Migrations and Movements of Atlantic Tarpon Revealed by Two Decades of Satellite Tagging

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Migrations and movements of Atlantic tarpon revealed by two decades of satellite tagging

Luo, Jiangang; Ault, Jerald Stephen; Ungar, Bruce T; et.al. https://scholarship.miami.edu/discovery/delivery/01UOML_INST:ResearchRepository/12378179450002976?l#13378179440002976

Luo, J., Ault, J. S., Ungar, B. T., Smith, S. G., Larkin, M., Davidson, T. N., Bryan, D., Farmer, N. A., Holt, S. A., Alford, A. S., Adams, A. J., Humston, R., Marton, A. S., Mangum, D., Kleppinger, R., Requejo, A., & Robertson, J. (2020). Migrations and movements of Atlantic tarpon revealed by two decades of satellite tagging. Fish and Fisheries (Oxford, England), 21(2), 290–318. https://doi.org/10.1111/faf.12430

Published Version: https://doi.org/10.1111/faf.12430

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Received: 1 October 2019 | Revised: 12 November 2019 | Accepted: 18 November 2019 DOI: 10.1111/faf.12430

ORIGINAL ARTICLE

Migrations and movements of Atlantic tarpon revealed by two decades of satellite tagging

1Department of Marine Ecosystems and Society, Rosenstiel School of Marine and Abstract Atmospheric Science, University of Miami, Understanding large-scale migratory behaviours, local movement patterns and Miami, Florida population connectivity are critical to determining the natural processes and an- 2National Marine Fisheries Service, St. Petersburg, Florida thropogenic stressors that influence population dynamics and for developing effec- 3Bonefish Tarpon Trust, Miami, Florida tive conservation plans. Atlantic tarpon occur over a broad geographic range in the 4 Pacific States Marine Fisheries Commission, Atlantic Ocean where they support valuable subsistence, commercial and recrea- Seattle, Washington tional fisheries. From 2001 through 2018, we deployed 292 satellite telemetry tags 5Port Aransas Marine Laboratory, University of Texas, Port Aransas, Texas on Atlantic tarpon in coastal waters off three continents to document: (a) seasonal 6 International Tarpon Conservation migrations and regional population connectivity; (b) freshwater and estuarine habitat Association, Houston, Texas utilization; (c) spawning locations; and (d) shark predation across the south-eastern 7Washington Lee University, Lexington, Virginia United States, Gulf of Mexico and northern Caribbean Sea. These results showed 8Fieldworkers Club, Three Oaks, Michigan that some mature tarpon make long seasonal migrations over thousands of kilome- 9 Robertson Foundation, New York, New tres crossing state and national jurisdictional borders. Others showed more local York movements and habitat use. The tag data also revealed potential spawning locations Correspondence consistent with those inferred in other studies from observations of early life stage Jerald S. Ault, Rosenstiel School of Marine and Atmospheric Science, University of tarpon leptocephalus larvae. Our analyses indicated that shark predation mortality Miami, 4600 Rickenbacker Causeway, on released tarpon is higher than previously estimated, especially at ocean passes, Miami, FL 33149, USA. Email: [email protected] river mouths and inlets to bays. To date, there has been no formal stock assessment of Atlantic tarpon, and regional fishery management plans do not exist. Our find- Funding information Division of Earth Sciences, Grant/Award ings will provide critical input to these important efforts and assist the multinational Number: EAR-1204752 community in the development of a stock-wide management information system to support informed decision-making for sustaining Atlantic tarpon fisheries.

KEYWORDS population connectivity, shark predation, spawning habitats, sportfish

1 | INTRODUCTION

1 INTRODUCTION 291 Atlantic tarpon (Megalops atlanticus, Megalopidae) has long been one 2 MATERIALS AND METHODS 291 of the most sought-after inshore marine game fishes with a long history in the United States as the focus of an important recreational 2.1 Tagging 291 fishery (Aflalo, 1907; Ault & Luo, 2013; Babcock, 1921a; Dimock & 2.2 Data analyses 292 Dimock, 1912; Holder, 1903; Pinckney, 1888; Spotte, 2016), includ- 3 RESULTS 293 ing participation by US Presidents, as recorded in many publications 3.1 Tag deployments 293 (Mares, 1999; Mill, 2010; Stilwell, 2011). The US fishery today has an 3.2 Seasonal migrations and distributions 294 annual economic impact of more than $6 billion, providing thousands 3.3 Estuarine and riverine utilization 295 of jobs (Ault, 2008; Steinback, Gentner, & Castle, 2004). Sport fish- 3.4 Spawning habitats 296 ing for tarpon is also very popular in other countries, where numer- 3.5 Predation by sharks 297 ous records have been recorded for large fish caught in Venezuela, 4 DISCUSSION 297 Sierra Leone, Guinea-Bissau, Mexico, Brazil, and Cuba (IGFA, 2018). 4.1 Tag deployments 297 While tarpon fishing in the United States is predominately catch- 4.2 Seasonal migrations and distributions 298 and-release, artisanal subsistence and commercial harvests of tar- 4.3 Estuarine and riverine utilization 299 pon occur in many countries outside the United States (Adams et al., 2014; Anyanwu & Kusemiju, 2008; Anyanwu et al., 2009). 4.4 Spawning habitats 299 Despite the history and importance of recreational catch-and- 4.5 Predation by sharks 300 release fishing for tarpon in the United States, the paucity of data 5 IMPLICATIONS FOR FISHERY MANAGEMENT 303 and models on population dynamics and migrations limits opportuni- ACKNOWLEDGEMENTS 304 ties for developing an effective regional management plan to sustain REFERENCES 305 this valuable resource (Ault et al., 2008). Atlantic tarpon are now SUPPORTING INFORMATION 308 threatened throughout their range by recreational fishing release APPENDIX 309 mortality, directed commercial harvests, intensive harvesting of key prey species and degradation of habitats for both predator and prey (Ault, 2008; Boesch et al., 1994; Polidoro et al., 2010; Waycott et al., 2009). A recent IUCN assessment classified Atlantic tarpon as Mississippi, Louisiana, Texas, North Carolina, South Carolina and vulnerable (Adams et al., 2014). Georgia), Mexico, Belize, Nicaragua and Trinidad (Table A1, Figure 1). A principal impediment to sustaining highly migratory marine The total of 292 deployed tags included the following: 199 Pop-up species is the lack of scientific understanding of the locations, at- Archival Transmitting (PAT) and 88 Smart Position and Temperature tributes and migratory connectivity of important habitats (Runge, (SPOT) tags from Wildlife Computers, Inc. (wildlifecomputers.com); Martin, Possingham, Willis, & Fuller, 2014; Webster, Marra, Haig, 1 PAT tag from Lotek Wireless, Inc. (lotek.com); and 4 PAT tags from Bensch, & Holmes, 2002). Without such knowledge, regional man- Microwave Telemetry, Inc. (microwavetelemetry.com). Each tag agement strategies are difficult to conceptualize, develop and imple- bore a fixed label containing our international toll-free phone num- ment (Hansson & Akesson, 2014). To improve this understanding, we ber, email address and a request for return of the tag. employed satellite telemetry technologies in an intensive multiyear Tarpon were captured with standard hook-and-line gears on (2001–2018) study of Atlantic tarpon over three continents with chartered recreational fishing vessels, using the heaviest tackle fea- focus on the coastal waters of the south-eastern United States, Gulf sible to minimize fight time. During the tagging process, tarpon at of Mexico and northern Caribbean Sea. Our specific objectives were boat side were kept completely in the water to prevent injury by to reveal (a) seasonal migrations and regional population connectiv- either: (a) guiding the fish into a specially designed sling or (b) using ity; (b) utilization of freshwater and estuarine habitats; (c) potential a lock-jaw gaff and tailer. Once secure, fork length (FL) and girth (G) spawning locations; and (d) predation mortality by sharks. were measured in centimetre. Body weight (kg) was computed with the algorithm of Ault and Luo (2013). Satellite tags were secured to tarpon via a tether (a 30-cm stainless steel wire encased in a med- 2 | MATERIALS AND METHODS ical grade Tygon tube) that was attached to a titanium anchor dart (Figure 1). Anchor darts were coated with an antibiotic cream. Two 2.1 | Tagging scales were removed from anterior of the first dorsal fin about one third of the distance to the head. The dart was then inserted approx- Electronic satellite telemetry tags from several manufacturers were imately 8 cm deep at a 45° angle towards the head into the tissue deployed on Atlantic tarpon from 2001 to 2018 in coastal waters where the scales had been removed. of the western central Atlantic Ocean, Gulf of Mexico (GOM) and Pop-up Archival Transmitting tags deployed on tarpon were Caribbean Sea including the United States (i.e. Florida, Alabama, pre-programmed to collect high-resolution data at either 1-, 292 | LUO et al.

FIGURE 1 Bathymetric map of the western central Atlantic, Gulf of Mexico and Caribbean Sea overlain with satellite tag deployment (red dots) and pop-off detachment locations (white crosses). Depths from 0 to 6,000 m are denoted by the colour bar. The photo inset shows PAT tag set-up and anchor dart placement location on tarpon

10- or 30-s intervals. Environmental variables collected were as 2.2 | Data analyses follows: water depth (via pressure); ambient water temperature; light level (for determining location of the tagged fish); salinity For PAT tags, light-level geolocation data were processed using (estimated from the wet/dry sensor (Luo & Ault, 2012)); and clock WC-DAP and WC-GPE2 software (Wildlife Computers), and a time, for deployment periods that ranged from 4 to 8 months. Tags sea surface temperature (SST) corrected Kalman filter (KF-SST; were also pre-programmed to release from the fish at specified Lam, Nielsen, & Sibert, 2008; Nielsen, Bigelow, Musyl, & Sibert, dates and times. Upon release, tags floated to the ocean surface 2006). A custom bathymetry filter was then applied to relo- where they transmitted their stored summary data to the orbiting cate any points that were on land or in shallow water, based on Argos satellite network, which were transmitted to us. Tag con- 2 × 2 min gridded ETOPO2 bathymetry data (Anon., 2006) and tact transmission time with orbiting satellites was restricted by the daily maximum depth recorded by the tag. For points where the tag's limited battery capacity and the quantity of data that maximum daily depth exceeded the bathymetric depth, all grid could actually be transmitted at any one time. This limitation was cells were selected along the longitude where bathymetric depth particularly evident with large datasets from tags that recorded was greater than the daily maximum depth within ±1 degree at short intervals for long monitoring periods. To contend with of latitude of the previous day's location. A final location was this problem, transmitted data were averaged and summarized at then assigned to a single cell selected randomly from that group 3-hr intervals into 14 user-defined temperature (from 20 to 32 at (Hoolihan & Luo, 2007). 1°C interval) and depth (from 0 to 50 at 5 m interval, then 50–75, Tag SST and ocean heat content (OHC) were estimated based 75–100 and >100 m) bins. This provided a constrained represen- on 3-hr binning of summarized profiles of depth and temperature tation of the animal's actual activities and minimized the potential (PDT), and depth and temperature histograms of all PAT tags. Daily loss of temporal blocks of data when transmission events failed or multiscale ultra-high-resolution (MUR, 0.01°) SST data were used to were otherwise corrupted. calculate mean SST of April and November 2008 (https://podaac.jpl. We were able to physically recover some tags by either travel- nasa.gov/) to demonstrate the position of thermal contours during ling to pop-up locations with a handheld Argos receiver that aided periods of known seasonal migrations. in finding tags, or through recovered tags returned to us by citizen Sea surface temperature corrected Kalman filter tracks were scientists, enabling us to obtain the entire dataset. SPOT tags did further refined with the fine-scale geolocation method of Luo et al. not archive data, but rather transmitted radio signals to the satellite (2015) which incorporated OHC estimated from Hybrid Coordinate network when a fish surfaced, exposing the wet-dry sensor to air. Ocean Model (HYCOM) and a genetic algorithm (GA-OHC). The con- The Argos System (www.argos-system.org) determined their geolo- cept of the OHC method, originally developed for use in hurricane cations by Doppler-shift calculations when two or more messages intensity forecasts, integrates the thermal energy from the depths were received during a limited time window of satellites passing associated with the 26°C isotherm (D26) to the ocean surface fol- overhead. lowing the equation of Palmen (1948). LUO et al. | 293

