera- tures bas leci to a decline in the reproductive success. and numbers of the peamouth chub (Mylocheilus caurinus) in a r.fontana tail water (~!ay and Huston 1979). The peamouth chub occurs and is occasionally creeled in the cold tailvaters belov Upper and Lover Salmon Falls dams, Bliss Dam, and C. J. Strike Dam in Idaho (Irving and Cuplin 1956). 309. The streamline chub (Hybopsis dissimilis) vas collected in. Barren Reservoir tailvater, Kentucky, during an electrofishing survey in March 1966 (J.P. Carter 1968a). The speckled chub (Hybonsis ~es.t ivalis) has disappeared ~om a cold tail water i n Texas, although it vas found in t he vicinity of the dam during: preimpoundJDent surveys and still survives in the river above the reservoir (Edwards 1978). Cold water temperatures and other .factors in the tailvater, caused by hypolimnetic releases, are most likely responsible for loss of this fish • 310. Reduced turbidity in tai~vaters may benefit some chub species. The turbidity-int olerant bigeye chub (Hybonsis amblons) is occasi onally found in Hoover Dam tailvater, Oh i o, perhaps due t o sl.i:glrtly lower tUl'bidi ty .levels .. _.This species is .seldom t:ound in the ·· "·"' .,.-"'"'" rest of the Big Wal nut Cr eek drainage (Cavender and Crunkil ton 1974) ·• · .. · ... 311. Bec·ause of 'the · lat-ge number of c hub · species and the wide .r·. · ~ange of their environmental requirements, generalizations on their success in tailwaters is difficult. Temperature changes and their effects on reproduction, along with changes in turbidity, appear to be ··most important in determining the p ·resence or absence of chubs. True minnows· 312. Habitat. The true minnows are a diverse group .vi th a variety . of habi tat preference. Many pre.fer lakes, ponds, or backwaters of 95 slow-moving streams with dense vegetation. Few species are found in reservoir tailwaters . They are tolerant of turbidit y as long as there is enough current to keep riffle areas free of silt. Many occur in . sc~ools n ear the river bottom, but they avoid-strong current.-- 313. ReProduction. Most minnovs spawn. between April and July. Some randomly spawn in shallow areas above gravelly or sandy riffles and others over silt bottoms; still others prepare nests in sand. or gravel areas which are guarded by the male. The following description for the bluntnose minnow (PimePhales notatus) taken from Pflieger (1975) is fairly typical for nest-building minnows. 314. The bluntnose minnow has a long spavning season, extending in Missouri fr::>m early May, when water ·temperatures are at least 20°C, to about mid-August. The peak of spawning is in late May and June. The eggs are deposited on almost any object (flat stones, boards, logs) that has a flat undersurface and lies on the bottom at depths ef 15 to 90 em . Usually sand or gravel, rather than a mud bottom, is selected for a nest site. Nest construction consists of excavating a small cavity beneat h the object selected and cleaning the undersurfac.es where the eggs will be deposited. In excavating ttie cavity, silt, fi'ne sand, and pebbles are svept away by violent motions of the tail fin, and larger objects are. pushed out with the tuberculate snout. The roof of the cavity is cleaned by the male with his mouth and the spongy pad on his back. · Only a sihgle male occupies a nest, but several males often nest in proximity beneath a single object. Several females may spawn in a single nest, vhich may ultimately contain more than 5000 eggs , Eggs hatch in 6 to 14 days, depending on water temperature, and. nevly hatched young are about 5 mm long . The male remains on the nest throughout the incubation period, driving away all other fish except females ready to spawn . 315. Food. The food of the mi ·nnovs is1 generalized and, includes both plants and animals; aquatic insects., small crustaceans, and algae predominate. 316. Age e.nd growth . Lengths of adult minnows generally range between 50and 75 mm, and maximum lengths betveen 75 and 100 mm. .Most average about 50 11111 long at end of first year of life and 75 mm at the end of the second. Few live longer than 3 years. Most are sexually mature in their second summer. 317. Males grow more rapidly and to a larger size than ·females for minnow species in which males guard the nest. Females are usually larger in species that do not guard the nest. True minnows in tailvaters 318. Seven genera of Cy}:)rinidae are commonly termed "minnows" (Bailey et al. 1970}. Two genera (Pimenhales and Phenacobi us) that reportedly occur in tailwaters are discussed in this section. 319. The fathead minnow ( Pimephales. nromelas} has a vide tempera- ture tolerance which makes it well-suited to the tailwater environment . It has increased in numbers in both the Navajo tailqter, New Mexico, and the Fontenelle tailwater, Wyoming (Mullai1 et al. 1976}. The fathead minnow is one of the remaining native species in cold Granby Dam tailwater, Colorado, where flows are only 11 percent of the historical average ('Weber 1959). The fathead minnow is the third most abundant species in the warm tailvate.r below Lake Carl Blackwell, Oklahoma,· and spawing activity was indicated in .mid-April when water tempera.tures reached 17°C (Cross 1950). The fathead minnow also occ·urs in Hoove'I' tailwater, Ohio . It apparently passed over the dam from the :reservoir above where it had been introduced as a bait fish (Cavender and Crunkilton 1974). This species vas also introduced into Dale Hollow tailvater, Tennessee, probably through its use as a bait fish (Bauer 1976). 320. The suc cess of the bluntnose minnow (Pimephales notatus) in cold tailvaters is varied . It is found in the cold Barr en Reservoir tail water, Ken t ucky (J. P. Carter 1968a) and is common in the drainage of Bi g Walnut Creek, Ohio, including Hoover Dam tail water (Cavender and Crunki lton 1974). The 'blunt nose minnow bas disappeared from the Grand River below Shand Dam, Ontario, presumably because of lack of spawn i ng ca.-used. by temperature reductions, since the spec.ies is com- monly found in the drainage above the reservoir (Spence and Hynes 197lb). 97 321. The bullhead minnow (Pimephales vigilax) vas present in the cold tailvaters below Canyon Dam, Texas, in 1976, but vas not collected from the river above the reservoir. No explanation vas apparent for the absence of this species in the 1976 collections above the reservoir since it had been collected here in othe·r years (Edwards 1978). 322. Adult suckerrnouth mi:mows ( P.henacobi us mi rabili s) we·re abun- d~lt in the river below Lake Carl Blackwell, Oklahoma, only during the · spawning season. Females in spawning condition occurred in pools below the dam in April at water temperatures. of lli to 25°C. Young of the year were abundant by mid~une in shallow· running water. By mid~uly, young suckermouth minnows had reached a length of 41 mm and vere only found in the s.wiftest flowing waters below the dam (Cro:ss 1950). 323. In three papers on tailwaters, minnows wer_e referred to in oniy general terms, without actual species being indicated. Pfitzer (1962) noted a general loss of minnow species in the cold tributary tailwaters of the Tennessee Valley. Diversity of minnow species de-· creased following the completion of a hypolimnetic release dam in Wisconsin in 1967. Although 8 to 10 minnow species were present at the three study stations belo.w the dam from 1965 to 1968, only 4 to 6 species vere found in 1969. Also, there vas a dec.rease in the overall abundance of minnows in this tailwater (Wirth et al. 1970). The authors ga:ve no reason for the observed minnow popltlation changes, although it would appear that cold va.ter temperatures were responsible. Mi.nnows were numerous in the warm tailvater below Elephant Butte Dam, New Mexico, 'W'he.re they made up 57 percent of samples collected in 1956 in seines and by electro shocking (Huntington and Navar:re 1957). Shiners 321!. Habitat. Shiners (Uotro"Ois) are the most abundant species vithin the Cyprinidae. They are a diverse group and the various species have a wide variety of habitat preferences. Many are found only in lakes, po·nds, or backwaters with little current, and others largely in small headwater streams. Relatively few shiner species are found in tail waters. s ·hiners in tailwate·rs prefer moderate to large- sized streams with relatively clear water throughout most of year, 98 ... . . ~- . ~ '· .· moderate or high gradients, and clean sand, gravel, .rubble and or boulder-strew bottoms. Some species are found in the deep, svif't riffles and in eddies and currents of pools immediately below such riffles. Other shiners prefer low to moderate current gradients and are mos.t abundant in shallow sandy pools. Some are found in schools in midwater or near the surface, and others near the bottom. 325. Reproduction. Shiners have varied spawning hab·its depend!.ng on the species. The following description for the common shiner taken ·from Scott and Grossman ( 1973) is generally applical;>le to most shiners. 326. Spawning usually begins when vater temperatures reach 15.6 to l8.3°C, usually in May or June. Spawning may occur over gravel beds in flowing vater; the fish may excavate shallow nests in gravel i.a .. . .. flowing 'Vater, or they may use the nests made by other fishes, even though such nests lack a current flow. They often spawn at the head or a gravelly riffle where the male establishes a territory. The females stay on the gravel below the males until ·ready .to spawn, then .move· upetrea,m and eventually spawn. The spawning act takes place in a · tractio~ of a second 'and is repeated many times, often· within a 'tf!!V ainutes, and there may be a constant succession of males and females . moving onto the spawning · site, spawning, and dropping back. Studies suggest that few eggs, probably not .mo·re than 50, are released at. each spawning. Tht! adhesive eo.~s drop to the bottom and become lodged between crevices of gravel. Eggs hatch in about 2-1/2 days at 21.1°C and larvae are about 5 mm long • 327. ~ Shiners have generalized food habits. They feed on aquatic and terrestrial insects, small crustaceans, arui algae. Larger individuals may prey on small fish. 328. Age and growth. Average length of adult shiners is variable . by species. Adults of some species range between 45 and 65 mm long and reach a maximum length of 70 mm; adults of other species range between 75 and 125 mm long and reach a maximum length of 175 mm. Males are the larger in some species, and females in others. ·Most shiners mature in their second or third summer; few live beyond age III. 9.9 ·. . ' Shiners in tailvaters 329. The shiners are a varied taxonomic group with many s~ecies. The majority discussed in this section are members of the g,enus NotroPis; however, two shiners from the genera Notemigonus and Richa=-dsonius are also included. ..· 330. The rosy1'ace· shiner (Notropis robellus) appears to thrive in ....... cold tailwaters. It is abunda:nt in Bull Shoals tailvater. Arkansas ~ -· (Brown et al. 1968; Hoffman and Kilambi 1970), and. is favored in· the tailve.ter environment below Hoover Reservoir, Ohio, by both the reduced temperature and the abundance of well-developed·riffles (Cavender and Crunkilton 1974). This species, togethe·r with the blackchin shiner (Notrop.fs heterodon) and redfin shiner (Notropis tDDbratilis), bas dis- appeared in the warm tailvaters below four hydropower facilities on the'Au Sable River, Michigan (Richards 1976). 331. The mimic shiner (Notropis volucellus) has become a common species below the same four bydropowe·r impoundments on the Au Sable· ·· Ri~er (Richards' 1976). This shiner has disappeared boca the cold :'~ Canyo~ Dam_ tailwater in Texas; although it is camnon in the river above !" the .. r~·servoir (Edvards·I978).' ·. · ' ··· ·: •:., '_-(;;:,:/;. ·: ;,. ·, ·• ... 332. The redside shiner (Richardsonius bait~at.u"s) does W ·ll. i~·. both varm and cold tailwaters. It thrives in water temperatures up to 26.7°C, which occur in tailvate.rs below dams on the 'Row River. Long Tom River, and Coast Fork Willamette River in Oregon (Hutchison et al. 1966). It i·s ·abundant in the cold tail waters below Owyhee Reservoir .and Antelope Reservoir • Oregon (Fortune and Thompson 1969). It also ~ppears to be maintaining its abundance in the cold Fontenell.e . . tailvater, Wyoming (Mullan et al. 1976). 333. Tne red shiner (NotroPis lutrensis) is fo~~d in both warm and cold tail waters. It is the most abundant species in Lake Carl ... Blackwell tailwater, Oklahoma (Cross 1950). Below Canyon Dam, Texas, it is the third most abundant species. Edwards (1978) believes that stabilization of va.te:r flows below this dam has increased red shiner abundance. 100 . . . -~ 334. The golden shiner (Notemigonus c·rysoleucas) and the spot tail shiner (Notropis hudsonius) are n.umero.us in. varm Holyoke Dam tailvater, Massachusetts (Jefferies 1974). In cold tailwaters, the golden shiner is found primarily due to its introduction as a bait fish. It is found in Dale Hollow tailwater, Tennessee, and appears to have moved from the reservoirs into both Hoover tailwater, Ohio, and Canyon Dam tail:water, Texas (Cavender and Crunkilton 1974; Bauer 1976; Edwards 1978). 335. Large .numbers of young eme::-ald shiners (Notropis atherinoides)-up to 800,000 in 24 hours--were lost in the discharge from Levis and Clark Lake, South Dakota (Walburg 1971). In the tailwater, this species is widely used a .s forage by game f'ish (Walburg et al. 1971). Thist species composed 16 perc.ent of the fish taken in Buckhorn tail water and vas also ·taken from Barren Reservoir tail water, Kentucky, in an electrofishing survey (Henley 1967; J. P. Carter 1968a). 336. The success of the spotfin ~•hiner (Notronis spilonteru:s) in cold tailwaters is varied. It occurs commonly in both Hoover tailwater, Ohio, and Barren Reservoir tailwater, Kentucky (J. P. Carter 1968a; ·Cavender and Crunkilton 1974). This shiner has disappeared below Sha.nd Dam in Ontario due to a delay in spavning cau.sed by cold water temper- atures (Spence and Hynes, 197lb). 337. A large number of other sl:Uner species occur in cold tailwaters. The duskystripe shiner (Nc•tropis Ei].sbryi) is the most abundant cyprinid in .Norfork tailwater, Arkansas. This species, along with the bigeye shiner (Notropis boons) and the whitetail shiner (Notropis galacturus), is abundant in Bull s·hoals tailwater, Arkansas (Brown et al. 1968; Hoffman and Kilambi 1970). In Hoover tailwater, Ohio, the str~ped sh.iner (Notropis cbrysocephalus) is abrmda:nt, while a species not found upstream, the sand shiner (Notroois stramineus), is common . Two other species, the rosefin shiner (Notroois ardens) and the silver shin.er (Notropis photogeni •s), are rare in Hoover tail water, although they are common. in the headwater streams above the reservoir (Cavender and Crunkil ton 1974). The silver shiner was also found in electroshocking samples in Barren Reservoir tailwater, and the common shiner ( Notropi s cornutus) was ta..'ten from both Barren 101 Reservoir and Nclin Reservoir tailwaters, Kentucky (J. P. Carter 1968a). Edwards (1978) found the blacktai.l shiner (Notronis ve.nustus) and Texas shiner (Notrop.is amabilis) in Canyon Dam tailvater, Texas. Though there was no change in abundance of the Texas shiner, the blacktail shiner '1."8.5 less abundant below the dam than in the stTeam above the reservoir, presumably because of colder temperatures. in the tailwater. Bauer (1976) identified the telescope shiner CNotropis telesconus) in fish samples collected from Dale Hollow tailvater, Tennessee. Stonerollers 338. Habitat. This fish is most abundant in streams having moderate or high gradients, well-defined gravel, rubble, or bedrock riffles, and permanent flow. It is generally found on riffles or in short, rocky pools where riffles and pools alternate in rapid succes- sion. It seems generally tolerant of turbidity. 339. Reproduction. The stoneroller spawns earlier in the spring than m0st other minnows. Spawning occurs on riffles where vater may be up to 30 em deep. The e.ggs are deposited in shallow pits dug by the males. Females remain in deeper vate;r near the spawning area, entering the pits indi...,.idually or in small groups to depo.sit their eggs. Spawn- ing occurs e.s one or more males press against the female,. The adhesive eggs become lodged in the gravel and are abandoned before they hatch. Eggs. hatch in 69-70 hours at 21°C. According to Smith (1935), nest building in Illinois started in mid-April, when vater temperature vas 12°C, and spawning continued until early June, whe.n temperatures were between 24 and 27°C . 340 . Food. The stoneroller lives in schools near the bottom.· It feeds primarily on algae and detritus that it scrapes from rocks, logs, and other submerged objects with the bladelike extension of its lower jaw. 341. As.e a,"1d growth. Adult stonerollers are c.ommonly 75 to 165 mm long, and maximum length is about 200 mm. In Missouri, tbey com- monly reach a length of about 35 to 60 mm ~J late August of the first summer of life (Pflieger 1975). Maturity is reached in the second or 102 · .. .. ~~ . " . ... _ .. ~ .. .. . :;o_ ··.~ .. .. .·~. .- ;•'' . . , .• third summer. Males grow more rapidly than females and attain a. much larger size. Stonerollers in tailwaters 342. The stoneroller is reported in a number of co·ld tailwaters. It. vas the most abundant species co•llected in Beaver tail water, Arkansas. It is also commonly found in the tailwaters of Bull Shoals Reservoir and. Norftlr.k Reservoir in Arkansas (Brown et al. 1968; Bacon e t &1. 1969; Hofflnan and Kilambi 1970). The stoneroller is abundant in Apalachia tailwater, Tennessee (Hill 1978), and is also found in other tailwaters of the Tennessee Valley system, whe·re it is used as forage ' by game ·fish (Pfitz~r 1962). Stonero·llers vere also taken in electro- fishing samples ~om Nolin tailvater, Kentucky (J. P. Carter l968a). The shallow gravel-bedrock substrate and 14. 4 to 21. 7°C tempe.ratures ba.ve favored the stoneroller in Hoover tailwater, Ohio. Large numbers : of stonerollers spavn in the shallow riffles below the da.m and both adults and young ar~ found in this tailwater throughout the year (Cavender and Crunkilton 1974). of< ··-~ ~ ..... ..; j il~·"-· , ~ ··;· ·. ~');· :)·;.··-:Dace a in tail waters ., -~-.;~:;~:,~:·:~~l~:r·: .1 ... t ~ Membe,rs of seven genera of cyprinids are referred 'to by the ·:~,; 1:' c011111l0tl name "dace." Only two genera, Phoxinus and Rhinichtbys, cQJD- prising four species, have been reported in tailwaters. 344. Tbe speckled dace (Rhinichthys osculus) appears to do well in some weste.rn coldwater tail water.s. It is abundant below Owyhee Dam . and Antelope Reservoir, Oregon (Fortune and Thompson 1969) . This species has survived below Blue Mesa Dam, Colorado, and has increased in Fontenelle tailwater, Wyoming (Kinnear 1967; Mullan et al. 1976) • •.. . ·~ ·~ 345. The longnose dace (Rhinichthys cata.ractae} has survived in , some cold tailwaters. It is commonly found below Owyhee Dam and has survived below Graney Dam, Colorado (Weber 1959; Fortune and Thompson 1969). ·It.has disappeared from the warm tailwaters below four sJD&l.l hydropower· dams ·on Michigan's Au Sable River (Richards 1976). 346. Two other species' of dace .bave been reported from cold ~-tailwaters. The southern redbel ly dac'e {Phoxinus erythrogaster) occurs in pools below Norfork Dam, Arkansas (Brown et &1. 1968) • and the 103 •' , .... ,· . ... -----;•-- ·, ----·-~-_.··----·-~ ~:-.·· --~ .. ;. .. -~ .. -.... --~-:------:"'--.:..~ blacknose dace (Rhinichthvs a.tratulus) in the headwa.ter streams of Big Walnut Creek, Ohio, and. occasionally downstream in the Hoover Dam tailwater (Cavender and Crunkilton 1974). Squaw1'ishes ·and chiselmouths in tailwaters · ... 347. Squawfishes and chiselmouths are found only in western North America . The northern squawfish thrives in the warm (up to 26.7°C) tailwaters below Cottage Grove Reservoir, Dorena Reservoir, and Fern Ridge Reservoir, Oregon (Hutchison et al. 1966). This species also ·appears to do well in most co~d tailwaters. It, together with the chiselmouth, is abundant below both Owyhee Dam and Antelope Reservoir in Oregon (Fortune and Thompson 1969). Both the northern squaw'f'ish and the chiselmouth are commonly creeled in a series of tailwaters on the Snake River, Idaho, including Upper Salmon Falls, Lower Salmon Falls, Bliss, and C. J. Strike (Irving and Cuplin 1956). A decrease in tenperature and a resultant 6-to 8-week delay in spawning is believed responsible for a reduction in northern squawfish numbers in a Montana tailwater (May and Huston 1979). 348. The Colorado squawfish appears to have been affected more . than the northern squaw'f'i sh by dam construction. The Colorado ,squawt'ish has disappeared "from a cold tailwater in Wyoming (Mullan et ~· 1976) and it is no longer fQund in a 105-km stretch of the Green River in Utah and Colorado (Vanicek et al. 1970). Tempe-rature reductions and alterations in normal flows may have eliminated it. It still. survives furt~r. downstream, but the alteration of seasonal temperature patterns since impoundment has reduced its growth rate {Vanicek a.nd Kramer 1969). Spawning migrations of Colorado squaw'f'ish in the Gunnison River in Colorado have been disrupted by lowered temperatures and reduced spring and summe r flows (Wiltzius 1978). Cyprinids in Russian tailwaters 349. Dam construction ·and ope:ration in the So·viet l,Jnion have affected many species of cyprinids. Historically, the bream vas the pr·edominant s,ec ies of cypri nid in the Volga River in the vicinity of the Kuibyshev Reservoir hydropcver dam (Diuzhikov 1961). Shortly after dam closure, the bream, together with the bleak, white· bream, and roach 104 I ·. ... (all cyprinids), were still nume-rous in the tailvater. However, un- favorable reproductive conditions in the tailvater caused by water temperature reductions, large diurnal water-level fluctuations, and a severe decline in spring flood flows, soon resulted in a decrease in abund.ance of all cyprinids { Sharonov 1963; Chikova 1968). Ide eggs, laid in the shallows, ·were lost through desiccation due to fluctuation in reservoir releases. Reduced temperatures in the tailvater delayed the spawning of bream, white bream, and zope. Between 24 and 50 percent of the females of t 'hese three species were found to be resorbing eggs in 1.963 and 1964 (Eliseev and Chikova. 1968). 350. Decreased. spring flood flows below the Volgograd hydroelec- tric facility have disrupted spawning conditions hundreds of kilometres downstream in the Volga Delta on the Caspian Sea. Spring spawning bream and carp have declined in abundance. However, the stabilizat:ion of flows throughout the year below this dam have improved the repro- ductive suc cess of the summer spawning white bream (Orlova and Popova 1976) .. 351. A reduction in mean ·water temperature of 4 to 5°C ha.'s f ·;.~ ~,':~·~·:· . :·affected many cyprinid species in the taUvater bel·ov Mingesbaur Hydroele.ctric Dam on the K·ura .River. Combined :f'yke net catches of gudgeon, bleak, podust, khramulya, and barbel {all c:yprinids) hav·e declined :from 73 percent before impoundment to only 40 percent after impoundment. In f&.ct, the barbel and the bystryanka, have completely disappeared. Thirty percent of the c.atch is nov composed of vobla, bream, and carp which have moved downstream into the .tailvater from the reservoir above. The al·~ered temperature regime has also shifted the spawning activity of the shemia from early summer to autumn. 