Survival and Growth Rate of Translocated Freshwater Mussels Lampsilis fasciola and Medionidus conradicus

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1 University of Tennessee, Knoxville Trace: Tennessee Research and Creative Exchange Masters Theses Graduate School Survival and Growth Rate of Translocated Freshwater Mussels Lampsilis fasciola and Medionidus conradicus Laura L. Pullum University of Tennessee - Knoxville, lpullum1@vols.utk.edu Recommended Citation Pullum, Laura L., "Survival and Growth Rate of Translocated Freshwater Mussels Lampsilis fasciola and Medionidus conradicus. " Master's Thesis, University of Tennessee, This Thesis is brought to you for free and open access by the Graduate School at Trace: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Masters Theses by an authorized administrator of Trace: Tennessee Research and Creative Exchange. For more information, please contact trace@utk.edu.

2 To the Graduate Council: I am submitting herewith a thesis written by Laura L. Pullum entitled "Survival and Growth Rate of Translocated Freshwater Mussels Lampsilis fasciola and Medionidus conradicus." I have examined the final electronic copy of this thesis for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Master of Science, with a major in Geology. We have read this thesis and recommend its acceptance: Colin Sumrall, Brian Alford (Original signatures are on file with official student records. Michael McKinney, Major Professor Accepted for the Council: Dixie L. Thompson Vice Provost and Dean of the Graduate School

3 !!!!! Survival(and(Growth(Rate(of(Translocated( Freshwater(Mussels(Lampsilis(fasciola(and( Medionidus(conradicus(!!!!!! A!Thesis!Presented!for!the! Master!of!Science! Degree! The!University!of!Tennessee,!Knoxville!!!!!!!!!! Laura!L.!Pullum! August!2015!

4 !!!!!!!!!!!!!! Copyright!!2015!by!Laura!L.!Pullum.!! All!rights!reserved.!!!!!!!!!!!!!!! ii!

5 ACKNOWLEDGEMENTS(! I!would!like!to!thank!my!advisor,!Dr.!Michael!McKinney,!for!his!introduction!to,!and! guidance!in,!the!field!of!freshwater!mussel!conservation.!comments!and! encouragement!from!my!committee!members,!dr.!colin!sumrall!and!dr.!brian! Alford,!have!been!greatly!appreciated.!In!the!field,!I!thank!Don!Hubbs!and!other! Tennessee!Wildlife!Resources!Agency!(TWRA!personnel!for!the!knowledge!they! passed!on!and!for!their!patience.!for!their!support!in!conducting!mussel!retrieval! and!measurement,!i!thank!dr.!mckinney s!undergraduate!ecology!students!(at!the! University!of!Tennessee.!!Thanks!to!Megan!Bradley!of!the!Aquatic!Wildlife! Conservation!Center!(AWCC!for!length!data!on!the!control!group!of!Lampsilis( fasciola.!!thanks!to!sarah!sheffield,!michael!lucas!and!dr.!josh!emery!for!pleasant! greetings!at!all!times.!thanks!to!angie!staley!in!the!environmental!and!planetary! Sciences!office!for!her!help!navigating!the!systems.!!My!sincere!appreciation!goes!to! Josh!Price!for!his!support!using!SAS!and!for!fruitful!discussions!on!mixed!model! ANOVA.!!Last,!though!not!least,!thanks!to!Jan!Pullum!for!hanging!in!there.!! iii!

6 ! ABSTRACT( Freshwater!mussels!(Family!Unionidae!and!Margaritiferidae!are!a!widely! threatened!group!of!bivalve!molluscs,!particularly!in!the!southeastern!united!states.!! Translocation!of!freshwater!mussels!is!an!increasingly!common!conservation! method.!however,!there!are!relatively!few!studies!that!quantitatively!investigate!the! factors!influencing!translocation!success!or!failure.!in!october!2013,!hundreds!of! Medionidus(conradicus!and!Lampsilis(fasciola!were!translocated!to!the!Pigeon!and! Nolichucky!Rivers!in!Tennessee,!with!an!interim!partial!survey!(June!2014!and!a! full!survey!(october!2014.!!in!this!study,!i!analyze!this!field\collected!data!to! determine!the!mechanism(s!that!currently!influence!the!outcomes!of!tennessee! mussel!translocation.!!!! My!recommendations!for!future!surveys!include!open!and!timely!data!sharing! between!investigators!and!the!scientific!community!at!large.!!given!these!data!and! associated!collection!methods,!a!better!understanding!of!freshwater!mussel! communities!and!restoration!success!factors!can!be!identified!at!lower!future!costs! and!facilitate!longer\term!research.!!my!research!recommendations!include!more! frequent,!complete!surveys,!and!quantitative!analyses!at!the!mussel!and!community! levels.! The!results!of!this!study!have!implications!for!conservation!translocation!efforts.!! My!results!indicate!that!both!L.(fasciola!and!M.(conradicus!can!be!successfully! translocated!to!the!pigeon!river,!if!1!they!are!translocated!to!the!pigeon!where!it! has!less!boulder,!cobble!and!exposed!bedrock!in!favor!of!more!coarse!and!fine! gravel!and!sand;!2!it!had!lower!peak!and!average!water!discharge!rates,!3!if!some! translocations!occurred!in!the!spring\early!summer,!and!4!if!the!translocated! mussels!are!initially!housed!in!cages!or!silos.!the!non\housed!mussels!were!not! recovered,!primarily!due!to!high!water!volumes!and!velocities!soon!after!the! beginning!of!the!study.!!the!housed!mussels!were!protected.!there!is!no!overall! iv!

7 predictability!of!the!water!discharge!timing!and!size!of!the!waterville!hydroelectric! Power!plant s!dam.!a!management!recommendation!is!for!incremental!releases!and! notification!to!conservation!authorities.!due!to!significant!mortality!in!the!first!8! months!of!this!study,!some!studies!should!start!in!the!spring\early!summer!rather! than!in!october!to!help!translocated!mussels!survive!their!first!over\winter!by! having!some!growth!and!habitat!acclimation!underway.!!! v!

8 TABLE(OF(CONTENTS!! CHAPTER!1!INTRODUCTION... 1! 1.1! The Rise and Fall of Freshwater Mussel Fauna... 1! 1.2! Habitat Factors that Influence Freshwater Mussel Mortality and Growth... 2! 1.3! Tennessee Efforts to Restore Freshwater Mussels... 5! 1.4! Research Study... 15! CHAPTER!2!SITE!AND!SPECIES!BACKGROUND... 16! 2.1! Sites... 16! 2.1.1! Clinch River at Kyles Ford... 18! 2.1.2! Nolichucky River... 18! 2.1.3! Pigeon River at Cosby Creek... 19! 2.1.4! Brood Stock from the VDGIF AWCC... 19! 2.2! Species... 20! 2.2.1! Lampsilis fasciola... 20! 2.2.2! Medionidus conradicus... 23! CHAPTER!3!MATERIALS!AND!METHODS... 26! 3.1! Overview... 26! 3.2! Translocation... 26! 3.3! Housing... 29! 3.3.1! Silo... 29! 3.3.2! Cage... 31! 3.3.3! Mesh Bag Release... 33! 3.4! Water Quality... 33! 3.4.1! Chlorophyll-A... 33! 3.4.2! Water Discharge Rate... 34! 3.5! Surveys... 37! 3.6! Data Collected and Analyzed... 37! 3.7! Recovery Rate and Effort... 37! 3.8! Analyzing Survival and Growth Rate... 38! CHAPTER!4!RESULTS... 42! 4.1! Interim Survey Results... 42! 4.2! Final Study Survey... 42! 4.3! Recovery Rate and Effort... 43! 4.4! Survival... 45! 4.4.1! Descriptive Statistics... 45! 4.4.2! Model-Based Analysis of Survival... 51! 4.5! Growth Rate... 53! 4.5.1! Descriptive Statistics... 53! 4.5.2! Model-Based Analysis of Growth Rate... 61! 4.6! Water Quality... 65! CHAPTER!5!DISCUSSION... 67! REFERENCES... 75! APPENDICES!!...!95! A!! Data!Collected!...!96! vi!

9 B! Mortality!Data!And!Analysis!Results!...!132! C! Growth!Rate!Data!And!Analysis!Results!...!137! VITA!...!169( vii!

10 viii LIST(OF(TABLES(!! ( Table!1\1.! TWRA!Freshwater!Mussel!Translocations,!2004!to!Present!...!8! Table!2\1.! Site!location,!altitude!and!substrate!...!16! Table!2\2.! Host!Fish!for!Introduced!Mussels!...!25! Table!3\1.! Source!&!Destination!Information!for!Tagged!Mussels!...!27! Table!3\2.! Length!of!translocated!mussels!and!AWCC!retained!mussels!...!28! Table!3\3.! Weight!of!translocated!mussels!...!28! Table!3\4.! Average!cage!and!silo!housing!occupants!\!Pigeon!River!mussels!...!32! Table!3\5.! Average!cage!&!silo!housing!occupants!\!Nolichucky!River!mussels!..!33! Table!3\6.! Study!Design!...!39! Table!4\1.! Summary!of!interim!survey,!6/10/2014.!Length!and!weight!are! measured!for!live!mussels!only!...!42! Table!4\2.!! Summary!of!final!survey,!9/23/2014.!Length!and!weight!are! measured!for!live!mussels!only!...!43! Table!4\3.! Recovery!rate!...!44! Table!4\4.! Catch!per!Unit!Effort!...!44! Table!4\5.! Full!Survival!Model!ANOVA!...!51! Table!4\6.!!! Species\Specific!Survival!Model!ANOVA!...!52! Table!4\7.! Summary!of!Absolute!Growth!Rate!...!54! Table!4\8.!!! Full!Absolute!Growth!Rate!Model!ANOVA!...!62! Table!4\9.!!! Full!Relative!Growth!Rate!Model!ANOVA!...!63! Table!4\10.!!! Species\Specific!Relative!Growth!Rate!Model!ANOVA!...!64! Table!A\1.! Initial!Length!and!Weight!of!Translocated!Mussels,!10/10/2013!...!96! Table!A\2.! Length!and!Weight!of!Translocated!Mussels!on!Interim!Survey,! 6/10/2014!...!119! Table!A\3.! Length!and!Weight!of!Translocated!Mussels!on!Final!Survey,! 9/23/2014!...!123! Table!B\1.! Survival!Rate!Full!Model,!Split\Split!Plot!Repeated!Measures!without! Replication!on!the!Arcsin!Transformed!Values,!Block!on!Destination!! Data!...!132! Table!B\2.! Survival!Rate!Full!Model,!Type!III!Tests!of!Fixed!Effects!...!132! Table!B\3.! Effect!=!Time,!Method!=!LSD!(P<0.05,!Set!=!4,!Survival!Rate!Full! Model!...!133! Table!B\4.! Effect!=!House*Time,!Method!=!LSD!(P<0.05,!Set!=!6,!Survival!Rate! Full!Model,!...!133! Table!B\5.! Full!Survival!Rate!Model!Assumptions!are!Met!...!135!

11 ix Table!B\6.! L.(fasciola!Survival!Rate!Model!Assumptions!are!Met!...!135! Table!B\7.! M.(conradicus!Survival!Rate!Model!Assumptions!are!Met!...!136! Table!B\8.! L.(fasciola!Survival!Model!\!Type!III!Tests!of!Fixed!Effects!...!136! Table!B\9.! M.(conradicus!Survival!Model!\!Type!III!Tests!of!Fixed!Effects!...!136! Table!C\1.! Absolute!Growth!Rate!Data!...!137! Table!C\2.! Absolute!Growth!Rate!Summary!Statistics!...!143! Table!C\3.! SAS!GLIMMIX!Procedure!Full!AGR30!Model!Information!...!144! Table!C\4.! Type!III!Tests!of!Fixed!Effects,!Full!AGR30!Model!...!144! Table!C\5.! Mean!separation!for!Rank!Absolute!Growth!Rate,!Full!AGR30!Model144! Table!C\6.! Mean!Separation!for!Rank!Absolute!Growth!Rate,!Effect=Destination,! Full!AGR30!Model!...!145! Table!C\7.! Mean!Separation!for!Rank!Absolute!Growth!Rate,!Effect=Species,!Full! AGR30!Model!...!145! Table!C\8.! Mean!Separation!for!Rank!Absolute!Growth!Rate,! Effect=Destination*Species,!Full!AGR30!Model!...!146! Table!C\9.! Mean!Separation!for!Rank!Absolute!Growth!Rate,!Effect!=!House,!Full! AGR30!Model!...!146! Table!C\10.! Mean!Separation!for!Rank!Absolute!Growth!Rate,! Effect=Destination*House,!Full!AGR30!Model!...!146! Table!C\11.! Mean!Separation!for!Rank!Absolute!Growth!Rate,! Effect=Species*House,!Full!AGR30!Model!...!146! Table!C\12.! Relative!Growth!Rate!Data.!...!154! Table!C\13.! Relative!Growth!Rate!Summary!Statistics!...!160! Table!C\14.! Type!III!Tests!of!Fixed!Effects!for!Relative!Growth!Rate,!Full!Model!160! Table!C\15.! Full!RGR30!Model!Assumptions!are!Met!...!161! Table!C\16.! Mean!Separation!for!Rank!Relative!Growth!Rate,!Effect!=Species,!Full! RGR30!Model!...!161! Table!C\17.! Mean!Separation!for!Rank!Relative!Growth!Rate,!Effect!=!House,!Full! RGR30!Model!...!162! Table!C\18.! Mean!Separation!for!Rank!Relative!Growth!Rate,! Effect=Destination*Species,!Full!RGR30!Model!...!162! Table!C\19.! Mean!Separation!for!Rank!Relative!Growth!Rate,! Effect=Destination*House,!Full!RGR30!Model!...!162! Table!C\20.! Mean!Separation!for!Rank!Relative!Growth!Rate,! Effect=Species*House,!Full!RGR30!Model!...!162! Table!C\21.! Tests!for!Normality,!L.(fasciola!RGR30!Model!...!163! Table!C\22.! Type!III!Tests!of!Fixed!Effects!for!L.(fasciola!RGR30!Model!...!163!

