¡. THESIS SUBMITTED TO THE FACUTTY OF GRADUATE STUDIES PARTIAT FULFILMENT OF T}iE RESUIREI\IENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY

lHE UNIVERS]TY OF MANTTOBA POPU],ATÏON PARANiETERS AND B]OENERGETIC DEMANDS OF VIALLEYE, Stizostedion vitreum vitreum (Mitchill), IN RETATION TO THEIR TROPHTC DYNAMIC ECOLOGY' WEST BLUE LAi{E, MANITOBA by JOHN R.M ¡ KELSO THESIS SUBMITTED TO THE FACUTTY OF GRADUATE STUDIES IN PARTIAT FULFILMENT OF T}iE RESUIREI\IENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY DEPARTIV]ENT OF ZOOLOGY WINNIPEG, MANITOBA FAT,L , Tg?I O i i,,ih i! ilì0 tJA o\ ko" DÅFtËj#S TAB],8 OF CONTENTS Page No Introduction Methods and Materials Vital Statistics of the Population Capture and Handling Mortality Trials 3 6 Estimation of "Abundance Natural MortalitY Grov¡th Biomass and Production 10 I,aboratory Food Studj-es Gapture and TransPort Holding Conditions 11 11 Natural Feeding L5 Calorimetry L7 l2 L2 Exper'imental Treatment Terminology and Statistical Analysis 18 2I Results Vital Statistics of the Population Mortality Trials Background to lvlark and Release Exp'ts Àbtindance Esti-nates 2L 2L 23 29 Natural MortalitY 34 Biomass and Produetion Estimates 49 Laboratory Studies Effect- of Ration Size on Conversion Conversion EfficiencY 52 54 5? 66 7o 7? Growth 36 Maintenance Requirements Availab1e Ration An Unaccounted for EnergY Excess Natural Feeding Caloric Condent of Walleye and Their Natu,ra1 \^Ial1e.ve Feeding Population ConsumPtion Food 79 79 B6 93 Discussion Vital Statístics of the Popula.tion ' Abu.ndance Estima-tes Natural I'{ortality Growth Biomass and Prod.uction T,aboratory Studies Natural Feed.ing Population Consumption 98 98 98 LOs LO7 110 113 L?? J-32 Summary L36 References r39 Appendix L]ST OF TABLES Page No Table I Numbers of vralleye captured, relea.sed, to June l9?O marked, and recaptured from \¡\ay L969 in West Blue lake, 22 Table II Effects of hand.ling and marking on walleye survival Table III Results of chi-square tests of the hypothesis that rates of recapture for fish age II to VI were simj-lar between adjacent capture periods and for the entire 1969f0 period Table IV Estimates of l¡/est Blue Lake walleye abundance excluding (Xf) and including recruitment (WZ), standard deviations indieated in parenthesis Table V Estimates of West Blue lake wall eye population * characteristics u.sing Jotly' s Oge 5) pro,c'edú.re, SymboJ-s in parenthesis column heads are those used throughou.t this study Figures in parenthesis are standard deviations Tab1e VI Survival and instantaneous mortality rates of walleye in trVest BIue Lake , Table VII Population characteristics of lVest Blue Lake walleye derived from a triple-ca-tch-tre11is analysis procedure du.ring periods of constant recruitment (May to September, 1969, and September L969 to May 1970) Figures i-n parenthesis are standard deviations Table VIII, Results of paired- "t" tests appl-i-ed to growth measLr.res for the period between lr'iay and- June L969 NS mea.ns not sígnificant Table IX Lengths (mm) at annulu.s forrnation of walleye in West Blue T,ake, 1969-70 N represents the number in each sample 24 26 30 32 33 35 ¿tl 45 Table X Rela.tive grorvth in length anci vreight (hĂ and hỗ,respectivelĂr) from previous annulớ, and instantalleous rates of growth in length and weight (gl and gG respec- tively) for vra.I1eye, 1969-?O , Table XI Percentage of annu.al growth in v¡aIleye by comparíson to size in May and to increment from adjacent annuli (in parenthesis) Table XII Estimates of biomass (E) and produ.ction (P) in rrVest Blue Lake u.sing the arithmetic approaeh v¡ith averà$e'lveight (w) for each age group and instantaneotls grovrth in weight (g) Estimates of N in