defined the core habitat utilization area (CHUA) as the areas of top 0 = c T − ◦ z 50% UD (da Anadón, D'Agrosa, Gondor, & Gerber, 2011; Graham OHC p z 26 d , (1) D26 et al., 2016). These metrics were calculated according to the equa- tions provided by Worton (1995) and plotted using Interactive Data where cp is the specific heat constant of water, ρ is seawater density, Languages (www.harris.com) software. and Tz is water temperature at depth z (Luo et al., 2015). In fisheries Rate of movement (ROM) was calculated at 12-hr intervals to applications, the 26°C isotherm can be modified to any water tem- determine whether a tarpon was migrating, or simply at residence. perature range, specifically by the lower bound of an animal's thermal To estimate the residence period of each tarpon track where days range, such as the 20°C isotherm that we used in this study for tarpon at liberty >10 days, we examined three ROM criteria (ROM < 5, 7, since 99% of temperatures recorded by the tags were above this iso- 10 km/day) for >3 consecutive days. To examine variability of mi- therm (Luo & Ault, 2012). gratory behaviour among individual tarpon, the proportion of time To improve location accuracy of SPOT tags, all Doppler-derived in residence was calculated as the residence period divided by total data were processed with the Argos system's Kalman filtering (KF) days at liberty. Distance from the tagging location was estimated method. Argos provided the following radius of error for each KF- using great-circle distance for each 12-hr location and these were derived location class (LC): LC 3, <250 m; LC 2, <500 m; and LC 1, plotted as a function of days at liberty. <1,000 m. Argos states that the median error for LC classes 0, A and Shark predation on tarpon was determined by two methods: (a) B ranges from 1 to 3 km (Bernard & Belbeoch, 2010). Class Z indi- from tooth marks observed on recovered PAT and SPOT tags and cates that the location process failed and estimates of position are (b) from data recorded by recovered PAT tags that showed reduced highly inaccurate. All transmitted locations were filtered to remove light levels with fluctuations of depth and temperature, similar to the positions with LC Z, those on land and those exceeding a speed of environment inside of sharks (Cosgrove, Arregui, Arrizabalaga, Goni, 2 m/s following Weng et al. (2005). & Neilson, 2015; Kerstetter, Polovina, & Graves, 2004). Data from To make SPOT location data compatible with the GA-OHC re- recovered PAT tags were also used to determine possible spawning fined PAT location data, and to further meet the requirements for activity based on deep dive behaviours around new and full moon kernel density metric estimates (De Solla, Bonduriansky, & Brooks, periods. 1999; Kie, 2013; Worton, 1989), the irregular sampling intervals of Tarpon estuarine and riverine habitat utilization were determined SPOT-derived raw data were standardized to a 3-hr interval. We for both SPOT tags and recovered PAT tags with archived wet/dry employed the piecewise Bézier interpolation method of Tremblay et data and track duration >10 days. For SPOT tags, this was done by al. (2006), but modified with the Lars Jenson algorithm (http://ljens counting the period when the locations occurred in inland waters. en.com/bezier/) to eliminate unnatural loops in the tracks. To mini- For PAT tags where the geolocation method was limited to ocean mize uncertainty, we did not interpolate track sections with tempo- waters, salinity was estimated from wet/dry sensor data using the ral data gaps that exceeded three days following Weng et al. (2008). Luo, Ault, Larkin, & Barbieri (2008) algorithm. We defined “rivers” as We employed utilization distributions (UD) to quantify the core inland waters that included lagoons, rivers and lakes where salinity regions of occupancy within an animal's home range or activity values were <25 PSU. The proportion of time in inland waters was space (Worton, 1995). UD were calculated for all tarpon using 12-hr calculated as the number of days in rivers divided by the number of interval (mid-night and noon) location data as the grouped species’ days at liberty. This proportion was then compared by tag types, re- habitat utilization for dry (Jan–Apr) and wet (May–Dec) seasons gions and seasons using t tests carried out with R statistical software. (Domeier & Nasby-Lucas, 2008; Hammerschlag, Luo, Luo, Irschick, & Ault, 2012; Weng et al., 2008). To avoid possible biases from short tracks, we excluded tarpon that were tracked for <10 days (Hoolihan, 3 | RESULTS Luo, Abascal, et al., 2011). To mitigate individual tarpon from biasing the results, we created a 0.5° × 0.5° spatial grid and counted the total 3.1 | Tag deployments number of 12-hr locations from all tarpon within each cell for each season and then multiplied that value by the number of unique tar- Detailed information on the 292 satellite tags (204 PAT and 88 pon within the cell. The resulting grid was used as a weighting factor SPOT) deployed from 2001 to 2018 on tarpon is presented in Table where the UD were weighted more heavily towards areas that were A1; tagging efforts varied annually (Figure 2a). The body weight frequented by many individuals. To achieve the final UD for each of tarpon tagged ranged from 14.9 to 99 kg, with a mode of 50 kg season, we set all portions that overlapped land to 0 and normalized (Figure 2b), with the majority (96.5%) ≥30 kg. Of the 204 total PAT all values so that the summation of all cells equalled 1 (Vaudo, Byrne, tags deployed, 19% (39) did not transmit any data to the Argos Wetherbee, Harvey, & Shivji, 2017). System (black bar in first column of Figure 2c), while 23 PAT and 33 Utilization distributions values are cumulated from the highest SPOT tags remained on fish for ≤10 days. These relatively short tag to lowest density areas to create kernel density contours. Thus, the deployments generally resulted from either tether failure, post-re - 25% contours represent areas of the top 25% highest densities; 50% lease mortality from failed resuscitation after tagging or predation contours represent areas of the top 50% highest densities, etc. We by sharks. There were 68 PAT and 28 SPOT tags that remained on 294 | LUO et al.

FIGURE 2 Numbers of satellite telemetry tags (PAT, dark bars; SPOT, open bars) deployed on Atlantic tarpon dependent on: (a) year; (b) tarpon weight; (c) days at liberty; and (d) geographic location. The black bar in panel (c) shows the number of deployed tags that never reported to us via the Argos System tarpon between 10 and 50 days due to pin breaks for PAT tags, or river mouths (Figure 4). Examples of individual tarpon tracks can fishing line entanglements for SPOT tags, and 73 PAT and 27 SPOT be found in Supporting Information. remained on tarpon over 50 days. Tag deployments and pop-off (or The monthly frequency distribution of 3-hr mean temperatures final for SPOT) locations are summarized by regions in Table 1 and from all PAT tags (Figure 5a) indicated that from November to April Figures 1 and 2d. Geographically, most of our tagging efforts (93 Atlantic tarpon spent most of their time in waters where tempera- PAT, 51 SPOT tags) occurred in waters around Florida, USA, mostly tures were between 20 and 26°C and from May to October above in the Florida Keys and Florida Bay (38 PAT, 9 SPOT), followed by 26°C. April and November are transition periods with temperatures Texas, USA (29 PAT, 14 SPOT), and Mexico (31 PAT). centred around 26°C, which has been reported by tarpon anglers as preferred migratory temperature (Babcock, 1951; Spotte, 2016; White & Brennan, 2010). The mean monthly temperature ranged 3.2 | Seasonal migrations and distribution from 22.5°C in December–January to 29.5°C in August (solid line Figure 5a). The overall minimum was 14.9°C, and maximum tempera- We were able to generate 154 (83 PAT, 71 SPOT) tarpon tracks ture was 35°C. However, in December and January, tarpon spent from the 292 deployed tags. Of those, 29 (19%) tarpon travelled 29% and 36% of their respective time at temperatures ranging from ≥600 km during their deployment periods, 81 (52%) tarpon trav- 16 to 20°C (Figure 5a). elled from 100 to 600 km and 44 (29%) tarpon travelled <100 km In April, as the water temperatures warm to 26°C, tarpon ag - (Table 2). The distance travelled from tagging location as a function gregate annually in Florida Bay and the Florida Keys (Luo, 2009; of days at liberty (Figure 3) indicated different modes of movement Mill, 2010). The spatial map of mean SST in April 2008 indicated (i.e. residence and migration). ROM were generated from 140 (78 that 26°C waters were south of Veracruz, Mexico in the western PAT, 62 SPOT) tarpon tracks that were >10 days. Distribution of GOM, and around the Florida Keys and south Florida in the east- tagged tarpon as function of per cent of time tarpon spent at resi- ern GOM (Figure 5b). Typically, as the 26°C isotherm moved north dence and ROM criteria are summarized in Table 3, which indicates along the US south-eastern and GOM coasts, tarpon followed. In most tarpon spent <50% time as residence for all ROM criteria the fall, as the 26°C isotherm moved south (Figure 5c), tarpon did tested. In addition, most residence locations (red dots) were near the same. LUO et al. | 295

TABLE 1 Summary of tag deployment Region deployed PAT SPOT Total Pop-off or end region region and pop-off or end locations Florida Bay (FB), FL 38 9 47 0 (7), FB (22), EG (4), SL (10), GA (1), SC (1), NC (1), VA (1) Everglades (EG), FL 9 8 17 0 (3),EG (10), FB (1), BT (2), NC (1) Boca Grande-Tampa (BT), FL 34 10 44 0 (10), BT (24), EG (1), FB (8), LA (1) Apalachicola (AP), FL 6 28 34 0 (1), AP (10), AL (4), LA (12), BT (4), FB (3) Miami (MI), FL 6 6 12 0 (2), MI (7), SL (2), GA (1) St. Lucie_Augustine (SL), FL 11 0 11 0 (1), SL (8), FB (2) Texas (TX) 29 14 43 0 (6), TX (28), MX (9) Louisiana (LA) 3 1 4 LA (3), FB (1) Alabama (AL) 1 0 1 FB (1) Georgia (GA) 1 0 1 SL (1) South Carolina (SC) 5 2 7 SC (5), MI (2) North Carolina (NC) 1 0 1 NC (1) Mexico (MX) 31 0 31 0 (8), MX (9), TX (10), LA (4) Belize (BZ) 10 4 14 BZ (14) Nicaragua (NI) 3 0 3 Caribbean Sea (3) Trinidad (TR) 14 4 18 TR (17), St Kitts & Nevis (1) Angola, Africa (AN) 4 0 4 AN (4) Total 206 86 292

Note: The abbreviation for each region is given in parentheses in the first column; the number of tags in pop-off or end regions is given in parentheses. A zero region indicates the number of tags that did not report. The Florida Bay (FB) region included waters in Florida Bay and around the Florida Keys from Key Largo to Key West. The Everglades (EG) region included waters of Everglades National Park and the nearshore Gulf of Mexico (GOM). The Boca Grande-Tampa (BT) region included waters in the estuaries and nearshore GOM from Naples to Tampa, FL. The Miami (MI) region included waters from Biscayne Bay to Ft. Lauderdale. The St. Lucie–Augustine (SL) region extended from St. Lucie to St. Augustine, FL. The regions of countries and other US states are defined by their names. Abbreviations: PAT, Pop-up Archival Transmitting; SPOT, Smart Position and Temperature.

Two tags (T118 and T187) deployed on tarpon in Florida popped border with Mexico (Figure 8a), reflected by the winter SST distri- up near Newfoundland (Figure 6). It is not known whether these re- bution (Figure 8b). During the wet season in east GOM, CHUAs ex- flect tarpon movements or post-detachment tag drifting. Based on panded north to the area between Boca Grande and Apalachicola reports in the literature (Banon et al., 2019; Twomey & Byrne, 1985), and west to the mouth of Mississippi River (Figure 8c), when the such movements would not be unprecedented (Figure 6). SST was >26°C for the entire GOM (Figure 8c insert). Similarly, in Core habitat utilization area is represented by the top 50% of west GOM, the majority of CHUAs expanded to US waters extend- kernel density area. During the dry season on the eastern US coast, ing from Port Isabel to Galveston, TX during the wet season. Only CHUAs were all located from south of Fort Lauderdale to the Florida small portions of CHUAs are in Mexican waters (Figure 8d). In the Keys/Everglades (Figure 7a), with some minor low kernel density north, CHUAs extended to the mouth of the Mississippi River, where areas north of Fort Lauderdale up to Cape Canaveral, FL. The CHUA they overlapped with CHUAs of tarpon tagged in the eastern GOM distribution is a reflection of the SST distribution during this period (Figure 8c). (Figure 7a inset). During the wet season, as the water temperatures warm (Figure 7b inset), tarpon habitats expanded northward to the Chesapeake Bay with a small portion of the CHUAs extending to 3.3 | Estuarine and riverine utilization South Carolina. The majority of CHUAs remained in the area from the Florida Keys to south of Cape Canaveral (Figure 7b). Majority of tagged tarpon (51.1% PAT, 59.6% SPOT) used estua- Similar seasonal habitat utilization patterns occurred in the GOM. rine and riverine habitat (Table 4). The mean proportion of time for During the dry season, in the eastern GOM most of the CHUAs are SPOT tags (0.5407) was significantly higher than that for PAT tags located in Florida Keys and western Everglades. For tarpon tagged (0.3979) at α = 0.05 (Table 4), which suggests that SPOT wags were in the western GOM, all the CHUAs were located south of the US better at discerning river use compared to PAT tags salinity-based 296 | LUO et al.

TABLE 2 Number of tags at maximum distance (km) from TABLE 3 Distribution of tagged tarpon as function of residence tagging location in different regions time and rates of movement (ROM) over three consecutive days

Maximum distance (km) Total Per cent of time as residence

<100 100–600 >600 Tags 0% 1%–49% 50%–99% 100% Speed limit in Mexico 0 4 10 14 3 days Number of tagged tarpon Texas 5 12 3 20 <5 km/day 30 88 17 5 NE GOM 10 15 4 29 <7 km/day 20 88 26 6 SE GOM 18 30 7 55 <10 km/day 12 87 33 8 Atlantic US 5 10 2 17 Note: Zero per cent of time as residence (0%) indicates a tarpon's Other countries 6 10 3 19 ROM was greater than the speed limits for the entire duration of the All regions 44 81 29 154 deployment; 100% means a tarpon's ROM was less than the speed limits for the entire duration of the deployment; 1%–49% and 50%–99% mean per cent of time when ROM was greater than the speed limits.