12 Table!C\23.! Table!C\24.! Table!C\25.! Table!C\26.! Table!C\27.! Table!C\28.! Table!C\29.! Table!C\30.! ( Mean!Separation!for!Rank!Relative!Growth!Rate,!Effect=Destination,!L.( fasciola!rgr30!model!...!164! Mean!Separation!for!Rank!Relative!Growth!Rate,! Effect=Destination*House,!L.(fasciola!RGR30!Model!...!165! M.(conradicus!RGR30!Model!Tests!for!Normality!...!165! M.(conradicus!RGR30!Model!Assumptions!are!Met!...!165! Type!III!Tests!of!Fixed!Effects!for!M.(conradicus!Relative!Growth!!!!! Rate!...!166! Mean!Separation!for!Rank!Relative!Growth!Rate,!Effect=Destination,! M.(conradicus!RGR30!Model!...!166! Mean!Separation!for!Rank!Relative!Growth!Rate,!Effect!=!House,!M.( conradicus!rgr30!model!...!167! Mean!Separation!for!Rank!Relative!Growth!Rate,! Effect=Destination*House,!M.(conradicus!RGR30!Model!...!168!!! x

13 !! Figure!1\1.! Figure!2\1.!! LIST(OF(FIGURES(! Freshwater!Mussel!Life!Cycle![Source:!Freshwater!Mussels!of!Iowa,! 2002,!life!cycle!diagram:!Mississippi!River,!Lower!St.!Croix!Team,! Wisconsin!Department!of!Natural!Resources]!...!3! Study!Site!Locations:!!Clinch!River!at!Kyle s!ford,!pigeon!river!at! Cosby!Creek,!Nolichucky!River!at!the!TWRA!Canoe!Launch.!!The! University!of!Tennessee!at!Knoxville!is!included!for!reference.!Map! courtesy!of!google!...!17! Figure!2\2.!!! L.(fasciola!US!range!((NatureServe,!2014a,!last!accessed! 05/12/2015!...!21! Figure!2\3.!!! Lampsilis(fasciola!(Encyclopedia!of!Life,! Figure!2\4.!!! M.(conradicus!US!range!((NatureServe,!2014b,!last!accessed! 05/12/2015!...!24! Figure!2\5.!!! Medionidus(conradicus!(Encyclopedia!of!Life,! Figure!3\1.!!! Mussel!length!measurement.!!Length!measurement!take!at!the!longest! dimension!(pictured!mussel!is!not!l.(fasciola!or!m.(conradicus!...!28! Figure!3\2.!! Source,!destination,!counts!and!housing!of!translocated!mussels.!...!30! Figure!3\3.!!! Silo.!Silo!width!is!27!cm,!height!12!cm,!metal!opening!at!top!is!8.6!cm! in!diameter.!the!interior!containing!the!mussels!is!4!cm!in!diameter! and!4!cm!in!height!...!30! Figure!3\4.!!! Cage.!Dimensions!of!the!cage!are!9!cm!in!height!and!6.6!cm!in! diameter!...!31! Figure!3\5.! Average!Monthly!Discharge!(m 3 /s!during!the!study!period!for!the! Clinch,!Pigeon!and!Nolichucky!Rivers.!Data!obtained!from! USGS!Site! !Clinch!River!above!Tazwell,!TN;!USGS!Site! !Nolichucky!River!at!Embreeville,!TN;!USGS!Site! ! Pigeon!River!at!Newport,!TN!...!35! Figure!3\6.! Maximum!Monthly!Discharge!(m 3 /s!during!the!study!period!for!the! Clinch,!Pigeon!and!Nolichucky!Rivers.!Data!obtained!from! USGS!Site! !Clinch!River!above!Tazwell,!TN;!USGS!Site! !Nolichucky!River!at!Embreeville,!TN;!USGS!Site! ! Pigeon!River!at!Newport,!TN!...!36! Figure!4\1.!!! Survival!by!Species!over!Time!...!45! Figure!4\2.!!! Survival!by!River!over!Time!...!46! Figure!4\3.!!! L.(fasciola!Survival!by!Destination!...!47! xi

14 Figure!4\4.! M.(conradicus!Survival!by!Destination!...!47! Figure!4\5.!!! Survival!by!Housing!...!48! Figure!4\6.!!! Survival!by!Housing!Condition!...!49! Figure!4\7.!!! Survival!by!Volume!per!Occupant!...!50! Figure!4\8.! L.(fasciola!30\day!Absolute!Growth!Rate.!The!growth!rate!density!is! shown!with!the!data s!normal!curve!...!53! Figure!4\9.! M.(conradicus!30\day!Absolute!Growth!Rate.!The!growth!rate!density! is!shown!with!the!data s!normal!curve!...!55! Figure!4\10.!!! Absolute!Growth!Rate!by!River!...!56! Figure!4\11.!!! Absolute!Growth!Rate!by!Housing!...!56! Figure!4\12.!!! 30\day!Absolute!Growth!Rate!by!Housing!Condition!...!57! Figure!4\13.!!! L.(fasciola!Relative!Growth!Rate!(30!d!...!58! Figure!4\14.!!! M.(conradicus!Relative!Growth!Rate!(30!d!...!58! Figure!4\15.!!! RGR30!by!River!...!59! Figure!4\16.!!! RGR30!by!Housing!...!60! Figure!4\17.!!! 30\day!Relative!Growth!Rate!by!Housing!Condition!...!60! Figure!4\18.!!! Housing!condition!by!Destination!River!...!61! Figure!4\19.!!! Pigeon!River!(USGS!site! !and!Walters!Dam!(USGS!site! !Mean!Daily!Discharge!(m 3 /s!1/1/2012!!9/30/ !66! Figure!B\1.!!! Least!Squared!Mean!(LSM!for!Survival!Rate,!Effect!=!Time,!Full!!!!! Model!...!133! Figure!B\2.!!! LSM!for!Effect!=!House*Time,!Survival!Rate!Full!Model!...!134! Figure!B\3.!!! Time*House!LSM!for!Arcsin_SR,!Survival!Rate!Full!Model!...!134! Figure!B\4.!!! Normality!of!Arcsin(Survival!Rate!Residuals,!Full!Model!...!135! Figure!C\1.!!! LSM!for!Destination,!Full!AGR30!Model!...!145! Figure!C\2.!!! Destination!untransformed!LSM!for!Absolute!Growth!Rate,!Full!AGR30! Model!...!147! Figure!C\3.!!! Check!on!Normality!for!Rank!Absolute!Growth!Rate,!Full!AGR30!!! Model!...!148! Figure!C\4.!!! Rank!Absolute!Growth!Rate!LSM!for!Species,!Full!AGR30!Model!...!149! Figure!C\5.!!! Rank!Absolute!Growth!Rate!LSM!for!Destination*Species,!Full!AGR30! Model!...!150! Figure!C\6.!!! Rank!Absolute!Growth!Rate!LSM!for!House,!Full!AGR30!Model!...!151! Figure!C\7.!!! Rank!Absolute!Growth!Rate!LSM!for!Destination*House,!Full!AGR30! Model!...!152! Figure!C\8.!!! Rank!Absolute!Growth!Rate!LSM!for!Species*House,!Full!AGR30!!! Model!...!152! Figure!C\9.!!! Conditional!Residuals!for!Rank!Absolute!Growth!Rate,!Full!Model...!153! xii

15 Figure!C\10.!!! Conditional!Residuals!for!Rank!Relative!Growth!Rate,!Full!Model!...!161! Figure!C\11.!! Residuals!for!Rank!Relative!Growth!Rate,!L.(fasciola!RGR30!Model!...!163! Figure!C\12.!!! Destination!effect!on!L.(fasciola!RGR30,!Least!Squared!Means!...!164! Figure!C\13.!!! Residuals!for!Rank!Relative!Growth!Rate!Model!for!M.(conradicus!...!166! Figure!C\14.!!! House!effect!on!M.(conradicus!RGR30,!Least!Squared!Means!...!167! xiii!

16 ! CHAPTER(1( INTRODUCTION( 1.1( The(Rise(and(Fall(of(Freshwater(Mussel(Fauna( Freshwater!mussels!are!an!important!indicator!of!ecosystem!health!and!biodiversity! (Neves!et!al.!1997;!Haag!&!Rypel,!2011;!Hubbs,!2014.!!North!America s!freshwater! mussels!are!bivalve!mollusks!of!the!order!unionida,!families!unionidae!and! Margaritiferidae.!!They!function!to!filter!water,!their!biodeposition!increases!food! availability!for!other!organisms!and!their!burrowing!releases!nutrients!from,!and! increases!the!oxygen!content!of,!the!sediment.!!freshwater!mussels!play!an! important!role!in!nutrient!cycling!by!reducing!the!amount!of!suspended!organic! matter!in!the!water!column,!which!otherwise!could!lead!to!eutrophication.!!in! addition,!they!are!a!food!source!for!fish,!aquatic!and!near\shore!mammals,!and! shore!birds.!(haag,!2012!!north!america!has!the!highest!freshwater!mussel! diversity,!with!estimates!of!around!300!species!(williams,!et!al.!1993;!graf!&! Cummings,!2007;!Vaughn,!et!al.!2004,!with!the!Tennessee!River!Basin!historically! home!to!102!species!(parmalee!&!bogan,!1998.!! Populations!of!some!freshwater!mussels!have!shown!severe!decline!due!to!a!series! of!human!engineering!projects!that!resulted!in!habitat!loss!and!degradation!of!river! and!lake!ecosystems!(graf!2007;!lyons!et!al.!2007.!thirty\five!species!of!freshwater! mussels!have!become!extinct!in!the!past!100!years!(neves!et!al.!1997.!! Environmental!factors!that!occur!at!large!spatial!scales!have!influenced!the!decline,! extirpation!and!extinction!of!freshwater!mussels!in!north!america,!including! systematic!habitat!destruction!by!dams!and!other!river!modification!projects,! pollution!(e.g.,!chemical!spills,!mine!and!agriculture!run!off,!and!flow!instability.! (Goudreau!et!al.!1993;!Diamond!&!Serveiss,!2001;!Havlik!&!Marking,!1987;!Aldridge! et!al.!1987;!neves!et!al.!1997.!! 1

17 Efforts!have!been!made!to!correct!habitat!destruction!at!large!spatial!scales!(e.g.,! within!the!tennessee!river!basin,!so!that!the!sites!may!be!viable!recruitment! grounds!for!mussel!population!enhancement.!for!example,!the!tennessee!valley! Authority!(TVA!has!implemented!several!methods!to!improve!the!dissolved!oxygen! concentrations!in!the!benthos!of!tailwater!sections!of!rivers!below!dams.!!these! methods!include!use!of!aerating!turbines,!surface\water!pumps,!oxygen!injection! systems!and!aerating!weirs!to!eliminate!hypoxia!by!reaching!dissolved!oxygen! concentration!targets!(daily!minimum!of!5.0!mg/l!necessary!for!benthic! invertebrate!and!fish!life!(brookshier!et!al.!1999.!!these!large\scale!restoration! efforts!are!necessary,!but!will!likely!be!insufficient!for!adequate!recovery!of!historic! mussel!populations.!!reintroduction!efforts!for!mussels!following!habitat! restoration!have!largely!been!unsuccessful.!!to!date,!however,!few!multi\factor! studies!have!been!conducted!to!understand!the!mechanism(s!influencing!the! success!of!tennessee!freshwater!mussel!translocation!efforts.!!!! 1.2( Habitat(Factors(that(Influence(Freshwater(Mussel( Mortality(and(Growth( Many!factors!influence!the!growth,!lifespan!and!abundance!of!freshwater!mussels,! including!the!environmental!factors!!habitat!destruction,!pollution!and!flow! instability.!!the!freshwater!mussel!life!cycle!and!habitat!present!additional!factors.!! For!instance,!the!mussel s!life!cycle!(figure!1\1!is!one!of!the!most!complex!of!any! group!!(us!fws!2006.!!the!male!mussel!releases!sperm!into!the!water.!the!water! and!sperm!are!siphoned!by!nearby!female!mussel!and!enter!the!female s!gills,!where! eggs!are!fertilized!and!develop!there!for!several!weeks.!!the!next!stage!of!the! lifecycle!is!the!emergence!of!microscopic!mussel!larvae,!called!glochidia.!the!female! releases!the!glochidia!(via!various!taxa\specific!methods!and!they!attach!to!a!host! to!develop.!! 2

18 !!! Figure(1L1.(Freshwater(Mussel(Life(Cycle([Source:(Freshwater(Mussels(of(Iowa,(2002,(life(cycle( diagram:(mississippi(river,(lower(st.(croix(team,(wisconsin(department(of(natural( Resources](!!! 3

19 Most!freshwater!mussels!prefer!specific!hosts!and!the!mussels!studied!herein!use! fish!as!hosts!(as!do!most!freshwater!mussels.!already!we!see!that!water!flow!and! host!availability!are!factors!in!mussel!survival!and!recruitment.!the!glochidia!attach! to!the!host s!gill!tissue,!which!provides!food!and!shelter!for!glochidia!development.!! The!larval!glochidium!develops!into!a!juvenile!mussel!within!several!weeks,!and! then!drops!off!the!host!to!the!substrate.!the!juvenile!continues!to!develop!to!a!sub\ adult!stage,!and!then!to!an!adult.!!a!mussel!is!considered!mature!or!adult!upon! reproductive!maturity.!in!this!latter!part!of!the!life!cycle,!we!note!additional!factors! affecting!the!survival!and!recruitment!of!mussels!including!host!fish!availability,!the! ability!to!attract!hosts,!and!the!availability!of!suitable!substrate.!!parasitic!life!cycle! and!selective!dependence!on!hosts!for!dispersal!to!suitable!habitats!are! complicating!factors!for!mussel!conservation!(neves!et!al.,!1997;!strayer!et!al.,!2004;! Zipper,!et!al.!2014.!! Predators!of!adult!freshwater!mussels!include:!Lutra(canadensis!(Otter,!Ondatra( zibethicus!(muskrat,!procyon(lotor!(raccoon,!mephitidae!(skunk,!laridae!(gull! and!scolopacidae!(shore!birds.!juveniles!are!eaten!by!platyhelminthes!(flatworm,! Hirudidae!(Leech,!and!Cambaridae!(Crayfish!(Strayer!&!Smith,!1996,!and! freshwater!fish!including!cyprinidae!(carp,!acipenseridae!(sturgeon,!ictaluridae! (Catfish,!and!Centrarchidae!(Sunfish!(Haag,!2012.!! Mortality!rates!and!survivorship!curves!are!unknown!for!most!mussel!species,! presenting!a!major!challenge!in!distinguishing!between!natural!and!human! influenced!population!fluctuations!(jones!&!neves,!2011.!however,!the!few!existing! studies!indicate!significant!variability!in!mortality,!longevity!and!growth!(jokela!&! Mutikainen,!1995;!Villella!et!al.,!2004,!Haag!&!Rypel,!2011.!In!general,!freshwater! mussels!produce!a!large!number!of!glochidia!from!which!a!relatively!few!survive!to! maturity.!mussels!are!typically!characterized!as!having!low!mortality!and!long!life! spans!(hart!et!al.,!2001.!!!! 4

20 Habitat!factors!such!as!dissolved!oxygen!(Strayer!&!Smith,!1996,!temperature! (Bogan,!2001;!Rodland!et!al.,!2009,!pH,!chemical!and!metal!concentrations,!and! sedimentation!disturbances!(ellis,!1936!have!been!suggested!(strayer!et!al.,!2004! as!additional!mussel!survival!factors.!!however,!subsequent!studies!showed!that! some!environmental!factors!had!almost!no!explanatory!power!(e.g.,!sediment!grain! size!and!current!velocity![strayer!&!ralley,!1993].!!availability!of!food!and!fish! hosts!for!mussel!glochidia!attachment!is!fundamental,!because!effective!recruitment! and!dispersal!are!critical!to!the!continuity!of!a!population.!!in!addition,!non\native! species!such!as!the!zebra!mussel,!dreissena(polymorpha,!reduce!food!and!habitat! available!for!native!mussels,!and!increased!parasite!densities!(e.g.,!trematodes! (Gangloff!et!al.,!2008!can!reach!a!level!that!reduces!mussel!reproductive!output.!! 1.3( Tennessee(Efforts(to(Restore(Freshwater(Mussels( Relocation!of!freshwater!mussels!from!one!location!containing!robust!mussel! population(s!to!another,!presumably!better,!location!has!been!used!as!a! conservation!strategy!for!freshwater!mussels!for!over!35!years!(cope!and!waller,! 1995.!In!an!effort!to!avoid!confusion!caused!by!all!of!the!terms!defining!relocation,! and!to!harmonize!the!freshwater!mussel!literature,!cope!(2003!proposed!the!term! relocation!to!refer!to!any!intentional!movement!by!humans!of!an!individual!or! population!from!one!location!to!another.!!however,!more!recently,!the!world! Conservation!Union!(IUCN,!2013!defined!the!following:! Translocation!\!the!human\mediated!movement!of!living!organisms!from! one!area,!with!release!in!another.!! Conservation(translocation!\!the!deliberate!movement!of!organisms!from! one!site!for!release!in!another.!it!must!be!intended!to!yield!a!measurable! conservation!benefit!at!the!levels!of!a!population,!species!or!ecosystem,!and! not!only!provide!benefit!to!translocated!individuals.! 5

