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Physical Factors Influencing the Distribution of a Top Predator in a Subtropical Oligotrophic Estuary

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Physical Factors Influencing the Distribution of a Top Predator in a Subtropical Oligotrophic Estuary Limnol. Oceanogr., 54(2), 2009, 472–482 E 2009, by the American Society of Limnology and Oceanography, Inc. Physical factors influencing the distribution of a top predator in a subtropical oligotrophic estuary Michael R. Heithaus1 and Bryan K. Delius Department of Biological Sciences, Florida International University, 3000 NE 151 St., North Miami, Florida 33181 Aaron J. Wirsing Department of Biological Sciences, Florida International University, 3000 NE 151 St., North Miami, Florida 33181; College of Forest Resources, University of Washington, Box 3052100, Seattle, Washington 98195 Meagan M. Dunphy-Daly Department of Biological Sciences, Florida International University, 3000 NE 151 St., North Miami, Florida 33181 Abstract We used longline fishing to determine the effects of distance from the ocean, season, and short-term variation in abiotic conditions on the abundance of juvenile bull sharks (Carcharhinus leucas) in an estuary of the Florida Everglades, U.S.A. Logistic regression revealed that young-of-the-year sharks were concentrated at a protected site 20 km upstream and were present in greater abundance when dissolved oxygen (DO) levels were high. For older juvenile sharks (age 1+), DO levels had the greatest influence on catch probabilities followed by distance from the ocean; they were most likely to be caught at sites with .3.5 mg L21 DO and on the main branch of the river 20 km upstream. Salinity had a relatively small effect on catch rates and there were no seasonal shifts in shark distribution. Our results highlight the importance of considering DO as a possible driver of top predator distributions in estuaries, even in the absence of hypoxia. In Everglades estuaries hydrological drivers that affect DO levels (e.g., groundwater discharge, modification of primary productivity through nutrient fluxes) will be important in determining shark distributions, and the effects of planned ecosystem restoration efforts on bull sharks will not simply be mediated by changing salinity regimes and the location of the oligohaline zone. More generally, variation in DO levels could structure the nature and spatiotemporal pattern of top predator effects in the coastal Everglades, and other tropical and subtropical estuaries, because of interspecific variation in reliance on DO within the top predator guild. Estuarine systems are vulnerable to human effects Studies of the movements and distribution of predators including eutrophication, habitat alteration and destruc- in estuarine systems have focused primarily on the most tion, degradation of water quality, increased hypoxia, and obvious changes in physical factors—salinity and water changes to freshwater inputs (Kennish 2002). In order to temperature. More recently, however, the role of chronic predict the consequences of anthropogenic modifications and/or acute hypoxic events that lead to mass movements for estuaries and to assess the likely effects of management or mortality of individuals have received greater attention alternatives, it is necessary to understand how variation in (Eby and Crowder 2002; Baird et al. 2004; Altieri and physical factors affects the distribution and abundance of Witman 2006). Most of these studies on the role of organisms within them. Of particular interest is how large dissolved oxygen levels in driving consumer distributions predators will respond to variation in physical conditions and mortality patterns have been conducted in temperate because these species can help to structure communities areas where enhanced primary productivity due to eutro- (Heithaus et al. 2008) and are often of commercial or phication combined with stratification of the water column recreational importance. can lead to hypoxia that can persist for considerable periods. For example, in the Neuse River Estuary of North Carolina, U.S.A., small mobile consumers (e.g., crabs, 1 Corresponding author ([email protected]). small teleosts) vary considerably in their responses to Acknowledgments chronic and episodic hypoxia and prolonged hypoxic Funding for this project was provided by National Science events may lead to enhanced interspecific competition or Foundation Grants DBI0620409 and DEB9910514 and Florida predator–prey interactions in shallow oxygenated waters International University’s Marine Sciences Program. We thank (Eby and Crowder 2002; Bell and Eggleston 2005) and the many volunteers who assisted with shark fishing, especially ultimately to a reduction in energy transfer to higher Derek Burkholder and Richard Chang. Thanks also to Joel trophic levels (Baird et al. 2004). However, species that can Trexler and Dan Childers who provided an introduction to the Everglades estuaries, funding, and logistical support for this tolerate low levels of dissolved oxygen may experience project. Research was conducted under Everglades National Park reduced predation risk from more oxygen-sensitive preda- permits EVER-2007-SCI-0025 and EVER-2005-SCI-0030 and tors (Eby and Crowder 2002; Altieri