'I'h:..-5 shift in activity has not affected s,avning success, howev~r; large numbers of shemia larvae and fry are still found in the tailvater ( Abdurakhmanov 1958) • 352. Reproduction of some cyprinid species has been disrupted by t 'he large flow fluctuati~:ms occurring below Narvskaya Hydroelectric Dam on the Narova Ri'ver. The spawning areas of the vimba have been sev:erely altered because of flow fluctuations, while the fry and eggs of the 105 ,. - f, I 1. golden shiner (European) have been destroyed through the dewatering of large sections of river bottom. Consequently, the numbers of both species have declined (Barannikova 1962). Catostomidae (Suckers) 353. The sucker family is largely restricted to North America. Suckers e.re one of the dominant groups of large fishes in fresh water, and in streams their total weight often exceeds that o-r all other fishes combined. In nUDber of species and individuals, they rank second only to the Cyprinidae. Each group of suckers has specific habitat prefer- ences, and most are bottom dwellers· vi.th similar but not i .dentical diets. All feed to some extent on larval and adult aquatic insects, small mollusks, small c·rustaceans, worms, and ~lgae .. All suc,kers spa\ltl in spring and none build a nest; eggs are scatte·reci in sui table 'habitat and abandoned. Preferred habitat of the various species .ranges from high, cold! mo\Ultain lakes and swift mountain stresms to warm, quiet ponds and lakes. 354. Most suckers are captured during spring spawning runs by use of gigs and snags; few are captured by book and line. Large numbers of some species are taken by commercial fishermen. The flesh of suckers has good flavor, but numerous sma11 bones detract from its value as food. Small suckers are an important source of forage for game fishes. 355. Those catostomids that commonly occur in tailwaters vil1 be discussed under three groups--buffaloes, suckers, and redhorse. Buffaloes. 356. Habitat. The three buffalo species, bigmouth, smallmouth, and black, have similar habitat requirements. They oc:cur primarily in the deeper pools of large streams, natural lowland lakes, and man-made impoundments. Buffaloes sometimes enter 'small streams to spawn, and the young rne.y remain here during their first stmiDer of life. Their distributional re.lation sugge,s.ts that the bigmoutb butt&J.o is 1110re tolerant of high turbidity than, the other two, and that the black buffa1o occurs most often in strong currents. lo6 .· ...... . : ·· .. •,•.: 357. Reproduction. The spavning habits of buffaloes are not well known. Buffaloes have been observed spawning in she.llow-water areas of rivers and reservoirs with a water temperature range of 15.6 to 18.3°C, between April and June. The• adhesive ·eggs are broadcast into the water, where they settle and adhere to the substrate-e.g., rocks and t'l.ooded vegetation. Spawning occurs in water so shallow that the backs of fish are often exposed. The eggs hatch in 9 to 10 days at a water temperature of about 16. 7°C. 358. Food. Studies by McCamish (1967) and others revealed that all ages of bigmouth buffalo feed principally on zooplank~on. The large, terminal mouth and numerous slender gill rakers are efficient devices for straining zooplankton from the water. 359. Zooplankton and attached algae were the principal foods found in smallmouth buffalo stomachs (McCamish 1967). Tt ~s species is primarily a bottom feeder, as indicated by the high frequency of insect larvae, attached algae, and associated detritus and sand in the stom- achs. The diet of the black buffalo is assumed to be similar to that of the smal1.mouth buf.falo. 360. Age and growth. T'.1e bigmouth is the largest 'bu:ffalo species. Adults are commonly 380 to 691 mm long and weigh 0.9 to 6.3 kg. Weights of 13.6 kg are no-t uncommon. Small.mouth and black buffaloes are somewhat smaller than the bigmoutb . 361. Buffaloes e.:re long-lived, many living more than 10 years. Females grow larger than males. According to Scho·ffman (1943) the average lengths and weights reached by the bigmouth buffalo in 'Reelfoot Lake, Tennessee, through the first eight summers of life were as follows: .. - 107 . ~ ....... ....... Ase in summers Lens!:b 1 mm 2 335 3 386 4 424 5 455 6 528 7 597 8 668 362. Growth in weight increases progressively eight summers, indica·ting that a fast-growing period of life. Buffaloes in taihraters fish in their eighth Weisht 1 k,s 0.7 1.0 1.1 1.4 2.4 3.4 6.2 dw-ing, the first summe:r are still 363. Buffaloes inhabit primarily lakes or large rivers and gen- erally are not important in tailwaters . In most instances vhere buffaloes a~·e found in tailvaters, they have migrated into tbe area from the downstream reservoir or river. in 364 .::;, Smallmouth and bigm~uth buffaloes are cOllllon SUlllller i.nbabi- tants of Levis and Clark Lake tailwaters in South Dakota and Nebraska. Fish collected in June had already spavned, and it vas assumed that their presence in the tailvater was related to feeding activity (Walburg et al. 1971). The increase of smallmouth b\lffal.oes in Dale Hollow tai'lwater, Tennessee, vas attributed to their migration upstream from Cordell Hull Reservoir (Bauer 1976). Buffaloes are important in t~e ~ler Rough River tailvater in Kentucky where they made up 9.3 percent of the fish catch by weight (Henley 1967). · This tailvater differs from most because it is essentially a long pool created by a mill dam 9.7 km downstream. These pondlike conditions closely dupli- cate the preferred lentic habitat of the buffaloes. 365 . The shallow, fast water in most tail waters, coupled vi.th a lack of recruitment, severely curtails or e·liminates buffalo popula- tions in most tailwat~rs. Black buffaloes were i!liportant in an Oklahoma tailvater shortly after impoundment. but were unable to sustain 108 ; . .. .. . :.·· : ....... .• . · .. themselves. This vas reflected by the rapid decline from 15 to 6 per- cent of total catch v.t'thin 2 years (Hall 1949; Hall and Latta 1951). 366. The food of buffaloes in tailvaters has not been studied extensively. Walburg ~~t al. (1971) found that bigmouth buffaloes from Levis and Clark Lake tailvater fed almost exclusively on zooplankton in the reservoir discharge, and smallmouth buffaloes fed on both zooplank- ton in the discharge and on attached algae and b~Jozoans found in the tailvater. Suckers 367. A number of sucker species occur in tailvaters. Most com- monly mentioned in the .literature are the northern hog sucker, spotted sucker, river carpsucker, quillback carpsucker, white sucker, longnose sucker, flannelmouth sucker, bluehead sucker, and largescale sucker. The first four species ar7 generally eastern or midwestern Unit-ed States in distribution, and the last four are extreme· northern or western. The white sucker is more videly distributed, except in the West and. South. Description of life history vill be limited to the ·northern hog sucker, spotted sucker, river and quillback ca.rpsuckers,···~ ... vhi te sucke.r, longnos,e sucker, and largescale sucker. Description of tail-water distribution includes all nine of the suckers mentioned above plus several other less common species. 368. Habitat. The northern hog sucker is an inhabitant of moderate-sized streams that have clean gravel or rock bottoms and ·permane~t flow. It is usually found on the stream bottom in riff les or in pools with noticeable current. The heavy bony head, slender tapering body, enlarged pectoral fins, and reduced svim o:adder permit it to maintain a position in svift currents vith little erfort. The northern hog sucker is nearly invisible on the stream bottom because of its strongly mottled and barred coloration. 369. The spotted sucker lives in lakes, over flow ponds, sloughs, oxbows, and clean s~uggish streams vith sandy, gravelly, or hard clay bottoms vi thout silt. It seems intolerant to turbidity, pollutants, and clay-silt bottoms . 109 370. Carpsuckers are common in large rivers, where they prefer deep, quiet pools and backwaters with moderate or low gradients. The river carpsucker prefers turbid waters with soft. bottcas, while the closely related quillback is found in clearer waters with finm bottoms. 371. White sucker hebitat is extremely varied, since the species occurs in both lakes and streams vi th low and high temperatures, lov and high turbidities, and fast and slow currents. The vhite sucker has fou."ld ma·n-made impoundments suitable, and bas become abundant in some. This species is especially characteristic of head.vater streams. 372. The longnose sucker occurs in the cold, clear water of both lakes and streams. Its occurrence in streams is usually related to spawning activity. 373. Le.!-gescale suckers live in lakes a:nd in large .rivers. They are often numerous in the veedy sho.reward areas of lakes, in backwaters, and in stream mouths. This species and the longnose s·ucker are often found together in the same general habitat. 374. Reproduction. The northern hog sucker spawns during spring near the heads of gravelly riffles in water 8 to 13 em deep, when water rl ... ·temperature reaches l5.6°C. Each female is attended by one or more· males. The demersal, nonadhesive eggs are deposited in a depression on the stream bottom and abandoned. 375. The spotted sucker spavn.s on riffles above large pools during the spring when water temperature ranges between 15 and 18°C. The eggs hatch in 7 to 12 days, depending on temperature. 376. Ca:rpsuckers are shallow-water, rand.ca. s ,pavners. They spawn in the spring when water temperatures reach about 21°C. Eggs are dispersed into the water column where they eventually settle to the stream bottom and adhere to the substrate. 377. White suckers spawn in the early spring. Adults usually migrate from lakes into gravelly st.reams when stream temperatures reach l0°C, but they are also known to spawn on .lake margins, or· quiet areas in the mouths of blocked streams and in tailwaters. Spawning sites 81"e usually in shallow water vi th a gravel bottom, but they may also spawn in rapids. No nest is built; eggs are scattered and adhere to the 110 ._.,.-. . · :.'; .. ··-.. '":• sravel or dri!'t· downstream and adhere to substrate in quieter areas. Eggs hatch in about 2 veeks, de,pending on temperature, and the young remain near the hatching site for about 2 weeks before moving to quiet areas along the stream bank or in a downstream lake . At this time, t .hey are 12 to 17 mm long. 378. Longnose suckers spawn in tbe early spr:i ng in streams where avatlable, but otherwise in shallow areas of lakes. They enter spawn- ing streams as soon as the water temperature exceeds 5°C. The spawning .run for this species reaches a peak several days bet"ore the run of white suckers into the same stream. Spawning often takes place in stream water 15.2 to 27.9 em deep, vith a current of 30 to 45 em/sec; and. a bottom of gravel 5 to 10 em in diameter. No nest is built; the · adhesive, demersal eggs are laid in small numbers and adhere to the gravel and substrate. Hatching and emergence of fry is similar to that reported for the white sucker. 379. Largescale suckers spawn in spring, usually in deeper sandy areas of streams vhere current is strong, but sometimes on gravelly or ·.sandy shoals in lake·s. '!'hey enter spawning streams when water· temper:..:-;~:: ature is 7.8 to 8.9°C and spawn a week or more later than the white sucker~ in ~he same streams. Spawning activity and hatching and' emer-· p gence of fry is similar to that reported for white-suckers. 380. ~ The northern hog sucker is an active feeder·, over- turning rocks and stirring up the bottom as it forages for immature aquatic insects and other bottom life vi th its fleshy, sucking lips. 381. The food of the s.potted sucker is said to consist mostly of mollusks and insect larvae. 382. Carpsuckers brovse extensively on attached filamentous algae: Other diet items include aquatic insects, worms, and mollusks. 383. White suckers have rather generalized food habits but subsist mos-tly on immature aquatic insects. 384. The diet of longnose suckers consists almost enti rely of algae, cbironomid larvae, amphipods, and other bottom organisms. Food o·f young fish includes illmature aquatic insects, copepods, c.ladocerans, and algae. lll ·----------="-----'----....!....------'L...--- . • .. - 385. Th~ di~t of the largescal~ suck~r consists almost entirely of bottom organisms such as aquatic insects, crustaceans, snails, and alga~. 386. Age and grovth. According to Pflieger (1975), northern hog suckers in Missouri streams reach a length of about 85 mm by the end of their first year of life and average 165, 246, 300, and 330 mm in their second through fifth years. Females grow mor~ ·rapidly than ~~s after. the fifth year and attain a larger maximum size. Males mature at age II and females at age III. 387. In Oklahoma. the spotted sucker attains a length of about 155 'IIDil in it.s first year and averages ·287, 338, 409, and 439 mm at the end of succeeding years. Maturity is reached at 3 years of age, and the maximum life span is about 5 years (Jackson 1957). 388. According to Pflieger (1975), river carps·uckers in Missouri average 81 mm in length by the end of their first year of life and 165, 229, 312, and 348 IIDil in succeeding years. Maximum life span is at l~ast 10 years. Average annual growth of the quillback carpsucker is .slightly gr~ater than that of the river carpsucker. . .. . . '. · · · 389. In Missouri, the white sucker averages 97 1IDD in length by the end of its first year of life and 173, 229, and 297 mm in succeeding years. Maximum length is about 508 mm. Fish mature when 3 or 4 years old; males mature a year earlier than females. 390. According to Brown (1971), longnose suckers in Montana average 76 mm by the end of the first year of lif~ and 140, 216, 267, 318·, apd 432 mm in succeeding years. The largest indi·vidual re.ported for Montana was 564 mm long and weighed 2.2 kg. Fi·sh .mature at 4 or 5 ..... -· years of age. 391. Grow-th of largescale suckers is generally slow . According to Brown (1971), grovth in Montana averages 51 nun by the end of the first yee.r of life and 89, 140, 190, and 254 mm in succeeding years. Specimens es old as 11 years have been reported. This suck~r matures when 4 or 5 years old . 112 .. ...... . . . ~ ·-·-~ .... Suckers in tail.waters 392. The effects of tailvaters on suckers have been varied, de- pending on the type o.f tailvater and the species of sucker. Taken as a group, suckers compose a significant segment of the fish population in many t~lwaters. 393 . The northern hog sucker is abundant. in many eastern coldwater tailvaters. Pierce (1969) found no change in the number s of hog suck.ers below Swmnersville Dam in West Virginia following c~osure. Hog suckers have remained extremely abundant in Chilhowee tailvater, Norris tailvater, and Apalachia tailvater in the Tennessee Valley, where young of the year are used as a food sourc e by game fish (Pfitzer 1962;. Hi ll 1978). The lover water temperatures (14.4-21.1°C), slightly reduced turbidity, mixed gravel-bedrock substrate, and higher dissolved oxygen level (always > 6. 0 mg/1) provide optimal conditions f o r pro- ducin g large numbers of hog suckers below Hoover Dam, Ohio (Cavend~r and Crunkilton 1974). Hog suckers apparently have 'been eliminated from a cold tailwater in Arkansas . They vere numerous here in 1950 vhen the ·project 1laS comple ted but had disappeared by 1959 (Baker 1959; Brown :~ .;'"-:"·~~-·· 1967). Reasons for the disappearance vere not specified. .'~----~:.,_,..._ -· _. · 394. The spotted sucke r is also common in some eastern tailvaters. It is one of the dominant species in Dale Hollow tallvater, Tennessee (Bauer 1976). It i .s also foW1d in moderate abundance in the cold tailwaters of Hoover Reservoir, Ohio, and East Lynn Lake, West Virginia ( Cave·nder and Crunk.il ton 1974; Goodno 1975). Occurrence in these tailwaters is mos t likely due to transport of f i sh over the dam from the reservoir above. Successful reproduction of spotted suckers occurs . . primarily in Hoover -Reservoir. Young of the year escape over the dam and are founci throughout-the length of lover Big Walnut Creek (Cavender and Crunkilton 1974). Adult spotted suckers are the fifth most numerous species lost ove·r the spillway of Little Grassy Lake in Illinois, com- posing 8 percent of total fish numbers (Louder 1958). 395 . The river carpsucker occurs in both col.d and varm tailvaters. It i s one· of t he major .r emaining native species found belov Canyon Reservoir, Texas (White 1969). It is also abundant in Levis and Clark 113- I I I tailwaters on the South Dakota-Nebraska border (Walburg 1971). The damming of Big Walnut Creek bas altered quillback distribution by blocking upstream migration and increasing spring concentrations in the tailwater {Cavender and Crunkilton 1974). 396. Ce.rpsuckers are among the most abundant fishes found below dams in the Rio Grande, New Mexico. They remain abundant in spite of the elimination of flows followi~g the irrigation season, which reduces the river to a series of isolated pools (Huntington and Navarre 1957). 397. The food of the ::-iver carpsucker in Levis and Clark Lake tailwate!' consisted primarily of zooplankton and algae (Walburg et al. 1971). 398. The white sucker has remained dominant in tailwaters below dams on many rivers where it was abun.d.ant before impoundment. The white sucker is adaptable to conditions in cold tailwaters which can closely resemble headwater stream habitats. It is abundant below Hoover . . . Dam and is one of three dominant species in Rocky Gorge· tailwater, Maryland ( Tsai 1972; Cavender an.d Crunldl ton 1974). In Twin Valley . Lake tailwater, Wisconsin, the carrying capacity of the white sucker increased threefold in the 3 years following impoundment · (Wirth et al. 1970). Even with a reduction of flows to only 11 percent of the historic ave::-age., white suckers have remained numerous in cold Granby tail water in Colorad.o (Weber 1959). They are also ab"..U'ldant . in the tailwater below Holyoke Dam, Massachusetts (Jefferi~s 1974). · ..... 399. On some rivers, the construction of reservoirs has allowed the white sucker to increase in numbers or become dominant in drainages where it was pre\~ously of only minor importance. Increased a~undance of the white sucker in Dale Hollow tailwaters, TeDDt!SSee, is attributed to the deeper, more lentic habitat found in the lower tailwater, caused by the downstreac impoundment of Cordell Hull Reservoir on the Cumberland River (Bauer 1976). 400. !n the Taylor and Gunnison rivers in Colorado, the reduction in temrera'ture and turbidity below some reservoirs has allowed the white sucker to outcompete most other native suckEr species, e.g., flannelmouth and bluehead suckers (Mullan et al. 1976; Wiltzius 1978). ... • r ... ~ .... The whit.e sucker is apparently able to reproduce in both of these reser- voirs and te.ilwaters, which accounts !"or its dominance over the native suckers in the Gunnison River drainage (Wiltzius 1978). Tailwater con- ditions apparently aid growth, since the condition factor (weight in relat,ion to length) of. white suckers increased in Blue Mesa tailwater following impoundlcent (Kinnear 1967). 401. The longnose suc.ker has become abundant in the tailvaters of two Colorado dams, and it has also displaced the nat:ive sucker species (Mullan et a.l. 1976; Wiltzius 1978). The reduced temperature and turbidity in these tailwaters, together with the longnose sucker's ability to reproduce successfully, may give this s.pecies a competitive advantage. 402. Longnose suckers also adapt well to reductions in flow below dams. They have survived in flows amounting to only 11 percent of the annual average (Weber 1959). Additionally, they remain in two Wyoming tailwaters, in spite of a seasonal reduction of discharge from 24.5 to 0.04 m3/sec and 8.8 to 0.01 m3/sec, respectively {Wesche 1974). The effects or reservoir construction and operation in· · .... ·-~.~- ·western rivers on the historically dominant flannelmoutb and bluehead .suckers bave 'been mixed. The flannelmoutb ~;;ucker remains tbe most .numerous native species below Glen Canyon Dam, Arizona, and the blue- head sucker is still present in Granby tailwaters {We.ber 1959; Mul.lan et al. 1976). Numbers of flannelmouth sucker have declined. in a Wyoming tailwater because of reduced water temperatures {Mullan et al • .. . 1976). Abundance of both the flannelmouth sucker and the b1uehead sucker has decreased in a New Mexico t .ailwater because of reduced temperatures and competition from trout (Mullan et a.l. 1976). However, 29 km downstream, wbe.re the river returns to 1 ts warmer, more turbid c'ondi tion, the flannelmouth sucker still predominates (Graves and Haines 1968, 1969). 404. The most serious reduction of bluebead and flannelmouth suckers has occurred below two Colorado tailwaters W'here cold water temperatures and reduced turbidity have allowed other species to ov.t- compete them. The result has been the disappeara.nce of both species 1.15 '·· .. - I r I I. o. from one of the ·tailvaters and their general decline. in the· other {Mullan et al. 1976; Wiltzius 1978). 405. The reduction of water temperatures in a Montana tailvater -· · bas delayed reproduction 6 to 8 weeks for both. the largesc.ale and long- nose suckers. Delay in spawning has decreased the abundance of large- scale suckers in this cold tiailwater {May and. Huston 1979). N-itrogen embolism 'below this site has caused sane mortality of largescale suckers. Largescale suckers are extremely abundant below Dorena Dam, Fern Ridge Reservoir, Lookout Point Dam, and Dex-ter Dam in Oregon, apparently thriving in the 21 to 27°C temperatures commonly attained in these warmvater tailwaters {Hutchison et al. 1966). 406. ·The effects of reservoir ope~ations on other less widely distributed sucker species have also been mixed. Lowered water temper- atures in a Wyoming tail water have eliminated the humpback suc-~er, ·wbi~e allowing the mountain sucker to stabilize or even increase its population (Mullan et al. 1976). 407. The blue sucker of the Missouri River is abundant in the ._::_-spring and fall ·in' t:he warm Le_wis and Clar.k Lake tailwaters on the ·.South Dakota-Nebraska border . Here it feeds heavily on zooplankton and algae and, COIIDDonly eats the bryozoans and aquatic insects that are· also available (Walburg et al. 1971) . . Red horse 408. Five species of redhorse are most often mentioned in litera- ture on-:tailvater fish populations. They are the black, golden, silver, ....... · .. s~_orthead, and river redhorse. Generally, little factual information is available on their spawning habits, age, growth, or other life history features. 409. Habitat. The various redhorses have slightly different habitat requirements. They usual ly inhabit clear streams having per- manent flov and clean gravelly or rocky bottoms. The black redhorse, golden redhorse, and river red horse, are most abundant in st:reams of medium size, whereas the shorthead redhorse and silver redhorse are ,most cOI!Dllon in larger rivers. The black redhorse is more abundant than the golden redhorse in the cooler and swifter streams. Where, the 116 . ·~ .. '· • tvo occur together, the black redhorse tends to predominate in short, rocky pools with current; whereas, the golden redhorse is most abundant in larger pools and backwaters without noticea'ble current . 