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Winter Feeding As an Overwintering Survival Strategy in Young-Of-The-Year Winter Flounder Richard J

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Winter Feeding As an Overwintering Survival Strategy in Young-Of-The-Year Winter Flounder Richard J This article was downloaded by: [Department Of Fisheries] On: 25 September 2012, At: 20:24 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Transactions of the American Fisheries Society Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/utaf20 Winter Feeding as an Overwintering Survival Strategy in Young-of-the-Year Winter Flounder Richard J. Bell a a Graduate School of Oceanography, University of Rhode Island, South Ferry Road, Narragansett, Rhode Island, 02882, USA Version of record first published: 13 Jun 2012. To cite this article: Richard J. Bell (2012): Winter Feeding as an Overwintering Survival Strategy in Young-of-the-Year Winter Flounder, Transactions of the American Fisheries Society, 141:4, 855-871 To link to this article: http://dx.doi.org/10.1080/00028487.2012.675896 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Transactions of the American Fisheries Society 141:855–871, 2012 C American Fisheries Society 2012 ISSN: 0002-8487 print / 1548-8659 online DOI: 10.1080/00028487.2012.675896 ARTICLE Winter Feeding as an Overwintering Survival Strategy in Young-of-the-Year Winter Flounder Richard J. Bell* Graduate School of Oceanography, University of Rhode Island, South Ferry Road, Narragansett, Rhode Island 02882, USA Abstract For fish, the first winter of life can be a period of high mortality prior to recruitment. When resources are limiting or when fish are unable to feed due to low temperatures, starvation can often lead to size-dependent overwintering mortality. Larger individuals are typically better able to survive starvation because they have a higher percentage of energy reserves and a lower metabolic rate per unit of body mass than small individuals. Alternatively, fish that can feed during the winter are able to maintain their lipid stores and reduce their chance of starvation. The aim of this study was to examine the overwintering mortality and physiology of young-of-the-year (age-0) winter flounder Pseudopleuronectes americanus in relation to body size. Size-dependent mortality was investigated with 17 years of length-frequency data. In addition, I sampled the diet and whole-body crude lipid content of 309 age-0 winter flounder over the course of 1 year. Samples were taken in three estuaries in the northeastern USA during October– April. Whole-body crude lipid content ranged from 4.7% to 12.4% of dry weight. Larger age-0 winter flounder did not have higher lipid stores, and the age-0 fish did not exhibit size-dependent overwintering mortality. Age-0 winter flounder fed on amphipods and polychaetes throughout the winter, and their whole-body crude lipid content was maintained through the fall and winter. The physiology data lack temporal replication, but the spatial coherence in results (i.e., consistency among the three estuaries, representing two different stocks) suggests that the consumption and energy allocation patterns are real and that age-0 winter flounder follow an alternative overwintering survival strategy. Juvenile fish in temperate latitudes face multiple sources Larger individuals typically have a higher proportion of of mortality during their first year of life. During the summer, lipids and (based on allometric scaling relationships) a lower predation is typically a major source of mortality, whereas phys- metabolic rate per unit mass than small individuals (Bochdan- iological shock and starvation are often cited as major sources of sky and Leggett 2001). In numerous overwintering starvation Downloaded by [Department Of Fisheries] at 20:24 25 September 2012 mortality during the winter (Conover and Present 1990; Sogard experiments, larger individuals within a year-class exhibited 1997; Schultz and Conover 1999; Hurst 2007). When fish are higher survival than smaller conspecifics, suggesting that size- faced with limited resources, energy allocation must be directed dependent overwintering mortality was an important source of toward the highest risk factors during different life stages. Dur- mortality prior to recruitment (Post and Evans 1989; Johnson ing the summer, energy is typically directed toward maximizing and Evans 1990; Pangle et al. 2004). These starvation experi- growth at the expense of lipid reserves to achieve large size and ments clearly demonstrated that when food was lacking or when to minimize predation. In the fall and early winter, juvenile fish fish were unable to feed, larger individuals had higher survival often switch from maximizing growth