1573 ARTICLE Temperature regimes, growth, and food consumption for female and male adult walleye in Lake Huron and Lake Erie: a bioenergetics analysis Charles P. Madenjian, Todd A. Hayden, Tyler B. Peat, Christopher S. Vandergoot, David G. Fielder, Ann Marie Gorman, Steven A. Pothoven, John M. Dettmers, Steven J. Cooke, Yingming Zhao, and Charles C. Krueger Abstract: Bioenergetics modeling was used to assess the relative importance of food availability and water temperature in determining walleye (Sander vitreus) growth. Temperature regimes experienced by both female and male adult walleye in three basins of Lake Huron and in Lake Erie were determined by use of surgically implanted temperature loggers and acoustic telemetry. Temperatures experienced by walleye were higher in Lake Erie than in Lake Huron. Walleye from Lake Erie grew at nearly double the rate of walleye from Lake Huron, and mass at age for adult females averaged about 50% greater than that for adult males in both lakes. Food consumption rate for an average adult walleye in Lake Erie was nearly twice as high as that in Lake Huron. Interbasin and interlake variability in temperature regimes accounted for a moderate degree of variability in walleye growth. We concluded that the driver for faster growth in Lake Erie compared with Lake Huron was higher food availability in Lake Erie compared with Lake Huron. The sex difference in temperature regimes explained 15% of the sex difference in Lake Erie walleye growth. Résumé : La modélisation de la bioénergétique a été utilisée pour évaluer l’importance relative de la disponibilité de nourriture et de la température de l’eau comme facteurs déterminants de la croissance des dorés jaunes (Sander vitreus). Les régimes thermiques auxquels sont exposés les dorés adultes femelles et mâles dans trois bassins du lac Huron et dans le lac Érié ont été déterminés en utilisant des enregistreurs de température implantés chirurgicalement et la télémétrie acoustique. Les tempéra- tures étaient plus élevées pour les dorés dans le lac Érié que pour ceux dans le lac Huron. Le taux de croissance des dorés du lac Érié était presque le double de celui des dorés du lac Huron, et le poids en fonction de l’âge pour les femelles adultes était en moyenne 50 % supérieur à celui des mâles adultes dans les deux lacs. Le taux de consommation de nourriture pour un doré adulte moyen dans le lac Érié était presque deux fois plus grand que dans le lac Huron. La variabilité des régimes thermiques entre bassins et entre lacs explique un degré modéré de variabilité de la croissance des dorés. Nous concluons que le facteur expliquant For personal use only. la croissance plus rapide dans le lac Érié que dans lac Huron est une plus grande disponibilité de nourriture dans le lac Érié. La différence entre les sexes sur le plan des régimes thermiques explique 15 % de la différence entre les sexes sur le plan de la croissance des dorés du lac Érié. [Traduit par la Rédaction] Introduction 1992; Jobling 1994; Quinn and Deriso 1999; Enberg et al. 2008). Thus, if the rate of food consumption is identical in two aquatic For a given species of fish, individual-level growth of fish is a ecosystems, and other factors affecting growth other than water function of food availability and the temperature regime experi- temperature are also identical across the two ecosystems, then enced by the fish (Hewett and Johnson 1992; Jobling 1994; Quinn fish growth would be expected to be faster in the aquatic ecosys- and Deriso 1999; Enberg et al. 2008). Food availability is a complex tem with the lower temperatures experienced by fish compared function of several factors, including prey abundance, prey size, with fish growth in the other aquatic ecosystem, owing to higher and handling times (Werner 1974; Persson and Greenberg 1990; metabolic costs in the warmer ecosystem (Hewett and Johnson Rennie and Venturelli 2015). Typically, metabolic costs for the fish 1992; Kao et al. 2015a, 2015b). increase with increasing water temperature, and metabolic costs Bioenergetics modeling has proven invaluable in assessing the represent a substantial energy loss for fish (Hewett and Johnson effects of various factors, including temperature regime, on fish Can. J. Fish. Aquat. Sci. Downloaded from www.nrcresearchpress.com by USGS LIBRARY on 09/27/18 Received 7 July 2017. Accepted 30 November 2017. C.P. Madenjian. US Geological Survey, Great Lakes Science Center, 1451 Green Road, Ann Arbor, MI 48105, USA. T.A. Hayden. Michigan State University, Department of Fisheries and Wildlife, Hammond Bay Biological Station, 11188 Ray Road Millersburg, MI 49759, USA. T.B. Peat and S.J. Cooke. Fish Ecology and Conservation Physiology Laboratory, Department of Biology, Carleton University, Ottawa, ON K1S 5B6, Canada. C.S. Vandergoot. US Geological Survey, Great Lakes Science Center, Lake Erie Biological Station, Sandusky, OH 44870, USA. D.G. Fielder. Michigan Department of Natural Resources, Lake Huron Research Station, 160 Fletcher Street, Alpena, MI 49707, USA. A. Gorman. Ohio