410. Tbe river redhorse seems less tolerant of turbidity, sil ta- tion, and. intermittent flow tba:n the other redhorse suckers. The golden redhorse is the most tolerant of both turbidity and intermittent flow, vhile ·the· silve·r redhorse avoids spring-fed streams having high gradients and those that have excessively high turbidity . The short head redhorse is the most adaptable in its habitat requirements. 411. Reproduction. Redhorses spawn in April or May when water temperatures reach about 13.3°C . Spawning occurs on riffles in :water 15.2 to 61 em deep and over a bottom of small rubble mixed with lesser amounts of small gravel and sand. No nest is buil t; eggs are scattered over the gravel and abandoned. Spawning is completed on any given riffle within about 4 days . 412. Food . Young-of-the-year redhorse feed in backwater areas on algae and small crustaceans . Older fish shift their feeding activ- . · · ities to riffles, where they forage for aquatic insect larvae and ····;;-.~·· ~·· ... : · othe·r small bott.om-dwelling invertebrate·s. The river redhorse has molarlike tee·th that are an adaptation for the crushing of mo l lusk shells. The· diet of this f i sh consists of mollusks, other inverte- brates, and plant material. 413. Age and growth .. Accord ing to Pflieger (1975), shorthead redhorse in Missouri streams reach a length of 107 mm at the end o f their first year and in succeeding years lengths '!l.re 193, 264, 305, an.d 335 mm. The maximum life span of most redhorse is 9 or more years, except for the golden redhorse which usually does not live beyond 6 or 7 years. 414. Age and growth of the redhorses are simi l ar except t .bat growth of the shorthead is more rapid than that of the golden and black but slightly slaver than that of the silver. The river redhorse usually grows slaver than the other species during the first few years of life, but eventually it overtakes and. surpasses them; it also .has a longer life span. 117 Redhorse in tailwaters 415 . The response ot: redhorse to tailvater condi'tions ia similar' to that of other suckers. Redhorse have increased in a number of' cold tailwater s in the Tennessee Valley (Pfitzer 1962). Redbo~se species were also abundant below Buckhorn Dam, Kentucky, where they numerically composed 49.8 percent of all fish present (Henley 1967). They were abundant.-in an Arkansas tailwater shortly after dam closure in 1950, but had disappeared by 1959 {Baker 1959; Brown et al. 1968). Cold water temperatures apparently eliminated the redhorse from this tail water. 416. The construction of Cordell Hull Reservoir has increased water depth in t 'he downstream section of Dal e Hollow t ail water, Tennessee. This alteration allowed the golden redhorse, a pool species, to become more abundant {Bauer 1976). In the shallower tailvate,rs below Hoover Re.servoir, Ohio, the golden redhorse is seldom seen despite being the most abundant redhorse in the drainage. (Cavender and Crunkilton 1974). Similarly, the golden redborse is of' only minor .'~ili.portance in terms of numbers and biomass· in East Lynn Lake tailvaters ;· ... tn we"st vt~sinia · (Goodno 1975>. · '· ·~:. :.i-::1 ~: :~:<·-::_·.:·:~ · ,~:. ·. ;•:·.:>.' 417. The distribution of silver redhorse in Big Walnut Creek was seriously influenced by construction of a reservoir . Although both young and adults were abundant in the reservoir headwaters prior to dam closure , they have subsequently disappeared. ~e presence of the dam has prevented the return of the silver redhorse to its former hz:?i t:S:t (Cavender and Crunkilton 197 4). Below Dale, Hollow Reservoir, however, a popul ation of silver redhorse has become est ablished and has been increasing (Bauer 1976). 418. Black redhorse are commo n in the t.ailwater below Hoover Dam in Ohio. Many were observed in shallow pools below the dam in early May, which suggests that they were attempting to spawn {Cavender and Crunkilton 1974) . . 419. Little has been reported on the ri'ver,·shorthead , e.nd gray redborses . The gray redhorse has survived in the cold tailvaters below Canyon Dam, Texas, and is one of the most numer ous native species 118 .. ., .. .. . ··~ . ·· .. ~~-~· .· ... :-... ·· ... -. •. '·- !' remaining (White 1969). The river reaborse bas increased in the lover tailvater of Dale Hollow Reservoir, Tennessee, since its inundation by Cordell Hull Reservoir (Bauer 1976). The shorthead redhorse is abundant in Levis and Clark Lake tailvate,r, South Dakota and Nebraska. Here it feeds primarily o.n zooplankton from the reservoir discharge, but ·algae, bryozoans, and aquatic insects are also C011Dftonly eaten (Walburg et al. 1971). Ictaluridae (Catfishes) 420. The catfish family includes 37 species, all generally re- stricted to 'North and South America. In the United States, they occur naturally in larger rivers, lakes, and slow-moving waters eas't , of the Continental Divide. Some of the larger species have been .widely int-ro- duced outside their natural range and now occur throughout th~ United States. Species commonly found in tailwaters are the channel catfish, flat-bead catfish, and, to a lesser extent, bullheads; the blue catfish :. ·· and the white catfish are important regionally. All a:r·e popular r~d and game fishes. Lesser known species which also occur in tailvaters '~:t ..... '.':"~ ·..,:_. ... are collectively referred to'as "madtoms ." They are rarely seen "be- cause of their small size and secretive habits. This discussion of ictalurids includes the three general groups--bullheads, catfishes·, and mad toms • Bullheads •' .. .. · ...... 421. Habitat. The black, brown, and yellow bullheads· may occur · in tailvaters, but they are essentially quiet-water fishes found in lakes, ponds, or sluggish streams. They occur in a variety of habitats, but are· most abundant in areas with turbid water, a silt bottom, a:1d D-' ~1 oticeable current O!" strong flow. Especially favorable babitats are the permanent pools of small intermittent creeks and muddy oxbows and backwaters of large streams. Black bu.1lheads usually inhabit the lover sections of small-to medi.um-sized streams of low gradient, .. ponds, backwaters of larger rivers, and silty, soft-bottomed areas of lakes and impoundments. They do not inhabit the areas in which brawn ll9 ... and yellow bullheads usually occur but seem to replace them if the habitat deteriorates. Yellow bullheads seem to prefer clearer water. 422. ReProduction. The reproductive habits of' the bullheads are similar. They spawn in late spring or early summer, vhen, water temper- atures reach about 21°C, in saucer-shaped nests fanned out by one or both parent fish. Nests are usually beneath some type of cover such as logs or objects elevated above the stream bottom. One of the parents remains continuously with the eggs until they hatch. f-1"innovs and sunfi s .h e.re often observed near the nest, rushing in to eat eggs when- . ev.er the opportunity arises. Eggs hatch in 6 to 9 _days. at 20.6 to 23.3°C. At hatching, the young are about 6 mm long; they remain in the nest until about the seventh day. The young bullheads move about in a compact school after leaving the nest, and continue to be accompanied by one or both adults. The young are abandoned by the adults when about 25 mm long but persist in schooling throughout the first summer of life. 423. Food. Bullheads are truly omnivorous and feed primarily from . the bottom on a variety of plant and animal material. Adults eat immature aquatic insects, clams, snails, crustaceans, plant material, leeches, and fish. Young up to 25 mm long feed almost exclusively on small crustaceans. Older young feed on chironomid larvae, clu.doc.erans, ostracods, ~phipods, and mayf~ies. 424. Age and growth. In Oklahoma vaters, the black bullhead averages 94 mm in length by the end of its first year of life and is about 170, 229, 274, 312, and 350 mm long at the end of succeeding years (Houser and Collins 1962). Growth in various ha:bitats is highly variable, however, being slowest in overpopulated ponds and streams and fastest in ne:w impoundments. Bullheads· may grow to 250 mm duri.ng the first year of life in new reservoirs, but may not reach this length until ~he fifth or sixth year in an overpopulated stream. Maximum life span is about 10 years, but few individuals live more t .han 5. Yellow and brown bullheads attain slightly larger size than the black bullhead. 120 -•: -J-o• I ~·. : .• ·. & ..... ·- •.· Bullheads in tailwaters 425. Bullheads may occur in tailwaters, but this habitat is generally unsuitable because• of reduced turbidity and strong currents. Bullheads found in tailwater•s are often produced in backwaters or in the upstream reservoir and c.arri.ed through the dam into the tail water. 426. In some warm tailwat~rs, bullhea.ds coming from the reservoir above are an important part of the fishery. A 1959 preimpoundment rotenone survey of Barren River, Kentucky, estimated bullheads to be 0.5 percent of the river fish community; few were taken by anglers (Carter 1969). In a 1968 creel survey at the Barren tailwater, it vas estimated that 33.8 percent of the anglers' catch vas bullheads •. Fishing success in the tailvater during 1968 was correlated with dam discharge; fishing was good when disc·harge ·W'as high and. poor when it vas lov (Charles and McLemore 1973). Good fishing success for bul.lheads during periods of high flov and lov historical bullhead populations in the ·· · river, indicate that the bullheads were probably produced in the res.er- voir and transported through the dam into the tailva.ter. The creel :. censu s in the Barren tailvater from 1968 through 1971 sho~ed a steady.·,::.·~ •·' · . ~ .. decline in the bullhead catch from 33.8 to 0.3 percent o .f the total. .harvest (Charles and McLemore 1973). This decline .not only indicates that the 'bull.head population vas unabl e to reproduce in the tailva.ter environment, but also reveals a lack of further recruitme~:+. into the tail water from the reservoir above. 427. Be,low Carlyle Dam, Illinois, bullheads made up 6.2 percent of the 1968 catc h by anglers. The bullhead harvest ·was 0.38 fish/hour in the reservoir compared with 0.08 fish/hour in the tailwater (Fritz 1969). These figures indicate a greater abundance of bullheads in the reservoir. Apparently bullheads favor the reservoir habitat over the florlng tailvater. Bullh.ead.s from Fern Rid!e Reservoir, Oregon, have established a fishery in the -warm tailwater (Hutchison et al. 1966). 'Hall and Latta (1951) found that 8 percent of the fish in the stilling basin. belov Wister Dem, Oklahoma, were black bullheads~ They believed that the fish found in the warm tailwater, including the bullheads, reflected the species composition of the reservoir above and not the 121 river downstream. The black bullhead is abundant in the Stillwater Creek drainage in Oklahoma, but occurs infrequently in the Lake Carl Blackwell tailvater. This is mo ;,t likely due to the strong currents in tailvaters, which do not provide suitable bullhead habitat (Cross 1950). 428 . Bullheads are found in tailwaters vith vide temperature ranges and lov dissolved oxygen concentrations. Electrofishing shoved that black bullheads vere the most abundant fish and ranked third in .. biomass in East Lynn tail water, \olest Virginia, where water temperatures ranged from 5 to 25°C and averaged 16.6°C (Goodno 1975). Pierce (1969) stated that bullheads became more abundant after construction of Summersville Dam, West Virgj.nia, even though tailvater temperatures did not exceed l5.6°C. Summers (1954) captured builheads by trap net below Tenkiller Dam, Oklahoma, in vater of 12.2 to l7.8°C, svi,ft cur.rents, and dissolved oxygen concentrations of less than 0.5 mg/1. Possible increased production of bullheads in these reservoirs and their export into the tailvaters may account ~or the large numbers of .bullheads · '. · ,; . _ -pbserved. . ~': . ": : ~::~~ . : . ·::-~ .. · ~ ,: , .,_ '.f/~~;.{ _1"·=· ' .. 429. Only tvo papers reported food habits of bullheads in · tailwaters. Olson (1965) stated that bullheads in Navaj4:> ttt.ilvater, Nev Mexico, ate a diet similar to that of rainbow trout, 'consisting of dipterans, plecopterans, ephemeropterans, algae, and fish; only gastropods (eaten by trout) vere excluded from the bullhead diet. Brown bullheads below Holyoke Dam, Massachusetts, ate (by volume) 37.4 percent detritus, 35.3 percent algae, 11.8 percent pelecypods, 11 percent fish, 2.2 percent bryozoans, and 2.4 percent other items (Jefferies 1974). Catfish 430. Habitat. Four catfish species have been collected .from tailwaters: channel catfish, flathead catfish, blue catfish, and vhite catfish. The last tvo species resemble the channel catfish and are regional in distribution. The white catfish is fo,und in Atlantic coastal streams from Nev Jersey south to Florida, and the blue catfish occurs only in the Missouri and Mississippi rive;s and their principal 122 ., .. , '· . ' . . ; ... ~; . ' tributaries. Tbil discuuion concerns only the cha.nnel catfish. The flathe•d catfish may occ·ur in ·tailvaters, but is 11 ttle I!1entioned. in tailwater literature. 431. The channel catfish is found in a variety of habitats but 11 most common. in large stream.s having low or moderate gradients. Though c·hannel catfish prefer currents, excessive current or insuft'ic.ient water· depth limits the.ir distribution. Adults are usually found in deep-water pools or near submerged logs and other cover. The young are commonly found in riffles or the shallower parts of pools. Adults are most active at night, when they move into shallow water to feed. 432. Reproduction. The channel catfish spawns in late spring or summer when water temperatures are between 24.0 and 29.4°C. Catfish spawn in nests selected by the male. Nest sites are natural cavities under submerged logs or debris, animal burrows, or undercut stream banks. The eggs hatch in about a week and the fry remain in the nest for 7 or 8 more days. The male guards the nest until the fry leave. Because of predation, the survival of catfish during their first-summer •; I . of life is· usually greater in turbid than in clear ponds or streams. ..-. · 433. Food. Channel catfish feed mostly on the bottom and locate food primarily by taste. Bailey and Harrison (1948), who conducted studies on the food habits of channel catfish in the Des Haines River, Iowa, found that fish less than 100 mm long fed almost entirely on small insects. Larger catfish had a more varied diet, including fish, insects, crayfish, mollusks, and plant material. 434. Age and growth. In the Salt River in Missouri, the channel catfish averages 65 mm in length at the end of its first year o~ life and is about 135, 206, 259, 297, 340, and 399 rnm long at the end of succeeding years (Purkett 1958a}. Studies by Barnickel and Starrett (1951) in the-Missouri-Illinois section of the Mississippi River showed that channel catfish mature when ' 4 or 5 years old at a length of 305 to 381 mm. Adults were commonly 305 to 815 aun long and weighed 0.4 to 6.8 kg. Life span is apparently dependent on growth rate. Slow-graving fish in Canada may live longer than 20 years; whereas, fast-grcwing fish from the southern United States may not live longer than 6 or 7 years, 123 ... I ~~ I I I i; r I Catfish in tailvaters 435. Channel catfish ere important game fish in ·some t&ilvaters, particularly in the southeastern United States. Change.s in stream habitat caused by dam construction have had varied effects' on channel catfish. They are often abundant in the warm tailwaters of. turbid main-stem or tributary rivers but are uncommon. or absent in clear, cold tail waters. 436. Channel catfish disappeared from the angler catch in a M1ssouri reservoir because of coldwater discharge (4.4-15.6°C) from an upstream reservoir {Hanson 1969). Below an Arizona dam, channel catfish were second in importance to rainbow trout until the vater cooled from 14. 8°C in 1967 to 10. 4°C in 1971 and. 1972. The cooling trend caused channel catfish to leave the tailvater (Mullan et al. 1976). 437. Channel catfish abundance below a Kentucky reservoir appar- ently decreased after dam construction ( Carte·r 1969). Water is re- ,leased from the hypolimnion to maintain a tail~ter trout fishery . C~el catfish c~posed 35.4 percent of the ~ngl~rs;' catch in the tailwater area before impoillldment 'in 1959, but never more than 2 per- cent after impouncbne.nt (Carter 1969; Charles and McLemore 1973). The change .. from hypolimnetic releases in 1968 and 1,969 to epilimnetic releases in 1970 and 1971 did not noticeably affect the channel catfish catch. Both release regimes resulted in water temperatures below those o.f th!Ot.historical monthly average for··tbe Barren River (Carter.l969). 438 .. Do'lll1stream from a cold tailwater, as local conditions moderate the dam discharge, catfish populations may inc·rease . Graves and Haines (1969) stated t hat channei catfish were the main game fish 29.0 km below a New Mexico dam vhere turbidity and water temperature had increased. Further upstream in the tail water, trout predomi,nated. 439. Turbid main-stem teilwaters and tributar y warmwater tailwaters often concentrate catfish and e reate a fishery. Cross (1950) believed that char~el catfish below Lake Carl Blackwell, Oklahoma, had escaped from the reservoir and remained in the tailwater because food was abundant and flows were stabilized. In June and July, an 124 -··.-.. :.-...·-.:-f· upstream migration of channel catfish in Stillwater Creek resulted in a large concentrati,on in the Blackwell tailwater. Channel catfish vere also most abundant in Levis and Clark tail water, South Dakota, during the SUIIIDer ('Walburg 1971). Channel catfish concentrate and provide a fishery below Lock and De.m Number 12 on the Mississippi River in Iowa (Gengerke 1978). This species is second in biomass and fi .fth in abun- dance in East Lynn Lake taih;ater, West Virginia. ( Goodno 1975). A comparison of the fish population in t .he stilling basin area before and after· closure of the warmwater Wister Dam, Oklahoma, showed a reduction in the channel catfish population from 34 percent of the total numbers of fish to 11 percent (Hall and Latta 1951). The reduc- tion reflected the change of a community associated vith a river to one 'IDIOre c0111Donly associat.ed vi th a reservoir. 440. Little information. is available on th.e fooC! and growth of ~hannel catfish .found in tailwaters. In Levis and Clurk tailwater, they ate fish, crayfish, and aquatic insects . In April and May, when Hexagenia was abundant. in the tailwater, it was a major food item, but ... as abundance of He.xagenia declined in the stDIIDer, catfish ate mo're · fish ··-~. (Walburg 1971). ··Catfish in Dale Hollow tailvater, Tennessee, ate pri-•· marily dipterans (Littj,e 1967}. · Channel catfish grew faster in Levis and Clark Lake than i.u the tailwater (Walburg 1971). Mad toms 441. Habitat . Had toms inhabit clear to moderately turbid streams having permanent flow and low or moderate gradients. Some species are also found in the shallows of lakes and their outlets. 'rhey generally occur on riffles over a gravelly or rocky bottom. Some species may ·· also be found over sandy bottoms in areas vi th fast current, and others, such as the brindledmadtom,in pools below riffles associated vith organic debris such as roots, leaves, tvigs, and logs. Species vary in their tolerance of current and turbidity. Madtoms a:re most active at night, often hiding by day beneath large rocks or other cover. 442. Reproduction. t;fadtoms spawn .from late spring into the sum- mer. Peak spavning occurs when water temperatures reach 25 .6 to 27.8°G. 125 ----......_. ______ _ .. Eggs are deposited as a compact cluster in a shallow depression exca- vated beneath a flat rock or other forms of protection such as tin cans, boards, or crockery. Nests are guarded by one of the parent fish. 1~e,dy hatched young are about 10 mm long. 443. Food. Madtoms are active at night, foraging over riffles and shal low pools. They feed primarily on insect larvae and small crustaceans that live on the bottom. Occasional small fish are also eaten. 444. Age and growth. Little information is available on gr-:>vth of madtoms. Carlson {1966), who studied the stonecat in the V~rmill ~~n River in South Dakota, found that they averaged 79 mm at the end of the first year of life and 99, 114, and 137 mm by the end of succeeding years. The largest specimen was 193 mm long and in its seventh year. 445. Other species of madtom are smaller than the stonecat and their total length at the end of their first year ranges from 36 to 64 mm. Adults of most species are 51 to 102 mm long and maximum length is about 127 mm.· Many mature in their second s-ummer and few live longer than 3 years. :)' MadtO..s in tail waters 446. There are few reports on madtoms in t ail waters. Carter {1969) found the brindled madtom in the Barren tailwater, Kentucky, in 1964 and 1965. Four species of madtoms--the slender, brindled~ freckled, and an undescribed species-were collected. in the Barren River during a preimpoundment survey in 1959. In the Owyhee River, Oregon, tadpole madtoms were commonly found in both the tailvater belov Owyhee Dam and in the reservoir above {F"ortune and Thompson 1969). Stonecats vere collected below four impoundments on the Au Sable Rive:-, Michigan, in the 1920's, but none ·were collected in a 1972 survey {Richards 1976). Percichthyidae (Temperate Basses) -'·· 447. These basses include several freshwater and marine genera. The white bass and the less common yellow bass 'bot'h occur naturally 126 .. · , . . . ".•. .. . . • J in the Miss.iss.ippi River drainage. The· striped bass was introduced into many reservoirs during the past 20 years .from anadromous stocks native to coastal rivers and estuaries in Virginia, North Carolina, and South Carolina. These plantings have been successful in some states, &ad. striped bass are nov relative·ly common in some river-reservoir systems. Examples of states vith successful striped bass fisheries in both reservoirs and tailvaters are Oklahoma and Tennessee. Reproduction of introduced striped bass has been limited, and the maintenance of fishable stocks is often dependent on annual plantings of hatchery- produced fish. The life history discussion of temperate basses is lici ted to the white bass; the tail water discussion incl udes both ·•·· white bass and striped bass. White bass 448. Habitat. White bass inhabit the· deeper pools of streams and the open waters of lakes and reservoirs. During the spring spavn- ing migrati.ons, large· numbers. enter tributary streams and are the. basis for an important seasonal fishery. The white bass ten ds to avoid .,.,.. . ' . '·,, .. .. vaters that are continuously turbid and. is most often found over a tina sand, gravel, or rocky bottom. It bas 'been introduced into many lakes ·. and reservoirs throughout the United States. 