Estuarine and habitat-related differences in growth rates of young-of-the-year winter flounder (Pseudopleuronectes americanus) and tautog (Tautoga onitis) in three northeastern US estuaries
Introduction
Estuaries along much of the northeastern coast of the United States provide nursery habitat for a wide variety of fish species (Able and Fahay, 1998), including those of commercial and recreational importance (Chambers, 1992). Within estuaries, a number of habitats, i.e. seagrasses, marshes, marsh creeks, have been identified as important fish nursery habitat. Seagrass habitats in the southeastern US (Hoss and Thayer, 1993, Heck et al., 1997) and marsh creek habitats (marsh surface, pools and creeks) along the east coast of the United States (Weinstein, 1979, Rountree and Able, 1992, Able et al., 1996) are considered high quality, in part because they support high abundances and densities of juvenile fish (Able, 1999). Seagrasses are known to provide refuge from predation (Heck and Orth, 1980, James and Heck, 1994) and food resources (Hoss and Thayer, 1993). The relative value of macroalgae, another macrophyte habitat in areas of the northeastern US (Short and Burdick, 1996, Timmons and Price, 1997), is somewhat debatable. Macroalgae are known to displace seagrasses (Short and Burdick, 1996), change the biogeochemistry of the water and sediment (Valiela et al., 1992), induce hypoxic conditions (D’Avanzo and Kremer, 1994) and alter benthic fauna (Valiela et al., 1997, Thiel et al., 1998). In laboratory experiments, a macroalgal (Ulva lactuca) exudate was toxic to invertebrates (Johnson and Welsh, 1985) and to larval winter flounder, Pseudopleuronectes americanus, but not to juvenile winter flounder (Johnson, 1980). In another study, Ulva-dominated habitats were comparable to eelgrass (Zostera marina) based on decapod crustacean abundance and were more important than areas lacking either type of vegetation (Sogard and Able, 1991). Ulva also provided a predator refuge for juvenile tautog, Tautoga onitis (Sogard and Able, 1991) and blue crabs, Callinectes sapidus (Wilson et al., 1990a, Wilson et al., 1990b).
High quality nursery habitats are those in which growth and survival of juvenile fish is enhanced and where ultimately reproductive potential is optimized (Houde, 1987, Gibson, 1994). Consequently, rapid growth can be used as one indicator of habitat quality because it implies that (1) there is sufficient food available, (2) individuals may achieve a size refuge from some predators, and (3) individuals may be larger at the end of the summer growing season, thus enhancing the chances of overwinter survival (Sogard, 1997, Able, 1999). Based on this, several studies have employed growth rates of juvenile fish in cages to evaluate habitat quality (Sogard, 1992, Guindon and Miller, 1995, Duffy-Anderson and Able, 1999, Able et al., 1999). Relative differences in growth rates for selected species held in cages in southern New Jersey estuarine habitats were generally maintained across experiments suggesting a consistency in foraging value (Sogard, 1992). In the heavily impacted Hudson River estuary, growth of caged young-of-the-year winter flounder and tautog has been used to assess the quality of man-made habitats (e.g. piers and pile fields). These studies have shown negative somatic growth under piers, which may be related to factors such as available light and food (Duffy-Anderson and Able, 1999, Able et al., 1999).
While previous research has concentrated on habitat comparisons within a single estuary, (Sogard, 1992, Guindon and Miller, 1995, Duffy-Anderson and Able, 1999, Able et al., 1999), our study compares growth of young-of-the-year winter flounder and tautog among different habitats within an estuary and among similar habitats in three different estuaries in the southern New England–New York Bight coastal watershed (Fig. 1). Winter flounder adults occur in estuarine and continental shelf waters in the northwest Atlantic (Bigelow and Schroeder, 1953), with the largest fisheries from Canada through southern New England. Young-of-the year winter flounder are known to occur across a variety of estuarine habitats (Sogard and Able, 1991, Able and Fahay, 1998). Tautog adults occur primarily on the inner continental shelf and the polyhaline portions of estuaries from Nova Scotia to South Carolina but is most abundant from Cape Cod to Delaware Bay (Bigelow and Schroeder, 1953). Young-of-the-year tautog occupies a variety of structured habitats and may be highly localized (Olla et al., 1979, Dixon, 1994, Able and Fahay, 1998). Both species, which are common in the study areas, have been shown to exhibit habitat related differences in growth in previous studies (Sogard, 1992, Duffy-Anderson and Able, 1999, Able et al., 1999). The nominal habitats studied, chosen for their emergent vegetation or proximity to vegetation and their common occurrence in each of the three estuaries, were evaluated for their ability to predict habitat growth both temporally and spatially. This study was part of a larger effort designed to examine several parameters that may be indicators of habitat quality within and among estuaries (Goldberg et al., 1993, Goldberg et al., 2000; Litvin et al., unpubl. data).
Section snippets
Study sites
Three estuaries (all within 250 km) were selected for comparison: (1) Great Bay–Little Egg Harbor in southern New Jersey, (2) the Navesink River in northern New Jersey, and (3) the Hammonasset River on the Connecticut coast of Long Island Sound (Fig. 1). Great Bay–Little Egg Harbor is part of the Jacques Cousteau Estuarine Research Reserve and is located within the Mullica River watershed and drains through Little Egg Inlet into the Atlantic Ocean. Compared with the other estuaries it is
Physical conditions
The physical parameters measured were similar in the three estuarine study sites. Temperature ranges in the three estuaries exhibited seasonal fluctuations typical of Mid-Atlantic and southern New England estuaries (Table 2, Table 3). Temperature was lower in early June, at the start of the winter flounder experiments, peaking in mid-summer at the end of the winter flounder and at the start of the tautog experiments before beginning to decline. Despite slight differences in the start date for
Artifacts of caging
Cages are a valuable tool for comparing the relative growth of small juvenile fish in different habitats but are also known to have certain limitations (Peterson and Black, 1994, Able, 1999). Cages confine fish to a small space, preventing movements to surrounding areas i.e. for food searching or to avoid areas of poor water quality. In general, this may be less of an issue for the target species in this study. Juvenile winter flounder are reported to make relatively limited movements (<100 m)
Acknowledgements
The authors thank Richard Trout of Rutgers University for advice on the statistical model used in the analysis and Alan Stoner, Tom Minello and Sam Wainright for reading the manuscript and providing helpful comment. Particular thanks go to John Manderson for all his insights and discussion. Jon Finn, Steve Fromm, Dave Packer and a host of others assisted in field data collection. Chris Zetlin assisted in the creation of Fig. 1. This research was supported in part by NOAA’s Coastal Ocean Program
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