21 Further,!conservation!translocations!consist!of!a!reinforcement!and!reintroduction! within(a!species!indigenous!range!(includes!augmentation,!and!b!conservation! introductions,!comprising!assisted!colonization!and!ecological!replacement,!outside( indigenous!range!(iucn,!2012.!!all!of!the!efforts!reported!herein!are!conservation! translocations,!and!will!be!referred!to!as!translocations.!!when!reporting!results! from!other!authors,!i!use!their!terminology!or!the!more!general!term,!relocation.!! The!Virginia!Department!of!Game!and!Inland!Fisheries!(VDGIF!Aquatic!Wildlife! Conservation!Center!(AWCC!cultivates!freshwater!mussels!for!augmentation! purposes.!!conservation!of!freshwater!mussels!in!the!tennessee!river!basin!has! used!both!approaches.!!! Cope!and!Waller!(1995!reviewed!33!papers!and!reports!on!freshwater!mussel! relocation!efforts.!!they!found!that!over!90,000!mussels!had!been!relocated!over!the! 37!projects!discussed!in!the!33!documents,!with!a!mean!mortality!rate!of!49%.!! Roughly!a!third!of!the!documents!reported!mortality!rates!of!over!70%.!!Half!of!the! relocations!occurred!in!the!southeastern!us!during!summer!months!(july!through! September.!!The!reasons!for!the!high!mortality!remain!unclear!because!of!lack!of! long\term!monitoring!and!inadequate!habitat!characterization!of!the!relocation! sites.!!!! A!major!obstacle!to!mussel!recovery!in!Tennessee!is!the!fragmentation!of!riverine! habitats!by!dams!and!impounded!waters!(twra,!2015.!in!tennessee,!habitat! conditions!in!some!rivers!that!once!supported!a!diverse!mussel!fauna!have!been! restored!sufficiently!to!again!support!mussels.!this!is!thought!to!be!the!case!for!the! Pigeon!River!site!included!in!this!study.!The!Tennessee!Valley!Authority!(TVA! began!a!reservoir!release!improvements!program!in!the!1990s!at!all!their!tributary! dams!to!provide!constant!minimum!water!flows!and!improve!water!quality!(e.g.,! increased!oxygen!content.!!! 6

22 The!Tennessee!Wildlife!Resources!Agency!(TWRA!is!the!regulatory!body!primarily! responsible!for!conservation!and!management!of!tennessee s!mussels!and!other! biological!resources!(twra,!2015.!cooperative!efforts!between!twra,!other! agencies!and!the!public!have!resulted!in!the!development!of!publications!to!help! guide!recovery!and!conservation!efforts!of!mussels.!!these!include! Tennessee!State!Wildlife!Action!Plan!(TWRA,!2005,!! Plan!for!the!Controlled!Propagation,!Augmentation,!and!Reintroduction!of! Freshwater!Mollusks!of!the!Cumberlandian!Region!(Cumberlandian!Region! Mollusk!Restoration!Committee,!CRMRC!2010,!and! Tennessee!Freshwater!Mollusk!Strategic!Plan!(Tennessee!Chapter!of!The! Nature!Conservancy,!2013.! TWRA s!aquatic!species!restoration!actions!follow!the!myers!rule!which!states!that! listed!endangered/threatened!species!can!be!stocked!into!tennessee! waters/locations!where!other!listed!species!are!extant!for!these!actions!do!not! change!the!regulatory!status!of!the!site!(twra,!2015.!!under!the!direction!of!don! Hubbs,!the!TWRA!Mussel!Restoration!Project!has!translocated!mussels!in!Tennessee! since!2004.!!table!1\1!(developed!from!data!provided!in!(twra,!2015!details!and! summarizes!those!efforts.!!using!6!distinct!source!rivers!and!14!distinct!destination! or!recipient!rivers,!the!twra!has!translocated!over!45,000!mussels!since!2004.!!in! 2014,!over!6,700!mussels!were!translocated.!!Of!those,!over!75%!were!federally! endangered!species.!! Communication!from!the!Pigeon!River!Relocation!Project!coordinator!(J.!Coombs,! UT!states!that!freshwater!mussels!were!released!in!the!Pigeon!River!(at!PRM!8.4,! near!the!mouth!of!english!creek!in!2000!(number!and!species!not!available.! Additional!translocations!occurred!in!August!2010!(50!L.(fasciola!by!the!North! Carolina!Wildlife!Resources!Commission!(NCWRC!above!Canton!mill!and!2011!(58! L.(fasciola!by!NCWRC!below!Canton!mill!and!upstream!of!the!confluence!with! Richland!Creek.!! 7

23 Table&1(1.&TWRA&Freshwater&Mussel&Translocations Source& Recipient& Species& VDGIF/! Tennessee! Diamond! Island! stock! Kyles!Ford! in!down> stream! end!of! pool!and! head!of! riffle! Lampsilis( abrupta( Year& 2004& 2005& 2006& 2007& 2008& 2009& 2010& 2011& 2012& 2013& 2014& Total& (( ((!!!!!!!!!!!! 300! 100! 113!!513!! River&Total(& 0& 0& 0& 0& 0& 0& 0& 0& 300& 100& 113& &513&& Ptycho2 branchus( subtentum( (( (( 211! 850! 239!!! 757! 693! 710! 602! 900!!4,962!!!! Kyles!Ford! includes! Clinch! stock! VDGIF! progeny! Milltown! Epioblasma( brevidens( (( ((!! 108! 76!!! 210! 458! 380! 599! 518!!2,349!! Cumber2 landia( (( ((!! 9! 1!!!!!!!!!!!!!!10!! monodonta( Epioblasma( triquetra( (( ((!!!!!!!!!! 330! 150! 27!!!!507!! L.(abrupta(!!!!!!!!! Cyprogenia( stegaria( (( ((!!!!!!!!!!!!! 8 298!!!298! 54! 45!!99!!!

24 Table&1(1.&TWRA&Freshwater&Mussel&Translocations.&Continued! Source& Recipient& Species& VDGIF/! Tennessee! Diamond! Island! stock! Saline! River! Arkansas! brood! stock/! Kansas! City!Zoo! progeny! Kyles! Ford,! Frost! Ford,!&! Wallen! Bend! Estill!Fork! of!paint! Rock! Littlelot! Hwy!230! Littlelot! Hwy!230! Slick! Shoals!and! Shelby> ville!dam! Venable! Spring! Year& 2004& 2005& 2006& 2007& 2008& 2009& 2010& 2011& 2012& 2013& 2014& Total& L.(abrupta( (( ((!!!!!!!!!!!!!! 121!!!!121!! Quadrula( fragosa( P.( subtentum( Toxolasma( cylindrellus( (( ((!!!!!!!!!!!!!! 103!!!!103!! 200! 115! 142!!!!!!!!!!!!!!!!!!457!! 200! 115! 88!!!!!!!!!!!!!!!!!!403!! River&Total(& 400& 230& 441& 967& 316& 0& 967& 1481&& 1240&& 1804&& 1463&& 9,309&&! & 9

25 Table&1(1.&TWRA&Freshwater&Mussel&Translocations.&Continued!!! Source& Recipient& Species& AABC/! Paint!Rock! stock! VDGIF/! Tennessee! Diamond! Island! stock! AABC/! Paint!Rock! stock! Kyles!Ford! includes! Clinch! stock! VDGIF! progeny! Elk!below! Harms!Mill! Winding! Stair!Bluff! Elk!below! Harms!Mill! Winding! Stair!Bluff! Winding! Stair!Bluff! Hwy!299! Bridge,! Oakdale! Lampsilis( virescens( River&Total(& 0& 0& 0& 0& 0& 0& 0& 0& 725& 938& 649& 2,312&&! 10 Year& 2004& 2005& 2006& 2007& 2008& 2009& 2010& 2011& 2012& 2013& 2014& Total&!!!!!!!!!!!! 430! 532! 500!!!!!!1,462!! L.(abrupta(!!!!!!!!!!!!!!!!!! 200!!!!200!! L.(virescens(!!!!!!!!!!!!!!!! 3,000!!!!!!!3,000!! River&Total& 0& 0& 0& 0& 0& 0& 430& 532& 3,500&& &200&& 0& 4,662&& Epioblasma( capsaeformis( Medionidus( conradicus(!!!!!!!!!!!!!!!! 525! 738! 149!!1,412!!!!!!!!!!!!!!!!!! 200! 200! 500!!900!!

26 Table&1(1.&TWRA&Freshwater&Mussel&Translocations.&Continued Source& Recipient& Species& Kyles!Ford! includes! Clinch! stock! VDGIF! progeny! Milltown! Island! E.( capsaeformis( Year& 2004& 2005& 2006& 2007& 2008& 2009& 2010& 2011& 2012& 2013& 2014& Total&!!!!!!!!!!!!!!!! 2,269!!!! 500!!2,769!! M.(conradicus(!!!!!!!!!!!!!!!! 800!!!!!!800!! River&Total& 0& 0& 0& 0& 0& 0& 0& 0& 3,069&& 0& &500&& 3,569&& Lemiox( rimosus(!!!!!! 212! 176!!!!!!!!!!!!!!388!! M.(conradicus( 201!!!!!!!!!!!!!!!!!!!!!!201!! Kyles!Ford,! Frost!Ford,! &!Wallen! Bend! Noli> upper!hale! Bridge! Actinonaias( pectorosa( 97!!!!!!!!!!!!!!!!!!!!!!97!! E.(brevidens(!!!!!! 44!!!!!!!!!!!!!!!!44!! E.( capsaeformis(!!!!!! 130! 240!!!!!!!!!!!!!!370!! P.(subtentum( 164! 120! 171! 238! 200!!!!!!!!!!!!!!893!!! Site&Total& 462& 120& 171& 624& 616& 0& 0& 0& 0& 0& 0& 1,993&&!! 11

27 Table&1(1.&TWRA&Freshwater&Mussel&Translocations.&Continued& Source& Recipient& Species& Year& 2004& 2005& 2006& 2007& 2008& 2009& 2010& 2011& 2012& 2013& 2014& Total& Kyles!Ford! includes! Clinch! stock! VDGIF! progeny! Milltown! VDGIF/! Tennessee! Diamond! Island! stock! Milltown! Noli> TWRA! canoe! launch! Wilton! Springs! E.(brevidens(!!!!!!!!!! 83! 200! 346! 380! 400! 485!!1,894!! E.( capsaeformis(!!!!!!!!!! 308! 799! 2,128!! 1,895!! 1,807!! 700!!7,637!! P.(subtentum(!!!!!!!!!! 637! 400! 700! 765! 675! 685!!3,862!! M.(conradicus(!!!!!!!!!!!! 506! 826! 500! 549! 500!!2,881!! L.(rimosus(!!!!!!!!!!!! 380! 562! 390! 330! 278!!1,940!! L.(abrupta(!!!!!!!!!!!!!!!!!!!! 130!!130!! Site&Total& 0& 0& 0& 0& 0& 1028&& 2285&& 4,562&& 3,930&& 3761&& 2778&& 18344&& River&Total& 462& 120& 171& 624& 616& 1028&& 2285&& 4,562&& 3,930&& 3761&& 2778&& 20337&& Cyclonaians(!!!!!!!!!!!!!! tuberculata( 217!!!!!!!!217!! Quadrula( pustulosa( Elliptio( dilitata(!!!!!!!!!!!!!!!!!!!!!!!!!!!! 59!!!!!!!!59!! 47!!!!!!!!47!!! 12

28 Table&1(1.&TWRA&Freshwater&Mussel&Translocations.&Continued Source& Recipient& Species& Tennessee! Diamond! Island! Q.(pustulosa( Actinonaias( ligamentina( Year& 2004& 2005& 2006& 2007& 2008& 2009& 2010& 2011& 2012& 2013& 2014& Total&!!!!!!!!!!!!!! 132!!!!!!!!132!!!!!!!!!!!!!!!!!!!! 372! 59!!431!! Kyles!Ford! Wilton! Springs! A.(pectorosa(!!!!!!!!!!!!!!!!!! 4! 199!!203!! C.(tuberculata(!!!!!!!!!!!!!!!!!! 88!!!!88!! E.(dilitata(!!!!!!!!!!!!!!!!!! 108! 36!!144!! Lampsilis(!!!!!!!!!!!!!! ovata(!!!! 5!!!!5!! Ptycho2!!!!!!!!!!!!!! branchus(!!!! 76! 51!!127!! fasciolaris( Villosa(iris(!!!!!!!!!!!!!! 100!!! 122!!!!222!! L.(fasciola(!!!!!!!!!!!!!! 100!!! 237!!!!337!! Q.(pustulosa(!!!!!!!!!!!!!!!!!! 17!!!!17!! M.(conradicus(!!!!!!!!!!!!!! 100!!! 284! 217!!601!! Site&Total& 0& 0& 0& 0& 0& 0& 0& 755& 0& 1313&& 562& 2,630&& 13

29 Table&1(1.&TWRA&Freshwater&Mussel&Translocations.&Continued& Year& Source& Recipient& Species& Total& 2004& 2005& 2006& 2007& 2008& 2009& 2010& 2011& 2012& 2013& 2014& Tennessee! C.(tuberculata(!!!!!!!!!!!!!!!! 909!!!!!!909!! Diamond! Island!!!!!!!!!!!!!!!!! Q.(pustulosa( 911!!!!!!911!! VDGIF! Propogat> ed/!clinch! stock! Kyles!Ford! Denton! L.(ovata(!!!!!!!!!!!!!!!! M.(conradicus(!!!!!!!!!!!!!!!! 200!!!!!!200!! 300!!!!!!300!! Site&Total& 0& 0& 0& 0& 0& 0& 0& 0& 2,320&& 0& 0& 2,320&& River&Total& 0& 0& 0& 0& 0& 0& 0& &755&& 2,320&& 1313&& &562&& 4,950&& Year&Total& 862&& 350&& 612&& 1591& 932&& 1028&& 3682&& 7,330&& 15084&& 8116&& 6065&& 45652&& 14

30 TheTWRAconductedmusseltranslocationintheNolichuckyRiversince2004and inthepigeonriversince2011.thestudyreportedhereinwasbasedon translocationsanddatacollectedaspartofthetwraeffortsduringtheoctober 2013 September2014timeframe. 1.4 ResearchStudy Theresearchpresentedhereinexaminesthesuccessofconservationeffortsattwo sites,oneofwhichhasbeenrecentlydeemedsufficientlyrestoredformussel reintroduction.bothtranslocationandaugmentationconservationstrategiesare examinedviataggingandsurveyingtwomusselspeciesoverapproximatelyone year.quantitativegrowthandsurvivaldatawerecollected,anddataonriver conditionswereobtainedbytestingwaterqualityandfromavailableusgswater qualitysites.thisresearchquantitativelyexaminestherivers,musselspecies,and housingfactorstodeterminetheirimpactonthesuccessofthetranslocationefforts oflampsilis(fasciolaandmedionidus(conradicusinthepigeonriverandthe NolichuckyRiver. 15

31 CHAPTER2 SITEANDSPECIESBACKGROUND InOctober2013,musselsweretranslocatedfromtheClinchRiveratKylesFord(in TennesseetothePigeonRiverattheconfluencewithCosbyCreekandinthe NolichuckyRiver,Tennessee.AdditionalmusselswerestockedfromtheVirginia DepartmentofGameandInlandFisheries (VDGIFAquaticWildlifeConservation Center(AWCCtothesamelocations.Thischapterprovidesbackgroundoneach siteandforthespeciestranslocatedtothosesites.thefollowingchapterwill describethematerialsandmethodsusedinthetranslocationandthesubsequent surveysandanalyses. 2.1 Sites Figure2W1providesamapofthestudysitelocations.TheseincludetheClinchRiver atkyle sford(36 25'30",83 23'54"NADV27,thePigeonRiveratCosbyCreek (35 57'38",83 10'28",andtheNolichuckyRiverattheTWRACanoeAccess (36 10'35",82 27'27".TheUniversityofTennesseeatKnoxville(UTKisincluded forreference.table2w1providesthelocationofeachsite,anditsaltitudeandabrief descriptionofitssubstrate. Table2C1.Sitelocation,altitudeandsubstrate Location Name Clinch River Pigeon River Nolichucky River Lat,Lon (NADV '30", 83 23'54" 35 57'38", 83 10'28" 36 10'35", 82 27'27" USGSSite Altitude (meters abovengvd 29 Drainage Area(sq. kilometers Substrate Cobblesand sand Boulders, cobbles,fine sediments Cobbles, sand,fine sediments 16

32 Ford Nolichucky CanoeLaunch Knoxville Figure2C1.StudySiteLocations:ClinchRiveratKyle sford,pigeonriveratcosbycreek, NolichuckyRiverattheTWRACanoeLaunch.TheUniversityofTennesseeatKnoxvilleis includedforreference.mapcourtesyofgoogle.!! ( 17