449. Reproduction. The white bass spawns in earl y spring and spawning is usually preceded by tht~ migration of mature adults into tributary streams. The prespavning schools are composed of only one sex. · Males move onto the spavniog grounds about a mont.h before the . females. The spavning ,Period throughout their range is from April ~ .. through June, when water temperature is about 15.6°C. Spavning occurs in midvater or near the surface, over a gravelly o.r rocky bottom, often in a current, and vi thout preparation of a nest. Spawning is generally completed at any given locality over a period of 5 to 1 0 days; there is no parental care of eggs or young. The eggs settle.to ·the bottom, where they hatch in about 2 days. Ne-wly ha.tched larv.ae are about 3 mm long. 450. Food. White bass usually feed in schools, appea·ring in ·large .numbers where food is abundant and moving on vhe.n the supply is 127 ---·-----...... ---~~ ----... ··---·-··-·-----·-.. _.,. ___ , lilt •. ..,.,::!);~ ,· ... ... exhausted. Most feed near the water surface in early morning and late evening, vhere they pursue forage fish, small crustaceans, and the emerging stages of aquatic insects. Zooplankton and aquatic insects s.re the most important diet items for young vhite bass. The diet of adults is largely composed of fish; however, zooplankton and insects are also important. 451. A~e and growth. Growth of the vhite bass is rapid; the life ... span is seldom more than 4 years in southern vaters, but may be 7 or 8 years in northern vaters. In Lake Wappapello, Missouri, tbis fi.sh reached a length of about 185 mm its first year and averaged 302, 338, and 358 mm by the end of succeeding years (Patriarche 19-53). Fev vhi te bass attain a length and veight of more than. 445 mm and 1. 2 kg .. Temuerate basses in tailvaters 452. The vhi te bass is a.n i.nportant. sport fish in many tailvater·s, especially during the spring. In the varm tailvaters of Clearwater Lake, Missouri, it made up 3.9 percent of the anglers' catch in 1961 and 14.1 percent in 1964 (Fry 1965; Hanson 1965). It composed 8 per- cent ~r the anglers' catch belov Pomme de Terre Reservoir, Missouri, during 1965-74 (Hanson 1977). 453. In tbe Tennessee Valley, white bass support important spring fisheries in cold tailvaters, which are primarily the result of spring migrations upstream from reservoirs (Pritzer 1962). The fish is a ·particularly importan·t game species belov the main-stem reservoirs; for example, in Watts Bar tailvater it canposed 18.8 percent of the anglers' catch in 1953 (Miller and Chance 1954). Large num:bers of vhite bass have also been observed migrating upstream out of Watts Bar Reservoir into Norris Dam tailvater (Escbneyer and Mange! 1945). 454. In Dale Hol1ov tailvater, Tennessee, vhite bass became abundant in the lover tailvater after the filling of Cordell Hull Reservoir (Bauer 1976). White bass are also important in the fishery belov Tenkiller Dam, Oklahoma, vhere they composed 8 percent of tbe catch immediately belcv the dam and 43 percent .at a point 12.8 km dovnstream (Deppert 1978). 128 J ;• .. . ·. • ... ~- ·-: . ...... -:., ... 455. Lake Taneycomo tailvater, Missouri, is influenced by the presence of Table Rock Reservoir upstream and Bull Shoals Reservoir downstream. Increased power production at the deep-release Table Rock Dam has converted Lake Taneycomo tailwater fr0m a varmvater to a coldwater tailvater, making conditions unfavorable for varmwater game fish. The magnitude of spring migration of vhite bass into Lake Taneycomo tailwater from Bull Shoals Reservoir, hovever, is most heavily influenced by the level of Bull Shoals Reservoir, vhich inundates Lake Taneycomo tailvater. Maximum spring migration of vhite bass intq the tailwater occurred vhen Bull Shoals Reservoir vas betveen 0.6 and 3.0 m belov paver pool. Above or belov these levels the migration rate declined proportionately (Fry 1965; Hanson 1969). 456. The food and reproductive habits of vhite bass in tailvaters are not vell .known. Of 165 stomachs of immature vhite bass collected. from Norri.s tailva.ter, Tennessee, in the vinter of 1942-43, 140 vere empty (Eschmeyer 1944). White bass in Levis and Clark tailvaters, . : -:·. .. .. ..: ... • .. ·. South Dakota, fed .. ~:n zooplankton. in the spring and tall, and pre~,;-.. ·.;·.;~~.iif;~ .. :>: :·~ · dominantly fish in SUlllllier (Walburg et al. 1971). Litt.le reproductive ·· .... ~ • 4J, : .. · success· was noted in either of these tailvaters · (Eschmey~; ~d ~ges -~ • ~1 '. '" ,. ·w • 1945; Walburg et al. 1971). 457. Striped bass have been stocked in a number of reservoirs and tailva.ters in an effort to control large gizzard shad populations and to enhance the sport fishery; hovever, they have been found to compete vi th trout in tail waters. Belov Davis Dam, ~izona, 24 trout <?00 to 230 m:m long vere found in 20 striped bass stomac~s examined (Arizona Game and Fish Department 1972). Additionally, food studies . on striped bass in Tenkiller tailvater, Oklahoma, shoved that rainbov . . trout composed 40 percent by number of the food items eaten by striped bass during the first veek after the trout vere stocked. Gizzard shad vas the prima1·y striped bass food in this tailvater, accounting for 56 percent of the total number of food items during the first veek after trout stocking and 75 percent at other times (Deppert 1978). Similar results vere noted in Keystone Dam tailvaters, Oklahoma, vhere gizzard shad made up 85 percent of the food eaten (Combs 1979). 129 ... .. .. : ... ·• '• ":~. . ,, ~- .C!: Centrarchidae (Sunfishes) 458. Fishes of this family are found in nearly all types of waters, and most are highly sought by sport fishermen. Their habi.ts and li'fe history are similar, differing only in detail. Ce~trarchids migrate little; oost remain in the sa."l!e stretch of stream .or shoreline through- out life. Feeding is by sight., and feeding activity peaks in early morning and late evening, accompanied by movement into. shallow water. All centrarchids construct nests for spawning, but only males partici- pate in this activity. They guard the eggs after they are deposited and remain until fry leave the nest. The centrarchids are discussed under three general groups--black basses, "true sunfi-shes," and crappies. Bla·ck basses 459. Three black bass species are commonly found in tailwaters-- tbe smallmouth bass, spotted or Kentucky bass, and largemouth bass. 460. Habitat. Smallmouth bass prefer clear; cool, permane·nt- f'loving streams vi th good gradient ( 1. 3 to 3. 8 m/km). Preferred SUIIIIIIer .-... vater .'temperature is about 21 °C. The fish generally occur, over a. ··:, : l -... ,·: • · silt-free rock or gravel bottom, near riffles, but not in the main current. Adults are most abundant near cover in the form of boulders, roots, or sunken trees. In large rivers vi tb navigation dams., they are usually restricted to the rocky shoals below dams, whe~e streamlike conditions still prevail. The smallmouth normally limits its. acti vi- t .ies to a single stream pool, but occasionally its home range includes several pools as much as 0.8 km apart. 461. Spotted bass inhabit flowing waters that are warmer and slightly more turbid than those favored by smallmouth bass. Preferred . summer temperature is about 24 °C. 'The habits of the spot ted bass are similar to those of the smallmouth, except that ·the spotted, bass is more migratory. 462. The largemouth bass is more widely distributed than the other basses and is the most abundant bass species in standing-water habitats. Its preferred summer temperature is about 27°C. It is 130 '· , .... commonly :round in lowland lakes, reservoirs, slow-flowing streams, and backwaters of large rivers. It is intolerant of exce;sive turbidity, and in streams with continuous strong flow it is rep:d.ced by one of the other basses. The . a.rgemouth displays more seasonal movement than either the smallmouth or spotted bass. 463. Reproduct.ion. The smallmouth bass begins nesting in the spring ~hen the water temperature exceeds l5.6°C. Nesting activity usual.ly peaks in late spring, but sometimes continues well into early summer. Renesting occurs if early nests are unsuccessful because of high water or low temperatures, and may occur even if the first nests are successful. Nests are in quiet water near shore or downstream from a boulder or other obstruction that breaks the force of the current. Water depth rarely exceeds. 1m over the nest. Smallmouth bass eggs are golden yellow and about 2. 5 .mrn in diameter. They are distinguished from those of the spotted bass and largemouth bass by' their larger size. Eggs hatch in 2 or 3 days and fry rel!lain on the gravel for about 6 days before leaving the· nest. 464. Spotted bass generally begin spawning several .days later.: b ·.•• •• • than smallmouth bass in the same stream. Nest-s are similar to those of_~."-.~· the smallmouth bass, and the development of eggs and fry is similar at · the same temperature. Fry of the spotted bass disperse from the nest 8 or 9 days after spawning if the water temperature is above 20°C. · 465. The largemouth bass begins spawning in the spring when water temperature reaches about l5.6°C and continues into late spring. Nests are constructed on almost any type of firm, silt-free bottom. Water ...... depth over nests may vary from 0. 3 m or less to 4. 6 m or more, being .deepest in the clear waters ·)f large impoundmen-ts. Nests are n.ever constructed where there is current or wave action. In streams, nests are located in the deeper and quieter parts of pools or in adjacent sloughs. Eggs are about the size of spotted bass eggs and are much ·smaller than thos·e of the sm.'lllmouth. They hatch in 3 or 4 days and fry leave the nest as a school when about 10 days old. Schools break up 26 to 31 days after hatching, when the young bass are slightly over 25 mm long. The male largemouth is a more attentive parent than any o:r 131 ,. ,. :, :· . - the other sunfishes, remaining with the schooling young for several weeks after they leave the nest. 466. Food. Midge larvae and zooplankton are the first foods of smallmouth bass. fry. According to Pflieger (1975), fry less than 25 mm long eat small fish, and fish .remain an important part of their diet throughout life. Crayfish and fish occur in about equal amounts in the diet of adult bass. Insects are taken frequently but are of only minor .. · ... importance .. 467. Immature stages of aquatic insects are the principal diet of spotted bass of all sizes. Insects are supplemented with small crusta- .. · c~ans in bass less than 75 mm·long, and with crayfish and fish 'in larger bass (Smith and Page 1969). 468. The first 'rood of young largemouth bass consists mostly of zooplankters, but these are supplemented by insects and their larvae as the young bass increase in size (Pflieger 1975). Adults feed princi- pally on fish,.crayfish, and large insects, along with an occasional .... frog, mouse, or ~ost any other animal that . swims or falls into the :.7 ·. ·_':.water. :In 'large re•se·rvoirs, the largemouth bass depends heavilY. on ' •• ,· • • • ; ' t " I gizzard shad as foo~, and there is a definite relation between the trends in abundance of the largemouth a .nd those of its principal. prey species. 469. Age and growth. In Missouri streams. the smallmouth bass averages 90 mm long when 1 year old, and attains lengths of about 170, 244, 290, 343, and 371 mm in succeeding years {Purkett 1 958b). As is coDJDon in fishe.s, growth is more rapid in larger streams than in headwater creeks. A Missouri smallmouth bass weighs. about 225 g at a length of 254 mm, and about 625 g at 356 mm . Smallmouth bass seldom exceed a length of 560 mm or a weight of 2.5 kg. Missouri smallmout'h bass bec.ome mature during their third or fourth summer of li.fe and . some live 10 or 12 years. 470. Growth of spotted bass is slightly faster than that of the smallmouth for the first 4 years of life but is slower thereafter. In Missouri streams, lengths attained are about 85 mm the .first year and about 183, 254, 292, 322, and 353 rmn in succeeding years (Purkett 132 .. 1958a). Growth appears to be more rapid in reservoirs than in streams. Most fish are mature when 3 or 4 years old. Few spotted bass live longer than 6 years or attain a weight much greater than 1.4 kg. 471. Growth of the largemouth bass is extremely variable, depend- ing on local conditions. In Lake Wappapello, Misso·ur.i, a length ot about 135 mm is attained the first year and lengths of 277, 338, 409, 460, and 498 mm are reached in succeeding years (Patriarche 1953). Growth rates are similar or faster in new, well-managed ponds, but much slower in highly turbid or overpopulated ponds, which may contain bass 4 years old or older that are still less than 254 mm long. Fish mature between the ages of II and IV, depending on growth rate. Under average conditions a 305-mm bass weighs about 340 g and a 560-mm bass about 2. 7 kg. Individuals weighing more than 3.6 kg are not uncommon. Few .largemouth bass live beyond 12 years. Black basses in tailvaters 472 . Black basses· are important game fish in. rive.rs and reservoirs in many areas of the United States. They occur in tailvaters if vater .. --:·.r· temperatures are suitable and cover is adequate. Black bass populations have be.en reduced below many hypolimnetic release dams constructed on varmwater streams because of low water temperatures, strong currents, and lack of instream cover. 473. The smallmouth bass fishery below Hoover Dam, Ohio, was lost due to hypolimnetic water discharge from the dam (Cavender and Crunkil ton 1974) • Pierce ( 1969) showed a reduction in smallmouth populations from 7. 24 to 4. 72 kg/ha in the Summersville tailvate '!*, West Virginia, based on preimpoundment and postimpoundment electrc- fishing studies. The maximum temperature in Summersvill e tailwater is 15.6°C, which is below the optimal range for smallmouth bass. l)endy and Stroud ( 1949) belie.ved that low water temperatures and low dis- solved oxygen following construction of Fontana Dam, North Carolina, adversely affected smallmouth bass for many miles downstream in the Little Tennessee River . The coldwater discharge from Table Rock Reservoir, Missouri, caused the loss of smallmouth and spotted bass downstream in Lake Taneycomo (Fry and Hanson 1968). 133 474. Cold tailvaters below two dams in Kentucky support fever black basses than occurred in the natural river before i~undment. Black basses ve.re estimated to be 47,9 and 19.1 percent ?f the angler catch in a 1959 creel survey from these tvo rivers (J. P. Carter 1968a). In 1968-71, after dam construction, black basses. composed only 0.1 to 2.6 percent of the angler catch (Charles and McLemore 1973). Apparently the tailwater habitat differed from the original varmvater stream habi- tat and did not provide the conditions necessary to sustain t.he bass .fishery. 475. Sroallmouth bass vere common in angler catches in the tailvater below Cherokee Dam, Tennessee, .for the first 8 years after impoundment. The fishery changed thereafter, and anglers had to fish 32 to 40 km do'Wt'lstream to catch smallmouth bass (Pfit:z.er 1962). Low vater temperatures apperently reduc·ed bass reproduction and numbers i ·n the upper 32 km of the tailvater. 476. Eschmeyer and Manges . (1945) shoved that the condition, factor ~ •. of'larg'em~·uth bass declined f~om aut~· 1942 'to ·autumn 1943 in a ··Tenne.ssee taii~at'e;~ They believed the fish ve;e experiencing lugh .. ::: ... • ••• ·.: ... ; J,. -· • • .. • • 'oo; -•• ~; • \ • .. .--:;i~.,.. ... ;.' natural mortality 'due to the harsh habitat in this 'cold tailvater."·•'-'• ~ ··· · •J 477. Some varm and cool tailvaters sustain black bass populations when water temperatures and cover are adequate. Many young-of-the-year smallmouth bass were captured by seine below a low-head dam on the Maquok:ta River, Iova. The large concentrations: of fish in this tailvater were believed to be a res-ult of high oxyge.n levels, blockage ··.. ,. of upstream movement, and habitat diversity (Paragamian 1979). Hutchison et al. (1966) found low to moderate smallmouth bass abundance below Cottage Grove Dam and Dorena Dam, Oregon. These flood control dams have caused an increase in water temperature in the tailvaters and have made them more sui table for smallmouth bas·s and le.ss sui table for trout and salmon. And.revs et al. (1974) compared the catch of largemouth bass in the river above and below an Ok1ahoma reservoir. Historically, largemouth bass vere abundant in the deep pools of the do'Wt'lstre&."D section where flows were more stable. After impoundment, 92 percent of the largemouth bass were ca.ught in the rive-r above the 134 ~-· ~ .. ... • ··· ... · ···•· ., .-··. ........ reeervoir, and they provided 35 pereent or the eet!mated eateh or all tilh epeeie1. Cool water dileharge and. tluetuating tlovs belov this main-stem hydropower dam apparently reduced habitat suitability for largemouth bass in the tail water . 478. Food of largemouth and smallmouth bass f'rom Holyoke Dam tailwater, Massachusetts, was studied in 1972 (Jefferies 1974). The frequency of occurrence. of fish, insects, and crayfish in largemouth bass stomachs was 88.9, 16.7, and 7.8 percent. Fish (spottail shiners) made up 93.4 percent and crayfish 5.3 percent of total food volume. The largemouth bass selected spottail shiners over Alosa s:pp. Smallmouth bass stomachs contained fish, insects, c.rustaceans, and pelecypods •.. Fish made up 45 percent of the stomach contents, crusta-. .. ceans 35 percent, and pelecypods 20 percent. In largem.outb bass ·:·collected belov Norris Dam, Tennessee, 42 of 53 stomachs examined vere empty. Apparently the cold tailvater did not provide good bass habitat since the fish exhibited no growth (Eschmeyer 1944, Eschmeyer and Manges 1945). 479. Reproduction of black basses in tailwaters is not vell docu- mented. There is no documented reproduction of smallmouth bass in South Holston tailwater, Tennessee. No juvenile bass were captured in this tailvater and many adult females from a November 1953 collection had resorbed. their eggs (Pfitzer 1962). Young-of-the-year largemouth bass were the most abundant centrarchid in collections taken in 1965 bel.ov Beaver Dam, Arkansas. However, it is not knovn whether these fish were produced in the tailwater or had moved out of the reservoir above. In 1966, no largemouth bass adults or juveniles were captured in the tailwater (Brown 1967). 480. Generally, cold tailwate.rs do aot provide good bass habitat. Limited bass fisheries have developed below some dams, but fish harve$t usually 'remains lov. Successful bass fisheries are usually found in warm tailwaters with backwaters or other sheltered areas. True sunfishes 481. The most common o f the "true sunfishes" (a term applied here to the Lepomis sp. of the family Centrarchidae) occurring in tailwaters 135 ..• ... &re the bluegill, gree.n sunfish,. and longear sunfish. Also included in thi.s section is the rock bass. Other true sunt'ishes may be locally abundant in tailvaters, but their life histories are generally 1Sim.ilar to those described. 482. Habitat.. The bluegill is common in the deeper pools and backwaters of' streams, and in lakes, ponds, and reservoirs. It is intolerant of continuous high turbidity and siltation and thrives best in warm, quiet-wat.er areas vi th some aquatic vegetati.on. 483. The green sunfish tolerates a vide range of conditions, but does best where few other sunfishes occur. It is adaptable f .or survival in fluctuating environments, since it tolerates extremes of t .urbidit;r, dissolved oxygen, temperature, and flov. 484. The longear sunfish is characteristic of clear streams vith sandy or rocky bottoms and permanent f'lov. It is more common in streams than in large rivers. Like other sunfishes, it avoids strong currents and is usually found in pools and backwaters adjacent to the stream channel. In most environments, when longear sunfish are camnon, ··.green' sunfish are rev. ·.~ · ~. 485. ·The rock bass CODIDonly occurs in streams. Permanent :flow, low turbidity, abundant cover, and silt-free bot toms are its basic requirements. It is usually found near boulders, subme·rged logs, and t ·ree roots· vhere there is a slight to mod.erate current. A deep rocky pool iDJmediately below e. ri.ffle is a f'avored spot. · 486. Reproduction. The true sunfishes appear to have similar spavning.habits. Fish begin nesting in the spring vhen vat.er tempera- tl;U'es are about 21 °C. Spavning reaches a peak in June but orte.n con- tinues into August. Nesting occurs on almost any t .ype of bott om.,· but gravel is preferred, Nests are usually in water 0.3 to 0.6 m deep and consist of roWldish depressions with a diameter about t .vice the length of t -he. male parent. Many nests are coDI!lonly close togetbe.r and in a limited area. The green SWlfish is less colonial than some of the other sunf'ishes in its nesting habits. The, male guards the nest unt.il the e.ggs hatch but does not guard. the fry once t .hey leave the. nest. 136 ·. Because. of their similar spawning habits, the various sunfish species often crossmate and produce hybrids. 487. The nesting season of the rock bass coincides with that of the smallmouth bass and precedes that of the sunfishes. Nests have been observed as early as the first week of April and as late as early June in Missouri, but in any given year the season seldom lasts more than one month (Pflieger 1975). Nesting begins when stream temperatures range between 12.8 and 15.6°C. The male rock bass fans out a saucer- shaped depression 200 to 250 mm in diameter over a bottom of coarse sand or gravel~ Nests arc in water from 0.3 to 1.5 m deep, usually near a boulder or other large object, and often where there is a slight current. The rock bass is a solitary nester, in contrast to the true sunfishes, which tend to nest in colonies. 488. Food. The diets of true sunfishes and rock bass are gener- ally similar. Young of the year feed on zooplankton and immature aquatic insects, and older fish on aquatic insects, supp.leme.nted with small fish, crayfish, and snails. Feeding is most intense during the 'early morning and in the evening. •··~·~.· 489. Age and grovth. Growth of the bluegill varies considerably . from one body of vater ~o another. Growth is usually slower in streams than in ponds, lakes, or reservoirs. In most Missouri waters, the bluegill reaches a length of 150 mm and a weight o:f about 70 g by the end of its third or fourth summer of life. A 215-mm bluegill weighs about 225 g. Bluegills commonly reach a length of 240 mm. and a weight of 340 g (Pflieger 1975). 490. The green sunfish attains lengths of about 43, 81, ll9, 150, and 193 mm at an age of 1 through 5 years, in the Salt River, Missouri (Purkett 1958a). A 150-mm green sunfish weighs about 85 g; few indi- viduals exceed .a length of 230 mm or a weight of 340 g. 491. In Missouri streams, the longear sunfish attains a length of· about 33 mm its first year and 64, 91, 109, 122, and 127 mm in succeed- ing years (Purke,tt 1958b). The maximum length and weight are, about 175 mm and 128 g. 137 . ~ . ·; ·. 492. Rock bass from Ozark streams in Mis,souri average 41 mm in length by the end of their first year of life and attain lengths of 86y 140y 178, 203, and 216 mm in succeeding years (Purkett l958a). Few live more thar. 5 or 6 years, but they commonly attain a length and veigh~ of up to 280 .mm and 454 g. 493. In s·ummary, growth of the true. sunfishes varies considerably from one water body to the next, and st\Ulting occurs in crowded popula- tions or vhere water is continuously turbid. Generally the bluegill, rock bass, and green S·Wlfish attain the largest size. T::-ue sun:'ishes in tailwaters 494. ··The abundance of sunfishes in tailwaters is variable and depends on recruitment and the available habitat. The occurrence of sunfishes in tailwaters below Tenness~e Vall7Y Authority storage reser- voirs was found t .o depend on fish present in the river be,fore impoWld- ment, fish entering the tailvater from the reservoir above, and migra- tion into the tailwater from tributary streams or reservoirs dovnst.ream (Pfitzer 1962). Cavender and Crunkilton (1974) reported bluegills and · ,, vhite 'crappies being carried over th!! spillway of Hoover Dam, Ohio, ·.