FIGURE 4 Composite of all 12-hr locations overlain on bathymetry and rivers. Orange dots indicate locations in dry season (Jan–Apr) for all regions; green + symbols indicate locations in wet season for tarpon tagged west of the Mississippi River; blue + symbols indicate locations in wet season for tarpon tagged east of Mississippi River. The red dots indicate residence locations with the rate of movement criteria <5 km/day for three consecutive days

method. For geographic comparison, we grouped the tags into four regions (Table 4), and analysis of variance indicated no significant difference of the mean proportion of time tarpon utilized rivers between any of the regions. For seasonal comparison, the mean proportion of time tarpon in rivers was significantly greater in the dry season than the wet season (Table 4). Examples of the detailed individual tarpon utilizing estuarine habitat can be found FIGURE 3 Distance from tarpon satellite tagging location as in Supporting Information. a function of days at liberty: (a) western Gulf of Mexico (GOM); tarpon tagged in Mexico (black line) and Texas (blue line); (b) eastern GOM; tarpon tagged in northeast GOM (black line) and 3.4 | Spawning habitats southeast GOM (blue line); (c) US east coast and elsewhere; tarpon tagged in US east coast (black line) and other countries (blue line). The red dots in all panels indicate residence time with the rate of Possible spawning activity during the known tarpon spawning sea- movement criteria <5 km/day for three consecutive days son (April–July) was inferred from deep diving behaviour identified LUO et al. | 297

FIGURE 5 (a) Three-hour mean sea surface temperatures (SST) recorded by all PAT tags (dots), associated monthly frequency distribution (red histogram bars) and monthly mean SST (solid line). Monthly sample sizes are shown. Mean SST maps for: (b) April 2008 and (c) November 2008. Colour scale bar indicates SST values from 14 to 30°C. The 26–27°C isotherm is the yellow portion of the spectrum

from individual depth data around new and full moon periods for 6 of the 25 PAT tags were ingested by sharks evidenced by re- of 90 recovered PAT tags (Figure 9). We compared the dates of these duced light-level values (Figure 12a,b), and these tags were trans- dives with migration track locations of each individual to determine ported inside of the shark for periods ranging from 2 to 27 days presumed spawning sites (Figure 10). Interestingly, most sites were (mean = 8.06, SD = 6.6 days) after which they were regurgitated. clustered in two locations: (a) northeast of Veracruz, Mexico (west- The other incidents of predation were based on observed tooth ern Gulf of Mexico) and (b) south of Florida Keys on the deep coral marks left on recovered PAT and SPOT tags (Figure 12c, d). It is reef track proximal to the Straits of Florida. Additionally, we had one well known that sharks regurgitate indigestible items by eversion PAT pop-off and two SPOT tagged tarpon in an area about 200 km of the stomach (Brunnschweiler, Andrews, Southall, Pickering, west of Boca Grande, Florida, in the GOM (Figure 10), which has & Sims, 2005). Eight PAT tags that were dislodged from tarpon been suggested as a spawning site based on the presence of early prematurely had teeth marks indicating shark attacks. Most life stage leptocephali larvae of the Atlantic tarpon (Crabtree, 1995; (>90%) shark attacks on tarpon occurred within 3 hr post-release. Crabtree, Cyr, Bishop, Falkenstein, & Dean, 1992; Smith, 1980). However, one tarpon was attacked by a shark 44 days after the Tarpon started the deep diving at dusk and the activity continued deployment (Figure 12e). until 4 a.m. (Figure 11). In addition, the temperature range from surface to 90 m depth was within 1°C (26.5–27.5°C). The presumed spawning sites described here are all continental slope habitats with 4 | DISCUSSION depths ranging from 100 to 200 m. The salinities at these sites were around 36 PSU. The surface temperatures were 26 ± 2°C with a 4.1 | Tag deployments mixed layer extending from 50 to 100 m. We deployed 292 tags from 2001 to 2018 averaging 30 tags/year, spanning three continents, six countries and eight US states, the 3.5 | Predation by sharks largest tagging effort for tarpon since the inception of satellite telemetry technology (Block, Dewar, Farwell, & Prince, 1998; Block Shark predation was evident on 25 of the 90 recovered PAT tags et al., 2016, 2011; Lutcavage, Brill, Skomal, Chase, & Howey, 1999; (27.8%), and 8 of the 29 recovered SPOT tags (27.6%). Seventeen Rooker et al., 2019; Wilson et al., 2015). The size distribution of 298 | LUO et al.

FIGURE 6 HCYOM derived SST on 20 August 2010, for the north Atlantic Ocean overlain with historic tarpon capture locations (solid white dots) (Banon et al., 2019), a PAT tag pop-off location (T118, pink dot) and a SPOT tag location (T187, grey dot). There were 11 tarpon captured in Long Island pound nets in the summer of 1981, one tarpon in Cork, Ireland, on 28 October 1981, 7 tarpon in Bay of Biscay from 1981 to 2003 and 4 tarpon in Azores from 1973 to 2011 tagged tarpon reflected our objective of tagging sexually ma- behaviour has been observed in many fishes (Chapman, Skov, et ture tarpon (>35 kg, Crabtree, Cyr, Chacón Chaverri, McLarney, al., 2012; Hansson & Akesson, 2014). Partial migration results from & Dean, 1997) which are believed capable of taking long seasonal individual differences in behaviour, that is, a kind of animal per- migrations (Crabtree et al., 1992). The amount of time that tags sonality (Nilsson, Bronmark, Hansson, & Chapman, 2014). Partial stayed on fish can be used as an index of tagging success. Causes migrations have been observed for many satellite-tagged spe- of tagging failure included equipment malfunctions (i.e. defective cies, for example yellowfin tuna (Thunnus albacares, Scombridae battery, pin or rigging, etc.), human error (inexperienced tagger) (Hoolihan et al., 2014)); bluefin tuna (Thunnus thynnus, Scombridae and post-release mortality. Hoolihan, Luo, Abascal, et al. (2011) (Block et al., 2005; Cermeño et al., 2015; Wilson et al., 2005); showed behavioural modifications from satellite tagging of large sailfish (Istiophorus platypterus, Istiophoridae (Hoolihan, Luo, pelagic fishes. We believe that tagging may have had similar ef- Goodyear, Luo, Goodyear, Orbesen, & Prince, 2011)); blue mar- fects on Atlantic tarpon, possibly making tagged fish more suscep- lin (Makaira nigricans, Istiophoridae (Goodyear et al., 2008)); white tible to predation. marlin (Kajikia albida, Istiophoridae (Hoolihan et al., 2015)); swordfish (Xiphias gladius, Xiphiidae (Dewar et al., 2011)); tiger shark (Galeocerdo cuvier, Carcharhinidae (Hammerschlag, Gallagher, 4.2 | Seasonal migrations and distributions Gallagher, Wester, Luo, & Ault, 2012)); and acoustically tracked common snook (Centropomus undecimalis, Centropomidae; Trotter, Previously, Atlantic tarpon seasonal migrations were largely un- Blewett, Taylor, & Stevens, 2012). Partial migration is one of the known or inferred from their seasonal distributions. In the central major factors that could potentially account for spatial and tem- western Atlantic, tarpon commonly range from Virginia to Florida, poral variations in population abundance and, hence, is a powerful Bermuda, Gulf of Mexico, the Caribbean Sea to Brazil (Wade, force shaping ecosystem dynamics and trophic effects (Chapman, 1962) and infrequently from Nova Scotia to Argentina. Holder Brönmark, Nilsson, & Hansson, 2011; Chapman, Hulthén, et al., (1903) had speculated on tarpon migrations, believing they moved 2012). Partial migration may confer some evolutionary benefits from Mexico to Texas and then to Louisiana in the western GOM. to species because it provides a natural buffer against extinction, In the eastern GOM, he postulated movements from Key West given that multiple contingents of individuals within the same up the Florida Keys and then bifurcating to both west and east population simultaneously use a range of different habitats and coasts of Florida. His early speculations were relatively consistent resources. Tarpon, in the evolutionary record for >100 million with our findings. In fall, we found that tarpon from east of the years (Carroll, 1988; Jordan, 1919; Nature, 1971; Sepkoski, 2002), Mississippi River migrated southeastward to Florida, and tarpon may have benefited from many advantageous aspects of partial from west of the Mississippi River migrated southwestward to migration. Mexico. Based on our data, we have not yet seen a tarpon from Utilization distribution (UD) estimates summarized the seasonality east of the Mississippi River migrate west of Louisiana, or vice of tarpon regional distributions, indicating that in the wet season the versa. range of tarpon distribution is twice as large as that in the dry season. In contrast to the 29 long-distance migrators, many of the tar- In the western GOM during the dry season, the distribution is almost pon in this study travelled moderate to relatively short distances. exclusively south of the USA–Mexico border, while along the eastern Termed “partial migration”, intraspecific variation in migration GOM and United States south-eastern coasts, their distribution was LUO et al. | 299

FIGURE 7 Seasonal tarpon habitat utilization maps as represented by kernel density contours calculated from locations of all tags from Florida Keys to Maryland: (a) January to April (dry season) with inset of Hybrid Coordinate Ocean Model (HYCOM) SST on 15 February and (b) May to December (wet season) with inset of HYCOM SST on 15 August. Coloured contours indicate cumulative kernel density values from high density to low density (i.e. 50% indicates the highest 50% of all kernel density values which denotes core habitat utilization areas, CHUA)

almost exclusively restricted to southern Florida. Our migration tracks showing that some adult tarpon used freshwaters extensively and UD maps clearly showed that tarpon migrated beyond state bor- (Supporting Information), while others rarely travelled to freshwa- ders along the USA east coast from Virginia to Florida, and along coast ter. This individual variation is similar to the concept of partial sea- of USA GOM from Florida to Texas and to Mexico. Genetic studies are sonal migration previously discussed. Matich et al. (2017) showed consistent with our research and have shown no genetic differences that food availability might govern tarpon distribution in the Shark among locations in the central western Atlantic and east Atlantic (Ault River estuary, Everglades National Park. Gillworms have been & Luo, 2013; Ward et al., 2004, 2008, 2005). found on both Indo-Pacific and Atlantic tarpon (Bunkley-Williams & Williams, 1994; Hutton & Sogandares-Bernal, 1960; Mendoza- Franco, Kritsky, Vidal-Martinez, Scholz, & Aguirre-Macedo, 2004; 4.3 | Estuarine and riverine utilization Williams & Jones, 1994), and rapid movements between freshwaters and ocean environments may reduce the infections of patho- Juvenile and adult life stages of tarpon inhabit estuarine bays and gens (Westerdahl et al., 2014). Babcock (1951) stated that tarpon freshwater rivers (Ault, 2008; Babcock, 1921a; Dimock & Dimock, might use the freshwater habitat to get rid of remoras and sea lice 1912; Goode, 1887; Pinckney, 1888). Tarpon are found in freshwater (Nerocila acuminate, Cymothoidae) since the remoras and sea lice Lake Nicaragua that sits some 195 km from its connection to the cannot survive the freshwater. ocean via Rio San Juan (Gill & Bransford, 1877; Koenig, Beatty, & Marinez, 1976; Simmons, 1900). Local Nicaraguans believe that tarpon in the Lake are a resident population, living and breeding in the 4.4 | Spawning habitats Lake. However, our 3 PAT tags deployed in the Rio San Juan, about 30 km east of Lake Nicaragua and 145 km west of the Caribbean Sea, To date, no one has directly observed tarpon spawning, other than all popped off in the Caribbean Sea. some ad hoc observations of presumed pre-spawning activities Despite the well-documented presence of mature adult tarpon (Baldwin & Snodgrass, 2008; Crabtree et al., 1992), and post hoc in freshwaters (Babcock, 1921a; Dimock & Dimock, 1912; Koenig predictions of spawning sites from the capture of leptocephalus et al., 1976), much of the scientific literature describes adult tar- larvae (Crabtree, 1995; Crabtree et al., 1992, 1997; Smith, 1980). pon habitat as mainly shallow coastal waters, bays and estuaries, In the early 20th century, Babcock (1921a) suggested that tarpon with only an occasional reference to freshwaters (Moyle & Cech, spawned in freshwater rivers, based on his observations of juve- 1988; Robins & Ray, 1986). A recent study based on elements and nile tarpon captured in those habitats. Subsequently as larval sam- isotopic ratios in tarpon scales (Seeley & Walther, 2018) showed ples were collected in ocean environments, it was determined that that tarpon spent on average 42 ± 34% of life histories within tarpon spawned in offshore waters (Babcock, 1951; Wade, 1962). oligohaline habitats, while another study using tarpon eye lens Smith (1980) and Crabtree et al. (1992) found young leptocephali showed 100% juvenile habitat in upper estuarine habitat (Kurth, offshore, further suggesting offshore spawning. Locations identi- Peebles, & Stallings, 2019). Our study reinforced this concept by fied as spawning areas from our study (Figure 10) matched spawning 300 | LUO et al.