33 2.1.1 ClinchRiveratKylesFord AbovetheNorrisReservoirisanunimpoundedportionoftheClinchRiverlocatedin northeasttennessee(figure2w1.itsupportsdiversehealthypopulationsofnative freshwatermussels.fiftywsixmusselspecieshavebeenreportedinthisregion (Stansbery,1973;Jonesetal.,2014,with10expectedtobeextinctorextirpatedin thenext50w100years.twentywfouroftheextantmusselspeciesareeitherfederally endangeredorproposedforfederallisting(hubbs,2014.thus,theclinchriverhas beendesignatedapriorityconservationareabytheusfishandwildlifeservice(us FWS,2012.AlthoughseveraloftheClinchmusselpopulationsareindecline,this siteremainsoneofthehighestintermsoffreshwaterbiodiversity(nevesetal. 1997;Parmalee&Bogan,1998.Infact,thissiteservesasasourceforsomeofthe translocationeffortsunderwaybecauseofitsconsistentabundanceofsomespecies. Inaddition,theglochidiaofsourceL.(fasciolafromtheClinchwerecollectedas broodstockforpropagationattheawccinva.harvestingbeginswithremovinga fewglochidiafromthefemaleandtestingtheirviability.viableglochidiawillsnap closewhenplacedintheproximity(e.g.,inasmalldishofafewgrainsofsalt.the saltisaproxyforthechlorideinfishblood,sotheclosingreactionindicatesthe glochidiaarereadytoinfestahostfish.viableglochidiaareremovedfromthe female,placedinanaeratedcontainerwithaknownhostfish,andtheinfestation begins.wheninfestationiscomplete,thefishandglochidiaaretransferredtotanks andcloselyobservedfor2weeksfortheappearanceofjuvenilemussels.(vdgif, NolichuckyRiver TheNolichuckyRiverinTennesseeatEvansIsland(theTWRACanoeAccessisthe secondtranslocationsite(figure2w1.in1913,thenolichuckydamwasbuilt, creatingdavycrocketlake.in1972,thedamwastakenoutofserviceandthe reservoirwasconvertedintoawildlifemanagementarea(hubbs,2014.a1980 survey(ahlstedt,1986identified21speciesoffreshwatermusselsina56km 18

34 stretchoftheriver,withonly2ofthespeciesexhibitingrecentrecruitment(hubbs 2014.Highconcentrationsofsandfrompastmicaandfeldsparmininginthe watershedcausedsevereimpactstoaquaticlifedownstream(tva1994.sincethe 1980survey,waterqualityconditionshaveimprovedsothatreintroductionof freshwatermusselscanbeconsidered PigeonRiveratCosbyCreek ThePigeonRiverofNorthCarolinaandTennesseehasalonghistoryofhabitat degradationfromalargedam,urbanandfarmrunoffandhighlevelsoftoxic effluentsfromapapermillincanton,northcarolina,whichwasconstructedin 1908.Thecombinedimpactsofthesedisturbancescausedtheextirpationofall nativemussel,andmanyfish,speciesdownstreamfromcanton,northcarolinato theconfluencewiththeriverintennessee,a101kmsectionofstream(bartlett, 1995.Overthelast20years,thepapermillhasmodernizedandgreatlyreduced wateruseandwasteproduction,therebyimprovingwaterqualityinthepigeon River(NCDENR,2008.Consequentlymanyhost(fishspecieshavereturnedtothe river,providingsupportfornativemusselpopulations.becauseofthese improvementsandtheidentificationofextirpatedmusselspeciesfromprevious archeologicalstudiesandobservationsbyexperts,thetwradecidedtoattempt musselrestorationviatranslocation(hubbs,2014inthepigeonriveratthecosby anddentonsites BroodStockfromtheVDGIFAWCC TheVDGIFAWCC,locatednearMarion,VA,wasestablishedin1998bytheVirginia DepartmentofGameandInlandFisheriestoactivelyrecoverVirginia sfreshwater mussels.theawcccultivatesfreshwatermusselsandprovidedl.(fasciolasubadult musselsforthetwratranslocationeffortssurveyedandanalyzedinthisresearch. Whenjuvenilemusselsreleasefromthehostfish,theyarecollectedviaanoutflow filterofthewaterrecirculationsystem.separatesystemsareusedforlargefishsuch 19

35 aslargemouthbass(micropterus(salmoides,andsmallerfishsuchasdarters (Percidae(AWCC2014.Collectedjuvenilemusselsarecountedandmeasured,and placedinarearingsystemallowingthemusselstogrowlargeenoughfor translocationoraugmentationandincreasingtheirchancesofsurvivalinthewild. Therearingsystemsusefilteredriverwatertoeliminatepredatorsthatmight consumesmalljuveniles,andkeepwaterchemistryasclosetonaturalaspossible. Throughthissystem,theAWCCstaffcancontrolthewatertemperatureandfood content.(awcc Species TranslocatedspeciesinthisstudyincludeLampsilis(fasciolaandMedionidus( conradicus.informationforthespeciesisprovidedbelow ! Lampsilis!fasciola! TheWavyrayedLampmussel,(L.(fasciola(Rafinesque,1820,isafreshwatermussel whoserangeintheunitedstatesandcanadianprovincesistheeasternusand Quebec,Canada(Figure2W2.InTennessee,itsconservationstatusis apparently secure (NatureServe,2014a.TheshortWtermtrendforL.(fasciolais relatively stabletodeclineof30%,wheremostofthedeclineisseenincanada,illinois,new YorkandMississippi(NatureServe,2014a.Lampsilis(fasciola(Figure2W3isasmall tomediumwriverspecies,andisfoundat1meterorlessdepth.thepreferred substrateofl.(fasciolaappearstobeamixtureofsand,cobbleandgravel(gordon andlayzer,1989.itoccursinsmallcreeksandmediumwsizedriversineastand MiddleTennessee,andinthemainstemCumberlandandTennesseeRiver reservoirs(parmalee&bogan,1998.itisextirpatedfromseveraltennessee 20

36 Figure2C2.L.!fasciolaUSrange((NatureServe,2014a,lastaccessed05/12/

37 Figure2C3.Lampsilis!fasciola(EncyclopediaofLife, lastaccessed4/3/

38 rivers,includingthewatagua,emory,frenchbroad,sequatchie,buffalo,andothers (Parmalee&Bogan,1998.Theshapeoftheshellisellipticalanditsthickness rangesfromthintothick/dense.thelengthofl.(fasciolaatmaturitycanreach90w 100mm(Parmalee&Bogan,1998.HaagandRypel(2011identifyLampsilinias shortwlivedthoughtheyfoundagreatrangeoflongevity(4w50years.scott(1994 statesamaximumageof32yearsformalesand24yearsforfemalel.(fasciola observedintheclinchriverinvirginia.thehostfishforl.(fasciolaarelistedin Table2W ! Medionidus!conradicus M.(conradicus(Lea,1834,orCumberlandMoccasinshell,isafreshwatermussel whoserangeisthesoutheasternus(figure2w4,endemictothetennesseeand CumberlandRiverdrainages.InTennessee,itsstatusis Vulnerable (NatureServe, 2014b.Itsshorttermtrendisadeclineof10W30%,withalongtermtrendof30W 50%decline(NatureServe,2014b.ThisspeciesisfoundinsmallWtomediumWsized streamssuchastheclinch,powell,holston,emory,watagua,andotherrivers (Tennessee.TheshellofM.(conradicusiselongateandelliptical,andrelativelythin (thoughitthickenswithage(parmalee&bogan,1998.thisisasmallspecies (Figure2W5,typicallyunder60mmlong. ItisfoundinmoderateWstrongcurrentsatdepthsoflessthan1meter.Medionidus( conradicushabitatissandandgravelsubstrate,thoughitisalsofoundinbedrock cracksandunderflatrocks(parmalee&bogan,1998.glochidiaarepresenton femalesinmidwseptemberandaredischargedinmidwlatemay(ortmann,1921. TherearefewdetailsonthelifehistoryandageofM.(conradicus,however,Scott (1994statesamaximumageof24yearsforspecimensintheClinchRiverin Virginia.ThehostfishforM.(conradicusarelistedinTable2W2. 23

39 Figure2C4.M.!conradicusUSrange((NatureServe,2014b,lastaccessed05/12/

40 Figure2C5.Medionidus!conradicus!(EncyclopediaofLife, Table2C2.HostFishforIntroducedMussels FreshwaterMussel L.(fasciola M.(conradicus *PrimaryHost HostFish(natural *SmallmouthBass(Micropterus(dolomieu(ZaleandNeves, 1982 BlackBass(Micropterus(spp(Jones&Neves,2011 LongearSunfish(Lepomis(megalotis(Wattersetal.,2009 *FantailDarter(Etheostoma(flabellare,*RedlineDarter (Nothonotus(rufilineatum(Zale&Neves,1982 RainbowDarter(Etheostoma(caeruleum,StripedDarter (Etheostoma(virgatum(Luo&Layzer,

41 CHAPTER3 MATERIALSANDMETHODS 3.1 Overview ThisstudyevaluatestheresultsoftranslocationofL.fasciolaandM.(conradicus,to tworiversites.musselgrowthandsurvivalareusedasproxiesforthenearterm successofthetranslocationefforts.additionalanalysesareconductedtoidentify factorsmostinfluencinggrowthandsurvival.mynullhypothesisisthatthereisno differenceingrowthorsurvivalregardlessofmusselspecies,destinationriver,or housingtype(cageorsilo.iexpect,however,thattheriverandspeciesinfluence boththegrowthandsurvivaloftranslocatedfreshwatermussels. Allmusselswereattachedwithacoloredtag(whitewithauniquenumeric identifier.thisallowedustocollectdataonindividualmusselsovertime. TranslocationoccurredonOctober10,2013withfollowWonsurveysofthe destinationsitesonjune10,2014andseptember23, Translocation Table3W1summarizesthesource,destination,species,quantity,andlifestageofthe translocatedmussels.thesourcesfortranslocatedm.(conradicusandl.(fasciolaare theclinchriveratkylesfordandtheawcc,respectively.onoctober10,2013, DonHubbs(TWRA,SteveAhlstedtandCraigWalkerreleased284adultM.( conradicusfromtheclinchrivertothepigeonrivercosbysiteand75adultm.( conradicusfromtheclinchrivertothenolichuckyriveratthetwracanoeaccess (Hubbs,2014.Atthatsametime,Dr.MichaelMcKinney(UTandstudents translocated237subwadult(27monthsl.(fasciola(awccpropagatestoboththe PigeonRiverCosbysiteandthesamenumbertotheNolichuckyRiverattheTWRA 26

42 Table3C1.Source&DestinationInformationforTaggedMussels(dataextractedfrom(Hubbs 2014 Source Destination Species Quantity VDGIFAWCC ClinchRiverat KylesFord(Hubbs 2014 VDGIFAWCC ClinchatKyles Ford PigeonRiver CosbySite PigeonRiver CosbySite Nolichuckyat TWRACanoe Access Nolichuckyat TWRACanoe Launch L.(fasciola( 237 M.( conradicus( Adult Lifestageat Introduction SubWadults (AgeW2years, 3months L.(fasciola( 237 SubWadults M.( conradicus( *AWCC=AWCCpropagatesfromClinchstock(Hubbs Adult CanoeAccess(Hubbs2014,hereafter PigeonRiver and NolichuckyRiver, respectively,unlessfurtherdistinctionisrequired.themethodoftransferwas madeviacoolersfilledwithriverwaterandincludingabatterywpoweredaeration pumptokeepthewateroxygenated,thusminimizingstress. Atotalof833musselsweretaggedandtranslocatedinthisstudy.TableAW1(in AppendixAprovidestheinitiallengthandweightforthetranslocatedmussels, withstatisticsforthesameprovidedintables3w2and3w3,respectively.theawcc retainedsomel.(fasciolafromthesamestockasthosetranslocated.thislengthis alsoreportedintable3w2,i.e.,l.(fasciola(retained.thelengthofamusselis measuredatthelongestpartoftheshellusingcalipers(figure3w1.thewetweight ofthemusselswasmeasuredusinganohauscs200scale.theawccdoesnot measuretheweightoflivemusselsbecauseoflikelyinconsistenciesinweightdue tovaryingamountsofwaterinandonamusselatdifferentmeasurementtimes.

43 Table3C2.LengthoftranslocatedmusselsandAWCCretainedmussels Length(mm Species Standard Average Deviation Min Median Max L.(fasciola( (translocated L.(fasciola( (retained M.(conradicus( Table3C3.Weightoftranslocatedmussels Weight(g Species Standard Average Deviation Min Median Max L.(fasciola( M.(conradicus( Figure3C1.Mussellengthmeasurement.Lengthmeasurementtakenatthelongestdimension (PicturedmusselisnotL.(fasciolaorM.(conradicus 28

44 3.3 Housing Atbothstudysites,translocatedmusselswerehousedincagesorsilos,orwere released10w15metersdownstreamofthecagesandsilos(nothoused.thesilos andcageswereusedtobothprotectthemusselsandtoenablesubsequentsurveys tofindthemusselsfordatacollection.thewaterdepthattimeofdeploymentofthe cagesandsiloswas0.9w1.5metersandthecagesandsilosweredeployedover3 metersfromeachshorebytwradivers.theseplacementparametersholdforboth thepigeonandnolichuckyriversitesandwererequiredtoreducetheriskof vandalism. Figure3W2identifiesthesource,destination,countsandhousingoftaggedand translocatedmussels.atotalof833musselsweretransferredinthisstudy;359m.( conradicusand474l.(fasciola.quantitiesof75and284m.(conradicuswere transferredtothenolichuckyandpigeonrivers,respectively.half(237ofthel.( fasciolaweretransferredtoeachdestinationriver.inthenolichuckyriver,64 musselswerecaged,80wereinsilos,and168weredistributedfrommeshbags.in thepigeonriver,64musselswerecaged,80wereinsilos,and377weredistributed frommeshbags Silo Silos(originaldesignbyChrisBarnhart,MissouriStateUniversityusedinthis study(figure3w3arecomposedofa10kgconcretedomeandapvcinner chamber.withintheinnerchamber,musselsarecontainedinstandard1.6wmm meshfiberglassbags.coveringtheinnerchamberisawiremeshwith1wcm 2 openings.oncethechamberisintheconcretedome,astrapispositionedoverthe PVCtokeepthechamberinthesilo.Thesilosweredesignedsuchthatwhenwater flowsoverthesilo,itcreatesabernoullieffect(bernoulli,1738;streeter,1966, drawingwaterupthroughthemusselenclosurethusprovidingasupplyofwater 29

45 All Tagged Individuals Clinch River! Medionidus conradicus AWCC! Lampsilis fasciola Nolichucky River Pigeon River Nolichucky River Pigeon River Cage Silo Not! Housed Cage Silo Not! Housed Cage Silo Not! Housed Cage Silo Not! Housed Figure3C2.Source,destination,countsandhousingoftranslocatedmussels Figure3C3.Silo.Silowidthis27cm,height12cm,metalopeningattopis8.6cmindiameter.The interiorcontainingthemusselsis4cmindiameterand4cminheight. 30

46 andnutrientstothemussels.thesilosalsoprotectthemusselsfrompredatorsand enablesurveyorstoeasilyretrievethemfordatacollection. Thevolumeofasilo,Vs=πr2h,whereristheradiusoftheinnerchamber,andhis itsheight.thesilosusedinthisstudyhaveavolumeof50.27cm3.thiswillbe usefulinformationtodeterminetheavailablevolumepermusselanalyzed Cage Cagesusedinthisstudy(Figure3W4areaPVCpipe(chamber.Withinthis chamber,musselsarecontainedinstandard1.6wmmmeshfiberglassbags.covering bothendsofthepipeisawiremeshwith1cm2openings.thecagesweredesigned tobeplacedparalleltotheground(layingonthepipe sside,suchthatwaterflows throughthecageprovidingasupplyofwaterandnutrientstotheenclosedmussels. Thecagesalsoprotectthemusselsfrompredatorsandenableeasyretrievalfor subsequentdatacollection. Figure3C4.Cage.Dimensionsofthecageare9cminheightand6.6cmdiameter. 31