~and e.stablishing small populations in the ·tailvater. '. Bluegills vere also periodically transported over spillways into the. tailvaters of Urieville Lake and Wye Lake, Maryland, ~nd r,;;rainfe Lake, Obi·~ ( Clal'k 1942; Elser 1960). 495. Sunfishes generally prefer areas vi th instream cover, lov currents_, and maximum water temperatures above 21 °C. SWl:fish popula- tions are usually depressed in tailvaters that are cold, or have high turbidity or little instream cover. Bl·uegills, longear sunfish, and green sunfish were collected in Norfork tailvatery Arkansas, in 1950 - ( rese:rvoir impounded in 1944), but were lac.k.ing in collections made in 1959 (Hoffman and Kilambi 1970). Brovn (1967) fO ·Wld several sunfishes in the same drainage below the never Beaver Dam in 1965 (reservoir impounded in 1961). Apparently several years are required before the reduction in temperature eliminates s\Ulfishes from these cold tail waters. Table Rock tailvater (Lake Taneycomo), Missow·i, vas con- verted from a warm to a cold tailwater when hypolimnetic discharges 138 .. .·.· : -.. . . , "J.·. , ..... .. ' -~ ~re begun in 1959. Bluegills were the only varmvater species to re- m&in abundant after the c.hange to cold water. Most of the blue gills were captured at the downstream end of Lake Taneycomo, where solar wa:rmlng and some thermal stratification occurred (Fry and Hanson 1968). Bluegills and longear sunfish composed over 50 percent of the total numbers of' fish collected by electrofishing in Nolin tail water, Kentuc.ky, in 1965 and 1966. The abundance of sunfishes in the tailvater was due primarily to export of fish from the reservoir above (J. P. Carte·r 1968b). A cha.:nge from epilimnetic to hypolimnetic release and the stocking of trout in the tailvater in 1970 and 1971 resulted in a reduc.tion of the sunfish harvest. The sunfish catch from 1968 to 1971 declined from 79.4 to 10.7 percent of' the total number of fish caug!Lt and .from 44.6 to 3.4 percent of the total fish weight (Charles and McLemore 1973). Reports of sunfishes in other cold ta.ilvaters are limited. Bluegills, green sunfish t and redear sunfish compo•sed only 5 perc.ent o·f the 1975 fish coi!IDuni ty in Dale RoLlow tail water, Tennessee (Bauer 1.976). 496. ·Studies on several va.rm tailvaters suggested that sunfishes ~..;.::.~ · .. can be _important in the fi.shery. Bluegills w:ere estimated to be '8, 12," 7 and 13 percent of the angler catch at Lake of the Ozark.s, Pomme de Terre, and Stockton tailwaters, Missouri. Both Lake of the Oza.rks and Pomme de Terre have epilimnetic discharges, resulting in ma.xi.mum· .;,a_ter' temperatures of about 29°C (Hanson 1974). A highly significant corre- lation betveen total annual discharge and annual average cate:h rate . '· W'a'S .found at Pomme de Terre. Hanson's findings agreed with the con- clusions of Moser and Hicks. (1970) that tailvater fisheries are sup- ported by fish from the reservoir (Hanson 1977). Carter (1969) stated that sunfishes in the Barren tailwat.er, Kentucky, were mo-re abundant . when water was released from the epilimnion rather than from the hypolimnion. The warm v:3.ter, in combination with the abundant instream · cover, accounts for the increase of sunfishes. Of the fish seined below Lake Carl Blackwell, Oklahoma, 15 percent were sunfishes (long eat", orangespotted, and green). The longear sunfish was most abundan.t in July when it was favored by reduced turbidity and stabilized flovs 139 . · .... belov the dam (Cross 1950) • Not all varm tailvaters :provide good sunfish habitat. Moser and Hicks (1970) found that sunfishes made up only 1. 5 percent of the fish biomass and 3. 0 percent of fish numbers in the stilling basin of an Oklahoma reservoir. Lack of cover may have ........ been responsible for the lov abundance of sunfishes. Fritz (1969) reported that bluegills made up 4.4 percent and green sunfish 2.6 per.- cent of the angler catch in Carlyle tailvater, Illinois. 497. Sunfishes are important to the fisheries in some"cool tailvaters. Longear sunfish vas the most common species taken by anglers in Broken Bow tailwate:-, Oklahoma ( 36 percent of the catch), and was second in biomass· ( 17 percent). However, more longear sunfish vere captured in the river above the reservoir than in the tailvater. Weekly temperature means averaged 3. 8°C lover in the tailvate.r than in the river upstream. Apparently the cool vater and fluctuating flovs. in- fluenced the harvest of sunfishes in this tailvat~ (Andrews et al. 19'74) . Cavender and Crunkil ton ( 1974) reported that a small conc·entra- tion of rock bass exists in Hoover tailvater, Ohio; the authors be- t lieved that the rock bass stay in the tailvater because of the abun- .. dalice of forage fish and crayfish. _ ... 498. The food habits of bluegills and longear _sunfish fro111 Wilson Dam tailvater, Tennessee, vere studied in the spring of 1977 (Warden and Hubert 1977). Fish eggs made up 67.2 percent of the total number of food items and 59.2 percent of the total volume in bluegills, and ... . .. 56.1 percent by number and 3.7 percent by volume in longear sunfish. Insects vere abundant in the stomachs of both species. Insects ac- counted for 27.1 percent by number and 32.1 percent by volume of food items in bluegills, and 23.0 percent by number and 22.2 percent by volume in longear sunfish. Insects eaten vere of the orders Diptera, Coleoptera, and Trichoptera and, of these, chironomid larvae and mayfly nymphs composed 90 percent of the tctal number and volume. Ot.ber items found in the stomachs vere decapods, larval fish, iaopods, mollusks, arachnids, and annelids. 499. Grovth of bluegills in Hartwell tailvater, South Carolina, did not vary from that of fish downstream or from those captured in &>n 140 .\ " I .. ·- .~. :- ·'· unimpounded. control stream. Apparently temperature fluctuati.ons and a temperature range of 6.1 to 16.8°C in the Hartwell tailwater did not adversely affect growth (Dudley and Golden 1974), although Fry and Hanson. (1968) stated that growth of warmwater fish (including bluegills) was reduced. in a cold Missouri tailvater (discharge temperature 4.4- 15.60C). Crappies 500. The white crappie and black crappie may both occur in tailwaters, and they have similar life histories. 501. Habitat. White crappies are found in ponds, lakes, reser- voirs, and slow-moving streams and rivers. In reservoirs, they are often found in areas having standing timber or other cover, and a·t other times they frequent dee,per va·ter, commonly occurring at depths of 4.6 m or more. Young crappies are often found over open vater of considerable depth. In streams, the white crappie is most abl.Uldant in the deeper pools or in 'backwater areas away from the main current. It avoids streams that are excessively turbid and those kept continuously ·cool ·.ay· tlov from spriligs. " ...... ~ · "':.;'"_'~'?'· .- . ; .,·~· 502. T.he black crappie requires habitat similar to that o·r the .. : · white crappi.e except that it is less tolerant of turbidity and silta- tion. In reservoirs, the black crappie is noticeably more abundan·t in embayments fed by the clearer streams. In streams, black crappies require clear water, absence of no·ticeable current, and abundant c.over. 503. Reproduction. Crappies begin spawning in April or May, when the vater temperature rises to about 15.6°C. In reservoirs, s.pavning occurs in shallow areas of coves protected from wave action; many nests are sometime•s concentrated in the same cove. Nests are prepared by the male on a variety of silt-free substrates in water 0.1 to 6.0 m deep. Sites with nearby logs or other large objects are favored locations for nests. The location of the nest is i 'ndicated only by the presence of tbe male. Eggs hatch in about 3 days and the try remain in the nest several more days. Fry do not school after leaving the nest. . 141 : : rl -· T' 504. Food. The diet of yoWlg crappies consists mainly o·f zoo- plankton, and that of adults includes zooplankton, aquatic insects, and small fish. The proportions of these · food. items in the adult diet vary with locality, season, and age of the fish. Small gizzard shad and threadfin shad are important foods of adult crappies in many reser- voirs. ·505. Age and grovth. According to Carlander (1977), the average calculated total lengths of white crappies at ages· I to VI from all are.as of the United States are 78, 158, 213, 257, 290, and 304 mm. Average weight of a 4-year-old white crappie is about 300 g. 506. Grovth in length of black crappies_is generally less than that of-white crappies in the same waters (Pflieger 1975). However, since the black crappie is heavier at any given length ·than the vhite crappie, grovth in weight differs little between the two species. Fev crappies live more than 3 or 4 years, but oceasional individuals live as long as 8 or 9 years. Maturity is reached during the second or third summ.er of 1i fe. ·Crappies in tailvaters ·-• •··-507. \-.'hi te crappies and black crappies are important in any · tailvater fisheries. When crappies are abundant in a reservoir, they are often carried through the dam and remain in the t .ailvater. Crappie abundance in a tailvater appears to be affected by water temperat.ure, season, and type of dam discharge. . .• • 0 ~ • .. .. • • 5'08. Crappies are often abundant in warmvater tai.lvaters and. can contribute substantially to the fishery. White crappies vere"estimated. to mak'!! up 56 percent of the angler catch at Lake of the Ozarks tailwater, Missouri, in 1965-74. A high correlation vas found between estimated number of fish caught and the number of days the flood gates were open at the dam (Hanson 1977). Crappies ·were estimated to be 41 and 54 pE.rcent of the m.UDber and 35 and 49 percent of the veight of the fish taken by sport fishermen during varmvater releases at Barren and Nolin tailvaters, Kentucky, in 1970 and. 1971 (J. P. Carter 196&; Carter 1969). P. Carter (1968a) reported that before reservoir construction in 1965, crappie populations in Barren and Nolin rivers 142 ' ' • ·.~ l '· .. I .. t :.,r •, .. .· ·'· .. _ ... . .. were low and. contdbu.ted little to the fishery. He attributed the increase in abund&Oce after impoundment to fish that vere produced. in the reservoirs and moved dovnstream through the dam. 509. Tb.e occurrence of fish in stilling bas.ins belov va.rmwater :release dams has been examined in tva studies: Pfitzer (1962) col- lected 120,000 crappies weighing 10,884 kg from 1.0 ha of water below Douglas Dam in Tennessee on October 30, 1953; Hall and Latta (1951) stated that 23 ,percent of the fish found in the stilling basin below Wister Dam ,, Oklahoma, in August were white crappies. 510. Several investigators have reported on the movement .and seasonal changes in abundance of white crappies in warm tailwaters. At Kentuclcy L&ke tailwater in Kentucky, 3552 white crappies were captured, tagged, and released fran January to December 1953'. Anglers • I • I ., recaptured_ 113 fish (3.2 percent) of which 95 (84 percent) were taken within 1.6 km of the release ·site (Carter 1955a·). A concurrent tagging, study in Kentucky Lake shoved little movement of white crappies (on the , .... basia. ~: recapture~ of 5 of 1752 marked fish) through. the navigati?I?- locks into the ta'ilwater (Carter 1955a). Anglers at Levis and Clark Reservoir and its1 tailvater, South Dakota and Nebraska, returned 42 tags fran 288 white crappies tagged in the reservoir. or these, 12 (28 percent) were tram fish captured in the tailwater (Walburg et al. ·· 1971). White ·crappies were abundant in the tailwater belov Lake Carl Blackwell, Okl.ahana, from October 1947 until March 1948 because flows were srtable and many fish escaped from the ·reservoir. Between Kovember and Janu&l"y, 139 white crappies were tagged and released in the tail waters. Most tag recoveries ·were reported during the •winter, 'axld we·re made in the tail waters. High water releases from the reser- voir during March 1948 apparently caused the crappies to leave the tailwater, since.,only tva tagged fish were captured during the subse-. .. quent spring and summer (Cross 1950). An increase in crappie abun- dance during. fall 1965 at Barren tailwater, Kentucky, was associated with the fall reservoir dra:wdo\ltl and the high water discharge into the· tailvat er (J. P. Carter l968b). There is a large populat ion of white crappies and black crappies in Hoover Reservoir, Ohio, and many young 143 ....... .... are carried over the spillway and into the tailwater (Cavender and Crunkilton 1974). 511. T.he change from hypolimnetic · (coldwater) to epilimnetic (warmwater) release (or vice versa) at some dams has affected the abundance of crappies in tailwaters. At Barren tailvater, Kentucky, crappies composed 13.4 and ll. 6 percent of the catch during hypolim- netic releases in 1968 and 1969, but 41 and 44 percent during epilim- netic releases in 1970 and 1971 (Charles and McLemore 1973}. The construction of Table Rock Dam, Missouri, changed the downstream Lake Taneycomo from a warmvater to a coldwater habitat. Test netting in Lak.e Taneycomo before and after the coldwater intrusion showed ·a reduction in white crappie abundance from 6.5 to 0.9 fish. per net day (Fry and Hanson 1968). Apparently the cold vater made the tailwater habitat 'less.suitable for the white crappie. 512. Small crappies are common forage for predatory fish. Com,bs ( 1979}, vho studied the diet of 164 adult striped ba.ss collected from · the tailva:ters of Keystone Dam, Oklahoma, in 1974 through 1976, found tb&t frequency of occurrence and percentage of total volume of ·vbite crappies in stomachs vas 10.4 and 5.8 percent, respectively. Walburg ~~ &1. (1971) reported tb~ occurrence of white crappies in stomachs of walleyes and saugers collected in Levis and Clark tailvater, South . . . Dakota and Nebraska. 5·13. Food of black crap'_;>ies from Holyoke Dam tailwa~er, Massa- chusett.s, in 1972 consisted mostly of spot tail shiners and insects (69.3 and 27.2 percent of total food volumes); the frequency of occur- rence of fish, insects, and zooplankton in stomachs vas 44, 68, and 42 percent, res9ectively (Jefferies 1974). 514. Studies of the growth of crappies in tailvate1#:has received little attention. Carter (1955b) reported that white cr~pies in Kentucky Lake grew faster than those in the tailvater. B;rore deep-vat er releases from a Missourireservoir converted the dovnstream Lake Taneycomo into a coldwater habitat in 1959, the average length of 4-year-old white crappies vas 338 mm, but after the change in 1963, it was only 211 mm (Fry and Hanson 1968). 144 ~ . . ~ .. i I .· •• t 515. In ·a report. on crappie gonadal development in tailvaters, 3 of 74 vhite crappies collected belov a Tennessee dam, betveen July 29 and September 1, 1941, vere immature.; the rest vere ripe, but none had spavned. Apparently the cold vater (< l0°C) had disrupted their repro- ducti.ve cycle (Eschmeyer and Smith 1943). Percidae (Perches) 516. Th~ perch family is one of the largest groups of North American freshvater fishes. Among them are three popular game fishes- valleyes, saugers, and yellov perch--and a large number of smaller fishes knovn collectively as darters. The three game fishes are repre- sented in Europe by the same or closely related species. The .darters are native only to North. America. 517. The closely related walleye and sauger both occur in rivers . and -are important in some tailvaters . The yellow perch is most oft en found in lakes but it is also abundant in backwaters of large rivers. ::V~~;t'!.·~~:-:,;:.~::.~:t~~-)·~_;:r;··It 'is seidom fo~d in small streams an.d usually does ~ot occur i n··:; ::o:-_·.:;.:-·.-r · , .. ''· .. :~'. ~ .. ,• .. a!;~·:t• •. ''r·· ., . ' . .. .,• . . tailwaters·ana, therefore, is not discussed further here. Darters are : adapted for life in svi:rt-floving sections of clear, rocky streams and . ·are common inhabitants of many tailwaters. The percids are discussed under two groups-(a) walleyes and saugers, and (b) darters • Walleyes and saugers 518. Habitat. Walleyes and sauge.rs inhabit the open water of large, shallow lakes on slow-nowing rivers . The habitat requirement s of the two species are similar, except that the sauger is m<:~re tolerant of high turbidity and is o:rten found in areas with strong current. The sauger is more common in habitats with silted bottoms, vhe1·~a~ valleyes prefer habitats with gravel, bedrock, and other types of firm bot~om. Tubb et al. (1965), vho studied fish distribution in the Sheyenne River in. North Dakota, found walleyes in pools 0 .9 to 5.5 ~ deep, but most commonly in pools dee·per than 2 . 4 m. The sauger vas taketl in only one pool, vhich vas 2.4 m deep. 145 .····II: a 519. The sauger feeds more actively during the day, vhereas tb~ walleye is more crepuscular. The walleye is light-sensitive and is usually found in deepwater pools during the day, especially when water is clear. Both generally occur in loose aggrega,tions of a fev to many individuals. They range over a vide area, rather than restricting activities to a definite home range. 520. ·Reproduction . Spavning occurs at night over a 2-wee:k period in the spring when water temperature exceeds 5.6°C. Spavning is com- monly preceded by move.ments out of larger rivers and reservoirs into tributaries, the males moving to the spawning grounds before the females. There is some evidence that these species tend to return to · a "home" spawning area in successive years. Spavning· occurs on ri ffle.s or rocky areas below dams in streams and along rocky wavesvept shorelines in lakes and reservoirs. Females are accompanied by several males during spawning, and eggs are scattered at random. Tbe adhesive eggs stick to the substrate, and tAtching occurs in 12 to 18 days, depending on water temperature. Newly hatched la.rvae are semi buoyant, and those produced in streams are therefore subject to dovnstream transport. ·' -. ' ::. ,.; ... ·., ... _~ .. -. 521. ~ Small crustaceans and insects are the food of walleye ·and sauger frJ. Insects are a significant food item throughout life, but .fish are the principal food of a~~ ts. They apl'&7.~ntly eat any specie.s •>f fish readily available to them. 522/ Age and growth~ The sauger grows more slowly than t ,he walleye and does not attain as large a size. Fish from the northern portion of' the range grow slower a;.1d live longer than those from more southern waters. Females attain greater lengths and 1i ve longer 'than males. Newly hatched larvae of both species are only a·oout 7 t ,o 8 Dill in length, but under ideal conditions may attain lengths up to 254 mm by the end of the first year. Pflieger ( 1975) .reported that the average length of walleyes frcn the C\lrrent River in Missouri is 200 11111 at the end of the fir .3t yes.r and 610 11111 at the end of the seventh year. The usual life span is 7 or 8 years, but muc·b older individuals are not uncommon. 146 I. . -•• ... .. -. '. , ..• 523. Vasey ( 1967) reporte.d that the saugers in the Mississippi Ri,·er in Iova reach a length of 145 mm in the first year and 515 mm after the seventh year. The usual life span of saugers in the South is 5 o.r 6 years, but some live to 12 or mo:re years in Canadian waters. W'a.lleyes and saugers in tailvater·s 524 •. 'Walleyes and saugecrs commonly occur in tailwaters below dams in many river systems. Their occurrence is often seasonal, caused by the blockage of upst.ream migration or passage downstream from the reser- voir above. Concentration of prey fishes attracts walleyes and saugers to tailvaters . . 525. 'Walleye numbers have increased belov a number of dams ill the years following construction. They are the second most nume.r ·ous species in East Lynn Lake tailwater, West Virginia (Pierce 1969). They have increased in abundance in SUDDDersville tailvater, 'West Virginia (Goodno 1975); below four hydropower impoundments on the Au Sable River, Michigan (Richards 1976); and in the tailwater belov Stockton hydropower dam, Missouri, where they composed 28 percent of the catch ··-by anglers in 1974 as compared with only 9 percent in 1 ,972 (Hanson ~:·. 1974). ~: .. ., 526. Saugers are highly migra·tory, moving up.stream as much as 380 km in 18 days, through the navigation locks in the Tennessee River main-stem dams (Cobb 1960). Blockage of upstream .migration has pro- vided significant winter and spring fisheries in most varm main-stem dam tailvaters and in some cold tributary dam tailwaters on the Tennessee River system (Pfitzer 1962). An estimated 88,703 saugers were caught between November 1959 and March 1960 in Pickwick Dam taillrate.r {Trenary 1962). Cobb (1960) reported there was no sauger fishery in the Tennessee River until the main-stem dams were con- structed. 527. The sauger fisheries in cold tributary tailvaters of the Tennessee Riyer system are generally smaller than those below the main-st em dams. Periodically som.e saugers migrate upstr-eam from 'Watts Bar Reservoir and congregate in the cold Norris tailwater (Eschmeyer 1944; Eschmeyer and Manges 1945). Large numbers of saugers vere also 147 I . .. i I. I .. .. caught in the upper 23 kn of the cold Chilhowee Dam tailwater on, the Little Te·nnessee River. In 1964 and 1965, most were taken from December to March and composed 16 percent of the creel. In the downstream portion of the tailwater, 24 to 46 km below the dam, saugers were less abundant than in the immediate tailvater, but constituted 80 · percent of ~he anglers' catch (Boles 1969). 528. Passage of fis.h over dams from re·servoirs upstream is also · impor tant in establishing walleye and sauger fisheries in tail waters. An estimated 19,102 walleyes passed into an Ohio taih:ater over a 5- year period (Armbruster 1962). A large percentage (42 percent) died . from broken backs and pressure damage while pasS'ing through or over the -: . . . dam, but 58 percent survived ,passage into the tai lwater. Of tag re- coveries from walleyes tagged in the reservoir, 30 percent c.ame from t.he tailvater. Studies on the Missouri River bave s ·hown that· large ·numbers of young-of-the-year walleyes and saugers--up to 700,000 in 24 hours--moved out of Lewis and Clark Lake, South Dakota and Nebraska , and into the tailwater (Walburg 1 971). Mark-and-recapture studies indica.ted that :some adult sauge-rs also move from the reservoir int o -:the tail:water (Walburg et al. 1971}. Additionally~ the tailwater walleye fisheries in Hoover tailwater, Ohio, and Canton Reservoi r taU water, Oltlahooaa, are the result of the export of fish from the reservoir (Moser and Hicks 1970; Cavender and Crunkilton 1 974). A related species, the Volga pike-perch, has increased in the Kuibyshe•t tailwater, U.S.S.R., after successfUl reproduction in the reservoir (Sbaronov 1963). 529. Other factors, including water depth and. temperature, aff~ct tailwater walleye and sauger fisheries. The in.cre.aae in water depth vith a probable increase i n water temperature in the l over sections of Dal e Hollow tailwa.ter, Tennessee, due to inundation by Cordell Hull Reser- voir, was foll owed by the appearance of both walleyes and aaugers (Bauer 1976). Spor t fishing success for walle yes in Lake Taneycomo tail water, Missouri, is dependent on the wate.r level of Bull Shoals Reservoir, which inundates the tailwater. Best catches occurred when Bull Shoals Reservoir was 0.6 to 3.0 m below power pool level. Catches 148 ' .. ~- .. ',J.