FIGURE 8 Seasonal tarpon habitat utilization maps as represented by kernel density contours calculated from locations of all tags in the Gulf of Mexico: (a) January to April (dry season); (b) HYCOM SST on 15 February 2008; (c) east Gulf of Mexico (GOM) from May to December (wet season) with inset of HYCOM SST on 15 August 2008; and (d) west GOM from May to December areas previously identified in the western GOM (Smith, 1980) and involves a rapid ascent from the substrate towards the surface using in the eastern GOM and Straits of Florida (Crabtree, 1995; Crabtree the change of hydrostatic pressure to force the release of gametes et al., 1992). Our observations of purported spawning dives occur- (Domeier & Colin, 1997; Graham & Castellanos, 2005; Whaylen, ring within a few days of new and full moons matched estimates of Pattengill-Semmens, Semmens, Bush, & Boardman, 2004). Similar spawning near full and new moons by Crabtree et al. (1997). Based rapid oscillatory dive patterns have also been observed for satel- on our tagging results and the occurrences of tarpon leptocephalus lite-tagged bluefin tuna in the GOM and Mediterranean Sea, and larvae, we think that tarpon spawning could occur around continen- these were considered to be spawning behaviours (Aranda, Abascal, tal slope areas where depths range from 100 to 200 m, temperatures Varela, & Medina, 2013; Cermeño et al., 2015; Teo, Boustany, & are near 26 ± 2°C, and salinity is around 36 PSU during spawning Block, 2007). A related species, bonefish (Albula vulpes, Albulidae), season, within 24 hr of travel distance from inshore coastal habitats. also spawns offshore at night near full moons and descends to ~60 m Our observations of deep diving behaviour corresponded with to spawn (Danylchuk, Lewis, Jud, Shenker, & Adams, 2019). seasonality of tarpon spawning (April–July in Florida; Crabtree et al., 1997) and with the likelihood of spawning in deep water based on egg characteristics (Babcock, 1921b). The deep diving behaviours 4.5 | Predation by sharks we observed were from May to June (Figure 9), well within the reported tarpon spawning season. Thus, we are very confident these Shark predation on tarpon has been well documented by anglers and observed deep diving behaviours were related to spawning activity. in the scientific literature (Ault, 2010; Ault et al., 2008; Churchill, Many studies have shown spawning behaviour of numerous fishes 1907; Dimock & Dimock, 1912; Guindon, 2011; Hammerschlag, Luo, LUO et al. | 301

TABLE 4 Comparison of tarpon estuarine and riverine utilization by tag types, tagging regions and seasons, showing the number (and percentage) of tagged tarpon that used rivers, the mean number of days spent in rivers and the mean proportion of time spent in rivers; SD is standard deviation

Used rivers Number of days in rivers Proportion of time in rivers

Comparison N (%) Mean SD Mean SD t p

Tag type SPOT 31 (59.6) 35.97 34.75 0.5407 0.2868 2.0615 .044* PAT 23 (51.1) 32.33 31.17 0.3979 0.2221 Region Atlantic US 15 (79.4) 27.53 22.96 0.4245 0.2589 1.3943 .1731 SE GOM 21 (56.8) 39.43 33.83 0.5481 0.2744 NE GOM 14 (56.0) 28.29 32.00 0.4362 0.2589 West GOM 4 (25.0) 62.25 56.06 0.5434 0.2469 Season Dry 12 (60.0) 47.00 42.97 0.6711 0.2298 3.186 .0047*** Wet 42 (54.5) 31.48 29.34 0.4252 0.2555

Note: The proportion of time in rivers is the total number of days spent in rivers divided by the days at liberty. Student's t tests were performed on the mean proportion of time in rivers for SPOT versus PAT, Atlantic US versus SE GOM, and Dry versus Wet. Abbreviations: PAT, Pop-up Archival Transmitting; SPOT, Smart Position and Temperature. *Significant at the α = .05 level. ***Significant at the α = .001 level.

FIGURE 9 Depth and temperature profiles from six recovered PAT tags showing deep diving (presumed spawning) behaviours of Atlantic tarpon: (a) T-30, (b) T-43 and (c) T-69 were tagged in Veracruz, Mexico on 11 May 2004, 28 May 2006 and 29 May 2007, respectively, and (d) T-53, (e) T-60 and (f) T-106 were tagged in Florida Bay, Florida on 11 June 2007, 29 April 2007 and 4 May 2008, respectively. The scale bar at the bottom represents the temperatures from 16 to 30 (°C). The arrows in each panel indicate dates of new and full moons 302 | LUO et al.

FIGURE 10 Bathymetric map of Gulf of Mexico (GOM) and Florida Straits overlaid with locations where deep diving behaviours were recorded by the recovered PAT tags (solid red dots) from Figure 9, and offshore movement locations reported from SPOT tags (open green triangles). The solid yellow line ellipses are predicted spawning areas based on the tagging data, and the dashed yellow line ellipses are other potential spawning habitat locations. The solid pink dots are presumed spawning locations from Smith (1980); the pink rectangular boxes are spawning areas noted by Crabtree (1995)

FIGURE 11 A close up of the deep dive from/T60 (Figure 9e) from 16:00 on 14 May to 04:00 on 15 May 2007 showing depth (a), light level (b) and temperature (c). The arrow in panel (b) indicates the time of sunset

et al., 2012; Tuma, 1976). Holder (1913) recorded his tarpon fishing those total mortalities, 8.3% were due to shark predation. Our study results in Aransas Pass, Texas, during June 1909, and reported 26.3% using recovered PAT and SPOT tags indicated a much higher shark of hooked tarpon mortality was due to shark predation. predation mortality: 27.8% and 27.6%, respectively. The fact that Guindon (2011), using acoustic tags, estimated a 13% rate for two different tag types produced nearly identical results gives us tarpon catch-and-release mortality, due to a variety of reasons. Of confidence that our mortality estimates are accurate. We believe LUO et al. | 303

FIGURE 12 Light-level (a) and depth (b) data recorded by PAT tag (T-117) deployed on a tarpon between 8 and 16 September 2007, indicating predation occurred about 2 hr after deployment. Examples of recovered PAT (c) and SPOT (d) tags that bore tell-tale teeth marks of shark predation. (e) Light-level data recorded by PAT tag (T-234) indicated predation occurred 44 days after deployment

that Guindon (2011) may have underestimated catch-and-release tagging process added handling time as compared to normal catch- mortality due to two key criteria used: (a) lack of acoustic tag move- and-release, thus may further weaken the tarpon when released. ment and (b) a relatively short time window (6 hr) of observation Since most of the shark predation in our study occurred near passes to make this determination. Using Guindon’s (2011) criteria on our and areas known with high shark density (Hammerschlag, Luo, et al., data, the 17 PATs inside shark stomachs would have been consid- 2012), this mortality estimate might not apply more broadly. ered alive, when in fact they were dead and consumed by the predator. Only the eight dislodged PAT tags of the 25 recovered, using Guindon's criteria, would have been considered dead resulting in an 5 | IMPLICATIONS FOR FISHERY 8.9% mortality estimate (8 out of 90). This is very close to Guindon's MANAGEMENT estimate of 8.3%. Our study showed that sharks preyed upon tarpon after 6 hr and up to 44 days after release. However, this post-release Improved understanding of tarpon movement ecology has significant mortality by sharks may not apply to tarpon released without a satel- resource management implications for fisheries and coastal habitats. lite tag. The electrical field from the tag may attract unwanted atten- Our results suggest a single, interconnected tarpon “unit stock” that tion from sharks, as their sensory systems are capable of detecting from a fishery perspective—a closed population comprised of the full weak electrical fields (Adair, Astumian, & Weaver, 1998; Kalmijn, life cycle of spawning adults, eggs, larvae and juveniles—may, at a 1971, 1982). Another tagging study (Kerstetter et al., 2004) showed minimum, stretch from the US Atlantic coast throughout the Gulf increased shark predation on satellite-tagged billfish. In addition, the of Mexico. 304 | LUO et al.

Overfishing and habitat loss are two obvious threats to sustain- Commission, International Commission for the Conservation of ability of the US Atlantic–Gulf of Mexico tarpon stock (Adams et Atlantic Tunas, International Scientific Committee for Tuna and Tuna- al., 2014; Ault et al., 2008; Spotte, 2016; Stilwell, 2011). Tarpon are like Species in the North Pacific Ocean). Atlantic tarpon should be fished throughout this geographic region and are targeted by both considered a highly migratory species managed federally and interna- recreational and commercial fisheries at different locations. In general, tionally like bluefin tuna and billfishes. With sufficient scientific infor- the United States has only recreational fisheries for tarpon (primar- mation, sustainable levels of fishing effort could be determined and ily catch-and-release), while commercial, subsistence and recreational allocated among individual states and Mexico. In contrast, interstate fisheries occur elsewhere (Ault et al., 2008; Ault & Luo, 2013; Browder, and international management of tarpon coastal habitats is almost in- Davis, & Sulliavan, 1981; Dailey & Landry, 2008; Spotte, 2016). conceivable at present. It is difficult to imagine, for example, a water Both harvest and catch-and-release fisheries incur some level of management or urban development project in sensitive tarpon habitat fishing mortality. Our tagging studies showed that the survival rate in one state being blocked by another state, let alone another country. of released tarpon is substantially lower than 100% due to a variety However, with sufficient ecological and resource economic informa- of factors, of which predation by sharks may also be a significant tion, the potential economic losses to the entire tarpon fishery result- unintended fishing mortality factor associated with a thriving recre- ing from coastal development projects could be quantified and the ational fishery. Thus, for the sake of tarpon conservation and fishery risks weighed against the economic benefits. Framing the trade-offs in sustainability, we highly recommend stopping tarpon fishing in areas economic terms may at least give the fish a fighting chance. where shark attacks on tarpon occur, and moving the angling effort to alternative fishing sites. ACKNOWLEDGMENTS Substantial loss of tarpon habitat, particularly mangrove wetlands, Funding was provided by the Bonefish & Tarpon Trust (www. has undoubtedly occurred along the south-eastern US and Gulf of tarbone.org), the Robertson Foundation, National Fish & Wildlife Mexico coastlines over the past 50 or more years due to coastal de- Foundation (www.nfwf.org), Buchanan Family Foundation, Wild velopment (Valiela, Bowen, & York, 2001). Loss of tarpon habitat is River Press, Baker Bishop, Sanctuary Friends of the Florida Keys perhaps a more insidious threat to sustainability than fishing mortal- Foundation (www.sffkf.org), Florida Fish and Wildlife Conservation ity. Adverse effects of overfishing on the reproductive capacity of the Commission, Texas Parks and Wildlife Department, Ocean Reef tarpon stock can be alleviated by relaxing fishing pressure. In contrast, Community Foundation, National Science Foundation under Grant loss of habitat area is irreversible and potentially limits overall stock No. EAR-1204752 and a number of other individual donors. This carrying capacity and, thus, maximum tarpon stock abundance. The research would not have been possible without the help of many impact of habitat loss may be especially damaging to tarpon because fishing clubs, professional captains and fishing guides, anglers of their use of coastal and riverine waters in all life stages. It is well and individuals who assisted us in providing vessels, catching and known that juvenile tarpon inhabit freshwater tributaries (Ault, 2008; tagging tarpon, and in tag recovery. Clubs include the following: Babcock, 1921a; Dimock & Dimock, 1912; Goode, 1887; Pinckney, Veracruz Yacht Club, Coatzacoalcos Yacht Club, Stuart Rod & 1888). Our study has shown that at times adult tarpon also inhabit Reel Fishing Club, Fieldworkers Club, Patagonia, Silver Kings TV, small freshwater creeks (see Supporting Information). Tarpon Lodge Pine Island, Tarpon River Lodge Nicaragua, Belize This study has shown that tarpon can travel long distances and River Lodge, El Pescador Lodge Belize, Rio Parismina Lodge Costa cross international borders; thus, in the long term, any local de- Rica, Green Reef Belize, and the Tarpon Tomorrow. Individuals in- pletion in the tarpon stock, via fishing mortality or habitat loss, clude: Dick Alario, Stu Apte, Mark Badzinski, Luiz Barbieri, Youssef will likely affect the tarpon population as a whole in the Atlantic Barquet, George Beckett, Jim Bohnsack, Bou Bosso, Curtis and Ocean. This spatially expansive view of the US Atlantic–Gulf of Andrew Bostick, Harold Brewer, Grant Brown, William Bryan, Mexico tarpon stock suggests that achieving long-term sustain- Charlie Causey, Felipe Fernandez Ceballos, Gustavo Cisneros, Roy ability will require a high degree of coordination among scientists Crabtree, Jack Curlett, Bill Curtis, Marty Dashiell, Alberto Pavon and managers from numerous Atlantic and Gulf coast states as David, Fuzzy Davis, Jay Decker, Don Demaria, Dave Denkert, well as Mexico. The most pressing need in the short term is to Jorge Diaz, Paul Dixon, Raymond Douglas, Jimmie Durham, Sally gather synoptic data to develop a stock-wide shared information and Jim Farley, Russ Fisher, Rob Fordyce, Frank Fowler, John system to support informed management decision-making focus- Frazier, Steve Fronterhouse, Gary Fungui, Chris Galvin, Heliodoro ing on three general areas: (a) fishery science and biological data Garza, Roger Gibson, Tom Gibson, Doug Hannon, Jeff Harkavy, for determining stock sustainability status; (b) ecological and map- Christine Harvey, Guy Harvey, Mike Heusner, Rick Hirsch, ping data for determining the location and amount of viable tarpon George Hommell, Brent Huggins, Ron Hunter, Dan Jacobs, Derek habitat; and (c) resource economics data for determining the mon- Jordan, Joel Kalman, Doug Kelly, Mitch Kiesler, Julian Lajornade, etary value of the tarpon stock. Bev Landstreet, Ken Lindeman, Paul Lubbers, Alberto Maderia, Controlling tarpon fishing mortality through coordinated manage- Behzad Mahmoudi, Erik Utrera-Lopez Marco, Jorde Martinez, Jim ment is at least conceivable and has precedent with intergovernmental McDuffie, Larry McKinney, Robert Meaher, Bill Michaelcheck, management organizations for highly migratory tunas and billfishes Andy Mill, Eduardo Perusquia Moran, Tom Morgan, Sandy Moret, in the Atlantic and Pacific Oceans (e.g. Inter-American Tropical Tuna Hunter Neblett, George Neugent, Iain Nicolson, Heidi Nute, Carlos LUO et al. | 305