47 Thevolumeofacage,Vc=πr 2 h,whereristheinteriorradiusofthepipe,andhisits height.thecagesusedinthisstudyhaveavolumeof307.91cm 3.Thiswillbeused todeterminetheavailablevolumepermussel. Thenumbersofinitialmusselsplacedinthecagesandsilos,alongwiththeirspecies areprovidedintables3w4and3w5forthepigeonandnolichuckyrivers, respectively.twocagesof24l.(fasciolaeachand2cagesof20m.(conradicuseach werelocatedinthepigeonriver.fivesilosof12l.(fasciolaeachand2silosof10m.( conradicuseachwereplacedinboththepigeonandnolichuckyrivers. Table3C4.AveragecageandsilohousingoccupantsCPigeonRivermussels House Type(CageorSilo InitialOccupants Species S1 Silo 10 M.(conradicus C1 Cage 24 L.(fasciola C2 Cage 24 L.(fasciola S2 Silo 12 L.(fasciola S3 Silo 12 L.(fasciola C3 Cage 20 M.(conradicus C4 Cage 20 M.(conradicus S4 Silo 12 L.(fasciola S5 Silo 12 L.(fasciola S6 Silo 12 L.(fasciola S7 Silo 10 M.(conradicus( Numberof Occupantsper Housing Cage: L.(fasciola:24 M.(conradicus:20 Silo: L.(fasciola:12 M.(conradicus:10 Totaloccupants =168 32

48 Table3C5.AveragecageandsilohousingoccupantsCNolichuckyRivermussels House Type(CageorSilo InitialOccupants Species S1N Silo 12 L.(fasciola C1N Cage 20 M.(conradicus S2N Silo 12 L.(fasciola C2N Cage 24 L.(fasciola S3N Silo 10 M.(conradicus S4N Silo 10 M.(conradicus S5N Silo 12 L.(fasciola S6N Silo 12 L.(fasciola C3N Cage 24 L.(fasciola S7N Silo 12 L.(fasciola C4N Cage 20 M.(conradicus( Numberof Occupantsper House Cage: L.(fasciola:24 M.(conradicus:20 Silo: L.(fasciola:12 M.(conradicus:10 Totaloccupants = MeshBagRelease Amajorityofthemusselstransferredwerereleasedfrommeshbagsdirectlyonto thesubstrate.thesewereplacedinshallow(lessthan1meterdeepwater,10w15 metersdownstreamofthecageandsilolocations.ofthetranslocatedmussels,168 weredistributedfrommeshbagsinthenolichuckyriverand377weredistributed thiswayinthepigeonriver. 3.4 WaterQuality ChlorophyllCA ChlorophyllWAisanindicatorofnutrientavailability.Weconductedpointwater samplingforchlorophyllatthepigeonriversiteonmay6,2014andthenolichucky RiversiteonMay15,2014.Two1Lsampleswereobtainedoffthebankateach location,wherethewaterdepthwasapproximately0.6meters.onceobtained,the samplewasplacedina4 Celsiuscoolerforstorageandtransportationbacktothe 33

49 laboratory.eachsamplewaslabelledwith ChlWA,sampledate,location,sample indicator(aorb,andthecollector sinitials. Uponarrivalatthelaboratory,wefollowedthetestingprotocolsoftheTurner10W AUFluorometerandrantriplicatesforeachsample.TheFluorometerprovides resultsinfluorescence,andweusedthefollowingequationtoconvertthevaluesto μg/ltocalculatetheconcentration. C=(F*Va/Vs wherec=chlorophyllwaconcentration(µg/l F=FluorescenceMeasured Va=Volumeofacetoneused Vs=Volumeofsamplefiltered WaterDischargeRate Waterdischargerate(cubicmeterspersecond,m 3 /secistheonlywaterquality datasetavailableforallsitesoverthestudyperiod.theaverageandmaximum waterdischargepermonth(figures3w5and3w6,respectivelyfortheclinch,pigeon andnolichuckyriversindicatehighlyvariabledischargesandmaximumdischarges exceeding400(clinchandpigeonriversand500(nolichuckyriverm 3 /s.for eachsite,themaximumlevelsoccurindecemberandarevariable,yetsimilartoone another,throughouttheyear. 34

50 140.0 AverageMonthlyDischarge(m 3 /s Figure3C5.AverageMonthlyDischarge(m 3 /sduringthestudyperiodfortheclinch,pigeon andnolichuckyrivers.dataobtainedfromhttp://waterdata.usgs.gov/tn/nwis/inventory; accessed6/7/2015;usgssite clinchriverabovetazwell,tn;usgssite NolichuckyRiveratEmbreeville,TN;USGSSite PigeonRiveratNewport,TN. Clinch Pigeon Nolichucky 35

51 600.0 MaximumMonthlyDischarge(m 3 /s Clinch Pigeon Nolichucky Figure3C6.MaximumMonthlyDischarge(m 3 /sduringthestudyperiodfortheclinch,pigeon andnolichuckyrivers.dataobtainedfromhttp://waterdata.usgs.gov/tn/nwis/inventory; accessed6/7/2015;usgssite clinchriverabovetazwell,tn;usgssite NolichuckyRiveratEmbreeville,TN;USGSSite PigeonRiveratNewport,TN. 36

52 3.5 Surveys FollowWonsurveyswereconductedonJune10,2014andSeptember23,2014.Wet suitsandbreathingapparatus(scubaorsnorkelwereusedinthesearchforthe cagesandsilos.thehousingswereplacedincoolerswithriverwaterandaerating pumps.cageswerebroughttoshoreindividuallyformeasurementtominimizethe timethemusselswereoutoftheirhabitat.aftermeasurement,eachhousingwas returnedtotheriverandplacedinitsoriginalposition. 3.6 DataCollectedandAnalyzed Datawerecollectedforeachmusselrecovered,including Tagnumber, River, Species(equatestosourceinthisstudy, Housing(cage,siloormeshbagrelease, Length(livemussels(mm, Weight(livemussels(g, Conditionofhousing(intermsofsilt,and Lifestatus(live,dead. Thefirstandthirdsurveyswerefullsurveys,whereasthesecondsurveyanalyzeda reducedsampleoftherelocatedmusselsaswedidnotwanttodisturballthe specimensbutwerejustcheckingonthegeneralstatusoftheexperiment.basic descriptivestatisticswerecalculated(usingmicrosoftexcel (2010foreach individualdatatypeandmorecomplexmodels(section3.8wererequiredto evaluatethesignificanceoftheeffectsofthesefactorsongrowthandsurvival. 3.7 RecoveryRateandEffort MembersoftheTWRA(TennesseeWildlifeResourcesAgencyandseveralofDr. MichaelMcKinney sundergraduatestudentsaccompaniedtheauthoronthefinal 37

53 survey.thecatchperuniteffort(cpueiscalculatedasfollows.therecoveryrate, R,isthenumberofmusselsrecoveredduringthei th survey(nidividedbythe numberofmusselsoriginallytransferredtothelocation(n0.thecpueisthe numberofmusselsrecoveredduringthei th survey(nidividedbytheeffort,e,(in personwhourstocapturethosemussels,measuredinmusselsperpersonwhour. ( R=ni/n0,0.0 R 1.0 CPUE=ni/e,CPUE AnalyzingSurvivalandGrowthRate Survivaliscalculatedasthenumberofmusselsaliveattheendofthestudydivided bythenumberofmusselstranslocated,i.e.,musselslive/musselstotal.bothabsolute growthrateandrelativegrowthrates(agrandrgr,respectivelyareevaluated. 30Wdayabsolutegrowthrate(AGR30iscalculatedasfollows: AGR30=!!"!!"%! 30 where m = number of days in the study, 348 for the current study, and l = length (mm AGR30unitsaremmper30days. RGR30isthegrowthraterelativetothemussel sinitialsize.itisafractionalvalue, whichifmultipliedby100providesthepercentrelativegrowthrate.rgr30is calculatedasfollows: 38

54 RGR30=!!"!!!!"%!!"%! 30 Amixedmodelanalysisofvariance(ANOVAisusedtotestforsignificanceof combinedeffectsofriversite,housingtypeandtimeonmusselsurvivalandgrowth rate.generalizedmixedmodels(saxton,2013wereimplementedusingthesas StatisticalSoftware,version9.4(SAS,2013GLIMMIXprocedure.Thetreatments analyzed(table3w6areriver,species,housingandtime.anexampletreatmentis: L.(fasciola(housedinaSilointhePigeonRivermeasuredon10/10/2013. Table3C6.StudyDesign Class Levels Values DestinationRiver 2 Pigeon Nolichucky Species 2 L.(fasciola( M.(conradicus Housing 2 Silo Cage Time 3 10/10/2013 6/10/2014 9/23/

55 Forsurvivalrate,arandomizedblockdesign(RBDisusedwiththedependent variable,survivalrate(sr.theeffectsofdestinationriver,species,housingand Timeareexaminedinthisfullmodel.Inaddition,separatemodelsweredeveloped foreachspecies.forthespecieswspecificmodels,thedependentvariableisthe survivalrate,andtheeffectsofdestinationriver,housingandtimeareexamined. TheSASPROCGLIMMIXwasusedforallmodelsinthisstudy.Otherspecificsofthe survivalmodelsinclude: Normalityoftheresidualsisassumed. Variance/covariancetype=vc(variancecomponentsisused.OntheRBD repeatedmeasuresdesigns,variance/covariancetype=ar(1didnotoffera significantenoughimprovementinaic(theakaikeinformationcriterion (Akaike,1973towarrantitsselection. KenwardRogerdegreesoffreedomadjustmentswereusedforthetypeIII testsforfixedeffects. BonferrnoniadjustedpWvalueswereusedintheLSWmeansposthoctests. Themodelingequationsregardingfixedandrandomeffectsfollowclassic techniquesforrbddesigns,splitplotandsplitwsplitplotdesigns. Waldcovarianceparametertestswereusedtodeterminewhetherthe covarianceparametersweredifferentfromzero. Forgrowthrate,acompleterandomizeddesign(CRDisusedwiththedependent variable30day[absolute Relative]GrowthRate.AsintheSRmodels,thefull modelexaminestheeffectsofthedestinationriver,species,housing,andtimeon growthratewithseparatemodelsdevelopedforeachspecies.thesasproc GLIMMIXwasusedforthegrowthratemodelsinthisstudy.Additionalspecificsof thegrowthratemodelsinclude: 40

56 Variance/covariancetype=vc(variancecomponentswasusedformost models. KenwardRogerdegreesoffreedomadjustmentswereusedforthetypeIII testsforfixedeffects. BonferrnoniadjustedpWvalueswereusedintheposthoctestswhere necessary. Themodelingequationsregardingfixedandrandomeffectsfollowclassic techniquesforcrddesigns,splitplotandsplitwsplitplotdesigns. TheWaldcovarianceparametertestswereusedtodeterminewhetherthe covarianceparametersweredifferentfromzero. 41

57 CHAPTER4 RESULTS 4.1 InterimSurveyResults TheinterimsurveyofthePigeonandNolichuckysitestookplaceonJune10,2014. Thepurposeofthissurveywastocheckthestatusofthemussels,examineasample, andcollectdataonthatsample.tableaw2providesthelengthandweightdata collectedintheinterimsurvey.table4w1summarizesthelength,weightand survivaldatacollectedonthe100livel.(fasciolaand37livem.(conradicustagged individualsrecoveredduringthissurvey. Table4C1.Summaryofinterimsurvey,6/10/2014.Lengthandweightaremeasuredforlive musselsonly. Species Length(mm Average StdDev Min Median Max L.(fasciola( M.(conradicus( Weight(g Average StdDev Min Median Max L.(fasciola( M.(conradicus( ( Survival Live Dead Total SurvivalRate L.(fasciola( M.(conradicus( FinalStudySurvey AfullsurveyofthePigeonandNolichuckysiteswasconductedonSeptember23, 2014.TableAW3providesthelengthandweightdatacollectedinthissurvey,while 42

58 Table4W2summarizesthelength,weightandsurvivaldatacollectedonthe168L.( fasciolaand120m.(conradicusfound. Table4C2.Summaryoffinalsurvey,9/23/2014.Lengthandweightaremeasuredforlivemussels only. Species Length(mm Average StdDev Min Median Max L.(fasciola( M.(conradicus( Weight(g Average StdDev Min Median Max L.(fasciola( M.(conradicus( ( Survival Live Dead Total SurvivalRate L.(fasciola( M.(conradicus( RecoveryRateandEffort MembersoftheTWRA(TennesseeWildlifeResourcesAgencyandUniversityof Tennessee(UTundergraduatestudentsofDr.MichaelMcKinneyaccompaniedthe authorontheseptember23,2014survey.overall,35%ofthe833totalrelocated musselswererecovered(table4w3.therecoveryrateforl.(fasciola(andm.( conradicus(were35%and33%,respectively,ofthetotalmusselsreleased.recovery ratebyriverwas28%and46%,respectively,forthepigeonandnolichuckyrivers. Noneofthemusselsreleasedfrombagswererecoveredandnosignificant concentrationofdeadshellsofl.(fasciola(orm.(conradicuswerefound.allrelocated musselsplacedincagesorsiloswererecoveredforanalysis. 43

59 Table4C3.Recoveryrate Species Site Housing NoliC L.! M.! Pigeon Bag chucky Cage Silo fasciola conradicus River Release River L.!fasciola! 0.35 Species M.!conrad icus Site Pigeon River Nolichucky River Housing Cage ( Silo ( Bag Release L.! fasciola! M.! conrad3 icus! Pigeon River Nolichucky River Pigeon River Nolichucky River Overall 0.35 Ofthoseparticipatinginthesurvey,4participantswereassignedtotherecovery effort,whileothersweighedandmeasuredtherecoveredindividuals.table4w4 describesthecatchperuniteffort(cpueofmusselsinthestudy,withanoverall CPUEof12.0,theNolichuckyat14.4andthePigeonwithaCPUEof10.3. Table4C4.CatchperUnitEffort Metric Pigeon Nolichucky Total Timespentonrecovery(hours No.personsonrecovery TotalpersonChours No.musselsrecovered CPUE UnitEffort=1personWhour 44

60 4.4 Survival DescriptiveStatistics Thesurvivalrate(NLive/MRecoveredofspeciesoverthestudytime(Figure4W1 showsasteeperslopeofdeclineform.(conradicus.thereisverylittledifference betweenthedestinationriversintermsofsurvivalrate(sr(figure4w SurvivalRate L.(fasciola( Figure4C1.SurvivalbySpeciesoverTime M.(conradicus( 45

61 SurvivalRate Pigeon Nolichucky Figure4C2.SurvivalbyRiveroverTime(valuesshownforNolichuckyRiver TheAWCCheldbackasampleofL.(fasciolajuvenilesofthesameageandstockas thoseusedinthisstudy.figure4w3presentsthesrovertimeforl.(fasciolainthe PigeonRiver,NolichuckyRiverandthoseheldbyAWCC.TheAWCCheldmussels hadthehighestsurvivalwithallmusselssurviving.thepigeonriverprovided slightlyhighersrforl.(fasciolathandidthenolichuckyriver.thesrform.( conradicuswashigherinthenolichucky(figure4w4. 46

62 SurvivalRate Pigeon Nollichucky AWCC Figure4C3.L.!fasciolaSurvivalbyDestination SurvivalRate Pigeon Nollichucky Figure4C4.M.!conradicusSurvivalbyDestination 47

63 IfweexamineSRandhousingtype(Figure4W5,weseelittledifferenceinSR betweencagesandsilosatthefinalsurvey.housingconditionwasnotedonthelast survey.icategorizehousingconditionbasedontheproportionofsiltinthehousing asfollows:clear/nosilt(0,lightsilt/lessthan10%silted(1,moderately silted/[10%<siltbyvolume<50%](2,andheavilysilted/morethan50%siltby volume(3.thosefewwithhousingcondition3were90w100%silted.srby housingcondition(figure4w6showsgreaterthan80%survivalforthosemussels inhousingofconditions0w2andlessthan20%survivalforthosemusselsinhousing ofcondition3(heavilysilted.thisfindingsupportsthefindingthatextremely highlysiltedhousing(cond3wasresponsibleforthedeathsofthemusselshoused therein SurvivalRate Cage Silo Figure4C5.SurvivalbyHousing 48