Department of Natural Resources, Fairport Harbor Fisheries Research Station, 1190 High Street, Fairport Harbor, OH 44077, USA. S.A. Pothoven. National Oceanic and Atmospheric Administration, Great Lakes Environmental Research Laboratory, Lake Michigan Field Station, 1431 Beach Street, Muskegon, MI 49441, USA. J.M. Dettmers. Great Lakes Fishery Commission, 2100 Commonwealth Boulevard, Suite 100, Ann Arbor, MI 48105, USA. Y. Zhao. Ontario Ministry of Natural Resources and Forestry, Aquatic Research and Monitoring Section, Lake Erie Fisheries Station, 320 Milo Road, Wheatley, ON N0P 2P0, Canada. C.C. Krueger. Michigan State University, Center for Systems Integration and Sustainability, 1405 South Harrison Road, East Lansing, MI 48823, USA. Corresponding author: Charles P. Madenjian (email: [email protected]). Copyright remains with the author(s) or their institution(s). Permission for reuse (free in most cases) can be obtained from RightsLink. Can. J. Fish. Aquat. Sci. 75: 1573–1586 (2018) dx.doi.org/10.1139/cjfas-2017-0280 Published at www.nrcresearchpress.com/cjfas on 6 December 2017. 1574 Can. J. Fish. Aquat. Sci. Vol. 75, 2018 growth (Chipps and Wahl 2008; Madenjian 2011; Deslauriers et al. reside in either the northern main basin or the southern main 2017). A fish bioenergetics model represents an energy budget for basin (Hayden et al. 2014). Walleye remaining in Saginaw Bay are a fish. Energy input via food consumption is equal to metabolic expected to experience warmer temperatures than those walleye losses, waste losses, and growth: out-migrating to the main basin, while food availability for wall- eye is presumably higher in Saginaw Bay than in the main basin C ϭ (R ϩ A ϩ SDA) ϩ (F ϩ U) ϩ G (Fielder et al. 2014; Hayden et al. 2014; Peat et al. 2015). The primary goals of our study were to compare walleye growth where C = consumption, R = standard metabolism, A = energy in Lake Huron with walleye growth in Lake Erie and to assess the expenditure due to activity, SDA = specific dynamic action (energy relative importance of food availability and temperature regime required to digest food), F = egestion, U = excretion, and G = growth. as determinants of walleye growth in the Lake Huron – Lake Erie Standard metabolism is typically modeled as an exponential func- system. A secondary goal of our study was to evaluate the effect of tion of water temperature. Bioenergetics model applications to sex differences in temperature regimes on sex differences in wall- evaluate the effects of various factors on fish growth are well eye growth. Specific objectives included (i) using temperature illustrated in the study by Hayward and Margraf (1987), who used loggers and acoustic telemetry to determine temperatures expe- bioenergetics modeling to show that faster growth of yellow rienced by female and male walleye in Saginaw Bay, the northern perch (Perca flavescens) in the central basin of Lake Erie compared main basin of Lake Huron, the southern main basin of Lake Hu- with the western basin of Lake Erie was due to an interbasin differ- ron, and Lake Erie, (ii) developing growth trajectories for female ence in food availability rather than an interbasin difference in tem- and male walleye in both Lake Huron and Lake Erie and then perature regime. compare growth between the two lakes, (iii) using bioenergetics Coupling archival temperature tag technology with acoustic modeling to estimate food consumption by female and male wall- telemetry has recently enabled researchers to record thermal his- eye in Saginaw Bay, northern main basin of Lake Huron, southern tories of fish within different basins of the same lake, as well as main basin of Lake Huron, and Lake Erie, (iv) using a bioenergetics within lakes connected by waterways (Cooke et al. 2013; Hayden modeling framework to exchange temperature regimes experienced et al. 2014; Peat et al. 2015). By constructing an appropriate acous- by walleye between basins and lakes to assess the effects of interba- tic receiver network, interbasin and (or) interlake movement of sin and interlake variability in temperature regimes experienced by fish can be detected. Implanting both an acoustic transmitter and walleye on walleye growth, and (v) again using a bioenergetics mod- an archival temperature tag in each fish, and subsequent moni- eling framework to exchange temperature regimes experienced toring of any interbasin and interlake movements of the fish us- by walleye between the sexes to evaluate the effect of sex differ- ing acoustic telemetry, can be used by researchers to determine ences in temperature regimes on sex differences in walleye temperature regimes experienced by fish while confined to a spe- growth. cific basin or lake. Materials and methods To date, the effects of both interbasin and interlake variability in temperatures experienced by fish on fish growth in a two-lake sys- Field methods tem, with a connecting waterway between the two lakes, have not Walleye in this study were captured and tagged as part of a large been investigated.