Partida, Billy Pate, Tom Pero, James Plaag, Stuyve Pierrepont, Aranda, G., Abascal, F. J., Varela, J. L., & Medina, A. (2013). Spawning behaviour and post-spawning migration patterns of Atlantic bluefin Marcus Poffenberger, Jesus Quijano, Gerardo Fernandez tuna (Thunnus thynnus) ascertained from satellite archival tags. PLoS Quijano, Fernandez Quijano, Jared Raskob, Mike Restivo, Jose ONE, 8(10), https://doi.org/10.1371/journal.pone.0076445 Manual Parada Rey, Michael Rich, Danny Romino, Gabriel Romo, Ault, J. S. (2008). Biology and management of the world's tarpon and bone- Nicky Runnels, Geoffrey Samuels, George Santry, Lance “Coon” fish fisheries. Boca Raton, FL: CRC Press/Taylor & Francis Group. Schouest, Jason Schratweiser, Jonathan Shenker, Dick Sherman, Ault, J. S. (2010). Silver King: A most perfect and ancient sport fish. In A. Mill (Ed.), A passion for tarpon (pp. 266–292). Mill Creek, WA: Wild Joe Skrumbellos, Curt Slonim, Chad Smith, Derke Snodgrass, River Press. Pedro Sors, Roe Stamps, Bob Stearns, Chris Sumers, Austin Tellam, Ault, J. S., Humston, R., Larkin, M. F., Perusquia, E., Farmer, N. A., Luo, J., Monty Trim, Colby Ulva, Felix Juan Malpica Valverde, Steve Venini, … Posada, J. (2008). Population dynamics and resource ecology of Pierre and Susie Villere, Rufus Wakeman, Natalia Zurcher, and Tim Atlantic tarpon and bonefish. In J. S. Ault (Ed.), Biology and management of the world’s tarpon and bonefish fisheries (pp. 217–258). Boca Choate. We also thank the Editor and two anonymous reviewers Raton, FL: CRC Press/Taylor & Francis Group. who provided insightful comments and suggestions that greatly Ault, J. S., & Luo, J. (2013). A reliable game fish weight estimation model improved this manuscript. This paper is dedicated to the indelible for Atlantic tarpon (Megalops atlanticus). Fisheries Research, 139, 110– memory of our friend and colleague, the late Capt. Bruce T. Ungar. 117. https://doi.org/10.1016/j.fishres.2012.10.004 Babcock, L. L. 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Marine Biology, 146, 409–423. https://doi.org/10.1007/ ics of tarpon (Megalops atlanticus): Microsatellite variation across the s00227-004-1445-0 distribution of the species. In J. S. Ault (Ed.), Biology and management Worton, B. J. (1989). Kernel methods for estimating the utilization distri- of the world's tarpon and bonefish fisheries (pp. 131–146). Boca Raton, bution in home-range studies. Ecology, 70(1), 164–168. https://doi. FL: CRC Press/Taylor & Francis Group. org/10.2307/1938423 Ward, R., Blandon, I. R., Vegal, R., Landry, A. M., Dailey, W., de Leon, F. Worton, B. J. (1995). Using Monte Carlo simulation to evaluate ker- J. G., … Needleman, D. S. (2005). Studies in conservation genetics of nel-based home range estimators. Journal of Wildlife Management, tarpon (Megalops atlanticus) - III. variation across the Gulf of Mexico 59(4), 794–800. https://doi.org/10.2307/3801959 in the nucleotide sequence of A 12S mitochondrial rRNA gene frag- ment. Contributions in Marine Science, 37, 45–59. Waycott, M., Duarte, C. M., Carruthers, T. J. B., Orth, R. J., Dennison, SUPPORTING INFORMATION W. C., Olyarnik, S., … Williams, S. L. (2009). Accelerating loss of sea- Additional supporting information may be found online in the grasses across the globe threatens coastal ecosystems. Proceedings Supporting Information section. of the National Academy of Sciences, 106(30), 12377. https://doi. org/10.1073/pnas.0905620106 Webster, M. S., Marra, P. P., Haig, S. M., Bensch, S., & Holmes, R. T. (2002). Luo J, Ault JS, Ungar BT, et al. Links between worlds: Unraveling migratory connectivity. Trends How to cite this article: in Ecology and Evolution, 17(2), 76–83. https://doi.org/10.1016/ Migrations and movements of Atlantic tarpon revealed by S0169-5347(01)02380-1 two decades of satellite tagging. Fish Fish. 2020;21:290–318. Weng, K. C., Castilho, P. C., Morrissette, J. M., Landeira-Fernandez, https://doi.org/10.1111/faf.12430 A. M., Holts, D. B., Schallert, R. J., … Block, B. A. (2005). Satellite LUO et al. | 309

7.99 7.36 9.01 9.95 0.39 0.00 0.79 8.03 4.71 5.39 4.07 0.03 2.96 27.92 29.77 11.72 12.47 30.88 Speed Speed (km/day) (Continues) 0 0 0 0 0 0 0 0 0 0 0 0 0.2 5.6 26 18.9 77 287.1 857.4 971.5 371.9 162.2 223.7 773.9 833.7 438 1,116.8 1,159.6 1,195.3 Distance Distance (km) 484.6

61 67 24 27 18 26 28 26 93 79 39 40 44 119 121 159 102 182 Days at liberty NO NO NO YES NO NO NO NO NO NO YES NO NO YES NO NO NO NO NO NO YES YES NO NO NO YES YES NO Tag recover NO NO

−76.551 −97.291 −97.463 −94.818 −96.788 −81.996 −96.873 −96.289 −96.617 −91.014 −92.246 −82.270 −90.840 −88.939 −80.630 −80.802 −80.500 −80.174 Lon (dd) ) 24.698 28.991 no loc 28.092 no loc 29.303 no loc no loc no loc 26.789 26.529 19.375 no loc 28.720 29.087 21.350 no loc no loc no loc no loc no loc 28.180 28.931 no loc 24.839 27.800 26.255 21.693 Reporting Lat-Lon Reporting 35.001 Lat (dd) 28.006 Megalops atlanticus Megalops 02/10/04 11/10/03 not heard 09/08/04 not heard 07/08/05 not heard not heard not heard 07/09/03 07/01/03 07/10/04 not heard 09/22/03 08/11/04 06/08/04 not heard not heard not heard not heard not heard 08/05/02 09/07/05 not heard 06/05/06 11/04/01 06/06/04 07/07/05 Reporting Reporting date 03/04/02 01/03/02 02/10/04 11/10/03 08/18/04 09/08/04 08/15/05 07/31/05 07/15/05 07/28/04 12/15/03 08/15/03 10/01/03 07/21/04 09/01/03 11/10/03 08/11/04 07/14/04 06/15/03 12/15/02 12/15/02 10/24/02 12/02/02 08/05/02 09/15/05 02/15/02 10/30/06 11/04/01 09/01/04 08/31/05 Expected Expected date pop-off 03/04/02 01/03/02 −76.533 −89.200 −89.204 −89.200 −96.289 −96.289 −96.290 −96.290 −96.294 −96.289 −96.256 −96.271 −96.289 −96.287 −96.274 −96.277 −96.274 −96.290 −81.080 −82.270 −82.270 −82.764 −82.765 −82.258 −80.868 −80.868 −80.771 −80.140 −80.267 Lon (dd) −80.749 19.377 19.377 19.360 19.360 19.328 19.377 19.318 19.320 19.377 19.341 19.366 27.605 27.607 19.323 19.366 19.360 27.001 24.798 24.798 24.842 26.699 26.699 31.702 28.945 28.960 26.699 35.049 28.962 25.333 32.331 Release Lat-Lon Release Lat (dd) 27.3 79.5 77.4 61.4 61.4 75.6 55.0 45.5 65.9 85.0 31.8 45.5 78.0 78.0 34.1 36.4 90.0 90.0 60.5 84.4 36.4 34.1 38.6 34.1 40.9 34.1 84.6 40.9 80.0 38.6 Fish, wt (kg) Venice, LA Venice, LA Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Boca Grande, FL Veracruz, Mexico Veracruz, Boca Grande, FL Venice, LA Veracruz, Mexico Veracruz, Tampa Bay, FL Bay, Tampa Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Tampa Bay, FL Bay, Tampa Boca Grande, FL Long FL Key, Long FL Key, Islamorada, FL Veracruz, Mexico Veracruz, Stuart, FL Ocean Reef, FL Savannah, GA Savannah, Release location Release Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Hilton Head, SC NC Oriental, 09/04/03 09/04/03 05/10/04 05/10/04 05/29/05 05/29/05 05/28/05 05/10/04 06/15/03 06/07/03 06/12/03 06/06/03 09/04/03 05/10/04 06/27/02 05/10/04 05/11/04 06/25/02 06/13/02 05/28/02 05/28/02 05/18/02 05/28/05 09/28/01 05/10/06 09/21/01 Tagging Tagging date 05/11/04 05/29/05 09/06/01 09/03/01 T-27 T-26 T-25 T-24 T-23 T-22 T-21 T-20 T-19 T-13 T-18 T-12 T-14 T-17 T-11 T-16 T-15 T-09 T-08 T-07 T-06 T-05 T-32 T-04 T-31 T-03 Tag name T-30 T-28 T-01 T-02 9 8 7 6 5 4 3 1 2 19 14 24 11 17 10 26 16 21 13 18 23 25 27 15 12 20 22 28 29 30 Tag count APPENDIX APPENDIX A1 DateTABLE and location of the start and and end SPOT of each tag PAT deployment on Atlantic Tarpon ( 310 | LUO et al.

9.65 9.59 5.36 4.08 5.47 1.49 0.02 0.43 0.90 0.43 2.33 1.17 8.79 8.05 3.19 0.43 6.30 3.48 3.02 11.35 14.04 24.83 16.70 13.88 12.67 Speed Speed (km/day) (Continues) 0 0 0 0 1.6 39.3 14 74.8 24.3 71.5 27 31.5 31.5 22.7 80.1 267.2 198.6 421.3 216.9 913.5 894.7 393.3 298.8 501.7 138.8 338.3 1,180.6 402.6 1,152.7 Distance Distance (km)

6 8 5 41 52 16 42 81 57 13 35 91 23 53 29 68 48 34 34 63 30 64 167 167 104 Days at liberty NO YES NO NO NO NO YES YES YES NO YES YES YES YES YES YES NO NO NO NO NO NO NO NO Tag recover NO YES YES YES YES