64 Survival Cond0 Cond1 Cond2 Cond3 HousingCondition Figure4C6.SurvivalbyHousingCondition IusedPearson scorrelationcoefficienttodetermineiftherewereanycorrelation betweenthevolumeavailablepermusselandsurvivalforthefollowing:within eachspecies,withineachriver,andwithineachcontainer.themaximum correlationwas0.36.figure4w7illustratessurvivalbyvolumeforeachspeciesand giventhelargestandarddeviationonmanypointsandthesmallnumberof replicates,moredataareneededtoexaminethisrelationshipfurther.this evaluationdoesnottakeintoaccountthereducedvolumeavailabletoeachmussel becauseofincreasedsiltation. 49

65 Survival VolumeperOccupant(cm 3 M.(conradicus( L.(fasciola( Figure4C7.SurvivalbyVolumeperOccupant. ( 50

66 4.4.2 ModelCBasedAnalysisofSurvival AsnotedintheMethods,IusedaMixedModelANOVA(splitWsplitplotforsurvival withmodelinputsprovidedintablebw1(appendixb.theassumptionsforthis modelaremet(seetablebw5withnormalityofthemodelresiduals(figurebw4 andnooutliersintheresiduals.thereisequalityofvariancebetweenthetreatment conditions. TheDestinationRiverwasshownnottobestatisticallysignificanttomussel survival,withpr>0.05(tablebw3.thismeansthat,inthisstudy,musselsurvivalis notinfluencedbyriver:thepigeonriverisnotdifferentthanthenolichuckyin termsofoverallmusselsurvival.inaddition,thecovariancefordestinationisnot differentfrom0.0,meaningthatallvariancesduetodestinationriverareexplained bythemodel.giventheseresults,thefullmodelisblockedonriver.thefullmodel ANOVAtableisshowninTable4W5. Table4C5.FullSurvivalModelANOVA Source Error Degreesof Freedom(DoF (numerator DoF (denominator Wp B*Wp 1 2 Sp B*Wp*Sp 1 2 SSp B*Wp*Sp*SSp 2 8 Wp*Sp B*Wp*Sp 1 2 Wp*SSp B*Wp*Sp*SSp 2 8 Sp*SSp B*Wp*Sp*SSp 2 8 Wp*Sp*SSp B*Wp*Sp*SSp 2 8 where B=Block,inthiscaseonDestination Wp=Wholeplot,Species 51

67 Sp=Splitplot,House SSp=SplitWsplitplot,Time ThefullmodelrevealsthattheTime(ofsurveyeffectonSurvivalisthemost significant(tablebw2.thetimeeffectisstatisticallysignificantandtheeffectsat eachsurveytimearestatisticallydifferent(tablebw3,figurebw1.themodelalso revealsthatthehouse*timecombinedeffectissignificant(tablebw4,figurebw2. ToprovideadditionalassurancethattheDestinationRiverisnotasignificantfactor (inthisstudyinmusselsurvival,iexaminesplitplotmodelsforeachspecies.that is,iprepared2additionalmodels oneforl.(fasciolaandtheeffectsonitssurvival, andaseparatemodelform.(conradicusandtheeffectsonitssurvival.thespeciesw specificmodelanovatableisshownintable4w6.themodelassumptions(ofno severeoutliers,normality,andequalityofvariancearemetforboththel.(fasciola andm.(conradicusmodels(tablesbw6andbw7.allvariabilityisaccountedforin eachspecieswspecificmodelwithoverallanddestinationcovariancesnotdifferent than0.thel.(fasciolamodelindicatesthattimeisaverysignificantfactoronl.( fasciolasurvivalandthehouse*timeinteractionissignificant(tablebw8.them.( conradicusmodelindicatesthattimeandthehouse*timeinteractionare significantfactorsonm.(conradicussurvival(tablebw9. Table4C6.SpeciesCSpecificSurvivalModelANOVA L.!fasciola! M.!conradicus! Source Error Num DoF Den DoF Num DoF Den DoF Wp B*Wp Sp B*Wp*Sp Wp*Sp B*Wp*Sp

68 where B=Block,onDestination Wp=Wholeplot,House Sp=Splitplot,Time 4.5 GrowthRate Thissectionpresentsresultsofanalyzingabsoluteandrelativegrowthrate.Note thatonlylivemusselsweremeasuredandusedintheseanalyses DescriptiveStatistics Absolute!Growth!Rate! The30Wdayabsolutegrowthrate(AGR30ofspeciesoverthestudytime(Table4W7 showsahigheragr30forl.(fasciola(figure4w8overm.(conradicus((figure4w9,at 0.80mmand0.29mm,respectively.AverageAGR30ishigherintheNolichucky althoughthepigeonrivermusselsexhibitthegreatestmaximumagr30. Figure4C8.L.!fasciola30CdayAbsoluteGrowthRate.Thegrowthratedensityisshownwiththe data snormalcurve. 53

69 Table4C7.SummaryofAbsoluteGrowthRate. AGR(mm/30days Species Average StdDev Min Median Max L.(fasciola( W M.(conradicus( W River Average StdDev Min Median Max Pigeon W Nolichucky W Species&River Average StdDev Min Median Max L.(fasciola( Pigeon W Nolichucky W AWCC M.(conradicus( Pigeon W Nolichucky W HousingType Average StdDev Min Median Max Silo W Cage W Species&House Average StdDev Min Median Max L.(fasciola( Silo W Cage M.(conradicus( Silo W Cage W

70 Figure4C9.M.!conradicus30CdayAbsoluteGrowthRate.Thegrowthratedensityisshownwith thedata snormalcurve. Asnoted,theAWCCheldbackasampleofL.(fasciolajuvenilesofthesameageand stockasthoseusedinthisstudyasacontrolgroup.table4w2includestheagr30for L.(fasciolainthePigeonRiver,NolichuckyRiverandheldbyAWCC.TheAWCCheld musselshadmoreconsistentgrowththanthoseinthepigeonandnolichucky.the NolichuckyRiverprovidedthehighestAGR30forbothspecies(Figure4W10.AGR30 byhousing(figure4w11showshigheragr30insilos,howeverthedifferencein averageagr30betweencagesandsilosis0.13mmper30days. 55

71 AGR W0.20 W M.(conradicus( L.(fasciola( Pigeon Nolichucky Figure4C10.AbsoluteGrowthRatebyRiver AGR W0.20 M.(conradicus( L.(fasciola( Cage Silo Figure4C11.AbsoluteGrowthRatebyHousing 56

72 Figure4W12depictsAGR30byhousingconditionandillustratesadifferentpattern thandoessurvivalbyhousingcondition.whereasthelowestsurvivalexistsfor musselsinhousingcondition3(heavilysilted,thosesamemusselsexhibitedthe highestgrowthrateat0.85mmper30dayperiod.inaddition,thereisno correlationbetweenagr30andavailablevolumepermusselinthehousingforthe timeperiodconsidered(348days AbsoluteGrowthRate(mm/30d Cond0 Cond1 Cond2 Cond3 HousingCondition Figure4C12.30CdayAbsoluteGrowthRatebyHousingCondition Relative!Growth!Rate! The30Wdayrelativegrowthrate(RGR30ofspeciesoverthestudytimeshowsa higherrgr30forl.(fasciola(figure4w13overm.(conradicus((figure4w14,at3.5% and0.96%,respectively.averagergr30ishigherinthenolichucky. 57

73 Figure4C13.L.!fasciolaRelativeGrowthRate(30d Figure4C14.M.!conradicusRelativeGrowthRate(30d 58

74 TheRGR30forL.(fasciolainthePigeonRiver,NolichuckyRiverandthoseheldby AWCCrevealsthattheAWCCheldmusselshadnonegativerelativegrowthrates, whereasthoseinthepigeonandnolichuckydid.thenolichuckyriverprovidedthe highestrgr30forl.(fasciola,wherebothrivershadthesamegrowthrateform.( conradicus(figure4w15.rgr30byhousing(figure4w16showshigherrgr30in silos,howeverthedifferenceinaveragergr30betweencagesandsilosislessthan 1.0%over30days.Figure4W17depictsRGR30byhousingconditionandillustratesa differentpatternthandoesabsolutegrowthratebyhousingconditionwith shrinkageexistingonlyintherelativegrowthrateformusselsincondition3 housing(heavilysilted.alloftheheavilywsiltedhousingswereinthenolichucky River.Inaddition,thereisnocorrelationbetweenRGR30andavailablevolumeper musselinthehousing.itisexpected,however,thattherewillbesuchacorrelation uponsubsequentsurveys,giventhemussels expectedpotentialgrowth. RGR W M.(conradicus( L.(fasciola( Figure4C15.RGR30byRiver Nolichucky Pigeon 59

75 RGR W0.10 M.(conradicus( L.(fasciola( Cage Silo Figure4C16.RGR30byHousing W % 0.55% 0.21% Cond0 Cond1 Cond2 Cond3 RelativeGrowthRate W0.1 W0.15 W0.2 W0.25 W0.3 W0.35 W0.4 W0.45 HousingCondition W40.12% Figure4C17.30CdayRelativeGrowthRatebyHousingCondition 60

76 Frequency HousingCondition(Siltation Pigeon Nolichucky Figure4C18.HousingConditionbyDestinationRiver ModelCBasedAnalysisofGrowthRate Absolute!Growth!Rate! Toanalyzethefactorsthatinfluencegrowthrate,IusedaMixedModelANOVA, (TableCW3(AppendixCwithmodeldatashowninTableCW1andsummarizedin TableCW2.ThedependentvariableisAbsoluteGrowthRateandIexaminedthe effectsofdestinationriver,species,andhousing(tablecw4.theassumptionsfor thismodelaremetwithnormalityofthemodelresiduals(figurecw3andthe conditionalresiduals(figurecw9.thefullmodelanovatableisshownintable4w 8. 61

77 Table4C8.FullAbsoluteGrowthRateModelANOVA DoF Source Error DoF(numerator (denominator Wp R(Wp 1 97 Sp Sp*R(Wp 1 41 SSp SSp*Sp*R(Wp 1 41 Wp*Sp Sp*R(Wp 1 85 Wp*SSp SSp*Sp*R(Wp 1 85 Sp*SSp SSp*Sp*R(Wp 1 85 Wp*Sp*SSp SSp*Sp*R(Wp 1 85 where Wp=Wholeplot,Destination Sp=Splitplot,Species SSp=SplitSplitplot,House R=Reps Themodeldeterminesthatall2Wwayeffectsarestatisticallysignificantforboth species(tablecw4.tablecw5providesthemeanseparationfor30wdayabsolute GrowthRateinmm(AGR30,whichisusedtoidentifystatisticallysignificanteffects. Themodelrevealsthatall2Wwayeffects,i.e.,Destination*Species, Destination*House,andSpecies*Housearestatisticallysignificant(TablesCW8,CW10, andcw11,andfigurescw5,cw7,andcw8,respectively.thatis,l.(fasciolainthe NolichuckyRiverhadthegreatestimpactonabsolutegrowthrate(fastestgrowth. TheNolichuckyRiver,regardlessofhousingtype,promotedthehighestabsolute growthrate.formusselsinthepigeonriver,thoseinsiloshadthehigherabsolute growthrate.l.(fasciolagrewfasterthanm.(conradicusregardlessofhousingtype, however,form.(conradicus,silospromotedgreaterabsolutegrowth. 62

78 Relative!Growth!Rate! Toanalyzethefactorsthatinfluencerelativegrowthrate,IusedaMixedModel ANOVA,withmodeldatashowninTableCW12andsummarizedinTableCW13.The dependentvariableisrelativegrowthrateandiexaminetheeffectsofdestination River,Species,andHousing(TableCW14.ThefullmodelANOVAtableisshownin Table4W9. Table4C9.FullRelativeGrowthRateModelANOVA DoF Source Error DoF(numerator (denominator Wp R(Wp 1 95 Sp Sp*R(Wp 1 36 SSp SSp*Sp*R(Wp 1 87 Wp*Sp Sp*R(Wp 1 36 Wp*SSp SSp*Sp*R(Wp 1 87 Sp*SSp SSp*Sp*R(Wp 1 87 Wp*Sp*SSp SSp*Sp*R(Wp 1 87 where Wp=Wholeplot,Destination Sp=Splitplot,Species SSp=SplitSplitplot,House R=Reps Theassumptionsforthismodelaremetwithnormalityofthemodelresiduals (FigureCW10,nosevereoutliersandequalityofvariance(TableCW15.Themodel revealsthatall2wwayeffects,i.e.,destination*species,destination*house,and Species*Housearestatisticallysignificant(TablesCW18,CW19,andCW20, respectively.intermsofthedestinationandspeciescombinedeffect,l.(fasciolain thenolichuckyrivergrewatthehighestrelativegrowthrate.forthecombined 63

79 DestinationandHousingeffect,theNolichuckyregardlessofhousingandthePigeon Rivermusselsincagesperformedbest.ForthecombinedSpeciesandHousing effect,thel.(fasciolaincagesperformedbest,thenl.(fasciolaandm.(conradicusin silosand,worst,m.(conradicusincages. Ialsodeveloped2mixedmodelANOVAforRelativeGrowthRatefortheindividual species.theanovatableforthespecieswspecificmodelsisshownintable4w10.for L.(fasciola,themodelassumptionsaremet(TableCW21andFigureCW11. Table4C10.SpeciesCSpecificRelativeGrowthRateModelANOVA L.!fasciola! M.!conradicus! Source Error Num DoF Den DoF Num DoF Den DoF Wp R(Wp Sp Sp*R(Wp Wp*Sp Sp*R(Wp where Wp=Wholeplot,Destination Sp=Splitplot,House TheeffectsonL.(fasciolaRelativeGrowthRatearedominatedbythedestination river(tablescw22,cw23,andfigurecw12,suchthatthenolichuckyriverprovides moregrowth.thecombinedeffectofdestination*house(tablecw22issignificant, inthatmusselsincagesinthenolichuckyhadthehighergrowth,thenthoseinsilos inthenolichucky. ForM.(conradicus,themodelassumptionsaremet(TablesCW25andCW26,andFigure CW13.WhiletheDestinationRiverisnotstatisticallysignificanttotheRelative 64

80 GrowthRateofM.(conradicus,theeffectsaredominatedbytheHousing(TableCW14 andfigurecw29andthedestination(tablecw28inwhichthehouseislocatedis statisticallysignificant.theyaredependentupononeanother(tablecw27andcw 30.Overall,musselsinSilosgrewfastest,whileSilosintheNolichuckypromoted thegreatestgrowth. 4.6 WaterQuality Iconductedpointwatersamplingforchlorophyllandprovidethoseresultsbelow. TheaverageChlorophyllWAconcentrationsatthePigeonRiversiteonMay6,2014 andthenolichuckyriversiteonmay15,2014were12.3µg/land18.5µg/l, respectively. Averageandpeakmonthlywaterdischargerates(m 3 /secfortheclinch,pigeon andnolichuckyrivers(figures3w19and3w20indicatehighlyvariabledischarges andmaximumdischargesexceeding400m 3 /sec.foreachsite,themaximumlevels arereachedindecember.iexaminedthemeandailydischargerateforthepigeon Riversite(USGSsite andthesitejustbelowWaltersDam(USGSsite ,for1/1/2012 9/30/2014(Figure4W19andfoundover98% correlationoverthetimeseries(95%confidencelevel,pwvalue<2.2ew16.though thiscorrelationdoesnotimplydirectcausation,itdoesindicatethatthewaterville powerplant(waltersdamwaterdischargeislikelythesourceofthepigeonriver site shighdischargerate. 65

81 MeanDailyDischarge(m 3 /s /1/12 3/1/12 5/1/12 7/1/12 9/1/12 11/1/12 1/1/13 3/1/13 5/1/13 7/1/13 9/1/13 11/1/13 1/1/14 3/1/14 5/1/14 7/1/14 9/1/14 11/1/14 WaltersDam PigeonatNewport Figure4C19.PigeonRiver(USGSsite andWaltersDam(USGSsite MeanDailyDischarge(m 3 /s1/1/2012 9/30/