parenthesis are recruited population, 4Z 48 50 Table XIII Effect of ration size (% body weight) on conversion (K1 and K2) for walleye at L6C Table XIV Changes in energy content of whole a.ge II+ walleye before and after growth determinati-ons (standard deviations are in parenthesis) Table XV Conversion efficieneies (standard deviations are in parenthesis) of age II+ walleye for experiments at different temperatures 55 6o 63 Maj-ntenance requirements expressed as ng/g/wk 68 V, and VI year old walleye at LZC Table XVII Experimental and derived (equations for cray- fish and emerald shiners derived by procedures in the text) relations between assimilation efficiency (E) and walleye weight (vi) for various natu.râl food organisms 76 Table XVIII Energy utilization by walleye for maintenance, routine metabóIism (fn) and tfre excess in energy ( T) above twice the routine level (r-Ztn) All measures are in calorj-es and figures in parenthesis are standard deviations ?B Table XVI of II, TV, Table XIX CaLs/g weight, moisture, and ash content of erayfish (0, virilis) collected in West Blu.e Lake, L969- ievlati-ons are in parenthesis Table XX Calories/gram, ash, and moisture of age perch, Sergg flavessens (mitctritf ), and older fish õollecteã TFom West BLue T,ake , I97o Fi-gures in parenthesis are standard deviations Table XXI Energy content (cals/g), percent moisture, and ash of common lvalleye diet items, West Blue T,ake, L9?O Stand.arcl deviatións are Í-n parenthesi-s, Table XXII Surnmary of wal] eye f eeding during the icefree period, L9?O, Table XXIII Contribution of the diet items found in walleye stomach to walleye nutrition, West Blue Lake, Lg7O, Table XXIV Biocontent and energy production at a hypothetieal temperatu.re (column 4) for vralleye in West Blue Lake Intake (n) for the period and daily ratj-ons v{ere to natu.ral calculated using derived conversj-on adapted situati-on ?O, Standãid B0 Bz 84 B? 9L 9l+ IIST OF FIGURES Page No Fig.-1 lfgst BIue Lake shovring location of gill netting sites Fig 2., T,ength frequency distributions of waI1e¡re captured by J.81, 6.35 and 8.8! cm giII nets in t¡/est Blue T,ake , L969-7O N represents number captu.red in each mesh, Fig Length frequ-ency dístributions of walleye in West Blue Lake captured during mark and release periods Lt 28 3? Fig Length increments (¿1,) of West Blue Ï,ake walleye during the growth year (from last week of June to one year later) 39 Fig 5, The relation between weight (tg) and total 42 leñgtñ (mm) for West BIue T,ake walleye (Glenn, MS 1969) Fig 6, Cunulative growth in walleye (¿c) in grams wet weight during the growth year, L969-70, in West Blue Lake blr Fig ? Effect of ra.tion size on conversion (K1) at 16c for walleye fed on emerald shiners 56 Fíg B Accumulation of weight (S) by walleye fed at l+% boãy weight at 2O, L6, and 16-BC 58 Fig Effect of experiment du-ration on K1 for vralleye net¿ at 20, L6, and 16-BC All fish were fed a.t a ration 6t of 4/" body weight Fig 10 Conversion efficiencies in three orders (Kt, 6t+ K?, and K3) as affected by size of experimental walleye Fig 11 Relationship between maintenance requirements (cãtoric values in paienthesis) and temperatu.re for walleye Fig 12, Effect of ration size (in calories ingested) on-the assimilation of age perch by walleye If+ (upper) and old.er (lower) at I6c Fig 13, Effect of fish size (g wet weight) on assirnilation efficiencies for walleye fed at 16C Fig 14 Change in energy content of walleye du.ring I97O in West Blue Lake Fie 15, Contribu.tion of all diet items in terms of welght-( ) and calories (-) in relation to body siàe of vìa1]eye in West Blue Lake, during the ice-free period, 69 73 75 B5 L970, BB Fig 16, Contribution, by