−97.464 −97.322 −97.265 −94.206 −94.548 −96.411 −96.634 −96.781 −81.449 −81.370 −81.109 −93.500 −81.178 −92.733 −63.271 −63.529 −80.157 −80.303 −80.297 −80.581 −80.125 −80.911 −80.504 −80.988 −80.535 Lon (dd) 18.194 28.404 no loc 13.316 12.696 18.016 27.177 27.466 29.666 27.227 21.676 28.373 25.953 27.352 24.788 27.502 no loc 28.225 no loc 24.392 29.020 no loc 24.864 28.122 Reporting Lat-Lon Reporting 24.709 29.196 24.736 Lat (dd) 28.448 25.387 11/26/06 11/30/06 not heard 09/26/06 09/18/06 11/22/06 05/29/07 11/20/06 09/09/06 10/22/06 11/21/06 07/01/07 07/15/07 11/11/06 07/01/07 11/11/06 not heard 05/26/07 08/13/06 not heard 05/20/06 07/01/06 no heard 06/07/06 10/12/06 Reporting Reporting date 08/28/05 07/16/07 05/28/07 07/02/07 01/22/07 01/08/07 05/02/07 04/02/07 02/05/07 02/12/07 06/16/07 03/12/07 11/15/06 01/29/07 02/26/07 07/01/07 07/15/07 11/06/06 07/01/07 11/10/06 11/03/06 06/15/07 01/08/07 10/20/06 10/01/06 10/04/06 06/30/07 10/01/06 12/25/06 Expected Expected date pop-off 09/30/05 07/16/07 07/03/07 07/02/07 −61.711 −61.714 −61.721 −94.436 −94.431 −94.436 −96.395 −94.913 −96.295 −96.294 −96.296 −96.295 −94.913 −96.291 −96.290 −96.290 −81.263 −81.263 −80.751 −80.208 −80.174 −80.753 −80.749 −80.746 −80.753 −80.290 −80.753 Lon (dd) −80.751 −80.751 19.369 19.370 19.344 19.371 19.347 27.196 27.165 19.360 19.360 29.176 29.176 24.837 24.843 24.841 24.858 24.852 24.658 10.689 10.686 10.678 24.658 24.852 24.837 24.837 18.187 18.187 18.182 28.406 25.276 Release Lat-Lon Release Lat (dd) 47.3 57.7 37.4 57.6 49.7 77.5 51.8 41.4 55.4 42.7 70.8 62.7 36.4 20.9 54.7 50.3 53.2 50.0 43.8 64.2 84.9 43.5 48.2 34.6 40.8 44.5 44.1 44.0 44.0 Fish, wt (kg) Mexico Mexico Mexico Port O'Connor, TX O'Connor, Port Coatzacoalcos, Trinidad, BWI Trinidad, Coatzacoalcos, Coatzacoalcos, Jensen Beach, FL Florida FL Bay, Trinidad, BWI Trinidad, BWI Trinidad, Florida FL Bay, Florida FL Bay, Jensen Beach, FL Veracruz, Mexico Veracruz, Coatzacoalcos, Coatzacoalcos, Florida FL Bay, Veracruz, Mexico Veracruz, Florida FL Bay, Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Galveston, Texas Bahia Honda, FL Veracruz, Mexico Veracruz, Bahia Honda, FL Key Largo, FL Veracruz, Mexico Veracruz, Galveston, Texas Release location Release Florida FL Bay, Veracruz, Mexico Veracruz, Florida FL Bay, Florida FL Bay, 09/10/06 09/30/06 08/16/06 09/30/06 10/16/06 05/13/07 08/15/06 08/15/06 05/21/07 06/11/07 10/16/06 05/28/06 09/30/06 05/14/07 05/28/06 05/13/07 05/28/06 05/28/06 08/05/06 05/15/06 05/27/06 05/15/06 06/01/06 05/30/07 08/05/06 Tagging Tagging date 05/14/07 05/29/05 04/29/07 04/29/07 T-48 T-49 T-47 T-50 T-54 T-55 T-45 T-46 T-56 T-53 T-52 T-44 T-51 T-57 T-43 T-58 T-42 T-41 T-40 T-35 T-39 T-36 T-38 T-34 T-37 Tag name T-59 T-33 T-60 T-61 51 41 47 52 55 42 59 45 49 37 31 57 35 32 39 56 53 50 36 58 33 46 48 54 38 43 40 34 44 Tag count TABLE A1 (Continued) TABLE LUO et al. | 311

7.45 5.19 1.97 1.17 1.29 4.94 1.18 0.19 0.19 3.86 3.76 2.87 2.34 6.06 6.65 2.11 2.64 2.03 8.92 11.88 10.04 10.52 16.99 13.48 16.36 15.20 12.91 12.07 12.97 Speed Speed (km/day) (Continues) 9.5 0 0 27.6 77.1 14.2 96.3 21.9 23.6 53.9 46.7 357.5 157.8 151.8 740 135.9 182.4 103.6 893.4 244.6 392.6 858.4 154.5 100.6 100.2 368.5 860.6 483.7 Distance Distance (km) 1,149.4 1,183.2 1,271.5

4 9 8 51 14 17 24 13 49 65 35 82 98 98 98 98 89 89 15 12 12 20 83 38 48 116 142 124 129 Days at liberty NO NO YES YES YES NO NO YES YES YES YES YES NO NO NO NO Tag recover NO NO NO YES NO YES NO NO YES NO YES NO NO NO YES

−97.492 −97.067 −97.406 −97.092 −97.064 −89.372 −95.223 −94.375 −96.343 −96.801 −96.730 −91.779 −81.036 −81.510 −81.414 −81.181 −93.562 −92.855 −62.261 −62.294 −82.531 −62.711 −90.151 −82.549 −83.586 −83.583 −80.980 −80.225 −80.019 Lon (dd) 11.352 26.776 10.811 25.385 28.419 24.459 27.835 28.106 29.017 29.716 28.158 27.563 18.702 10.780 no loc 20.631 16.998 Reporting Lat-Lon Reporting 18.551 29.552 24.749 28.414 11.763 Lat (dd) 25.951 20.888 9.922 30.224 27.244 28.898 27.990 no loc 25.698 11/05/07 06/30/08 02/24/09 12/03/07 10/08/07 11/01/07 09/26/08 09/12/07 08/21/07 08/17/07 09/17/07 09/16/07 09/05/07 Reporting Reporting date 03/02/09 not heard 01/28/08 11/06/07 09/05/07 08/27/07 09/13/07 09/05/07 08/29/07 10/23/07 01/15/08 05/18/09 09/10/07 09/10/07 09/05/07 07/05/07 not heard 07/11/07 12/15/07 11/15/08 07/20/09 01/10/08 12/06/07 02/20/08 03/01/09 03/15/08 11/26/07 11/26/07 01/15/08 01/15/08 09/05/07 Expected Expected date pop-off 07/20/09 08/25/09 01/30/08 02/11/08 09/05/07 09/05/07 09/13/07 09/05/07 02/04/08 02/05/08 01/15/08 07/20/09 09/10/07 09/10/07 09/05/07 07/31/07 07/03/07 08/29/07 −97.145 −87.982 −61.705 −61.759 −61.759 −95.014 −94.993 −94.988 −96.478 −96.607 −96.628 −96.626 −96.283 −96.384 −96.292 −96.296 −96.283 −96.596 −96.288 −81.289 −81.638 −81.290 −82.263 −84.476 −84.463 −84.474 −80.771 Lon (dd) −80.166 −80.180 −80.749 −80.771 19.378 19.368 19.375 19.378 19.370 27.205 27.169 29.096 29.091 29.097 11.041 11.031 11.038 24.656 24.844 24.652 24.841 24.844 10.681 10.697 10.697 26.712 26.067 25.897 28.293 28.559 28.568 28.571 28.335 28.587 30.565 Release Lat-Lon Release Lat (dd) 47.3 47.4 37.4 49.6 51.3 51.2 76.3 55.1 55.7 52.0 52.0 32.4 56.7 50.3 50.2 33.1 66.0 22.9 60.3 46.8 43.5 46.2 40.9 30.5 43.5 40.8 40.9 48.5 48.5 44.0 44.5 Fish, wt (kg) Boca Grande, FL San Juan, Nicaragua Trinidad, BWI Trinidad, Port O'Connor, TX O'Connor, Port Mobile AL Bay, Port Isabel, TX Galveston, TX Port O'Connor, TX O'Connor, Port TX O'Connor, Port Galveston, TX Port O'Connor, TX O'Connor, Port Bahia Honda, FL Galveston, TX Port O'Connor, TX O'Connor, Port Veracruz, Mexico Veracruz, Release location Release San Juan, Nicaragua Trinidad, BWI Trinidad, Trinidad, BWI Trinidad, Port O'Connor, TX O'Connor, Port San Juan, Nicaragua Veracruz, Mexico Veracruz, Veracruz, Mexico Veracruz, Hutchinson Island, FL Island, Hutchinson Florida FL Bay, Mexico Veracruz, Bahia Honda, FL Naples, FL Florida FL Bay, Florida FL Bay, St Lucie, FL Mexico Veracruz, 06/16/08 01/20/09 08/15/07 09/08/07 10/20/07 09/05/07 08/04/07 09/08/07 09/08/07 08/04/07 09/06/08 05/27/09 08/04/07 09/08/07 05/30/07 Tagging Tagging date 01/23/09 08/15/07 08/14/07 09/08/07 01/22/09 05/30/07 05/30/07 07/24/07 05/12/07 05/30/07 06/11/07 10/11/07 04/29/07 05/21/07 07/23/07 05/30/07 T-92 T-91 T-77 T-90 T-78 T-76 T-75 T-80 T-79 T-73 T-89 T-81 T-74 T-82 T-72 Tag name T-88 T-83 T-84 T-86 T-87 T-70 T-71 T-64 T-68 T-69 T-62 T-85 T-63 T-66 T-65 T-67 74 76 61 75 67 71 69 87 81 62 73 78 72 65 70 79 82 89 85 90 86 77 66 68 83 88 63 60 80 84 64 Tag count TABLE A1 (Continued) TABLE 312 | LUO et al.

7.04 9.74 5.00 4.70 1.96 0.05 0.01 0.25 0.10 0.00 0.20 0.00 0.25 0.33 0.38 2.82 2.40 8.87 19.75 10.51 10.37 10.95 16.36 Speed Speed (km/day) 38.63 (Continues) 0 0 0 0 0 0 0 0 5.4 5.4 4.1 1.2 0.2 0.9 8.4 79.3 11.5 16.2 30 169.3 237 135.5 726.1 936.7 866.7 630.4 1,145.7 1,391.1 2,977.2 Distance Distance (km) 1,188.4 2086

6 0 6 24 69 21 70 35 89 12 22 33 22 60 54 43 43 182 182 109 127 133 183 134 134 Days at liberty NO YES YES YES YES NO NO NO YES YES NO NO YES Tag recover YES NO YES YES YES YES YES YES YES YES NO NO NO YES YES YES NO NO

13.152 13.181 13.152 12.502 −79.271 −79.998 −76.910 −97.109 −75.821 −96.293 −81.148 −81.324 −81.024 −81.230 −81.512 −81.186 −93.779 −92.511 −54.501 −82.259 −82.252 −82.259 −82.263 −80.650 −80.660 Lon (dd) 29.406 26.760 40.400 −9.352 19.369 25.938 21.883 24.800 −9.024 36.615 24.642 −9.415 26.761 Reporting Lat-Lon Reporting 26.760 no loc no loc 25.294 34.999 33.168 24.973 30.500 Lat (dd) 26.712 25.857 −9.356 no loc no loc 29.490 24.705 32.611 no loc no loc 09/11/08 07/01/08 09/15/09 04/08/09 06/28/09 10/19/09 11/29/09 05/10/08 03/28/08 09/15/08 07/01/08 09/30/09 06/15/09 Reporting Reporting date 07/01/08 not heard not heard 09/15/09 09/15/08 09/15/08 06/17/08 09/16/08 06/30/08 07/13/08 04/17/08 not heard not heard 08/14/08 06/24/08 08/07/08 not heard not heard 09/25/08 09/25/08 09/15/09 03/03/09 09/15/09 02/15/10 03/02/10 09/10/08 05/26/08 09/15/08 09/15/08 09/21/09 09/15/09 Expected Expected date pop-off 09/25/08 09/15/08 09/15/08 09/15/09 09/15/08 09/15/08 09/20/08 09/15/08 09/25/08 09/15/08 05/26/08 09/15/08 03/02/08 03/02/09 09/15/08 09/15/08 09/15/08 09/25/08 13.149 13.125 13.152 13.125 −94.926 −96.288 −96.292 −96.363 −81.014 −81.287 −81.287 −81.016 −81.287 −82.263 −82.252 −82.263 −82.263 −82.263 −82.263 −80.152 −80.750 −80.750 −80.968 −80.750 −80.750 −80.750 −80.750 Lon (dd) −80.750 −80.119 −80.750 −80.750 19.370 19.368 27.171 −9.342 −9.383 −9.345 −9.383 29.171 24.853 24.853 24.658 24.658 24.853 24.853 24.853 24.853 24.853 24.658 24.853 24.853 26.712 26.761 26.712 26.712 26.712 26.712 25.292 25.231 25.291 25.897 28.371 Release Lat-Lon Release Lat (dd) 67.6 37.9 37.9 57.6 57.0 49.5 77.4 79.3 39.3 61.6 41.9 75.0 55.0 52.6 52.0 31.0 70.0 70.7 32.3 50.2 58.3 54.5 54.4 63.6 63.5 60.2 46.2 40.8 38.0 40.8 44.6 Fish, wt (kg) Boca Grande, FL Angola, West Africa St. Lucie, FL Veracruz, Mexico Veracruz, Florida FL Bay, Bahia Honda, FL Whitewater FL Bay, Veracruz, Mexico Veracruz, Boca Grande, FL Port O'Connor, TX O'Connor, Port Boca Grande, FL Angola, West Africa Bahia Honda, FL Florida FL Bay, Angola, West Africa Release location Release Florida FL Bay, Florida FL Bay, Florida FL Bay, Florida FL Bay, Boca Grande, FL Florida FL Bay, Boca Grande, FL South Beach, FL Whitewater FL Bay, Bahia Honda, FL Whitewater FL Bay, Angola, West Africa Florida FL Bay, Florida FL Bay, Galveston, TX Boca Grande, FL 06/09/08 04/02/09 08/20/09 05/25/08 05/04/08 05/19/08 03/17/09 06/04/09 06/15/09 09/20/09 06/09/08 02/24/08 05/19/08 05/11/08 04/01/09 Tagging Tagging date 05/05/08 05/04/08 05/05/08 05/05/08 06/09/08 05/05/08 06/16/08 01/19/08 03/13/09 05/20/08 03/16/09 02/23/08 05/10/08 05/11/08 08/02/08 06/09/08 T-100 T-120 T-124 T-99 T-98 T-117 T-118 T-123 T-122 T-125 T-101 T-97 T-115 T-116 T-119 Tag name T-106 T-105 T-107 T-103 T-102 T-104 T-110 T-96 T-114 T-111 T-121 T-94 T-109 T-113 T-112 T-108 96 95 94 97 91 98 93 92 99 114 119 111 117 110 116 113 118 107 115 112 101 102 105 121 103 106 109 108 120 104 Tag count 100 TABLE A1 (Continued) TABLE LUO et al. | 313