82 CHAPTER5 DISCUSSION Thepurposeofthisresearchistoprovidequantitativeinsightsintothefactors affectingfreshwatermusseltranslocationefforts,inparticularforl.(fasciolaandm.( conradicusinthepigeonandnolichuckyriversofeasttennessee.iexaminedthe factorsoftherivertowhichthemusselsweretranslocated(thedestinationsite,its nutrientlevel,waterdischargelevels,sedimentandsiltation,themusselenclosure orhousing(cage,siloandbagreleased,thestudytimeandthespeciesthemselves. UsingfieldobservationsofthesefactorsandstatisticalandmodelWbasedanalyses, severalinsightswerequantitativelyillustrated. Oneunfortunateaspectofthestudyisthatnomusselsreleaseddirectlyontothe substrate(bagreleasedweresubsequentlyfound.hence,nostatementonsurvival orgrowthofbagwreleased,translocatedmusselscanbemade.anumberoffactors mayhavecontributedtotheinabilitytorecoverthenonwhousedmussels,including predation,burrowing,andwaterdischargevolume.thoughpredationmayhave accountedforsomeoftheunrecoveredmussels,wesawnoevidenceofspoilpiles, brokenshellsorunbrokenandemptyshellsatornearthetranslocationsitesfor eitherspecies. L.(fasciolaandM.(conradicusareburrowingmusselsandourinabilitytorecoverthe bagwreleasedmusselsmaybedueinparttothisbehavior.wedidnotdigintothe substratetolocatepotentiallyburrowedmusselsbecausedoingsowoulddisturb thehabitat.copeandwaller(1995surveyedmusseltranslocationeffortsand determinedanoverallrecoveryrateof43%.forriversintennessee,therecovery ratewas12.2%overallwhenanestimatewasprovidedintheliteraturebefore1995 (Cope&Waller,

83 ThePigeonRiversite(consistingofboulders,cobblesandexposedbedrock,with somefinesedimentsmayhavepreventedburrowing.theclinchandnolichucky Rivershavefinesedimentsthataremoresuitableforburrowing.TheNolichucky RiversitehadthehighestpeakdischargeWover500m 3 /sduringthestudyperiod (December,2013.Dischargewasalsohigh(andalsohighestinDecember,2013in thepigeonriverandthesourceclinchriver,bothover400m 3 /s.giventheinitial weightsofthemusselsunder5g,thetranslocatedmusselsmayhavebeenwashed awaybyhighwatervelocities. Alltranslocatedmusselsplacedincagesorsiloswererecoveredandanalyzed.The cageandsilohousingsareeffectiveinprotectingmusselsfrompredatorsandhigh velocity,thusenablingtheirrecovery.oftherecoveredmussels,thesurvivalrates were59%and63%forcagesandsilos,respectively,muchhigherthanthesurvival rateforriversintennesseeincopeandwaller s(1995survey(11.4%. TheL.(fasciolainthisstudyweresubWadultswhenreleasedinOctober,2013. HanlonandNeves(2006notethatjuvenileL.(fasciolareleasedinJunehadgreater survivalratesthanthosereleasedinseptemberormarch.thosereleasedin SeptemberandMarchexperiencedhighmortalitywithinthefirstmonthofrelease. TheyalsofindthatoverwintersurvivalexhibitedasizeWdependentrelationship.Due tothelimitedsamplesizeofmyinterimsurvey,astatisticalevaluationofsizew dependentoverwwintersurvivalwasnotpossible. ComparingthesurvivalrateforL.(fasciolainthePigeonRiver(72%,Nolichucky River(66%andthoseheldbyAWCC(100%showsthattheAWCCWheldL.(fasciola hadthehighestsurvivalrate.thisislikelyduetothe1consistentavailabilityof nutrients,2lackofsiltationand3lowwatervelocitiesandlowvelocityvariability forthel.(fasciolaraisedbyawcc.thesurvivalratesforcageandsilowhoused musselswerequiteclose.however,ifwelookattheconditionofthehousingin relationtosurvival,weseethatthosemusselsinthemostheavilysilted(~90% 68

84 siltedhousessufferedthehighestmortality(81%,althoughthosehousedwith50w 70%silthadthehighestsurvival(100%.Thismayhaveoccurredbecauseof potentialdifferencesintimingofsiltationofthehouses,withearliersiltation resultinginamorecompletelysiltedenclosureandhighermortality.thosewith thesecondhighestlevelofsiltationmayhavebeenabletobestsurvivebecauseof theavailabilityofenoughroomandfinesedimentformanyofthosemusselsto partiallyburrowwithintheenclosure. UsingmixedmodelANOVAtoidentifythemostsignificanteffectsonsurvivalofthe musselsinthisstudy,thereareseveralfindings.timehasthemostsignificanteffect ontranslocatedmusselsurvival.specifically,survivaldecreaseswithtimeforboth species,asexpected.whatisunexpectedistherateatwhichsurvivaldecreases.it isunknownwhatmayhavecausedthisdecline,butinitialhandlingand translocationstressesmayhaveplayedarole.inaddition,ifindtheeffecton survivalofthetimeandhouseinteractionissignificant.survivaldecreaseswith TimeandHouse.Thedifferenceoccursattheinterimsurveywheresilossupported highersurvivalthancages.thismaybeduetothebernoullieffectinsilosproviding morenutrientsandoxygenatedwaterortounidentifiedsamplingbiasattheinterim survey.contrarytomyexpectations,thedestinationriverwasnotstatistically significanttotranslocatedmusselsurvival. SpeciesWspecificmodelsrevealedthesameresultsasthefullmodel,withTimebeing verysignificanttol.(fasciolasurvival.thesimilarityofresultsinthefulland specieswspecificmodelsmaybeduetothesimilarityinsurvivalrateforbothspecies. FollowWonsurveysofthesemusselsorlongerWtermstudiesofthesespeciesare recommendedtodetermineiftheseresultsholdwithadditionaltime. HanlonandNeves(2006reportedjuvenileL.(fasciolagrowthobservations(in additiontosurvival.thosereleasedinjunehadgreatergrowthratesthanthose releasedinseptemberandmarch.thosereleasedinseptemberormarchexhibited 69

85 poorgrowthinthecoolseasonalwaterconditions.theaverage30dayrelative growthrateforl.(fasciolaatthefinalsurveyforthisstudywas3timesthatinthe interimsurvey.thisislikelyduetoslowedgrowthduringthefallandwinter seasons. A2004W2008studyinClinchRiverinTennessee(Jones&Neves,2011foundL.( fasciolamaximumageandsizetobe45yearsand91.3mmformales,and24years and79.8mmforfemales.the2014twrasurvey(twra,2014foundthelengthof M.(conradicuscollectedintheClinchRiverrangedfrom25W55mm.IntheNolichucky River,M.(conradicuslengthrangedfrom15W45mm(notincludingM.(conradicusin thisstudy.ageswerenotreported.thelengthofawccwheldl.(fasciolaranged from15.7w24.2mm(average20.9mm.thetranslocatedl.(fasciolaofthisstudy rangedinsizefrom19.08w58.97(average31.87mm,andthem.(conradicussize rangedfrom25.31w50.92mm(average40.56mmattheendofthestudy.the minimumlengthsofbothspeciesarewithinreportednormalranges.theminimum lengthofthestudyl.(fasciolawaslessthanthatoftheawccwheldl.(fasciolabecause ofshellshrinkageinsomeofthetranslocatedmussels.shrinkageiscommonin musselsthatarestressed(haag,2012. Themaximumlengthsofmusselsinthisstudyaresmallerthanthemaximum lengthsrecordedinthewildbecausethetranslocatedmusselswerelessthan5 yearsoldwhenmeasuredatthefinalsurvey,muchyoungerthantheirmaximum reportedage.themaximumlengthofthestudyl.(fasciolawasgreaterthanthatof theawccwheldl.(fasciola.thismaybeduetoincreasednutrificationfrom agriculturalrunwoff.however,alongwiththenitrogeninagriculturalrunwoffcomes otherpollutantssuchaspesticidesandfinesedimentsthatservetosuffocatethe mussels.thesepollutantsandotherchemicaltoxinsdecreasesurvivalofmussels withincreasedexposureandconcentrations(hariharanetal

86 RelativegrowthrateforL.(fasciolawasgreaterthanthatforM.(conradicus.While bothspeciesexhibitedshellshrinkage,m.(conradicussufferedgreatershrinkage.in particular,thosemusselsinheavilysiltedhousingsexhibitedanegativeaverage relativegrowthrate. UsingmixedmodelANOVAtoidentifythemostsignificanteffectsonrelativegrowth rateofthemusselsinthisstudy,ifindthatall2wwayeffects,i.e., Destination*Species,Destination*House,andSpecies*House,arestatistically significantforbothspecies.l.(fasciolagrewfasterthanm.(conradicusandsilos promotedfastergrowththancages.intermsofthedestinationandspecies combinedeffect,l.(fasciolainthenolichuckyrivergrewatthehighestrelative growthrate.forthecombineddestinationandhousingeffect,thenolichuckyriver, regardlessofhousing,andthepigeonrivermusselsincagesperformedbest.for thecombinedspeciesandhousingeffect,thel.(fasciolaincagesperformedbest, thenl.(fasciolaandm.(conradicusinsilosand,worst,m.(conradicusincages. TheL.(fasciolaWspecificgrowthratemodelrevealedthattheDestinationRiver dominatestheeffectsonl.(fasciolarelativegrowthrate,withthenolichuckyriver promotingfastergrowthofl.(fasciola.inaddition,thecombinedeffectof DestinationandHouseissignificant.Thatis,ifthedestinationistheNolichucky, Silospromotefastergrowth.However,inthePigeonRiver,therewasnosignificant differenceingrowthratesforeitherhousing. TheM.(conradicusWspecificgrowthratemodelrevealedthattheeffectsonrelative growthrateofm.(conradicusaredominatedbyhousing,withsilospromotingfaster growth.thedestinationinwhichthehouseislocatedissignificant,wheresilosin thepigeonriverpromotethefastestgrowth.weseeadifferencebetweenspecies withrespecttodestinationand/orhousing.followwonsurveysofthesemusselsand longerwtermstudiesofthesespeciesarerecommendedtodetermineiftheseresults holdwithadditionaltime. 71

87 Averagemonthlywaterdischargerates(m 3 /secfortheclinch,pigeonand NolichuckyRiversindicatehighlyvariabledischargesandmaximumdischarges exceeding400m 3 /sec.foreachsite,themaximumlevelsarereachedindecember. ThemeandailydischargerateforthePigeonRiversiteandthesitejustbelow WaltersDam,for1/1/2012 9/30/2014haveover98%correlation.Thoughthis correlationdoesnotimplydirectcausation,itdoesindicatethatthewaterville powerplantwaterdischargeislikelythesourceofthepigeonriversite shigh dischargerateandanegativefactorinthesurvivalandgrowthoffreshwater mussels. ThroughthisstudyIsoughttodeterminethemechanism(sthatinfluencethe outcomesofrecenttennesseefreshwatermusseltranslocations.theabilityto housethetranslocatedmusselsisimportanttotheabilitytorecoverandanalyzethe mussels.whentheconditionsoftheseenclosuresbecomesignificantlysilted(over 90%,however,themusselscansuffershellshrinkageandincreasedmortality.One mayconsidermorefrequentsurveysandreleaseofhousedmusselswhentheir housingconditiondeteriorates(orrinsingthehousingofsediment.atsomepoint thehousedmusselswillgrowtoasizeatwhichthevolumeofthehousinglimits growthandpossibly,survival. AsurprisingresultofthemultiWfactormodelingisthatthedestinationriverwasnot asignificanteffectinthesurvivalofthetranslocatedmussels.thenolichuckyriver wasthesoledestinationwithhousingsufferingsignificantsiltation.onewonders whetherfindingsomeofthereleasedmusselsmayhaveaddedsupporttoor contradictedthisfinding.thedestinationriverwassignificantonlyintherelative growthrateofl.(fasciola,withthenolichuckyriverpromotingfastergrowth. 72

88 Recommendations(for(future(research:( Severalsmall,singleWinvestigatorfreshwatermusselstudieshavebeenandcontinue tobeconductedintennesseerivers.giventhepaucityoffundingdirectedtoward freshwatermusselcommunityrestoration,isuggestopenandtimelydatasharing betweenthoseinvestigatorsandwiththescientificcommunityatlarge.giventhis collectionofdata,alongwiththeassociateddatacollectionmethods,abetter understandingoffreshwatermusselcommunitiesandrestorationsuccessfactors canbediscernedatlowerfuturecosts.havingthisdataavailablecanalsofacilitate longerwtermresearch. IsuggestfutureresearchtakeperiodicandquantitativeobservationsovermultiW yearstudies.factorsshouldincludewaterquality(chemical,geological,and physicalcharacteristics;sedimentspecifics;othermussels,hostfish,andpredators inthecommunity;musselbiologyandnutrientavailability. Implications(for(conservation(efforts:( TheresultsofthisstudyindicatethatbothL.(fasciolaandM.(conradicuscanbe successfullytranslocatedtothepigeonriver,if1themusselsweretranslocated wherethepigeonriverhaslessboulder,cobbleandexposedbedrockinfavorof morecoarseandfinegravelandsand,2thepigeonriverhadlowerpeakand averagedischargerates,particularlyatcriticaltimesinthemussellifecycle,3some translocationsoccurredinthespringwearlysummer,and4thetranslocated musselsareinitiallyhousedincagesorsilos.thesiloandcageenclosureswere helpfulinrecoveringmusselsforanalysis.noneofthenonwhousedmusselswere recovered,primarilyduetohighwatervolumesandvelocitiesinthefirstthirdof thestudy.thehousedmusselswereprotectedfromtheseissues.however,if musselsareinitiallytranslocatedintheheavysilosandrockwwedgedcages,atsome pointthosemusselswillbereleased.thoughthemusselsinthisstudywereprew adultandwerenotsexuallymature,thehighwatervelocityinthepigeonriverwill likelyreducethetimemalemusselspermarespatiallyaccessibletolocalfemale 73

89 mussels.thisreducestheopportunityforrecruitment.duetosignificantmortality inthefirstthirdofthisstudy,somestudiesshouldstartinthespringwearlysummer ratherthaninseptember/octobertohelptranslocatedmusselssurvivetheirfirst overwwinterbyhavingsomegrowthandhabitatacclimationunderway.inaddition, siltationoftheenclosuresandmusselgrowthshouldbemonitoredtoensurethe volumeavailabletothemusselsissufficient. 74

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110 APPENDICES 95

111 APPENDIXA:RawData TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013 TagNum Species Length(mm Weight(g 0 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

112 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 35 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

113 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 73 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

114 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 112 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

115 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 150 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

116 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 188 M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola

117 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 227 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola

118 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 265 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

119 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 303 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

120 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 340 M.conradicus L.fasciola M.conradicus L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

121 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 377 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

122 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 415 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

123 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 454 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

124 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 492 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

125 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 531 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

126 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 569 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

127 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 606! M.conradicus! 29.6! 2! 607! M.conradicus! 39.31! 5! 608 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

128 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 645! M.conradicus! 43.77! 9! 646! M.conradicus! 41.62! 8! 647 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

129 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 683! M.conradicus! 42.02! 8! 684! M.conradicus! 28! 3! 685 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus

130 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 721! M.conradicus! 36.78! 4! 722! M.conradicus! 45.2! 10! 723 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus L.fasciola M.conradicus M.conradicus

131 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 759 M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

132 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 797 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

133 TableA41.InitialLengthandWeightofTranslocatedMussels,10/10/2013.Continued TagNum Species Length(mm Weight(g 835 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

134 TableA42.LengthandWeightofTranslocatedMusselsonInterimSurvey,6/10/2014 TagNum Species Length(mm Weight(g* 1 L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 110 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus M.conradicus

135 TableA42.LengthandWeightofTranslocatedMusselsonInterimSurvey,6/10/2014. Continued TagNum Species Length(mm Weight(g* 124 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus M.conradicus L.fasciola M.conradicus L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola 247 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

136 TableA42.LengthandWeightofTranslocatedMusselsonInterimSurvey,6/10/2014. Continued TagNum Species Length(mm Weight(g* 273 L.fasciola L.fasciola L.fasciola 289 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus 568 M.conradicus M.conradicus

137 TableA42.LengthandWeightofTranslocatedMusselsonInterimSurvey,6/10/2014. Continued TagNum Species Length(mm Weight(g* 570 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus 713 M.conradicus M.conradicus 743 M.conradicus 762 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 805 L.fasciola L.fasciola L.fasciola L.fasciola *Deadmusselswerenotweighed.Musselswithablankweightcellweredeadatthe secondsurvey. 122