·'t . .-. .. -,. ~ . _ ... ,.,. .. _ progressively declined when water levels were either above or below this range (Hanson 1969) • 530. Water temperatures. in some col~ tailvate.rs have had a nega- tive effect on vall eye populations. Lowered vater temperature in a 'Korth. Carolina tailwater eliminated the walleye fishery (Dendy and Stroud 1949). Lov water temperature in Table Rock tailwater, Missouri~ has also affected the walleye catch. A rapid temperature increase of 5.6°C in the tailvater due to flood. flows spilling over the dam re- sulted in an immediate increase in'feeding activity and consequent increase in wall.eye catch (Fry 1965). 531. Reproduction of saugers has been adversely affected in some cold tailvaters. Saugers in a Tenness ·ee tailvater have sbovn signs of .resorbi.ng eggs. This vas attributed to lov water temperatures~ which ve·re generally less than 10.0°C (Escbmeyer and Smith 1943). 532. Flow regulation can influence walleye reproduction hundreds of kilometres downstream. Reduced winter flov·s from Bennett Dam. on the Peace-Athabasca River in Canada allowed the inlet to Lake Richardson ' to · t'reez~., 'thereby delaying access of walleyes to spawning area• in the :~:­ spring (Peace-Atbabasca Delta Project Group 1972; Geen 1974). A similar . situation was reported below the Volgograd li;,•droelectric Dam, U.S.S.R. Reduced spring f'lovs caused a deterioration of pik~-perch spawning . habitat far downstream in the Volga River Delta on the Caspian Sea (Orlova and Popova 1976) • 533. Water-level fluctuations in tailvaters have a negative in- fluence on sauger reproduction. Year-class strength in Fort Rand all · Dam tailvaters, South Dakota, vas 15 times greater in years when vate.r levels fluctuated only 0.8 m/day than in years when fluctuations were 1.4 m/day (Nelson 1968). Apparently, reduced water-level fluctuat ion · resulted in greater survival of eggs and larvae. To increase sauger abundance, Walburg (1972) recommended that water releases from Fort Randall Dam be not less than 566m3 /sec during the spawning and egg incubation period. 534 . Walleyes have been stocked in some tail waters where natural reproduction does not occur. A 26-km stretc h of river be,low the . ' -. I I. I i ,. I I Boysen Unit Dam, Wyoming, has provided a good wal~eye fishery as a result of stocking (U. S. Bureau of Sports Fisheries and Wildlife 1969}. Hicks (1964} recormnended stocking walleye fi,ngerlings instead of walleye fry in Tenk.iller tailwater, Oklahoma, because-··of·"Wlsuitable zooplankton supplies caused by intermittent water releases. The recom-............ mended stocking of fingerlings was appar ently successful, since both walleyes and saugers nov occur in the tailvater-(Deppert 1978"). 535. Food habits and grovth of walleyes and saugers in tailvaters ·· are not well documented. An examination of six walleyes collected from ..: Holyoke Darn tailvater, Massachusetts, shoved that fish made up · 96 per- · cent of the total volume of food in stomachs (Jefferies 1974). Saugers in Chilhowee tailwater, Tennessee, preyed heavily on stocked rainbow trout during the spring spawning run (Boles 1969). Food of walleyes and. saugers from Levis and Clark tailvaters, South Dakota and Nebraska, consisted primarily of gizzard shad, emerald .shiners, yellow perch, vhi.te bass, and white crappies (Wa.lburg et al. 1971). The grovth of .wa11eyes and saugers in Levis and Clark Lake tailvater vas superior ·: to that ·in t~e reservoir; for fish of si~la.r lenCtbs, ·the weights o~ · · _;-·' . _, •. ~ . . .· .. . · .. ~; ,··j. .,. . . .. walleyes and saugers vere respectively 7 and.l2 percent greater in the tailvater. Darters 536. According to Bailey et al. (1970), 109 species of darters are found in the United States and Canada. Comparatively .few ar·e ·.· mentioned in the tailwater literature. A general description Qf. I darter life history is presented because of the large number of species. 537. Habitat . Most darters are found in c l ear, smal·l -to medium- sized streams vi th permanent flow and clean, gravelly·, or rocky bottoms. They are most often found in the deeper sec tions of ri.ffles ,, but e.lso occur in rocky pools having no perceptible current. Some spe cies are .f more tolerant to turbidity than other s. The young of most species can be found in quiet-water areas associated vith leaves,·sticks, and organic debris. 150 ..-; :-!' ·~ .... .;. ••• :.:: ; ~. .· , ..... . ...... . , .... , ·····.'"·· ....... ..... .. 538. Darters are adapted for life in svift-floving streams. They sink immediately to the bottom vhen they stop svimming, and the press of the current against their enlarged pectoral fins tends to hold them in place. Darters are usually found beneath or betveen rocks and are thus afforded protection from the direct action of the current. When moving from place to place, they proceed by a series of short darts. 539. Reproduction. Most darter's usually spavn in lat~ spring over a sand or gravel bottom in vater about 0.3 m deep having moderate current. Eggs are laid and fertilized in a depression on the stream bottom, vhere they hatch in about 21 days at 21°C. Eggs and larvae receive no parental protection. 540. Some darter species attach eggs to strands of filamentous . algae or aquatic mosses and males establish territories. Breeding males of still other species seek out and occupy cavities beneath · rocks. Ripe females enter the cavity and deposit their eggs, vhere they adhere to the underside of the rock. The male stays vith the • ~ -~~ ·,;· .. : .. 1 .•• • • .t:~;.:·~~.:~. ;~,hit.'~ .. eggs until. they hatch. ..,.., .. · ,;· . 541. Food. . Darters are carnivorous, feeding principally on insects and other small aquatic invertebrates. 542. Age and growth. Lengths of adult darters other than logperch usually range from 64 to 89 mm, vith a maximum of about 100 mm. Males grov more rapidly and attain a larger size than females. Most are mature in the first spring after hatching and fev live longer than 3 year~. The logperch, the largest darter, usually ranges in length from 102 to 152 mm, but sometimes attains 178 mm. Darters in tailvaters 543. Darters have not been studied extensively in tailvaters. They flourish in a variety of.environments, and some have become established in cold tailvaters. Many darter species, particularly the orangethroat darter, rainbov darter, and logperch, are abundant in Beaver tail vater, Arkansas ( Brovn et al. 1968; Bacon e·t al. 1969; · Hoftman and Kilambi 1970) . Their abundance may be due to the uns·table composition of the fish population in this relatively nev tailvater. 151 -------· --------... ·-. -·----------.... ..---.... -------·-·--·--...___,...,. ____ ...,... ..... -...... -··-. --- ... Darter numbers are reduced in older tailwaters in the same drainage (Brown et al. 1968; Hoffman and Kilambi 1970). 544. The relatively low water temperatures (14.4-21. 7°C), low turbidities, mixed gravel-bedrock substrates, and high dissolved oxygen levels in Hoover tail water, Ohio, provide excellent habitat .for some _darter species. The logperch, greens ide ·darter, rainbow-dart-er.., and -•banded darter ·are all abundant, and the blacks ide darter, johnny darter, and fantail darter also occur there (Cavender and Crunkilton 1974). 545. The logperch, gilt darter, and band.ed darter were all abun- dant in the cold Chilhowee tailwater in Tennessee. The logperch was also nwmerous in the cold Norris tailwater (Hill 1978). The tessellated darter was dominant in the cold Rocky Gorge Dam tail water, Maryland (Tsai 1972). The orangethroat darter was the seco.nd most abun4ant s .pecies in the cold tailwater below Canyon Dam, Texas. The reduced temperatures in this taUwater appear to have extended the winter and spring breeding season of this species, and reproduction now occurs throughout the year (Edwards 1978). 546. Only one report on the age and. growth of darters in • _ . tailvaters was found·. Tsai (1972), who studied the tessellated darter in Rocky Gorge Dam tailwater in 1967, found that the mean. standard lengths of females at ages I, II, and III vere 35, 47, and 55 DID; males were 37 mm long at age I and 50 .mm at age -II. Sciaenidae (Drums) 547. The drum family contains many important marine fishes; only one is a freshwater species, the freshwater drum. Freshwater drum 548. Habitat. This fish is found in large, shallow lakes and large, slow-moving r:l.vers. It is usually found in the larger pools of streams and in lakes and reservoirs at depths of 9.0 m or more. The freshwater drum avoids strong current, 1 s usually found near the bottom, and is tolerant of high turbidity. It is particularly common in the ~tissouri and Mississippi rivers and t-he dovnstream sections of their 1 .52 I ~.'";. ' i"-., major' tributaries . It is also c011111on in Lake Erie and in many reser- voirs. 549. Reproduction. Spawning of the freshwater drum occ·urs in late spring or early summer, vhen water temperatures reach about 18°C. Spawning occurs over a period of about 6 veeks (Svedberg and Walburg 1970). .Eggs are fertilized in the open water and :t"loet until hatcbinf). They hatch in about 36 hours at 21 °C; nevly hatched larvae are 3. 2 mm long. The larvae are semipelagic until they are at least 15 mm long. The vide distribution of the freshwater drum in floving-vater systems is related to the pelagic state of their eggs and larvae. 550 . Food. The diet of freshwater drum consists mainly of fish, crayfish, and immature aquatic insects; mollusks are eaten if available. ·Young of the year eat mostly zooplankton and chironomids; as fish in- crease in size, larger aquatic insects become important. 551. .Age and growth. In Missouri streams, freshwater drum average 112 mm in length by the end of the first year of life and 2o6, 269, 315, 353, and 378 mm in succeeding years (Purkett 1958b). On the average, a· 330-mm · t1 sh weighs about 4 50 g and a 400-mm fish about · 900 "t;;.· .. g. Most drum caught by fishermen veigh 900 g or less, but individuals weighi.ng up to 1~.9 lt.g are occasionally taken. The mAximum life span is at least 13 years. Freshwater drum in tailwaters 552. The role of freshwater drum in tailvaters has not bee.n studied extensively. Several reports deal vith their occ·urrence in warm and cold tail waters. They are relatively common in the warm Keystone Dam tailvater, Oklahoma, v~ere they are .eaten by striped bass (Combs 1979). They are abundant in Levis and Clark Lake, a main-stem Missouri River reservoir on the South Dakota-Nebraska border, and large numbers of young of the year less than 25 mm long pass into the tailwater vith the discharge during the summer (Walburg 1.971). · In spite of the large numbers of young lost from the reservoir, adults are relatively uncommon in this warm tailwater (Walburg et a.l. 1971). Apparently fev of the young carried downstream in the river flow later return to the tailwater. Freshwater drum made up 10 percent 153 of the total sport catch in the cool Pomme de Terre tailvate:r, Missouri, from 1965 through 1974 (Hanson 1977). They were also common in the cold Tenkiller Dam tailwater, Oklahoma (Deppert 1978). The numbers of freshll!lter drum in a Mi ·ssouri tailvater have declined in 'recent years because water temperatures have decreased since completion of the upstream dBJII (Hanson 1969). Cottidae (Sculpins) 551 The sculpins are bottom-living. primarily marine fishes, of arctic and temperate seas; several genera are found in fresh waters of the northern hemisphere. This discussion vill be limited to the mottled and banded sculpins, vhich occur in some streams. Scul"Dins 554. Habitat. The tvo sculpin species often occur together because tbeir requirements are similar. The mottled sculpin is usually found in streams vitb clear, cold water, in both riffles and pools vith bottom types ranging from silt to gravel and. rock. Generally, it is most abundant in cover such as coarse rock or thick srovths or wa~er­ cress. The banded sculpin tolerates higher temperatures than the mottled sculpin and is the more abundant of the tvo in larger and va.""Dler ' streams. Sculpins live on the bottom, spending considerable time ly,ing motionless in one spot and moving in short, quick dashes. .... .555.. Rel:lroduction. The mottled sculpin spawns in the spring vhen vater temperature reaches about l~C. The adhesive eggs are deposited in clusters of about 200 on the undersides of stones. Tbe incubation period is 3 to 4 veeks and the male remains near the nest until the fry dispe!"se. 556. Food. Larval aquatic insects are the main diet of sculpins. Cottids are predaceous, but they do not feed extensively on eggs and young of trout, as is sometimes claimed (Pflieger 1975). 557. Arz.e and growth. The mottled sculpin is 28 to 36 mm lang vhen one year old. It probably does not mature until its third or fourth summer of life. Adults are commonly 60 to 90 mm long and the 1 54 .· . · ' . ·. . . . _. .... ,. ... 111&XiDru:m is about 115 mm. The banded scupli n is somewhat larger. Ad:ul t a are co.aonly 65 to 130 mm long and the I:I&Ximum is 185 mm or more. Sculpins i n tailvaters 558. Sculpins have not been extensively studied in tailvaters, although they are numerous in s001e cold tailvaters. Pfitzer (1962) noted that soulpins became important as forage in many cold tailvaters of the Tennessee Valley whe.n the number of minnow species declined. Both the banded sculpin and mottled sculpin have become numerous in Chilhovee tailvater, Norris ta.ilvater, and Apalachia tailvater (Hill 1978). The mottled sculpi'n is the most abundant species in both the Nor~ork and Bull Shoals tailwaters in Arkansas (Brown 1967; Hoffman and ICilambi 1970). Sculpins are also abundant in the McKenzie River. System. Oregon, vhere they are found in the cold tailvaters of four hydropower fa.cilities and one flood-control dam (Hutchison et. al. 1966). 559. Cottids are not found in all tailwaters. The mottled . sculpin has, disappeared below four hydropower dams on the Au Sabl e River, Michigan (Richards 1976), and were rarely collected during 1966 studies in the cold Beaver tailwater, Arkansas (Brown 1967) . 560 . Sculpins are highly susceptible to stranding during lar.ge water fluctuations because of their sedentary behavior. A total of 55 sculpins vere found ·stranded in three 0.82-m2 sections of the tailwater below a Wyoming dam. It was recommended that flow dec.reases not exceed 2.8 m3 /sec/day , to allow for fish migration out of the area (Kroger 1973). 561. Overall, reductions in flow do· not appear to affect sculpin survival. The piute sculpin (formerly eagle sculpin) is one of the· surviving native species in the Granby Dam tailwater, Colorado, in spite of large flow reductions. 562. Sculpins appear to reproduce suc cessfully in a number of tailvaters . The banded sculpin vas the only fish speci es able to revroduce in the cold Dale Hollov tailvater, Tennessee; no young of the year other than sculpins were observed (Little 1967). The partial inundation of a tailvater by a dovnstream reservoir beginning in about 155 1970 seriously reduced the abundance of the banded sculpin. and the ·' species is nov infrequently collected (Bauer 1976). 563. The food babi ts of sculpin& in tail:vaters are not well known. The common bullhead, a cottid which occurs in Cov Green tailvater on t .be 'l'ees River, United Kingdom, exhibited a feeding shift following impoundment of the reservoir. The reduction of Plecoptera caused the adults to begin feeding on mollusks • and t he fry shifted to Diptera and Epbemeroptera (Crisp et al. 1978), The f~ ot the banded sculpin in Dale Hollow t-ailvater, prior to inundat~ion by the downstream Cordell Hull Reservoir, cons.isted of Diptera, Coleoptera, Isopoda, c.rayfish, and small trout. The sculpin& did not eat Cladocera,. vbich vere abundant in the reservoir discharge (Little 1967) • . ' ·. -· 156 . · ~.· . -·"' . .. : ·,· . : . PART VIII: CONCLUSIONS 564. The construction or an impoundment alters the biological, chemical, and physical characteristics of the stream environment be,lov the reservo·ir. Many or the biological changes are a direct result or dam construction, and include blockage of upstream fish migration, inundation of spawning grounds, and the interruption of downstream invertebrate drift. In addition, the tailvater biota is influenced by the characteristics of the impoundment and the f 'aunal and geomorphic characteristics of the preimpoundment stream. 565. Physical and chemical characteristics in the tailwater are primarily determined. by the volume and t i.ming of water released and by the depth from vbich water is withdrawn from the reservoir. The effects of these releases are further modified within the tailwater by inflows from downstream tributaries and groundwater, riparian vegetation, atmospheric conditions, and physica1 characterist.ics of the streambed. The tailvater biota reflects interactions between the nati,ve or intro- duced organisms and the physical and chemical cond.i tions .in t.he , -' -, '. ·;·:·';"'<,. tailwater. . . ~· ,. : . • • • -. '•' ' : :~1'1. .... Effects of HyP<?limnetic Release on Downstream Biota 566. The depth of the discharge is of primary importance in deter- mining the tailwater environment below stratified reservoirs. The release depth affects vater temperatures, dissolved gas concentrations, nutrients, turbidity, and the presence of toxic concent.rations of some dissolved substances in the tailwater. These factors have a profound effect on the tailwater biota. 567. Maximum and average water temperatures are generally colder -----.in the tailwater below a hypolimnetic release ~serv~ir than in the -~ unimpounded stream. The effects of these coldwater releases are similar ----- on both warmwater and coldwater streams but are more severe for the . wa.rmvater streEW.S . The reduced wat er temperatures may fall bel ov t he tolerance levels of c.ertain native species of invertebrates and fish. 157 Lowered water temperature can increase the competition between native species and introducfl!d organisms adapted to the colder environment. The result ma~ be the loss of some native species from the tailwater. 568. A change in the seasonal water tempe,rature pattern also occurs in tailvaters. Water in hypoli.mnetic discharges is colder than that in the unimpounded stream during the sUDIIler and warmer during, the winter. Delays in spring warming because of cold hypolimnetic releases may e.l ter the .. reproduction, hatching, emergence, and developnent of many inve!"tebrates and fish. The altered tempe.rature· reg.i•es may disrupt the life cycles of sCDe insect spedes and. cause t .hem to emerge during the winter or prevent them t'rCD hatching in the spring. Some -fishes may not reproduce because the cold water disrupts their physio- logical developnent and eliminates the temperature stimul.us to spawn. Some may spawn several weeks late, retarding egg and larval ile.velopment. The smaller young are subject to more intense interspecific competition and reduced over-winter survival. 569. The temperature of cold tailwat.ers below bypolimnetic release dams built on wa!"mVater s t reams eventual.l.y returns to ambient as waters .·,· proce~d downstream. The biota in the downstream section closely re-.. sembles that in the natural unimpounded stream. A transitional zone may exist between the cold tail water and. the varm dovnstream river that may not be readily inhabited by ei'ther coldwater or varmvater organisms. This transitional zone is often larger than the immediate• tailvater (Hulsey 1959). ( 510. Hypolimnetic releases into coldva.ter streams generally do mot have e. drastic effect on the stream biota. \ilater temperat.ures in these tail waters generally remain vi thin the tolerance le·vels of the coldwater organisms that inhabited the original stream. Temperatures may sometices be reduced below the tolerance level of certain organisms vhich may disappear from the iirDDediate tailvater. This si tuatitm may also result in some redistribution of i nsects and fish. For example, chironomids and simuliids have replaced most other insect specie:s, and brook trout have replaced rainbow a .nd brown trout in some tailvaters. 158 : .. ... . . , ___ •;~··· .. . r·· 571. The volume of cold, hypolimnetic vater stored in the reser- voir and the loss of daily tempe·rature :f'luc.tuations in the tailvater atfects downstream biota. Some reservoirs lack sufficient sto·rage capacity of cold, bypoli.mnetic vater to maintain coldwater releases throughout the summ.er and f'e.ll.. Inadequate storage. capac.i ty may .result in the change from a coldwater to a varmvater tailvater during the latter part of the sUIIIIIier. This change significantly affects the tailvater biota, since coldwater organisms cannot survive in the varm vaters of late summer and ve.rmvate.r organisms cannot reproauce or grov in the cold waters vhic!l occurred earlier in the year. 572. The loss of diurnal fluctuations in vater temperature .may remove the temperature -stimulus necessary for normal progression of invertebrate life processes. Some invertebrates may disappear but other better adapted species usually replace them. Invertebr'ate popu-') lations in these stressed environments often display lov diversi 't,ies I and high densities. 573. Lov dissolved oxygen concentrations in tailvaters belov St·ratified deep-release reservoirs may cause physiological Stress in the aquatic coomrunity and limit fish and invert.ebrate diversity. Bio- logical decomposition in the hypolimnion of some reservoirs during the summer eliminates most of the dissolved oxygen and results in the re- lease of deoxygenated vater. If insufficien·t reaeration occurs in the outlet vorks, invertebrates may enter the drift and fish may actively migrate downstream. In extreme cases, lack of adequate dissolved oxygen has been responsible for the die-off of fish in tailvaters. In most tailvaters, however, turbulent flow over riffles rapidly inc reases the· dissolved oxyge.n concentration as the vater proceeds dovnstresm. 574. Nutrients vhic·h enter a reservoir may be used by the reser- voir phytoplankton or may settle into the hypolimnion. The dissolved nutrients vhich accumulate in the hypolimnion, either as a result of the decomposition of organic matter or directly from the wa'~ershed, are flushed into the tailwater during release of hypolimnet i c w~~c!r and may enhance primary productivity in the tailvaters. The additional nut!"i- ents may increase periphytic algal production in the tailvater and, 159 consequently, increase the numbers of invertebrates reeding on or living in the algae. 575. Toxic levels of' reduced substances, including iron, manga- nese, hydrogen sulfide, and anmonia, may be formed in the hypolimnion of a reservoir during low oxygen conditions. It is possible for these substances to be released into the tailvater at levels which may stress both the invertebrate and fish populations. 576. Turbidity is usually reduced in tail waters belov deep-release dams and organisms vhich are adapted to turbid waters may be at a competitive disadvantage. For example, reduced turbidity favors trout and other clear-water species over rough fish. In some instances, however, turbid inflows move through the reservoir as a density current and are released into the tailwater. Effects of EPilimnetic Release on Downstream Biota 577. Reservoirs with epilimnetic releases are generally less . ' disruptive t'o tail water "!:liota than are those vi th bypolimnetic releases • • <u• \ · ·· Warmvater str~ams are· subject to only minor temperature .