Winter Flounder Movements in the Southern Gulf of Maine. ICES CM
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INTERNATIONAL COUNCIL FOR ICES CM 2009/B:16 THE EXPLORATION OF THE SEA Beyond Geolocation: Inferring and explaining the behavior of tagged fish Winter Flounder Movements in the Southern Gulf of Maine Gregory R. DeCelles and Steven X. Cadrin NOAA/Umass Cooperative Marine Education and Research Program School for Marine Science and Technology A two-year acoustic tagging experiment was conducted in the Plymouth Bay estuary to examine seasonal movement patterns of adult winter flounder. During the first year of the study, 47 individuals were fitted with acoustic transmitters and tracked using an array of 15 Vemco receivers. During the second year of the study, the passive array was expanded from 15 to 30 receivers and an additional 25 flounder were tagged within the array, allowing more direct tests of alternative hypotheses about spawning migrations. Telemetry data suggests that seasonal movements within the estuary were strongly associated with water temperature. A proportion of tagged flounder migrated into the estuary in the spring as water temperature increased, while others remained in coastal waters. Of those individuals that entered the estuary in the spring, the majority emigrated to coastal waters in June, when water temperatures in the estuary exceeded 15oC. A small portion (n=5) of tagged fish were detected reentering the estuary in autumn, as water temperatures decreased to less than 15 oC. Tagged flounder exhibited spatially and temporally diverse movements, and it appears that sympatric, contingent spawning groups of winter flounder may occur within this relatively small geographic area. Keywords: winter flounder, Pseudopleuronectes americanus, acoustic telemetry, contingent Contact Author: Greg DeCelles, NOAA/Umass CMER Program, School for Marine Science and Technology, 200 Mill Rd. Suite 325, Fairhaven, MA 02719 U.S.A. [tel: +1 508 910 6393 email: [email protected]] 1 Introduction Knowledge of seasonal changes in the distribution and behavior of individuals within populations is a crucial step in understanding population structure (Metcalfe, 2006). Acoustic telemetry offers researchers the ability to better understand fish behavior and migration, and ultimately allows for contingent groups within populations to be recognized (Secor, 1999). Acoustic tags also allow unprecedented opportunities to understand the amount of connectivity between estuarine and coastal populations (Able, 2005). Another advantage of this technology is that data acquisition is not dependent upon the recapture of tagged individuals by the fishery. In this study we sought to understand the migratory patterns of winter flounder in the southern Gulf of Maine using acoustic telemetry. The miniaturization of acoustic transmitters has allowed for the study of flatfish biology and migration. Telemetry has been successfully used to study many flatfish species including; plaice (Greer Walker et al., 1978), summer flounder (Sackett et al., 2007), and sole (Greer Walker et al., 1980). More specifically, this technology has been used to investigate winter flounder spawning habitat (Pereira et al., 1994), movement patterns (John Manderson, NOAA, personal communication) and habitat use by juveniles (Elizabeth Fairchild, University of New Hampshire, personal communication). In the coastal waters of the U.S., winter flounder are managed as three stocks; Gulf of Maine, Southern New England/Mid-Atlantic (SNE/MA) and Georges Bank. Individuals in the Gulf of Maine and SNE/MA stocks are resident in coastal and estuarine waters. Flounder in the Georges Bank stock remain offshore throughout their entire life cycle. The seasonal distribution of winter flounder in the SNE/MA stock have been studied using a series of mark-recapture experiments (e.g. Perlmutter, 1947; Saila, 1961; Powell, 1989), and this stock is considered to be dependent upon estuarine habitats (Able and Fahay, 1998). There has been far less research on winter flounder in the Gulf of Maine, relative to the SNE/MA stock. As a result, there is considerable uncertainty regarding the movement and spawning patterns of winter flounder in the Gulf of Maine stock. Traditionally, winter flounder in the Gulf of Maine were presumed to spawn exclusively in estuarine habitats (Bigelow and Schroeder, 1953). However, this paradigm may have arisen due to observations in the SNE/MA stock area, or due to the relative ease of sampling flounder in estuaries, rather than coastal habitats. More recent data suggests that some flounder in the Gulf of Maine are using coastal waters as spawning grounds, rather than estuaries. Due to the demersal and adhesive nature of winter flounder eggs (Pereira et al., 1999), dispersal of eggs is limited, and ichthyoplankton tows capturing their eggs can be used to infer spawning locations. Winter flounder eggs have been sampled in coastal locations of Massachusetts Bay and Cape Cod Bay (Lux and Kelly, 1982). The ages of eggs sampled in Plymouth Bay, Massachusetts suggested that a substantial proportion were produced by coastal