138 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014 TagNum Species Length(mm Weight(g* 0 L.fasciola 1 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 35 L.fasciola 38 L.fasciola M.conradicus 42 L.fasciola 44 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 60 L.fasciola L.fasciola L.fasciola 65 L.fasciola L.fasciola L.fasciola

139 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014.Continued TagNum Species Length(mm Weight(g* 69 L.fasciola L.fasciola L.fasciola 74 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 88 M.conradicus 91 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 102 L.fasciola L.fasciola 107 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 125 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus 124

140 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014.Continued TagNum Species Length(mm Weight(g* 137 L.fasciola L.fasciola L.fasciola 144 L.fasciola L.fasciola 147 L.fasciola 149 L.fasciola 152 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 175 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola M.conradicus 192 L.fasciola L.fasciola M.conradicus M.conradicus L.fasciola 206 L.fasciola M.conradicus L.fasciola L.fasciola

141 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014.Continued TagNum Species Length(mm Weight(g* 219 L.fasciola L.fasciola 222 L.fasciola M.conradicus L.fasciola 226 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 245 L.fasciola L.fasciola 248 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 259 L.fasciola 260 L.fasciola M.conradicus 264 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 289 L.fasciola L.fasciola L.fasciola L.fasciola

142 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014.Continued TagNum Species Length(mm Weight(g* 300 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 342 L.fasciola 344 L.fasciola L.fasciola 350 L.fasciola L.fasciola L.fasciola L.fasciola 356 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola

143 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014.Continued TagNum Species Length(mm Weight(g* 388 L.fasciola 390 L.fasciola L.fasciola L.fasciola M.conradicus M.conradicus M.conradicus 419 M.conradicus 422 M.conradicus M.conradicus M.conradicus 429 M.conradicus M.conradicus M.conradicus M.conradicus 447 M.conradicus 448 M.conradicus M.conradicus M.conradicus 452 M.conradicus M.conradicus 461 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus 509 M.conradicus 515 M.conradicus M.conradicus 517 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus 128

144 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014.Continued TagNum Species Length(mm Weight(g* 550 M.conradicus M.conradicus 556 M.conradicus 557 M.conradicus 560 M.conradicus M.conradicus M.conradicus M.conradicus 569 M.conradicus 570 M.conradicus M.conradicus 572 M.conradicus M.conradicus M.conradicus M.conradicus 576 M.conradicus 577 M.conradicus M.conradicus M.conradicus 580 M.conradicus M.conradicus 585 M.conradicus M.conradicus M.conradicus 593 M.conradicus 596 M.conradicus M.conradicus 607 M.conradicus 612 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus 661 M.conradicus M.conradicus M.conradicus

145 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014.Continued TagNum Species Length(mm Weight(g* 671 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus 700 M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus M.conradicus 743 M.conradicus 761 L.fasciola L.fasciola L.fasciola 770 L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola L.fasciola 805 L.fasciola L.fasciola L.fasciola L.fasciola

146 TableA43.LengthandWeightofTranslocatedMusselsonFinalSurvey,9/23/2014.Continued TagNum Species Length(mm Weight(g* 836 L.fasciola L.fasciola *Deadmusselswerenotweighed.Musselswithablankweightcellweredeadatthe finalsurvey. 131

147 AppendixB.AnalysisofSurvivalRateEffects TableB41.SurvivalRateFullModel,RBDSplit4SplitPlotRepeatedMeasureswithout ReplicationontheArcsinTransformedValues,BlockonDestination4Data Obs Destin4 ation Species House Time Survived Trials Survival_ Rate 132 Arcsin_SR TableB42.SurvivalRateFullModel4TypeIIITestsofFixedEffects Effect NumDF DenDF FValue Pr>F Species House Species*House Time <.0001 Species*Time House*Time Species*House*Time

148 TableB43.Effect=Time,Method=LSD(P<0.05,Set=4,SurvivalRateFullModel Obs Species House Time Esti4 mate Standard Error Mean Standard Error ofmean Letter Group A B C FigureB41.LeastSquaredMeans(LSMforSurvivalRate,Effect=Time,FullModel TableB44.Effect=House*Time,Method=LSD(P<0.05,Set=6,SurvivalRateFullModel LSMArcSin(SR Obs Spec4 ies T1 T2 T3 Time House Time Estimate Standard Error Mean Standard Error ofmean Letter Group 18 _ A 19 _ C 20 _ C 21 _ A 22 _ B 23 _ C 133

149 A B C Connected(Lines(are(Statistically(the(Same 0.63%%%%%%%%%0.64%%%%%%%%%0.78%%%%%%%%%1.19%%%%%%%%%1.57%%%%%%%%%1.57 FigureB42.LSMforEffect=House*Time,SurvivalRateFullModel FigureB43.Time*HouseLSMforArcsin_SR,SurvivalRateFullModel 134

150 FigureB44.NormalityofArcsin(SurvivalRateResiduals,FullModel TableB45.FullSurvivalRateModelAssumptionsareMet ***DiagnosticSummary*** Nosevereoutliersorinfluentialpointsfound Normalityshouldbesatisfactory Equalvarianceshouldbesatisfactory TableB46.L.fasciolaSurvivalRateModelAssumptionsareMet ***DiagnosticSummary*** Nosevereoutliersorinfluentialpointsfound Normalityshouldbesatisfactory Equalvarianceshouldbesatisfactory 135

151 TableB47.M.conradicusSurvivalRateModelAssumptionsareMet ***DiagnosticSummary*** Nosevereoutliersorinfluentialpointsfound Normalitymightbeanissue,KS>0.10; ShapiroTWilkestestindicatesNormality Equalvarianceshouldbesatisfactory TableB48.L.fasciolaSurvivalModel4TypeIIITestsofFixedEffects Effect NumDF DenDF FValue Pr>F House Time <.0001 House*Time TableB49.M.conradicusSurvivalModel4TypeIIITestsofFixedEffects Effect NumDF DenDF FValue Pr>F House Time House*Time

152 AppendixC.AnalysisofGrowthRateEffects TableC41.AbsoluteGrowthRateData TagNum Destination Species House 30dayAbsolute GrowthRate

153 TableC41.AbsoluteGrowthRateData.Continued TagNum Destination Species House 30dayAbsolute GrowthRate T

154 TableC41.AbsoluteGrowthRateData.Continued TagNum Destination Species House 30dayAbsolute GrowthRate T T T T T

155 TableC41.AbsoluteGrowthRateData.Continued TagNum Destination Species House 30dayAbsolute GrowthRate T

156 TableC41.AbsoluteGrowthRateData.Continued TagNum Destination Species House 30dayAbsolute GrowthRate T T T

157 TableC41.AbsoluteGrowthRateData.Continued TagNum Destination Species House 30dayAbsolute GrowthRate T

158 TableC42.AbsoluteGrowthRateSummaryStatistics Destination Species House NObs Mean StdDev Min Max T T T T T Key Obs Observationnumber TagNum TagNumberofmussel Length Lengthofmussel(mm LifeStatus Live(1orDead(0 SCStatus SiloorCageStatus: 0 NotSilted 1 LightlySilted 2 ModeratelySilted 3 HeavilySilted Destination Locationmusseltranslocatedto 0 PigeonRiver 1 NolichuckyRiver Species MusselSpecies 0 Lampsilisfasciola 1 Medionidusconradicus House Housingmusselsplacedin 0 Cage 1 Silo Time ObservationTime 1 9/10/ /10/ /23/2014 Replication Replicationnumberofthe (Destination,Species,House,Time instance 143

159 TableC43.SASGLIMMIXProcedureFullAGR 30ModelInformation DataSet ResponseVariable ResponseDistribution LinkFunction VarianceFunction VarianceMatrix EstimationTechnique DegreesofFreedomMethod TableC44.TypeIIITestsofFixedEffects,FullAGR 30Model WORKS.RANKS RankGrowthRate Gaussian Identity Default Notblocked RestrictedMaximumLikelihood Containment Effect NumDF DenDF FValue Pr>F Destination <.0001 Species <.0001 Destination*Species <.0001 House <.0001 Destination*House <.0001 Species*House <.0001 Destination*Species*House NumDF:Demoninatordegreesoffreedom DenDF:Numeratordegreesoffreedom Pr>F(pTvalue<α,α=0.05 TableC45.MeanseparationforRankAbsoluteGrowthRateinFullAGR 30Model Set AverageSig DiffValue MinimumSig DiffValue MaximumSig DiffValue

160 FigureC41.LSMforDestination,FullAGR 30Model TableC46.MeanSeparationforRankAbsoluteGrowthRate,Effect=Destination,FullAGR 30 Model Obs AGR30(mm Destin4 ation 0 1 Destination Method=Bonferroni(P<0.05Set=1 Estimate Standard Mean Standard Error Error ofmean B A Letter Group TableC47.MeanSeparationforRankAbsoluteGrowthRate,Effect=Species,FullAGR 30Model Obs Species Estimate Standard Error Method=Bonferroni(P<0.05Set=2 Mean 145 Standard Error ofmean A B Letter Group

161 TableC48.MeanSeparationforRankAbsoluteGrowthRate,Effect=Destination*Species,Full AGR 30Model Method=Bonferroni(P<0.05Set=3 Obs Destin4 ation Species Estimate Standard Error Mean StandardError ofmean Letter Group B B A B TableC49.MeanSeparationforRankAbsoluteGrowthRate,Effect=House,FullAGR 30Model Method=Bonferroni(P<0.05Set=4 Obs House Estimate Standard Error Mean Standard Error ofmean B A Letter Group TableC410.MeanSeparationforRankAbsoluteGrowthRate,Effect=Destination*House,Full AGR 30Model Obs Destin4 ation Method=Bonferroni(P<0.05Set=5 House Estimate Standard Error Mean StandardError ofmean C B A A Letter Group TableC411.MeanSeparationforRankAbsoluteGrowthRate,Effect=Species*House,FullAGR 30 Model Method=Bonferroni(P<0.05Set=6 Obs Species House Estimate Standard Error Mean StandardError ofmean Letter Group A A B A 146

162 FigureC42.DestinationuntransformedLSMforAbsoluteGrowthRate,FullAGR 30Model 147

163 FigureC43.CheckonNormalityforRankAbsoluteGrowthRate,FullAGR 30Model 148

164 0.72mm 0.52mm FigureC44.RankAbsoluteGrowthRateLSMforSpecies,FullAGR 30Model 149

165 1.14mm 0.56mm 0.49mm 0.49mm FigureC45.RankAbsoluteGrowthRateLSMforDestination*Species,FullAGR 30Model 150

166 0.66mm 0.55mm FigureC46.RankAbsoluteGrowthRateLSMforHouse,FullAGR 30Model 151

167 0.80mm 0.81mm 0.54mm 0.44mm FigureC47.RankAbsoluteGrowthRateLSMforDestination*House,FullAGR 30Model AGR30(mm House Species0 Species1 FigureC48.AbsoluteGrowthRateLSMforSpecies*House,FullAGR 30Model 152

168 FigureC49.ConditionalResidualsforRankAbsoluteGrowthRate,FullModel 153

169 TableC412.RelativeGrowthRateData. TagNum Destination Species House 30dayRelative GrowthRate

170 TableC412.RelativeGrowthRateData.Continued TagNum Destination Species House 30dayRelative GrowthRate T

171 TableC412.RelativeGrowthRateData.Continued TagNum Destination Species House 30dayRelative GrowthRate T T T T T

172 TableC412.RelativeGrowthRateData.Continued TagNum Destination Species House 30dayRelative GrowthRate T

173 TableC412.RelativeGrowthRateData.Continued TagNum Destination Species House 30dayRelative GrowthRate T T T5.85ET

174 TableC412.RelativeGrowthRateData.Continued TagNum Destination Species House 30dayRelative GrowthRate T

175 TableC413.RelativeGrowthRateSummaryStatistics Destination Species House NObs Mean StdDev Min Max T T T T T TableC414.TypeIIITestsofFixedEffectsforRelativeGrowthRate,FullModel Effect NumDF DenDF FValue Pr>F Destination Species <.0001 Destination*Species <.0001 House <.0001 Destination*House <.0001 Species*House <.0001 Destination*Species*House NumDF:Demoninatordegreesoffreedom DenDF:Numeratordegreesoffreedom Pr>F(pTvalue<α,α=

176 FigureC410.ConditionalResidualsforRankRelativeGrowthRate,FullModel TableC415.FullRGR 30ModelAssumptionsareMet ***DiagnosticSummary*** Nosevereoutliersorinfluentialpointsfound Normalitymightbeanissue,KS>0.10 Equalvarianceshouldbesatisfactory TableC416.MeanSeparationforRankRelativeGrowthRate,Effect=Species,FullRGR 30Model Obs Species Estimate Standard Error Method=Bonferroni(P<0.05 Mean 161 Standard Error ofmean A B Letter Group

177 TableC417.MeanSeparationforRankRelativeGrowthRate,Effect=House,FullRGR 30Model Obs House Estimate Standard Error Method=Bonferroni(P<0.05 Mean Standard Error ofmean B A Letter Group TableC418.MeanSeparationforRankRelativeGrowthRate,Effect=Destination*Species,Full RGR 30Model Obs Destina4 tion Method=Bonferroni(P<0.05 Species Estimate Standard Mean Error Standard Error ofmean B B A B Letter Group TableC419.MeanSeparationforRankRelativeGrowthRate,Effect=Destination*House,Full RGR 30Model Obs Destina4 tion Method=Bonferroni(P<0.05 House Estimate Standard Mean Error Standard Error ofmean B A A A Letter Group TableC420.MeanSeparationforRankRelativeGrowthRate,Effect=Species*House,FullRGR 30 Model Method=Bonferroni(P<0.05 Obs Species House Estimate Standard Error Mean Standard Error ofmean A AB C B Letter Group 162

178 TableC421.TestsforNormality,L.fasciolaRGR 30Model Test Statistic pvalue Shapiro4Wilk W Pr<W < Kolmogorov4Smirnov D Pr>D < Cramer4wonMises WTSq Pr>WTSq < Anderson4Darling ATSq Pr>ATSq < FigureC411.ResidualsforRankRelativeGrowthRate,L.fasciolaRGR 30Model TableC422.TypeIIITestsofFixedEffectsforL.fasciolaRGR 30Model Effect NumDF DenDF FValue Pr>F Destination <.0001 House Destination*House

179 TableC423.MeanSeparationforRankRelativeGrowthRate,Effect=Destination,L.fasciola RGR 30Model Obs Destina4 tion Method=Bonferroni(P<0.05 House Estimate Standard Mean Error Standard Error ofmean 1 0 T B 2 1 T A Letter Group FigureC412.DestinationeffectonL.fasciolaRGR 30,LeastSquaredMeans 164

180 TableC424.MeanSeparationforRankRelativeGrowthRate,Effect=Destination*House,L. fasciolargr 30Model Obs Destina4 tion Method=Bonferroni(P<0.05 House Estimate Standard Mean Error Standard Error ofmean C C A B TableC425.M.conradicusRGR 30ModelTestsforNormality Test Statistic pvalue Shapiro4Wilk W Pr<W Kolmogorov4Smirnov D Pr>D Cramer4wonMises WTSq Pr>WTSq Anderson4Darling ATSq Pr>ATSq TableC426.M.conradicusRGR 30ModelAssumptionsareMet ***DiagnosticSummary*** Nosevereoutliersorinfluentialpointsfound Normalitymightbeanissue,KS>0.10 Equalvarianceshouldbesatisfactory Letter Group 165

181 FigureC413.ResidualsforRankRelativeGrowthRateModelforM.conradicus TableC427.TypeIIITestsofFixedEffectsforM.conradicusRelativeGrowthRate Effect NumDF DenDF FValue Pr>F Destination House <.0001 Destination*House <.0001 TableC428.MeanSeparationforRankRelativeGrowthRate,Effect=Destination,M.conradicus RGR 30Model Method=Bonferroni(P<0.05 Obs Destina4 tion House Estimate Standard Error 166 Mean Standard Error ofmean 1 0 T A 2 1 T A Letter Group