taxon, of resident organisms to the diet of walleye in lVest Blue Lake , ]-970, 9o ACKNOV¡LËDGEMENTS The kind help and guidance of Dr F.J I¡/ard financíal support of the Manitoba Provincial Government made this investigation possible Dr T,;J Johnson's aid in establishing priorities in the laboratory was most helpfut The remaj-ning committee members, Drs K, Doan and R, Green, also provided advice and 'guidance The help of graduate students in the field, and my wife in the laboratory was critical to this stud.y and the ABSTRACT During 1969-ZO, 569 walleye age II and older were marked in West Blue Lake of which J6 were recaptured No short term mortality was found in conjunction with mark and release periods As well, distribution of marked fish was essentially random, and sirnilar segments (by age) of the population v¡ere undm examination at all times Frequency distributions a.nd * tests ind.icate that net selection was not by age and unlikely to be by size The May 1969 population, 1090 walleye, decreased to 819 individuals in May 19?0, but was augmented by 2100 Rew recruits in September, 1969, Petersen estimates, the Jolly (irg65) approach, and the triple-catch trellis provid.ed similar descriptions of the population Mortality on a per day basi-s was small, mean i = 0.0045, and was greatest ,during faIl and winter .Growth, and consequently production, was greatest between kg, was stable from year to year, and production, l4O kg, was primarily from younger fish June and September Biomass, approxima.tely 800 conversion; K1 (##) and K2 (uffi), was affected by neither ration size nor temperature, but decreased with (#hUT) was affected only by temperature Walleye assirnilation efficiency was d.ependent upon diet type (1east efficient for invertebrates, and most efficient for fish), and fish size Maintenance per unit walleye was independent of size but was affected increasing fish size , K3 conversion by tempe.rature Maintenance requirements, all converted to zoc for various agg walleye approxirnates vlinberg's J (Lg56) T = J w' for routine metabolisrn ¡, séasonal cycle in caloric content of whole walleye (Iess,.gonads) occurred and was also evident in perch No in invertebrates examined Greatest energy eontribution to the nutrition of West Blue Lake wâI1eye was by pereh and stieklebacks Greatest numêrÍưaI contribution was by amphipods 4nd nayflies but both provided less energy such cycle was apparient Laboratory conversion and assimilation efficiency was applied to the natural diet of walleye for an estimation of population intake The resident population required from 40 to 1860 K ca¡/kg/day for production, and the intake depended upon season and population structure t INTRODUCT]ON Investigations into quantitative relationships of fish and their food follow three general approaches: defining biomass of food and fish; determining turnover in inorganic materials, such as nitrogen; and establishing energy transformations Allan 0.95]-).and Johnson Ogee) utilized maintenance requirements and gross conversion in the defi.nition of population requi-rements In addition Gerking (1-962) ¡ras described the turnover of nitrogen in the bluegill sunfish Mann (1965) tocated the enerỵgy transfor:matiorr in a river using procedures originally pointed out by winberg (L956), of this study were twofold: a) to determíne chara.cteristics of growth, number, survi-val and ultimately biomass and production of an unexploj.ted population of walleye, Stizostedion v vitreum (wlite¡rifl), and b) to describe under controlled conditions the effect of temperature, ration, and walleye size on growth, rnaj-ntenance, and assimilation The latter aspect of this investigation was based on theoretical