~hanges as a result of the construction of an epilimnetic .release reservoir. The water temperature of the discharge is usually within the tolerance limits of the native va.rmwater stre8ln species and does not affect their survival. Add.:i.tionally, scme species of fish and invertebrates -may be ... transported out of the reservoir and added to the taflvater bi:9~a. Fish common in the reservoir often predominate in the tailwater. EpiUmnetic discharge from a dam built. on a coldwater stream usually has a higher summer and autumn temperature than the original stream. The varmer discharges may cause changes in the biota of the tailwater. Coldwater species ( plecopterans and trout) may be eliminated by temperatures exceeding thei.r tolerance levels. .E.levated temperatures also may allow rough fish (carp and suckers) to outCOl'lPete ,many of the native or i.ntro- duced species (trout). 578. Low dissolved oxygen concentrations rarely limit the biota below surface-release reservoirs. High levels of photosynthesis in 160 ... ~-. . ' •I ' .. - the reservoir epilimnion, gas exchange during passage of water from the reservoir, and downstream turbulent flow all increase concentra- tions of dissolved oxygen in the tailvater. Toxic concentrations of iron, manganese, and. hydrogen sulfide are rarely encountered since these materials are products of anoxic conditions in the hypolimnion. 579. Spillway releases from high dams, particularly in years of high fl.ov, can cause gas supersaturation in tailvaters and result in mortality to fish and invertebrates. Lov dovnstream temperatures and a laminar flow inhibits the dissipation of the dissolved gases back to the atmosphere. Gas supersaturation has been noted in the tail- vater belov a dam on a varmvater stream in the central United States, but the problem is most common at high dams on the larger rivers of · the Pacific Northwest. Nitrogen supersaturation and nitrogen em- ·.bolism generally do not occur at lowhead dams. 580. Epilimnetic discharges are generally low in dissolved · nutrients, since particles (algae, suspended material) containing ...... or sorbing nutrients settle into the hypolimnion. This loss of nutrientft result~ to ·~~duced' .primary produc~ivity i~ t .he immediate tailwater. However, the export of insects, phytoplankton and zoo- , ·:· ·· .. :~.\.,;.::-: .. , plankton from the reservoir often compensates for this reduction. Tlie exported organisms are often used as food and may increase the numbers of fish congregating below the dam. Many of the inverte- brates. may flourish below surface-release reservoirs because of the export or plankton and other suspended organic matter from the reservoir. -· .... 581. Many reservoir fish move into the tailwater either through the turbines or over the spillway. Tailwater populations of shad, . sunfishes, suckers , pikes, and some percids may be maintained through export fram the reservoir. In addition, native stream fishes often concentrate here, and consequently sport fishing in tailwaters is sometimes excellent. Tailvater fisheries are most successful in spring &nd early summer because of migrations related to spawning. Sustained low flovs adversely affect fishing success. 161 .. Effe~ts of Water Release Patterns on Downstream Biota 582. The timing and volume of vater released from a dam may severely limit or enhance the tailvater biot~:: Changes in the flow patterns after dam construction include seasonally stabilized flows, sustained high flows (vith reduced peak flows) during h~gh water ..... ., periods, minimum flows during dry periods, and diel fluctuating flows below hydropower facilities. The effect of these flow changes on the· biota may, however, be masked by other factors such as temperature, dissolved oxygen, and toxic levels of reduced substances. 583. ·The tailvaters below most nonhydropowe.r reservoirs ba.ve a more stable annual flow regime than that in unimpounded streams. The stabilized flows that result from both a reduction in the intensity or floods and t :he maintenance of flows during low-water periods provide a less variable habitat. Dense algal mats, often associated with stabi- lized flows, may inhibit the production of some native invertebrate species that prefer rock substrates. However, the additional habitat and food supply provided by the algal mats generally attract nev inTer- tebrate taxa.that become an important part of the topd web~ but species feeding on allochthonous material become less abundant. Nesting'fishes may reproduce more successfully in a stabilized tailwater. Stable flows may, however, be detrimental to fish that require moderate· flov variation to initiate spawning activities. 584. The effects of high flows (floods) in natural streams have been described by several authors (Tarzwell 1938; Seegrist and Gard 1972; Ryck 1976). High flows in tailwaters are generally less intense than in unaltered streams but are continued over a longer time. The effects of high flows on tailwater fishes are partly dependent on the tailwater physiography. If the tailwater has deep pools, sufficient cov~r. or backwater areas for fish shelter, high flows are less detri- mental to fish populations than if the tailvater has little physical variability. High flows during the reproductive per.iod of fish can scour the stream bottom and destroy the eggs and larvae and··reduce 162 .· ·. .. ,._·. i:-"•• ... ... .... ~····· ..... i. . -····. •• ! . .. •. ., ... ~ ~- ,, .... reproductive success, especially in unsheltered areas. Increased discharges may ca.use catastrophic drift of benthos, causing a sub- stan,tial decrease in benthic standing crops. High flows can be bene- ficial in some tailvaters as they flush sediment frcm the interstices of the rubble substrate and supply food and oxygen "':.o benthic invertebrates. 585. Fall dravdovn of flood control reservoirs increases flows into the tail water. Thi s drawdovn usually occurs after reserroir de- stratification, when fish and other organisms a r e more evenly distri- buted vi thin the reservoir. Many young-of-the-year and older fish may be lost from the reservoir during this time. causing an increased abundance in the ta ~ ,_vaters. The effects of the increased flows on the ·· resident stream fi.shes in the tailvater, both during the rese!'"'roir dravdovn and for sustained periods after flood flows, are largely unknown •. The increased flows may initially produce catastrophic effects on stream inverteb.rates, but t 'he ben'thic community gradually stabilizes ·. as the remaining flow-tolerant species adjust to the prevailing condi- tions ,' ·Higher flows extend the influence of the reservoir discharg~ '· .·.: · ·downstream. 586. The subject of mini'mum flovs is one of the most videly studied as.pects of regulated s treams. During the dry season of late summer and fall, mi nimum flows are often maintained below dams to provide aquatic habitat for the survival of invertebrates and fish . The habitat that remains, however, is usually of dec.reased quality and •; ., .. , .. ··· · ·· quantity. Reduced . .habitat inc·r eases interspec i fic competition. for apace and food ·among fish and among invertebrates. Both groups experi- ence physiological stress and reduced production during low flows . ... ~ flovs· in tailvaters reduce water velocities and associated detrital material, and thus food and oxygen for benthic organisms are also reduced . Lov flows allow silt and detritus to accumulate in the t&ilvater frOD1 streamside rtmoff . This material may be beneficial to the productivity i'n the tailvater if not present in excess i ve quanti- ties. In coldwater streams, minimum flows from deep-release reservoirs .·· ·• often maintain vate.r temperatures vi thin tolerance levels for coldwater species (e.g., trout.). 587. Irrigation storage reservoirs impound winter and spring runoff, vith a consequent reduction in tailvater flo•vs. Winter de- watering of the tailvater reduces overwinter survival of many organisms by limiting habitat and exposins them to harsh winter conditions. Poor survival of trout has been documented in streams devatered in winter. Dewatering in the spring probably affects the reproduction of some fishes by reducing both the stimulus to spawn and the availability of .. spawning habitat. 588. Below hydropower dams, large diurnal·flow fluctuations most often have a destructive influence on the tailwater biota and create an unstable, highly variable downstream habitat. Tbe extreme variation in flo,y SCO'.lrS the tailvater and displaces both fauna and flora. Species vith narrowly defined envirorunental requirements are eliminated from these tailwaters. During power generation, high water velocities cause streambed and bank instability and habitat degradation. The stream may .. be subject to increased. turbidity, and algal and macrophytic growth is · discouraged. The widely fluctuating flaws discourage the establishment of streamside vegetation and other aquatic plants .. The benthic !ood. base is radically reduced and some. species may be e1iminated. A "zone of fluctuation" is permanently establishe.d where no production takes place because of periodic streambed exposure during nonpower cycles. 589 .. Fluctuating flows disrupt the spawning and reproductive success of some fish species by destroying .nests and sweeping avay unsheltered eggs and fry. Only fish adapted :to high vater velocities are able to sustain their populations in tailvaters below hydropower dams. Stranding and desiccation of many species of invertebrates, fish eggs, salmonid fry, and sculpins have been ·reported,. Invertebrates located in 1luctuating tailwaters may attain community equilibrium, provided they are able to adapt to the vr.riati.ons in flow. 590. The release of large volumes ot cold bypolimnetic water during power generation maintains a cold tailwater environment below some southern reservoirs. However, during nongenerating periods, the 164 ' .. ,.._ ... . ' • " \,o'', '" ....... : small volume of water released may warm rapidly due to solar radiation and exposure to warm. air temperatures. The thermal tolerance of some tish and invertebrates may be exceeded during these periods, and the organisms either move out of the tailwater or die. The increased water temperatures may be beneficial to some species, such as carp and smellmouth bass, that compete with coldwater fishes. Despite the fluc- tuating flows encountered below hydropower dams, many excellent trout tisher:ies have developed in these waters. The quality of the fishery depends on the habitat suitability for trout, which includes cold water, adequate flow, plentiful cover and food. Past Tailwater Research and Suggestions for Future Study 591. .Review of the available literature revealed that the present understanding of biological problems in tailwaters is far from complete. CUrrent research being funded or conducted by various groups, including the U. S. Army Corps of Engineers, Tennessee Valley Authority, U. S. Water and Power Resources Service (formerly Bureau of Reclamation)~ and ·.·:·.:·.: ' • ,A._ the U. S. Fish and Wildlife Service, may provide the necessary infor-."' ; mat.ion. to overcome the inadequacy of the literature. 592. Most tailwater research has been narrow in scope. Usual!y, t .he study of the biota in a tail water has been l.im.i ted to the compila- tion of lists of invertebrate species and fish species or creel census . It has sometimes been assumed that a major change in one physical factor (e.g., temperature, flow, etc.) has caused a change in the tailwater biota. However, the more comprehensive investigations needed to confirm these assumptions have rarely been conducted. 593. The fev studies that have been directed toward. determining the causes for observed changes in tailwater biota have generally been the result of acute short-term problems (e.g., fish kills cau~ed by gas supersaturation or reduced dissolved oxygen). The mo ::-e subt].e changes resulting in the disappearance of a species or change o:: spe- ~ies composition in tailwaters have general.ly not been dete"rmined. As an example, the loss of a fish species in cold tailwaters has been 165 attributed to the reduction in temperature. What is usually not known is whi c h stages of the fish's life cycle were affected by the lowered temperature. Reduced temperatures may have had any of several effects or combinations of effects: (a) prevented the initiation of spawning activity, (b) inhibited the hatching of eggs, (c) impeded the growth of fry, (d) destroyed a required food. source, (e) provided other species with a competitive advantage, or {f) simply be.en below the tolerance limits of the affected species. If the "weak link" in the life cycle of the species was known, it might be possible to release water of a more favorable temperature during the critical period (e.g., releasing water of a wa_~er temperature until egg hatching is completed). 594. It is improbable that c'hanges in species composition were due solely to one factor, such as a reduction in temperature. Changes in the tailwater biota are more likely due to the alteration of a number of factors such as temperature, flow, ha:bita.t·availability, food abundance, and the levels of turbidity, dissolve.d gases, and certain chemicals. Additionally, the degree to which these &l.terations . affect · ·. ·, the''biota· in" each tail water may be highly variable. Th~ bi~t~ i~· 't¥o .' ., .. ~·t a,p·p~~ntly. s~~ilar tailwater~ may. "re~ct diffe~ntly t'o·i~i~ii~--~~~~5- in chemical and physical factors because of differences in project location, construction, and operation. Indeed, it is possible that in some tailwaters the assumed cause of the faunal changes (e.g., t~per­ ature ·redu~tion) may not have had any real effect, but simply masked the actual causative factors. 595. Studies are needed to determin~ the tailwater chemical and physical properties that cause changes in diversity and abundances in the biotic community. Such studies should investigate all parts of the tailwater ecosystem and must include continuous (i.e., daily) monitoring of major chemical and physical variables and periodic sampling of the tailwater biota (i.e., periphyton, plankton, benthos, fish). Once sufficient information on the chemical and phy-sical environment of a tailwater is obtained, it should be possible to relate this information to observed changes in the biotic community with appropriate statistical analyses. 166 ~. ··-.;:~~·. ), - •. . . . .. ...... .-.. •, ... . . ~ . 596. In addition to determining which chemical and physical factors act to alter the biotic community, further more intensive, limited studies will be needed to di scover how these factors act on the most seriously affected members of the biota. Only by determining how the affected tailwater organisms are influ.enced will it be possible to suggest improvements in the current tailwater management plan; for example, how a valuable varmwater fish species is influenced below a bypolimnetic release, hydropower dam. It may be determined that the disappearance of this species is most closely correlated with diurnal flow and tempe.rature fluctua·tions and delayed seasonal warming. More narrow·ly defined studies may d etermine that fish spawning has been delayed because of a lack of adequate thermal stimulus and that fish egg hatching and invertebrate food production have declined because of the periodic drying-of spawning beds and riffle areas ~ Ultimat~ly, it will be up to the managing agency to determine if any of the suggestions .. might be implemented within the context of an overall reservoir manage- m~nt plan . 597. The type of intensive study just discussed is only useful for understanding the problems of an individual tailvater being investi.gated . It would be helpful in the· future to have a generalized conceptual model of each major tailwater type. to assist in the recognition of tail~ter problems without resorting to large-scale intensi-ve studies. Such a conceptual model mu s t provide a clear understanding of how the major chemical, physical, and biologi.cal variables relate to and interact with each other. · ... -· ... . ~ .. .. ..... ......... .. • . ... 167 ': · REFERENCES Abduraltbmanov, Y. A. 1958. The effect ot' regulation on the flow of tb.e Kura River on the behavior and· abundance of fishes in the region below the Mingechaur Hydroelectric Station. Rybn. Khoz. ····· 3-4(12) :13-15. (Fish. Res. Board Can., Transl. Ser. 258). Aggus, L. R., D: I. Morais, and R. F. 'Baker. 1979. Evaluation o.f the trout fishery in the tailvater of Bull Shoals Reservoir, Arkansas, 1971-1973. Proc. Annu. Conf. Southeast. Assoc . Fi~h Wildl. Agencies. 31:565-573 . Allee, K. R. 1959. The distribution of stream bottom fauna. N. z. Ecol. Soc. Proc'. 6 :5-8. AmbUhl, H. von. 1959. The significance of flow as an ecological factor. Schveiz. z. Hydrol. 21(2):133-270. (Transl. by John Devitt, Humboldt State College, Arcata, Calif., U.S.A.). Anderson, K. R. 1972. Report to the Federal Power Commi.ssion on the fish and wildlife aspects of the relicensing of the Potter Valley Hydroelectric• Project (F .P.C. Projec.t No. 77), Lake and Mendocino Counties, California. Calif. Dep. Fis~ G8Jile. 59 pp. (Mimeogr.). · · Anderson, N. H., and K . W. Cummins. 1979. life histories of aquatic insects. ,; . 36( 3) :335-342 •. Influences of diet on the J. Fish. Res. Bo.ard Can. ·;c.·' • · 'Andrews, A; K., G. A. Earls, and R. C. Summerfelt. 1974. Rec ~eational use of an Ok.laboma' scenic river b.isected "by a mainstream bydro- e.lectric impoundment. Okla. Dep. Wildl~ Conserv., Fed. Aid ·. Proj. 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Some factors associated with fluctuation in year-class strength of sa:uger, Levis and Clark Lake, South Dakota. Trans. Am. Fish. Soc. 101(2):311-316. Walburg, C. H., G. L. Kaiser, and P. L. Hudson. 1971. Levis and Clark Lake tailwater biota and some relations of the tailwater and r~servoir fish populations. Pages 449-467 in G. E. Hall, ed. Reservoir Fisheries and Limnology. Am. Fisb. Soc. Spec. Publ. 8. Walburg, c. H., J .. F. Novotny, K. E . Jacobs,.T. M. Campbell, and w. D. Swink. 1980. Water quality,·macroinvertebrates, and fisheries in tail waters and streams -an annotated bibliography. National Reservoir Research Program, U. S. Fish Wildl. Serv. Contract Report to Vicksburg, U. S. Army Engineer Waterways Experiment Station. 199 pp. Wallace, J. B., J. R. Webster, and W. R. Woodall. 1977. ·.The role of filter feeders in flowing waters. Arch. Hydrobiol. 79(4): 506-532. Ward, J. v. 1974 •. A temperature-~tressed ~tream ecosystem below a bypolimnial release mountain reservoir. Arch. Hydrobiol. 74(2) :247-275. 187 -···. Ward, J. V. 1975. Dm•nstream fate of zooplankton from a hy'polimnial release mountain rese~voir. Verb. Int. Verein. Limnol. 19:1798-1804. Ward, J. V. 1976a. Effects of flow patterns below large dams on stream benthos: a review. Pages 235-253 in J. F. Orsborn and C. H. Allman, eds. I~.lstream Flow N'eeds, Vol. 2. Am. Fish. Soc., Washington, D. C. Ward, J. V. l976b. Comparative limnology or differentially regulated sections of a Colorado mountain river. Arch. Hydrobiol. 78(3): 319-342. Ward, J. V. l976c. Effects of thermal constancy and seasonal tempera- ture displacement on coumunity structure of stream macroin:verte- brates. Pages 302-307 in G. W. Escb and R. W. McFarlane, eds. Thermal Ecology II. PrO'Ceedings o·f a symposium held at Augusta, Georgia, April 25, 1975. Energy Res. and Dev. Admin., · Washington, D. C. Ward~ J. V., and R. A. Short. 1978. Macroinvertebrate community structure of f 'our special lotic habitats in Colorado, U.S.A. Verb. Int. Verein. Limnol. 20:1382-1387 ... Ward, J. V., and J. A. Stanford. 1979. Ecological factors controlling stream zoobenthos with emphasis on thermal modification of regulated streams. Pages 35-56 in J. V. Ward and J. A. Stanford., 'eds •. }'h~ Ec.ology of Regulated Streams. Plenum Press, Rev York. ·~_Warden, R. L., Jr.; and W. A. Hubert. 1977. · ·•· · · Lepomis from the Wilson Dam tailvater. 179-183. Spring stomach contents of J. Ala; Acad. 'Sci. 48(4): Waters;T. F. drift. 1964. Recolonization of denuded stream bottom areas by Trans. Am. Fish. Soc. 93(3):311-315. Waters, T. F. 1969. Invertebrate drift -ecology and significance to stream fishes. Pages 121-134 in T. G. Hortbcote, ed. Symposium on Salmon and Trout in Streams-.-H. R. MacMillan Lectures in Fisheries. Univ. B.C., Vancouver. Waters, T. F. 1972. The drift of stream insects. Annu. Rev. Entomol. 17:253-272. Weber, D. T. 1959. Effects of reduced stream flows ou the trout fishery below Granby Dam, Colorado. M.S. thesis, Colo. State Univ., Fort Collins. 149 pp. Webster, J. R., E. F. Benfield, and J. Cairns, Jr. 1979. Model pre- dictions of effects of impoundment on particulate organic matter transport in a river system. Pages 339-364 in J. V. Ward and J. A. Stanford, eds. The Ecology of Regulated Streams. Plenum Press, New York. Weitkamp, D. E., and M. Katz. sat.ur.ation literature. 1980. A review of dissolved gas super- Trans. Am. Fish. Soc. 109(6):659-702. 188 ,. :• Welch, E. B. 1961. Investigation of ~ish age and growth and food abundance in Tiber Reservoir and t 'he river below. Mont. Fish Game Dep., Fed. Aid Proj. F-5-R-10. 18 pp. Wesche, T. A. 1974. Relationship of discharge reductions to available trout habitat for recommending suitable stream flows. Water Resour. Ser. 53. Water Resour. Res. In.st., Univ. Wyo., Laramie. 73 pp. Wetzel, R. G. 1975. Limnology. W. B. Saunders Co., Philadelphia. 743 pp. White, R. L. 1969. Evaluation of catchable rainbow trout fishery. Tex. Parks Wildl. Dep., Fed. Aid Proj. F-2-R-16. 18 pp. Whitford, L. A. 1960. The current ef'fect and growth of fresh-water algae. Trans . .Am._Microsc. Soc. 79(3):302-309. \ol"bit .ford, L. A. , and G. J. Schumacher. 1961. Effect of current en mineral uptake and respiration by a fresh-water alga. Limnol. Oceanogr. 6:423-425. Williams, R. D., and R.· N. Winget. 1979. Macroinvertebrate response, to flow manipulation in the Strawberry River, Utah (U.S.A.). Pages 365-376 in J. V. Ward and J. A. Stanford, eds. The Ecology o.f Regulated Streams. Plenum Press, New York. Wiltzius, w. J. 1978. Some factors hi.storically affecting the di-stri- bution and abundance of fishes in the Gunnison River. ·Colo.···· Di.v. Wildl., Final· Report. 202 P.P· Wirth, T. L., R. C. Dunst, P. D. Uttormark, and W. Hilsenhof.f. 1970. Manipulati~n of reservoir waters for improved quality and fish population response. Wis. Dep. Nat. Re·sour., Res. Rep. 62. 23 pp. Wright, J. C. 1968. Effect of impoundments on productivity, water chemistry, and heat budgets of rivers. Pages 188-199 in Reservoir Fishery Resources SYlllposium. Sout'hern Div., .Am. Fish. Soc. Yang, C. T. 1971. Formation of ri.ffles and pools. Water Res our. Res. 7(6) :1567-1574. Young, W. C., D. H. Kent, and B. G. Whiteside. 1976. The infiuen~e of a deep storage reservoir on the species diversity of benthic macroin.vertebrate commu."li ties of the Guadalupe River, T.exas. Tex. J. Sci. 27(1):213-224. 189 .. . ·- •,. 