and Witman 2006). Florida International University Institutional Animal Care and Less appreciated is the potential for variation in dissolved Use Committee authorizations. oxygen levels over short temporal and small spatial scales 472 Abiotic drivers of bull shark abundance 473 to structure the distribution of large consumers, especially the three sites farthest downstream (mouth, SRS4, and in subtropical and tropical estuaries that are less prone to SRS5) occurred near long-term environmental sampling the seasonal hypoxia characteristic of temperate estuaries. platforms maintained by the Florida Coastal Everglades The subtropical estuaries of the coastal Everglades, long-term ecological research program (www.fcelter.fiu. Florida, U.S.A., which constitute the largest mangrove edu). Sampling near the mouth of the estuary and at SRS5 ecosystem in the Western Hemisphere, are human-dominat- occurred in broad (,50–100 m wide) mangrove-lined ed systems that exhibit marked temporal and spatial channels with water depths of ,1.2–3.0 m. SRS4 and variation in physical factors including dissolved oxygen Otter Creek were located at upstream portions of a broad levels. Most obviously, there is distinct seasonal variation in open area, Tarpon Bay (Fig. 1). Sampling in Tarpon Bay precipitation and freshwater delivery to these estuaries. occurred where water depths ranged between 1.5 m and Although the timing and magnitude of wet and dry seasons 2.5 m in both channels (Otter Creek and SRS4) and more within the Everglades vary interannually, salinities tend to be open waters where several branches of the slough converge lowest from December to May and highest from June to (SRS4). Sampling in upstream sites occurred in channels November (Childers et al. 2006). The overall delivery of similar to those of SRS5 and the mouth, but these channels freshwater to the estuaries, however, is low compared to often were narrower (,20–50-m wide) and rarely exceeded historical levels due to the construction of water manage- 2 m depth, but were at least 1.2 m deep. ment canals and levees over the past 100 yr and the diversion of much of the freshwater that historically flowed to the Field methods—From May 2005 to February 2008 we Everglades (Davis et al. 2005). This modification to the sampled bull sharks using a ,500-m longline fitted with 42 system’s hydrology has almost certainly had major effects on 6 6.7 SE gangions. Each gangion consisted of a heavy-duty mobile predators in the coastal estuaries of the Everglades, clip with swivel attached to ,2 m of 400-kg monofilament as will the planned attempts to restore water flow to a more line. A second swivel was positioned in the middle of the natural level (Comprehensive Everglades Restoration Plan line, which terminated in a 12/0–15/0 Mustad tuna circle [CERP] 1999). Therefore, studies of the factors influencing hook with offset point that was baited with mullet (Mugil the distribution and abundance of top estuarine predators sp.). Pilot fishing efforts found no effect of hook size on size are important for predicting how ecosystem engineering will of sharks captured or shark catch probabilities. Lines were influence their use of the Everglades estuaries. set between dawn and dusk and were allowed to soak for Juvenile bull sharks (Carcharhinus leucas) are a euryhaline 1 h after the last gangion was deployed before being upper trophic-level predator in many estuaries of the retrieved. After the longline was set, we recorded environ- subtropics including those of the Everglades (Pillans and mental conditions including water temperature, salinity, Franklin 2004; Pillans et al. 2005; Wiley and Simpfendorfer and dissolved oxygen using a YSI85 at depth of 1–1.5 m. 2007). Thus, studies of the factors influencing the distribu- Upon retrieval, we recorded the presence or absence of tion and abundance of bull sharks are potentially important bait at every hook. Sharks were brought alongside the boat, for understanding the dynamics of Everglades estuaries and sexed based on the presence or absence of claspers, predicting the consequences of proposed ecosystem man- measured (precaudal, fork, total, and stretched total agement strategies. The objective of this study was to lengths) to the nearest centimeter, tagged using a plastic determine the factors influencing the distribution and roto tag affixed through the first dorsal fin, freed from the abundance of juvenile bull sharks in the estuary of the hook, and then released. Shark River Slough of Everglades National Park, and in particular to evaluate the influence of temporal and spatial Data analysis—Because shark responses to environmen- variation in abiotic factors (water temperature, salinity, tal conditions may vary with age (e.g., Simpfendorfer et al. dissolved oxygen, distance from the Gulf of Mexico). 2005) we analyzed data for individuals ,1 yr old (young of the year [YOY]) and sharks .1 yr old (juveniles) separate- Methods ly. Consistent with previous studies (Simpfendorfer et al. 2005; Heupel and Simpfendorfer 2008), we considered all Study site—Juvenile bull sharks are one of the largest individuals ,96 cm stretched total length (STL) as young bodied predators in the Shark River Estuary (Loftus 2000; of the year.
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