to maximizing energy than smaller fish. However, fish of some species feed during stores to prevent starvation during the food-limited winter (Post the winter, potentially reducing the risk of starvation and alter- and Parkinson 2001; Hurst and Conover 2003; Biro et al. 2005; ing the energy allocation dynamics that occur prior to the onset Heintz and Vollenweider 2005). of winter. With a reduced risk of starvation, age-0 fish would *E-mail: [email protected] Received March 25, 2011; accepted January 4, 2012 Published online June 13, 2012 855 856 BELL not need to switch their energy allocation from growth to lipid METHODS storage, and the overwintering survival advantage of larger in- dividuals may not exist. Assuming that prey is available and Sample Collection that all size-classes are able to feed equally, the greater energy The length-frequency data for determining overwintering reserves in larger fish would not result in higher survival. Lipid mortality of age-0 winter flounder were collected during the levels would (1) remain relatively constant through the winter, RIDEM seine survey in October and the spring (April) trawl (2) vary with the availability of resources, or (3) both (Bystrom¨ survey from 1988 to 2005. A gear change in the spring trawl et al. 2006). survey prevented the inclusion of data from later years of the Winter flounder Pseudopleuronectes americanus are cold- survey. Age-0 winter flounder of the two coastal stocks remain adapted fish that are able to survive low temperatures by pro- in the nearshore areas and are susceptible to seining until the ducing antifreeze proteins (Fletcher 1981). Adults of the two fall (Saucerman and Deegan 1991); thereafter, they move into coastal stocks (Gulf of Maine and southern New England–Mid deeper water and are then susceptible to trawling (Dominion Atlantic Bight) move into estuaries and breed during the coldest Resources Services 2007). The seine survey initially sampled 15 part of the year at temperatures as low as −1◦C (Collette and stations throughout Narragansett Bay, increasing to 18 stations Klein-MacPhee 2002). However, the winter habitat of young-of- in subsequent years. Each station was sampled with a single the-year (age-0) winter flounder is not well known. It has been sweep of a seine, covering an area of approximately 500 m2 widely assumed that age-0 winter flounder move into deeper wa- from the shore to the shallow subtidal zone. The majority of ter within their native estuary and burrow into the mud during winter flounder caught were age 0; individuals greater than the coldest parts of the year (McCracken 1963). Most poten- 150 mm total length were excluded based on a visual inspection tial predators of winter flounder (predators larger than 30 mm) of the length-frequency distribution and previous studies (With- in the northeast leave the estuaries during the coldest months, erell and Burnett 1993). Samples from all stations were pooled thus limiting predation mortality during the winter (Collette and and divided into 5-mm size-classes. The abundance in each Klein-MacPhee 2002). Prey resources may be limiting, how- size-class was converted to individuals per 500 m2 (Delong et al. ever, which suggests that starvation could be important. It is 2001). unknown whether age-0 fish are feeding at these temperatures, The RIDEM spring trawl survey was a random stratified sur- living off energy reserves, or displaying some combination of vey that included 26 stations in Narragansett Bay. The RIDEM both strategies. Winter flounder primarily store energy as lipid vessel towed a bottom trawl at 4.63 km/h (2.5 knots) for 20 min. and utilize it during periods of starvation (Maddock and Bur- The net had a headrope length of 13.7 m and a footrope length ton 1994). If age-0 winter flounder are relying solely on stored of 18.3 m. The width was assumed to be 60% of the footrope energy, lipid levels would be expected to increase in the late (Laura Lee, RIDEM, personal communication), resulting in a fall and to become depleted through the winter. Larger individ- total area swept of 16,946 m2. The length-frequency data for uals would have a higher proportion of lipids and would most all winter flounder caught during the spring survey in each year likely exhibit higher survival. If winter flounder are meeting were separated into cohorts with NORMSEP software (Hassel- their daily requirements through feeding, lipid levels would not blad 1966; Gayanilo et al. 1995). NORMSEP assumes that the increase in the late fall but would remain relatively constant length-frequency distribution of a single cohort can be repre- through the winter; therefore, size-dependent starvation would sented by a normal distribution.
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