'It ~· . . ~ . . .. . .. ··!" .. . .... .. . . .. .. ., \. ·•'· · . . .,.1. . :_ .APPEI.'lDIX A: ALPHABETICAL LIST OF THE 113 TAILWATERS MENTIONED .. ":. ,: . . IN THE TEXT WITH LOCATION BY RIVER AND STATE, PROVINCE, OR COUI'l'RY > 1\) Antelope Apalachin Augusta Barren Deaver Bennett Berlin Big Bend Bliss Blue Lake Tail water 'Blue Mesa Boulder (Hoover) Boysen Unit Broken Bow Buckhorn Bull Shoals Caballo Calderwood Canton Canyon Cape Horn Diversion Carl Blackwell Carlyle Center Hill River Jordan Creek Hiwasse Kennebec Barren White Peace, Athabasca Mahonlng Missouri Snake Sawmill Creek Gunnison Colorado Wind Mountain Fork Kentucky White. Rio Grande Little Tennessee North Canadian Guadelupe Eel Stillwater Creek. Kaskaskia taney Fork (Continued) State(s), Province(s) or Country Oregon Tennessee .Maine. Kentucky Arkansas Alberta, British Columbia Ohio South Dakota Idaho Alaska Colorado Nevada, Arizona Wyoming Oklahoma Kentucky Arkansas New Mexico Tennessee Oklahoma Texas California Oklahoma Illinois Tennessee > w Cherokee· Chilhowee Tailvater C. J. Strike Clark Canyon Clearwater Conowingo Cooke Cordell Hull Cottage Grove Cow Green · Coyote Cumberland Dale Hollow Davis Dexter Diversion (unnamed) Dorena Douglas East Lynn Elephant Butte Ennis Fern Ridge Five Channels Flaming Gorge Fontana Fontenelle Foote Fort Randall ·., .· . River Holston Little Tennessee Snake Beaverhead 'alack Susquehanna Au Sable Cumberland Coast Fork Willamet te Tees Russian Cumberland Obey Colo·rad.o Middle Fork Willamette Blacktail Creek Row French Broad Tveh•epol e Cr eek Rio Grande Madison Long, Tom Au Sable Green Little Tennessee Green Au Sable M1ssouri (Continued) State(e), Province(s) or Country Tennessee Tennessee Idaho Montana Missouri .Maryland Michigan Tennessee Oregon United Kingdom California Kentucky Tennessee Arizona, Nevada Oregon Montana Oregon Tennessee West Vi rginia New Mexico Montana Oregon Michigan Utah, Colorado North Carolina Wyoming Michigan South Dakota Tail water Glen Canyon Granby Hartwell Hebgen Hog Park Holyoke Hoover Jackson Lak.e Kentucky · Keystone Kuibyshev Lake of the Ozarks (Bagnell) Lewis and Clark (Gavins Point) Libby (Koocanusa) Little Goose Little Grassy Lock and Dam 12 Lookout Point Loramie Low head (unnamed) Lower Salmon Falls McNary Mingeshaur Mio Morrow Point N~vskaya ·'· j.-· .··:·.·~- ~.. !'I '-. River Colorado , Colorado Savannah · Madison Hog Park Creek Connecticut Big Walnut Creek Snake Tennessee Arkansas Volga Osage Missouri. Kootenai Snake Crab Orchard Creek Mi.ssissippi Middle Fork Willamette Loramie. Maquoketa · Snake ColtDDbia · Kura Au Sable ~' ~~· Gunnison : Narova (Continued) State(s), Province(s) or Co\Ultry Arizona Colorado Georgia, South Carolina Montana Wyoming Massachusetts Ohio Wyoming Kentucky Oklahoma U.S .S.R. Missouri Nebraska, South Dakota Montana Washington Illinois Illiqoi~, Iowa Oregon · Ohio Iowa Idaho Oregon, Washington U.S.S.R. Michigan Colorado U.S.S.R. .. Tail water Navajo Nolin Norfork Norris Oahe Owyhee Perc.ha Diversion Pickvick. PoDIIle de Terre Roanoke Rapids Rob Roy · Rocky Gorge (Triadelphia) Rough Shand Shasta South Holston Stockto.n SUJJIIIersvilJ.e Table Rock Taneycomo Taylor Park TenkUler Tiber Twin Valley Upper Salmon Falls Urievllle •state owned pondu. RiYer San Juan Nolin .' North Fork Cli.nch Missouri Owyhee Rio Grande Tennessee Po11111e de Te.rre Roanoke .Douglas Creek Patuxent Rough Grand Sacramento South Fork Holston Sac Ga.uley White White Taylor Illinois Mari.as Mill Creek Snake • (Continued) State(s), Province(s) or Country New Mexico Kentucky Arkansas Tennessee South Dakota Oregon New Mexico Tennessee Missouri North Carolina Wyoming Maryland Kentucky Ontario California Tennessee Missouri. West Virginia Missouri Missour.i Colorado Oklahoma Montana Wisconsin Idaho Maryland > 0\ Vir Volgograd Watauga Watts Bar Wilson Wister Wye Tail water Wyman Yellowtail -state owned ponds. •. ., . Svratka Volga Watauga Tennessee Tennessee Poteau • Kennebec Bighorn .· River State(~). Province(s) or Country · C7.echoslovakia U.S.S.R. Tennessee Tennessee Alabwna Oklahoma Maryland Maine Monta:na ----~------------------------------------------------------------------------------------------- ... ·· APPENDIX B: ·LOCATION OF 105 RESERVOIR TAILWATERS ., IN THE UNITED STATES MENTIONED IN THE TEXT ..... 6-1 ~ . ., .... ~··· w 1\) .. .. . "'-· .": . ; . Figure Bl ' . •-. . . · . • . APPENDIX C: . COMMJN AND SCIENTIFIC NAMES OF FISHES MENTIONED IN THE TEXT • ARRANGED BY FAMILY t-\ •' .... ' .. . ; ' Part I: Fishes from North American Tailwaters ······ . Polyodont i dae · . Paddlefish Clupeidae Skipjack herring American shad Gizzard shad Th:::-eadfin shad Salmonidae ..... -· Coho salmon Cutthroat trout Rainbow trout Brown trout BTook trout Esocida.e Grass pickerel Northern pike Muskellunge Chain pickerel CyJ>rinidae Chiselmouth Stoneroller Carp Humpback chub Bonytail Roundtail chub Speckled chub Bigeye chub Streamline chub Peamouth Hornyhead chub River chub Golden shiner Texas shiner Rosefin shiner Emerald shiner Polyodon spathula (Walbaum) Alosa c sochloris (Rafinesque) Alosa sanidissima Wilson) no;;soma cepedianum (Lesueur) Dorosom.a pete·nense (GUnther) Oncorhynchus kisut.ch (Walbaum) Salmo clarki Ri.chardson ~ gairdneri Richardson Salmo trutta Linnaeus ~linus fontinalis (Mi tchill) Esox americanus vermicul.atus Esox lucius Linnaeus Es·ox masouinone;y Mitchill Esox niger Lesueur. , ... ,., Lesueur Acrocheilus a1utaceus Agassiz·and · Pickering Campostoma anomalum (Rafinesque) Cyprinus carpio Linnaeus Gila cypha Miller Gila elegans Baird and Girard Gila robusta Baird and Girard HYbQpsis aestivalis (Girard) Hybopsi s amblops ( Rafi.nesque) Hybopsis dissimilis (Kirtland) Mylo e heilus caurinus (Richardson) Nocomi.s biguttat.us (Kirtland) .Nocomis micropogon (Cope) Notemigonus crysoleucas (Mitchill) Notropis amabilis (Girard) Notropis ardens (Cope) Notropis atherinoides Rafinesque (Continued) C2 :•:. •,_ .•. ;·. ·-;-. ,· Cyprinidae (continued) Bigeye shiner Striped shiner Common shiner Whitetail shiner Blackchin shiner Spottail shiner Red shiner Silver shiner Dusky!ftripe shiner Rosyt'ace shiner Spotfin shiner Sand shiner Teles~ope shiner Redt'in shiner Blacktail shiner Mimic shiner Suckermouth minnow Southern redbelly dace Bluntnose minnow . Fathead minnow . ....... , . Bullhead minnow : · · · Colorado · squaWrish Northern squawfish Blacknose dace · Longnose dace Speckled dace Redside shiner . Creek chub Catostomidae River carpsucker Quill back Longnose sucker White .sucker Bluehead sucker Flannelmouth sucker Largescale sucker Mountain sucker Blue sucker ·. . Northern hog sucker SmalJ.mouth buffalo Bigmouth buffalo Black buffalo .. ; ~. ~ .... ,J -•• Notropis boops Gilbert Notropis ehrysocepbalus (Rafinesque) Notr opis cornutus (Mitchill) Notropis galacturus (Cope) Notr opi s heterodon (Cope) Notropis hudsonius (Clinton) Notropis lutrensi s (.Baird and Girard) Not ropis photogenis (Cope) Notropi s pilsbryi Fowler Notropis rubell us (Agassiz) Notropis spilopterus (Cope) Notropis stramineus (Cope) Not roois t elescop1ls (Cope) Notropis umbratili s (Girard} Notroni s venustu s (Girard) Notropis volucellus (Cope) Phena c obius mira.bilis (Girard) Phoxinus erythrogaster (Rafinesque) Pime phales notatus (Ra.t'inesque) Pimephales oromela.s Rafinesque Pimepha.les vie;ilax (.Baird and Girard) : Ptychacheilus lucius G~rard · -: :· · .;..; · Ptychocheilus oregonensis (Richardson) Rbinichthys atratulus (Hermann) · .Rhinicbtbys cat a.ractae · (Valenciennes) Rhi n i cht hys osc.ulus (Girard ) Ricbardson ius bal teat us (Richardson) Semotilus atromaculatus (Mitchill) Caryiodes carpio ( Ra.finesque) · Carpi odes cyprinus (Lesueur) .. Catostomus catostomus (Forster) Cat osto.mus commerson i (Lacepede)' Cato stomus discobolus Cope Catostomus latipinnis Baird and Gi rard Catostomus m.acrocheilus Girard Catos tomus olatyrbyn.chus (Cope) Cyele;ptus elongatus {Lesueur) Hypentel ium nigr.icans (Lesueur) Ict.iobus bubalus (Rafi oesque) Ictiobus . cypr'inellus (Valenciennes) Ictiobus niger (Rafinesque) (Cont i nued) C3 Catostomidae (continued) Spotted sucker Silver redhorse River redborse· Gray redhorse Black redhorse Golden redhorse Shorthead redhorse Humpback sucker Ictal uridae White catfish Blue catfish Black bullhead Yellow bullhead Brown bullhead Ch.annel catfish Slender madtan Sto·necat Tadpole madtom Brindled madtom Freckled madtom Flathead catfish · Percichthyid&e White bass Yellow bass Striped bass Centrarchidae Rock bass Green sunfish Ora.ngespotted sunfish ······'Bluegill Longear sunfish Redea:r sunfish Smallmouth bass Spotted bass Largemouth bass White crappie Black crappie Miny-trema melanops (Rafinesque) Moxostoma anisurum (Rafinesque) Moxos:toma. carina tum (Cope) Moxostoma congestum { Bail1 and Girard) M'oxostoma duouesnei (Lesueur) Mox.ost.oma erythrurum {Rafinesque) Moxostoma macrolepidotum (Lesueur) Xyraucben texanus (Abbott) Ictalurus catus (Linnaeus) Ictalurus ~tus (Lesueur) Ictalurus me~as (Rafinesque) Ictalurus iiit8i'is {Lesueur) Ictalurus nebulosus (Lesueur) Ictalurus punctatus (Rafinesque) Noturus exilis Nelson Noturus flavus Rafinesque Noturus e;:rrinus {Mitchill) Not u:rus . miurus Jordan Noturus nocturnus Jordan and Gilbert Pylod.ictis olivaris (Rafineaque) Moron.e chrysops (Rafi'nesque) Morone mississippie·nsis Jordan. and Eigenmann Marone saxatilis (Walbaum) Amblonlites rupestris {Rafinesque) Lenomis cyanellus Rafinesque Lepomis humilis (Girard) .Lepomis macrochi rus Rafinesque Lenomis megalotis (Rafinesque) Leopmi s microlophus (Giinther) Micropterus dolo~eui Lacepede Micropterus punctulatus (Rafines que) Micropt erus .s a lmoides (Lacepede) Pomoxis annul.aris Rafinesque Pomoxis nigroma c ulatus (Lesueur) {Continued) c4 . ... Perc.idae Greenside darter Rainbow darter Fantail darter Johnny darter Tessellated darter Orangethroat darter Banded darter YelJ.ov perch Logperch Gilt darter Blacks.ide darter Sauger Walleye Sciaenidae Freshwater drum Cottidae Mottled sculpin Piute sculpin .Banded sculpin •• J ......• Etheostoma blennioides Rafinesque Etheostoma caeruleum Sto.rer Etheostoma tlabel.lare Ra~inesque Etheostoma n.igrum Rafinesque Etheostoma ol.mstedi Storer Etheostoma s ectabile ( A8assiz) Eth.eostoma. zonale Cope ) Perea flavescens (Mitchill) ~na caprodes (Rafinesque ) Percina evides (Jordan and Copeland) Percina maculata (Girard) Stizosted.ion canadense (Smith) Stizostedion vitreum. vitreum (Mitchill) Aplodinotus grunniens Rafinesque Cottus bairdi Girard Cottus beldingi Eigenmann and Eigenmann Cottus carolinae (Gill) C5 . ...... . I I ,, Part II: Fishes from Europea.n Tail waters Salmonidae Baltic salmon Cyprinidae Zope Bream Golden shiner (European) Bystr:ranka Bl:eak Barbel White bream Shemaia Nase (Podust) Carp Gudgeon Ide Roach or Vobla IChramulya Vimba Percidae Volga pike-perch Cottidae Common bullhead (sculpin) ~ salar Linnaeus Abramis bal~erus (Linnaeus) Abramis ~ (Linnaeus) Abramis sp. Alburnoides bipunctatus (Bloch) A1 burn us a1 bum us { Li nnaeus ) Barbus be:rbus (Linnaeus) BUcca bjoerkna (Linnaeus) Chalcalburnus chalcoides (~Uldenstadt) Chondrostama nasus (Linnaeus) Cyprinus caroio Linnaeus ~ gobio ~Linnaeus) Leuciscus idus ( Linnaeus) Rutilus rutuUs (Linnaeus) Varicorhinus capoeta (~Uldenstadt) Vi mba ~ ( Linna.eus) Lucioperca volgensis (Gmelin) Cottus gobio Linnaeus C6 .I ·. . ,· .•. APPENDIX D: LIFE HISTORY INFORMATION FOR THE M:>ST COMK>N FISH GROUPS MENTIONED IN THE TEXT :D-1 Fish group, occurrence Gizzard shad Rivers, lakes, and reservoirs; most found eastern U.S. Trout Rivera, lakes, and reservoirs; common throughout U.S. Carp Rivers, lakes, and reservoirs; common throughout U.S. Suckers Category (forage, F; game, G; rough, R) F G .R Rivera, lakes, and reservoirs; various species round through- out U.S. R '· Table Dl -Fish Groups, General Characteristics Life span years 4-6 3-5 5-6 5-6 First Age years 2-3 2-4 .; 2-4 maturit;y: Length nn Habitat 250-350 Pools, backwaters 250-380 Riffles, pools 300-450 Backwater pools 300-400 Pools and· riffles (Continued) 0 ~ Pre terence's Temperat'ure oc 22-24 10-15 20-22 14-20 ., Water clarlt;y: No preference Clear :Turbid Clear to . turbid '• ' (Sheet 1 or 3) t1 w . . Category (forage, F; Fish group, game, G; occurrence rough, R) Channei catfish Most common in G large rivers but also found in some reservoirs; intro- duced throughout the U.S. White bass Lakes, rivers, and G reservoirs; intro- duced throughout the U.S. Black basses* Found throughout G the U.S . in lakes, reservoirs, and rivers :;· . J. :-. \ 0 • ' Table Dl (Continued) Life First maturit;y: span Age Length years . ;y:ears IIDil --- 6-10 4-6 330-560 4-6 2-3 250-300 4-7 3-4 250-320 (Continued) • Preferences Teaperature Vater Habitat oc clarit;y: Deep pools 27-30 Semi turbid Deep pools 23-25 Clear Pools 21-27 Clear *Three species are c~mmon--largemouth,'smallmoutt.~ and spotted--and eac~ has different requirements. .· (Sheet 2 of 3) Table Dl (Continued) Category (forage, F; Life First maturitz PreJ:erences Fish group, game, G· , span Age Length Temperature · Water occurrence rO!!ejh 1 R) zears , zears nun llabi tat oc clarity Sunfishes Lakes, rivers, and F,G 3-4 2-3 100-150 Pools 22-25 Clear to reservoirs; m.ost turbid common eastern two-thir ds U.S . Crappies Lakes, rivers, and F,G 3-4 2-3 150-200 Pools 22-24 Clear t o res ervoirs; most slightly t;l .e:-common easter n turbid half U.S. Walleze Lakes, rivers, and G 5-7 3-4 350-450 Open water, 18-21 Clear reservoirs; most pools COftlllon eastern half U.S. (Sheet 3 of 3) ( Fish sroup Gizzard shad Trout d VI Carp Suckers Table Fish Groups, Spawning and Spawning Season Spring, early Winter, spring Spring, early summer Late winter, spring Temperature oc 17-?3 4-16 17-20 13-23 Location Shallow, protected areas (scattered) Riffle (nest) Flooded shallows (scattered) Shoal areas (scattered) D2 Eating Characteristics IncUbation period dazs 4 20-Bo 4-8 4-14 Food Youna Adults Plankton Pl ankton Plankton Insects, fish Plankton Insects, plant material Plankton Insects, algae (Continued) . .. f: • Remarks Young ooDJDon in taflwaters dur- ing fall and winter; impor- tant forage. The rainbow is the most common trout stocked in cold tallvaters. Adaptable to· many habitats; considered a nuisance because of destruct! ve habits. About 12 species ot suckers, in- cluding redhorse s uckers, occur in tailwaters; 11 fe history varies by species. (Sheet 1 o~ 3) . Table D2 (Continued) SP,!vnins Incubation Temperature period Food Fish sroul2 Season oc Location dals Youns Adults Remarks Channel catfish Late 24-29 Natural 5-10 Plankton, Insects, Highly sought as spring, cava ties insects fish food fish, early (nest) particularly swrmer southeastern u.s. White bass Spring, 14-21 Midwater 2 Plankton Insects, Important tail- early over hard. .fi.sh vater sport fish swmner bottom .. in spring and 8'-. ( scatte.red) summer. Black basses Spring, 16-18 Sand-2-4 Plankton, Insects, Important aport early gravel insects fish fish. Small- summer (nest) mouth and spotte.d basses most common in streams; large- mouth in lakes. Sunfishes Spring, 20-22 Firm 3-5 Plankton Insects Important sport early bottom fi.shes. Blue- summer (nest) gill , green and longear sun- fishes most com- mon in tailvaters. ,'(f (Continued) (Shee·t 2 or 3) Fish group Season Crappies Spring Walleye Spring ... ... , .·: ........ ,· Table D2 {Continued) Spawning Temperature oc 6-8 Locat·ion Firm bottom (nest) Gravel, rubble (scattered) Incubation period days 3 12-18 Food Young Adults Plankt.on • Insects • insects fish Plankton, Fiah insects • • Remarks Important sport fishes. Adults may be common in tailvaters dur- ing spring and young during fall and winter. Important sport f:lsb in tail- waters of some large rivers. (Sheet 3 of 3) .. · . • -.... . ·' : .. -!· '· .~' ,· ... ,r.;. _:! .. ·: ·.: '.t· ......... .,. . APPENDIX E: ··. GLOSSARY .,JI• . , :-: .-r,. ~.~: ·• .. -~· ... • • £-1 Algae: primitive plants, one-celled to many-celled. Alloc.hthonous: materials such as leaves and detritus that originate from outside a l~e or stream. Amphipods: group of crustaceans that includes the freshwater forms Hyalella and Gammarus. Anadromous fish: fish that spend most of their lives in the sea or lakes but ascend rivers to spawn. Anaerobic organisms: microorganisms that thrive in the absence of oxygen. Annelids : earthworms and leeches . Anoxia: state of having too little oxygen in tissues for normal met.abol:!.sm. Arachnids : spiders and water mites. Armoring: acctDDulation of coarse particles on a stream bottom through loss of finer materials to the current; the formation of a firm layer on the streambed that is resistant to further degradation. Arthropods: group of invertebrate animals that 'includes . crustaceans, insects, and spiders. Autochthonous: materials such as algae, macrophytes, and their decom- posi t .ion products that originate vi thin a lake or stream. Auto~ropby: type of nutrition in which an organism manufactures its own· food from inorganic compounds. Benthic org~isms (benthos): immature aquatic insects, stream bottom. aquatic invertebrates such as mollusks,. and crustaceans that live on or in the Biomass: total weight of a particular species or of all organisms in a particular· habitat. Biota: all living organisms in a region. Bryozoans: small bottom organisms that make up part of the benthos. Carnivore: any animal partly or wholly dependent on catching other animals for its 'food. Chironomids: family of insec~s of the Order Diptera; large group that includes the non biting, mosqui tolike midges. Cladocerans: freshwater crustaceans; includes such zooplankton genera as Daphnia, Chydorus, and Alona. E2 .. ' • ~t • . : ···.•· .... • .. ·. Copepoda: freshwater crustaceans; includes such zooplankton genera as Diaptomus and, Cyclops. Crustaceans: includes certain zooplankters (copepods and cladocerans), ampbi,pods, dec a pods, 1 sopods , and, ostracods. Daphnids: any member or the cladoceran genus Daphnia. 'Oecapods: freshwater shrimp and crayfish . Detritivores: organisms 'that ingest detritus • . Detritus: fine particulate debris or organic or inorganic origin. Diatoms: class or r:a.lgae having silicified skeletons. Dipterans: order of insects that includes true flies. Drift: . aquatic or terrestrial invertebrates that move or float with ·· the current. Ecology: science or the interrelations between living organisms and the.ir environment. Encystment.: formation of a resi·stant cyst by c.ertain microorganisms, especially under un.!avorable environmental conditions. Ephemeropterans: order or insects that .inc~udes the mayflies. Epilimnion: upper stratum of more or less uniformly warm circulating water that forms in lakes a.nd reservoirs during periods or stratifi- cation and extends from the surface to the metalimnion or thermocline. Excystment: portion of the life cycle of an organism when it e111erges from its cyst stage and resumes normal. metabolic activity. Fingerling: immature fish, from a length or about 25 mm (or size at disap,pearance of yolk sac) to the end of first year of life. Fry: life stage· or fish between hatching o'f the egg and assumption of adult c.b&racteristics (usually at a length or about 25 DID). Gastropods: snails. . Habitat: Herbivore: place where a particular plant or animal lives . organism that feeds on plant material. Heterotrophy: type of nutrition in which an organism depends on organic matter for food. -...... ·.Hydraulic residence time: time (usually days) required for a volume of · vater equal to the reservoir capacity to move. through 'the reservoir 'and. be discharged downstream. ··.·' E3 . .. . ...... , •' !. I· ·. Hypolimnion : lover stratum of cold and relatively undisturbed vater that forms in lakes and reservoirs during periods of stratification and extends from the bottom up to the metalimnion or thermocline. Instar: any one of the successive stages in the life history of an insect. Isopods: freshvater c.rustaceans (.Asellus), vhich are similar to terrestrial sow bugs. Laminar flow: smooth, low-velocity flow, vith parallel layers of water shearing over one another, and vi th li.ttle or no mixing of layers. Lentic: standing vaters such as lakes an.d ponds. Lepidopterans: order of insects that includes the moths and butter- flies. Limnology: study of the physical, chemical, and biological conditions in fresh vaters. Lotic: running waters such as streams and rivers. Macrophytes: macroscopic or large forms of vegetation. Metalimnion: stratum betveen the epilimn·ion and the hypolimnion in stratified lakes and reservoirs; exhibits marked thermal discon- tinuity; temperature changes at least 1°C per metre throughout this .stratum. -Mollusks: soft-bodied animals usually enclosed in a shell and having a muscular foot; freshwater forms include snails and clams. Nymphs: one of a serie.s of immature stages in certain insects . Oligochaetes: earthworms and their aquatic representatives. Om.."livore: any animal that eats a variety of living and dead plants and animals. Ostracods: small crustaceans enclosed in bivalve shells; resemble small clams. Periphyton·: ~:~.ssociation of aquatic organisms attached or clinging to stems and leaves of rooted plants or other surfaces projecting above the stream bottom. pH: the negative logarithm of the effective hydrogen-ion concentration . Used to express both acidity and basicity on a scale of 0 to 14; 1 represents neutrality, numbers less than 1 increasing acidity, and numbers greater than 1 increasing basicity. E4 • ·.· ~. . ··~- ' ·-· .. · •·· t· : ·. =·:· ~-.. ·: .. . .•. . .... -;~:. · . . . .. ·~ ... --.: . ... . • Photosynthesis: complex of processes involved in the . formation of carbohydrates from carbon dioxide and water in living plants in the presence of light and chlorophyll. Phytoplankton: small plants (algae) that live unattached in the water. Piscivorous: feeding on fishes. Plankton: organisms. of relatively small size, mostl y microscopic, that drift with the water current; some have vea.k powers of locomotion. Plecopterans : order of insects that includes the .stoneflies and salmonflies. Pool: portion of a stream that is dee·p and quiet relative to the main current .• Redd: type of fish spawning area (usually a cleared circular or oblo.ng depression) in running wat er with a gravel bottom. Redox potential: oxidation-reduction potential; a m~asure of the . · oxidizing or reducing intensity of a solution. Riffle: shallow rapids in an open stream, where the water surface is broken into waves by obstructions wholly or partly submerged. Run : stretch of relatively deep, fast-flowing water with the surface essentially nonturbulent. ' Seston: :· living or nonliTing bodies of plants or animals that float ·or . swim ·in tbe .water. · . Simuliids: family of insects of the Orde.r Diptera; includes black flies and buffalo gnats . Spate: a sudden freshet or flood. Stenothermal: refers to an. organism that can maintain itself only · over a relatively narrov range of temperature. Strati~ication : separation of and nonmixing between the s urface ep1limnetic water and the deep bypolU!znetic water because of density differences between the tva layers • Tail.Ma ter: channel o r stream below a dam. Thermocline : see metalimnion . ·· Trichopterans : order of insects that includes caddis fli e s. Trophic level : refers to the position occupied by an organism in a simplified food c.hain • E5 -····----.. -... ·--··----..------··-•.. --·-_ ....... ___ ..... ----·~ . .. .... .... ··-... -· .... ---··· .... --·--·-· ___ .. __ _ Turbellarians: free-11 ving flatworms. Turbidity: cloudiness of water caused by the presence of suspended matter.·· Turbulent flov: flov vi th secondary, heterogeneous eddies supe:r- i=POsed on the main forward flow, accompanied by considerable mixing of components. Water quality: a term used to describe the chemical, physical, and biological characteristics of water in respect to its suitability for a pa-rticular use. · Zooplankton: animal microorganisms that 11 ve \Ulattached in the water • . . . E6