reviews of growth anrl metabolism (paloheimo and Dicki.e , ]1965, 1966a and b) The stated otrjectives were fundamental eomponents in understanding production processes in natural systems and were consequently extended to describe seasonal differences in intake requíred to produce observed produeti.on in an unexpl-oited population This study util-ized the calorie to describe two The objec'cives LI+2 1969 Seasonal rates of growth within a population of walLeye Stizostedion vitreum vitreum 1¡¡titcnitl ), in west- Biïffiffi'Itỉa,-õrđfl 1966-67 M.Sc Thesis, University of Manitoba, Irlinnlpeg, Manitoba 63 p G1enn, C .L !S Go11ey, F B 1961 Enỗrgy.valqes of ecologcaI Ecology, lr2(3 materíal-s 5ù-58b Greenberg, B I9l+7 Some reilatlons between territorlr social hf-erarchy, and leadership in the gre€n sunfish ' (Lepomis llrneli-us) Phyèio1 Zoo1., 2Oz 269-299 Gross, W L., E, W Roelofs and P, O Fromm L965 fnfluence of photoperiod on grov{th of green sunfish, Lepomj-s cvairellu3 J Flsñ Res Bd; canada , zz(61ffiD86, Hathaway, ): E S L927 The relation of temperature to the quantity of food consumed by fishes EcoJ,., 13: h'23-l+34'' Hfle, R 1941 Age and growth of the rock bass, ArnblopLi,tes rqpgstltg (Rafinesgue ), in Nebish Lake',-WIsEonsïn, ffisc.Acad Sci.,Arts¡andI,etúerb,33lI89-337 R ].95l+ Fluctuations in growth and yearaclass strength rH1le, ' of walleye in Saginaw Bay Fish BuL1., U S Fish and lrli1d1, Serv., 561 7-59 of Uunt, B P.' 1960 Digestion rate and food consumption'Trans Florida Bârr warmouth and largemouth bass Amer Fiéh.-Soc., 89(2): 206-2L0 Job, S V 1960 Growth and calorific approxÍmation in the speckl-ed trout fnd J Fish , z 129-136 Johnson, L J 1966 Experimental determination of food j' J Fisñ Res BA Canada , Zl (l-0): ].L+95-l-5O5 Jo11y, G M, l-1965 Explicit estimates from capture-recapture data with both death and imnigration stochastic model Biometrika, 522 225-247 Keast, A Ì968 Feeding of some Great Lakes fishes at 1ow A.' I95l+ Growth, maturity, and mortality in the relatively unexploÍted lake tqgut of Great Slave Lake, J Fish Res BA Canada, 11(6): 827-852 Kinne, 1960 Growth intake and food conversion in a euryplastic fish exposed to dlfferent temperatures andsa1initios.Physiol.ZooI.,33l288-3l7 Keinnedy; W L43 Lawler, G L{ .196|.- Th-e food- of t4e plke, Esox rucÍus, in HgryiF kEg,-Maniroba J Fisrr: ; m:-rãáada, 22(61| 1357-1377 Mannl K 7?62 transforrnatíons by a population of _En"Igy ftsh ln the River Thames J Aniir E'coi., 3t+zZjZ-ZTj I,ỵccann, J A and K D carrander- rg?o Mark and recovery estimates of fish populaüions in cl-ear Lake, rowä r-958 and 1959 roiva' ftàüe j: sõi.;- 1116l-;- j6g:ioi Menzel, Dn q 1960 Utilizatíon' of food bf a Berrruda reef fÍsh, - ' Ep¿¡gøgfus euttatus.: J Cons Þem int Explor, G 19n studÍes on the feeding habits of fishes Ecology, 2?(ll z 9I-96 MÖore, G A - ]-gl+b The retj-nae of two North American üel_easÈs with specÍal, reference to their tapeta lucida Jour Comp Neur., 80(3 ): 369-j79.MoyJ-e, J B !2io Gflr nets for sampl-ing fish popurations in lvllnnesota waters Trans Àmér FÍsir.Moore, w 195-2Or+ Soc., T9: Molnar, G T and r To1g 1962 Rel-atj.on between water temperature and gastric digestion of largemouth bass (4{iõropterus salñoides Lacõpede) .1 Fïsh Res sFGtraÇlqf6lmõs-iol| Muir, B s.- 1963 vital statistics of muskie in Nogies creek f T.g loslr nrortalÍty due to tagging, anã the estiqate of exploitatlon J Fisñ ñés Bd canada, 20(51: tzt3-Lz3o Northcote,' T G and H Ìf, Lorz 1966 seasonal and diel changeb in'food of adult kökanee (oncorhvnctrus:l nerka ) in lrllcora Lake, British colffiõffiTish Ondonera, K \962 some aspects of behavior infl-uencing production fN Biologícal Basís of Freshwatei Fish ProductÍon S l Geit