7.08 1.12 5.18 1.76 1.92 1.45 5.30 4.44 0.79 0.13 0.18 4.49 0.50 0.22 0.00 3.46 0.40 6.18 0.04 0.25 2.49 8.55 2.69 14.60 18.98 Speed Speed (km/day) (Continues) 9.6 0 2 0 0 0 0 0 0 4.4 4.6 2.1 67.4 13 55.9 10.6 31.9 31.1 25.4 22.8 22.6 34.6 359.3 151.8 171.9 116.8 255.9 106.2 220 288.8 302.6 Distance Distance (km)

8 5 7 8 2 11 11 11 10 16 69 42 49 95 13 26 49 29 15 50 22 112 101 105 164 Days at liberty NO YES YES YES YES YES YES NO YES NO YES YES NO NO YES NO NO YES Tag recover NO YES NO YES NO YES NO NO NO NO YES NO NO

−97.196 −97.370 −97.108 −81.154 −81.570 −81.379 −82.565 −82.200 −82.083 −82.537 −82.146 −82.360 −82.366 −63.644 −80.900 −80.254 −80.312 −88.216 −80.801 −88.249 −88.304 −80.797 −80.752 −80.986 Lon (dd) −88.136 12.150 17.516 27.629 25.453 17.534 24.800 17.231 no loc 27.623 24.901 27.433 no loc 27.207 26.704 26.500 no loc 26.612 24.820 Reporting Lat-Lon Reporting no loc 26.916 26.404 Lat (dd) 24.837 26.500 24.843 24.759 27.269 24.627 no loc 24.663 no loc 17.647 09/22/09 04/12/10 05/29/10 05/11/10 04/06/10 05/31/10 04/07/10 not heard 09/28/09 01/25/10 09/11/09 not heard 08/18/09 09/08/10 06/20/10 not heard 06/08/10 Reporting Reporting date 07/08/10 not heard 11/07/09 06/10/10 09/29/10 06/10/10 06/20/10 09/19/10 09/16/07 05/15/09 not heard 01/08/11 not heard 10/29/10 01/25/10 09/01/10 09/01/10 09/01/10 08/01/10 09/01/10 08/15/10 02/15/10 01/10/10 01/25/10 12/01/09 07/27/10 02/09/10 09/08/10 08/25/10 02/15/10 09/01/10 Expected Expected date pop-off 09/29/10 02/15/10 02/15/10 09/15/10 09/29/10 09/15/10 09/29/10 09/15/10 SPOT5 SPOT5 02/15/10 01/20/11 02/15/11 02/01/11 −61.698 −96.524 −96.363 −96.362 −96.461 −96.517 −96.607 −96.462 −81.016 −81.016 −81.016 −81.287 −82.108 −82.211 −82.205 −82.205 −82.065 −82.186 −82.060 −88.231 −88.231 −88.237 −80.169 −80.181 −80.758 −80.168 −80.758 Lon (dd) −80.752 −80.752 −80.752 −88.063 17.528 17.528 17.535 17.604 27.169 27.171 27.167 24.828 24.828 24.843 24.843 24.843 24.672 10.707 26.906 26.616 26.514 26.514 26.410 26.956 26.937 25.291 25.291 25.291 28.269 28.371 28.371 28.394 28.262 28.559 28.395 Release Lat-Lon Release Lat (dd) 47.8 67.0 37.9 57.6 49.8 79.8 74.5 41.1 52.0 52.6 65.3 21.3 56.4 28.2 56.9 36.5 56.4 56.4 50.8 50.1 54.8 66.4 83.9 60.9 46.6 60.8 34.1 46.2 30.9 43.5 43.0 Fish, wt (kg) Whitewater FL Bay, Trinidad, BWI Trinidad, Whitewater FL Bay, Port O'Connor, TX O'Connor, Port Belize River, Belize Whitewater FL Bay, Belize River, Belize St. Lucie, FL St. Lucie, FL Florida FL Bay, Belize River, Belize Boca Grande, FL Boca Grande, FL Port O'Connor, TX O'Connor, Port St. Lucie, FL Boca Grande, FL Port O'Connor, TX O'Connor, Port Port O'Connor, TX O'Connor, Port Release location Release Florida FL Bay, Port O'Connor, TX O'Connor, Port Florida FL Bay, Boca Grande, FL Florida FL Bay, Boca Grande, FL Florida FL Bay, Port O'Connor, TX O'Connor, Port Bahia Honda, FL TX O'Connor, Port Long Cay, Belize Hog Island, FL Peace River, FL 04/10/10 08/11/09 04/11/10 09/20/09 03/27/10 04/12/10 03/26/10 08/14/09 08/20/09 06/28/10 03/28/10 05/26/10 05/25/10 09/20/09 08/13/09 05/26/10 09/20/09 08/30/09 Tagging Tagging date 06/27/10 08/29/09 06/09/10 06/03/10 06/09/10 05/26/10 06/10/10 09/08/07 05/13/09 08/29/09 09/10/10 10/05/10 10/05/10 T-142 T-134 T-143 T-139 T-141 T-140 T-133 T-135 T-137 T-144 T-138 T-146 T-145 T-132 T-136 T-147 T-131 T-130 Tag name T-152 T-129 T-150 T-149 T-151 T-148 T-153 T-126 T-127 T-128 T-154 T-155 T-156 141 147 151 142 149 145 131 137 135 152 124 132 139 146 143 148 136 140 127 126 133 128 138 123 129 144 130 134 125 150 122 Tag count TABLE A1 (Continued) TABLE 314 | LUO et al.

7.54 9.53 9.51 1.37 1.70 4.10 0.65 1.82 0.50 0.03 0.00 3.85 0.79 0.48 2.59 2.65 2.04 2.22 19.63 57.75 39.28 39.20 13.75 12.01 58.15 Speed Speed (km/day) (Continues) 0 0 1 0 0 0 0 0.5 8.2 59.5 29.9 85.8 15.4 18.7 15.8 32.8 12.1 249.2 414.5 251.6 116.3 115.5 235.7 252.2 262.2 235.2 685 522.6 136.6 1,138.4 Distance Distance (km)

9 6 6 2 1 2 4 6 8 1 11 75 55 95 21 72 23 58 25 20 46 38 40 118 123 Days at liberty 122 YES YES NO YES NO YES YES YES YES NO YES NO NO YES NO NO NO NO NO NO YES Tag recover YES NO YES NO YES YES YES YES YES

−89.847 −81.550 −81.596 −81.313 −81.016 −81.267 −81.553 −81.301 −81.539 −81.287 −81.924 −81.461 −92.932 −82.002 −82.126 −82.050 −80.327 −80.884 −80.683 −80.676 −80.666 −80.869 −80.074 −88.237 −80.210 Lon (dd) 31.400 27.617 25.857 24.900 26.980 26.940 24.883 24.687 25.882 26.591 25.291 25.883 no loc 25.299 18.608 17.525 29.385 no loc 24.914 24.687 no loc 29.945 25.864 24.658 Reporting Lat-Lon Reporting no loc 24.666 24.644 27.353 Lat (dd) 27.250 no loc 07/08/11 05/19/11 03/28/11 05/29/11 07/31/11 08/29/11 04/08/11 05/02/11 04/04/11 08/25/11 03/16/11 04/04/11 not heard 03/05/11 12/12/14 04/27/11 07/11/11 not heard 05/29/11 12/13/10 not heard 11/11/10 Reporting Reporting date 07/31/11 05/17/11 not heard 01/10/11 01/31/11 02/11/11 02/23/11 01/24/11 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 01/15/12 SPOT5 SPOT5 SPOT5 01/15/12 SPOT5 SPOT5 09/12/11 09/12/11 12/12/14 09/19/11 SPOT5 09/12/11 SPOT5 12/13/10 02/15/12 03/01/11 Expected Expected date pop-off SPOT5 SPOT5 01/10/12 01/10/11 01/17/11 04/05/11 04/05/11 01/24/11 −79.139 −96.365 −81.016 −81.011 −81.130 −81.130 −81.015 −81.287 −81.287 −82.237 −82.158 −82.258 −82.258 −82.237 −82.103 −82.110 −82.060 −82.186 −80.260 −80.260 −80.750 −80.751 −80.751 −84.478 −80.749 −88.229 −88.063 −88.063 −80.234 Lon (dd) −80.233 17.520 17.604 17.604 27.348 27.357 29.899 24.841 24.847 24.847 24.841 24.658 24.658 26.632 26.750 26.711 26.711 26.631 26.892 26.898 26.937 26.956 25.352 25.352 25.291 25.293 25.370 25.360 25.290 28.339 33.181 Release Lat-Lon Release Lat (dd) 47.9 47.9 47.1 87.4 59.5 59.0 49.1 79.8 39.4 89.9 14.9 24.3 45.0 45.6 65.0 45.5 35.4 35.0 70.8 36.0 20.1 53.2 36.5 50.2 54.5 60.5 60.2 43.2 40.9 44.8 Fish, wt (kg) FL FL Broad FL Key, Broad FL Key, Boca Grande, FL Boca Grande, FL Boca Grande, FL Florida FL Bay, Florida FL Bay, Whitewater FL Bay, Boca Grande, FL Florida FL Bay, Florida FL Bay, Ponce de Leon Bay, Ponce de Leon Bay, Belize River, Belize Whitewater FL Bay, Whitewater FL Bay, Port O'Connor, TX O'Connor, Port Apalachicola, FL Apalachicola, Boca Grande, FL Charlotte Harbor, FL Bahia Honda, FL Bahia Honda, FL Charlotte Harbor, FL Long Cay, Belize Peace River, FL Georgetown, SC Release location Release St. Lucie, FL Long Cay, Belize Hog Island, FL St. Lucie, FL 04/28/11 04/27/11 06/15/11 05/23/11 08/21/11 04/09/11 04/02/11 03/19/11 08/14/11 04/02/11 04/02/11 03/01/11 03/01/11 03/18/11 03/15/11 02/28/11 10/15/14 06/03/11 05/23/11 09/09/10 05/17/11 05/16/11 09/10/10 08/17/11 10/05/10 09/17/10 Tagging Tagging date 01/29/11 08/23/11 09/23/10 01/22/11 T-180 T-179 T-181 T-182 T-183 T-176 T-177 T-178 T-171 T-175 T-174 T-170 T-169 T-172 T-173 T-168 T-166 T-187 T-186 T-159 T-184 T-185 T-160 T-165 T-161 T-164 Tag name T-157 T-158 T-162 T-163 174 176 161 175 171 167 169 173 178 172 170 162 179 165 181 159 155 177 182 166 157 168 163 156 160 158 153 164 180 Tag count 154 TABLE A1 (Continued) TABLE LUO et al. | 315