spawning events (Chau and Pearce, 1977). Howe and Coates (1975) hypothesized that coastal spawning groups may exist based upon their observations of adult fish captured in coastal and offshore waters during the spawning season. In addition, the Massachusetts Division of Marine Fisheries Industry-Based Survey commonly captured adult winter flounder in coastal waters in March and April in 2 2005, 2005 and 2007 (Hoffman et al., 2007; Figure 1). The presence of adult fish in coastal waters in March and April is significant, because these are considered the peak spawning months for winter flounder in the Gulf of Maine (DeCelles and Cadrin, 2007). Contingents are defined as groups of fish that exhibit divergent migratory patterns or habitat use (Secor, 1999). In this paper, we refer to contingents as adults that spawn in either coastal or estuarine waters. The presence of contingent structure within the Gulf of Maine stock will have important ramifications for population structure and the identification of Essential Fish Habitat. Spatial structure may provide fish populations with enhanced stability and resilience due to variable survival of early life history stages in different habitats (Kerr et al., 2009). For winter flounder, which spawn demersal eggs, the conditions encountered by the early life history stages are largely determined by the locations where adults spawn. The progeny of estuarine and coastal spawning flounder will likely exhibit different larval transport trajectories and survival. In this study we tracked the movements of adult winter flounder using acoustic transmitters and a moored array of acoustic receivers. Plymouth Bay and the adjacent Plymouth Harbor, Duxbury Bay, Kingston Bay estuary (hereafter Plymouth estuary) in the southern Gulf of Maine were chosen as the study site. Winter flounder is a dominant member of the groundfish community in the Plymouth estuary (Lawton et al., 1984), and the area serves as an important nursery habitat for juvenile winter flounder. The Plymouth estuary is a known spawning area for winter flounder (Normandeau Associates, 2001) with peak spawning typically observed in March and April. In addition, the coastal waters of Plymouth Bay have been identified as an area where coastal spawning likely occurs (Chau and Pearce, 1977; Marine Research Inc., 1986). Furthermore, the geography of the Plymouth estuary allowed us to achieve adequate sampling coverage using relatively few acoustic receivers. The goal of this study was to characterize the seasonal movement patterns of adult winter flounder in Plymouth Bay and the Plymouth estuary using acoustic telemetry. In order to investigate the hypothesis of coastal spawning, we sought to identify probable spawning habitats using the observed movements of tagged individuals during the spawning season. Gaining a better understanding of the spawning habits of winter flounder in the Gulf of Maine area may improve management of this species, aid in conservation efforts and allow further insight into the population and stock structure of this species. This information is critical in light of Essential Fish Habitat requirements. Materials and Methods Study Site The Plymouth estuary and the adjacent Plymouth Bay are located in the southern portion of the Gulf of Maine (Figure 2). The Plymouth estuary is bordered on its seaward side by two barrier beaches. Tidal exchange between the Plymouth Estuary and Plymouth Bay occurs through a 2,020m opening between Saquish Head and the northern extremity of Plymouth Beach. Approximately 66% of the water in the Plymouth estuary is replaced during each tidal cycle (Iwanowicz et al., 1974). Extensive sand and mud flats are exposed at mean low water when the surface area of Plymouth estuary is reduced 3 from 10,057 acres to 5,465acres (Iwanowicz et al., 1974). Plymouth Bay is bordered on its eastern extent by Cape Cod Bay. The average depth within Plymouth estuary is 3.3m at mean high water and 2.1m feet at mean low water, and the average tidal amplitude is 2.9m (Iwanowicz et al., 1974). A deeper channel is present between Saquish Head and Plymouth Beach, where depths reach nearly 26m. The major sources of freshwater input to the Plymouth Estuary are; Back River, Bluefish River, Jones River, and Eel River. Acoustic Telemetry In total, 72 adult winter flounder were fitted externally with acoustic transmitters (Model V92L, 69kHz, 9mm X 21mm, Vemco Ltd.) between 2007 and 2009 (Figure 3). The transmitters had an expected battery life of 384 days (Matthew Holland, Vemco Ltd., personal communication), allowing each fish to be tracked for approximately one year. Prior to the first year of tagging in 2007, we planned to release 47 tags in two discrete batches. The first group of tags (n=24) were to be deployed in the coastal waters of Plymouth Bay during the winter of 2007, prior to the spawning season. We planned to release the second batch of tags (n=23) on spawning fish inside the Plymouth estuary during the months of March and April, 2008. In November of 2007, 24 pre-spawning flounder were tagged in Plymouth Bay, approximately 5km from the mouth of the Plymouth estuary (Figure 4).