7.85 9.29 5.28 4.27 1.81 0.12 0.02 5.12 0.04 5.50 0.36 0.31 1.77 2.06 0.10 0.24 6.57 1.28 0.53 0.18 0.43 3.00 8.95 59.60 11.01 13.51 18.88 20.04 Speed Speed (km/day) (Continues) 0 4.9 1.3 0.2 5.5 2.8 17.8 19.5 51.2 10.9 20 35.8 18.1 23.8 30 43.3 38.9 48.6 137.3 351.3 811.8 418.3 795.3 281.9 298 539 222.3 1,090.8 440.8 Distance Distance (km) 5 4 0 1 5 10 11 71 10 10 26 55 26 81 45 57 56 29 58 66 22 38 40 43 38 216 139 121 108 Days at liberty NO YES NO NO NO NO NO NO YES NO YES YES NO YES NO YES NO NO YES YES NO YES NO NO Tag recover NO YES NO NO YES −87.181 −89.550 −61.864 −95.049 −61.376 −61.696 −61.533 −61.799 −61.433 −61.723 −61.550 −81.412 −81.829 −81.191 −81.655 −82.068 −82.746 −83.356 −80.118 −80.748 −80.104 −88.233 −80.129 −80.543 −80.112 −88.121 −88.141 −80.136 −80.621 Lon (dd) 25.945 26.934 25.774 24.635 18.615 25.172 17.523 25.812 17.517 26.094 10.644 25.948 28.499 29.944 26.081 30.367 10.458 10.687 10.633 26.689 26.686 29.093 26.761 30.097 10.713 Reporting Lat-Lon Reporting 10.758 10.763 Lat (dd) 28.399 10.233 01/21/12 10/18/12 04/15/14 11/13/12 03/03/15 09/04/12 04/05/12 10/04/12 03/30/13 01/27/12 02/24/12 11/10/11 11/25/11 08/12/11 04/10/12 08/18/11 09/22/11 06/08/12 10/31/11 08/08/12 06/03/11 05/24/13 09/08/12 11/07/11 12/19/11 Reporting Reporting date 11/12/11 09/28/11 06/13/11 10/24/11 03/15/12 04/15/13 02/15/13 04/15/13 03/03/15 09/10/12 08/30/12 04/15/13 09/16/13 02/15/12 05/15/12 03/01/12 02/15/12 02/15/12 08/15/12 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 09/30/11 SPOT5 04/15/13 SPOT5 04/15/12 Expected Expected date pop-off 03/15/12 03/15/12 SPOT5 02/15/12 −79.147 −79.147 −79.147 −79.147 −89.181 −61.709 −61.628 −61.675 −61.628 −61.628 −61.628 −61.628 −61.628 −96.365 −82.109 −82.116 −82.262 −82.108 −82.264 −82.235 −82.233 −80.109 −80.107 −88.403 −88.121 −84.257 −84.257 −80.108 Lon (dd) −80.129 17.534 17.515 29.901 29.901 10.694 10.663 10.702 10.663 10.663 10.663 10.663 10.663 26.909 26.908 26.633 26.876 25.902 25.767 26.667 26.630 26.733 28.339 25.901 25.758 28.960 33.190 33.190 33.190 33.190 Release Lat-Lon Release Lat (dd) 67.1 79.2 77.2 79.2 71.0 45.2 62.7 42.5 72.9 72.9 50.0 50.2 66.0 33.6 58.9 53.3 60.7 33.6 60.7 46.6 83.8 54.3 40.0 54.5 34.6 38.9 64.2 48.5 38.3 Fish, wt (kg) FL FL Miami beach, FL Charlotte Harbor, FL North Miami Beach, Port O'Connor, TX O'Connor, Port Charlotte Harbor, FL Charlotte Harbor, FL Boca Grande, FL Belize River, Belize Belize River, Belize Georgetown, SC Trinidad, BWI Trinidad, North Miami Beach, Georgetown, SC Apalachicola, FL Apalachicola, Georgetown, SC Trinidad, BWI Trinidad, Apalachicola, FL Apalachicola, Trinidad, BWI Trinidad, Trinidad, BWI Trinidad, Boca Grande, FL Venice, LA Boca Grande, FL Boca Grande, FL Georgetown, SC Trinidad, BWI Trinidad, Trinidad, BWI Trinidad, Release location Release Miami, FL Trinidad, BWI Trinidad, Trinidad, BWI Trinidad, 02/03/14 09/08/12 01/20/12 10/15/14 09/08/12 09/08/12 05/19/12 03/26/12 03/25/13 09/28/11 12/29/11 02/14/12 09/28/11 07/21/11 09/28/11 05/10/12 07/23/11 09/18/11 09/16/11 05/19/12 10/20/12 05/24/11 09/08/12 09/30/11 12/08/11 09/18/11 Tagging Tagging date 06/08/11 09/18/11 09/16/11 T-210 T-211 T-213 T-212 T-209 T-207 T-208 T-206 T-205 T-204 T-200 T-202 T-203 T-201 T-199 T-197 T-196 T-198 T-195 T-194 T-215 T-193 T-214 T-216 T-192 T-191 Tag name T-188 T-190 T-189 195 196 194 197 191 198 211 193 192 199 210 187 190 207 189 185 201 186 183 205 188 202 206 208 204 203 200 184 209 Tag count TABLE A1 (Continued) TABLE 316 | LUO et al.

7.45 4.42 5.21 5.31 4.20 0.66 0.00 3.10 3.82 6.19 0.07 2.11 0.20 3.54 8.55 2.56 3.17 6.48 49.71 39.18 11.67 22.90 46.37 Speed Speed (km/day) (Continues) 9.3 0 0 0 0 0 5 0 0 0.1 8.6 29.4 94 53 28.2 20.8 36.5 22.9 43.6 417.3 447.4 195.9 521.7 105.4 501.8 563.8 634.6 346.6 460.1 440.4 Distance Distance (km)

9 3 7 9 5 1 7 0 0 5.875 11 11 67 13 87 70 56 12 50 63 83 43 145 166 221 Days at liberty NO NO YES YES NO YES YES NO NO YES NO NO NO NO NO YES NO NO YES NO NO NO Tag recover NO YES NO YES NO NO NO NO

−79.972 −97.157 −89.114 −89.420 −89.269 −89.945 −61.979 −95.488 −61.761 −61.761 −96.370 −81.493 −81.277 −85.001 −82.181 −82.011 −86.315 −83.980 −80.141 −84.326 −84.606 −88.826 −80.109 −80.130 −88.270 Lon (dd) 30.019 28.685 24.795 26.884 29.743 25.908 25.772 no loc 28.379 27.674 26.792 no loc no loc 29.669 10.639 no loc 30.149 no loc 26.089 29.093 30.312 29.611 17.330 29.271 Reporting Lat-Lon Reporting Lat (dd) 29.858 27.071 29.900 29.270 10.691 10.691 06/30/15 10/02/12 11/27/12 09/20/12 07/04/12 03/22/14 09/16/12 not heard 09/10/12 11/10/12 06/26/15 not heard not heard 06/28/12 06/03/12 not heard 06/30/12 not heard 01/25/12 07/27/12 06/19/12 06/21/12 08/16/14 Reporting Reporting date 12/07/12 10/05/13 04/17/13 10/24/12 07/17/12 12/29/11 12/08/11 01/10/16 02/05/13 SPOT5 SPOT5 SPOT5 SPOT5 09/15/12 03/15/13 03/15/13 03/15/13 11/15/15 12/15/12 12/15/12 SPOT5 SPOT5 01/10/13 SPOT5 09/15/12 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 Expected Expected date pop-off SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 −79.176 −61.726 −61.761 −61.761 −96.365 −96.380 −96.390 −96.380 −82.098 −82.146 −82.233 −82.233 −84.503 −84.318 −84.481 −84.503 −80.088 −80.142 −84.481 −84.329 −84.481 −84.328 −84.334 −84.446 −80.117 −80.109 −88.235 −84.503 −84.599 Lon (dd) −80.757 17.533 29.871 29.915 29.896 29.871 29.896 29.947 29.896 29.921 29.910 29.908 29.871 29.861 24.856 10.689 10.691 10.691 26.861 26.875 26.733 26.733 25.884 25.773 25.896 25.902 28.339 28.326 28.297 28.326 33.168 Release Lat-Lon Release Lat (dd) 19.3 37.8 87.3 69.4 49.7 49.7 49.9 51.1 61.8 41.2 55.8 52.6 81.3 52.5 62.9 62.2 58.9 36.9 63.8 66.0 43.1 43.7 30.1 46.4 43.7 43.2 43.0 48.1 48.2 Fish, wt (kg) 44.0 FL FL Port O'Connor, TX O'Connor, Port Charlotte Harbor, FL Apalachicola, FL Apalachicola, Port O'Connor, TX O'Connor, Port FL Apalachicola, Port O'Connor, TX O'Connor, Port Port O'Connor, TX O'Connor, Port Apalachicola, FL Apalachicola, Miami Beach, FL Miami, FL Boca Grande, FL Apalachicola, FL Apalachicola, Boca Grande, FL North Miami Beach, Apalachicola, FL Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Trinidad, BWI Trinidad, North Miami Beach, Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Belize River, Belize Release location Release Boca Grande, FL Georgetown, SC Florida FL Bay, Apalachicola, FL Apalachicola, BWI Trinidad, Apalachicola, FL Apalachicola, Trinidad, BWI Trinidad, 09/21/12 09/07/12 06/21/15 09/21/12 06/14/12 09/07/12 09/21/12 06/22/12 01/14/14 07/15/12 09/08/12 05/01/15 09/08/12 03/23/12 06/14/12 06/21/12 06/21/12 05/23/12 01/19/12 06/14/12 06/14/12 06/14/12 08/15/14 Tagging Tagging date 09/08/12 09/15/12 07/29/12 05/13/13 12/29/11 05/08/12 12/08/11 T-239 T-240 T-238 T-236 T-241 T-237 T-235 T-242 T-243 T-232 T-233 T-234 T-231 T-229 T-228 T-246 T-247 T-227 T-224 T-225 T-226 T-244 T-245 Tag name T-219 T-223 T-218 T-217 T-222 T-220 T-221 241 219 214 217 216 213 218 231 237 224 235 232 215 212 226 227 239 236 221 240 233 228 223 229 234 230 238 225 222 220 Tag count TABLE A1 (Continued) TABLE LUO et al. | 317

9.43 5.94 0.21 1.44 5.94 1.43 0.50 4.34 5.74 1.33 0.11 1.30 0.36 0.16 0.77 0.53 0.40 0.41 6.95 2.95 2.76 2.14 11.91 10.81 10.33 16.25 16.37 13.20 Speed Speed (km/day) (Continues) 9.3 0 0 4.8 0.5 5.4 6.5 3.9 87.7 39.6 18 15 13.8 26.6 20.9 86.5 33.1 38.9 451.6 129.8 439.9 361.4 491 278 178.7 677 216.9 160 498.9 422.6 Distance Distance (km)

0 1 8 3 5 7 3 7 74 61 23 26 94 35 15 23 23 58 58 20 63 84 30 40 44 111 118 112 104 Days at liberty NO YES YES NO YES NO NO YES YES NO NO NO NO NO NO NO NO Tag recover YES YES YES NO NO NO NO YES YES NO YES YES NO

−97.209 −89.552 −89.515 −89.152 −89.404 −81.023 −81.172 −81.086 −96.290 −96.419 −96.411 −96.375 −96.522 −81.223 −81.471 −81.797 −85.372 −85.408 −82.942 −82.041 −82.472 −86.153 −80.995 −80.339 −80.083 −88.948 −88.871 −88.075 −88.262 Lon (dd) 25.413 27.321 25.270 25.281 29.002 29.686 29.807 26.929 26.504 30.446 25.366 no loc 29.071 29.206 29.680 17.443 29.492 17.424 Reporting Lat-Lon Reporting 25.296 Lat (dd) 26.252 27.834 28.964 25.839 28.444 28.402 28.299 26.687 28.361 26.522 28.377 04/30/13 06/06/14 04/09/13 04/11/13 07/24/15 06/26/15 07/23/13 06/23/14 07/01/13 05/09/13 06/04/13 not heard 08/07/14 07/18/14 06/04/15 10/15/15 09/05/13 Reporting Reporting date 07/17/13 04/30/13 02/06/14 08/28/13 10/22/13 10/21/12 09/22/14 09/30/12 09/14/12 12/11/12 09/24/12 10/14/12 09/28/12 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 12/15/15 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 Expected Expected date pop-off SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 −81.172 −81.081 −96.365 −96.380 −96.381 −96.380 −96.380 −96.380 −81.087 −82.643 −82.683 −82.098 −82.111 −80.977 −80.755 −84.503 −84.503 −84.503 −84.503 −84.411 −84.503 −84.503 −80.972 −84.503 −84.503 −88.056 −88.056 −88.121 −80.994 Lon (dd) −80.113 17.630 17.630 17.514 27.578 27.566 29.871 29.871 29.871 29.871 29.890 29.871 29.871 29.871 29.871 24.852 25.423 25.270 25.282 26.861 26.896 25.351 25.421 25.415 25.897 28.339 28.326 28.340 28.326 28.326 28.326 Release Lat-Lon Release Lat (dd) 59.0 59.0 59.1 59.1 59.1 49.1 49.1 49.6 49.5 77.3 51.2 51.1 24.1 55.8 45.4 52.6 45.3 52.7 62.8 62.5 36.9 36.3 36.4 54.5 54.2 64.2 43.9 54.8 64.2 Fish, wt (kg) 44.0 FL FL Tarpon FL Bay, Florida FL Bay, Ponce de Leon Bay, Mud FL Bay, Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Shark River, FL Tarpon FL Bay, Apalachicola, FL Apalachicola, Long Cay, Belize Long Cay, Belize Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Belize River, Belize Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Release location Release Tarpon FL Bay, North Miami Beach, Boca Grande, FL Port O'Connor, TX O'Connor, Port Tampa, FL Tampa, Tampa, FL Tampa, Port O'Connor, TX O'Connor, Port Boca Grande, FL TX O'Connor, Port Port O'Connor, TX O'Connor, Port Port O'Connor, TX O'Connor, Port TX O'Connor, Port 04/07/13 04/27/14 04/09/13 04/10/13 05/01/15 06/18/15 05/19/14 03/11/13 05/23/13 04/07/13 04/24/13 08/18/15 08/21/15 05/25/14 06/18/14 03/28/13 05/09/15 05/24/13 Tagging Tagging date 04/10/13 02/03/14 09/07/12 09/17/14 06/26/13 06/26/13 09/07/12 09/08/12 09/07/12 09/21/12 09/21/12 09/21/12 T-272 T-273 T-271 T-270 T-269 T-268 T-265 T-267 T-264 T-266 T-263 T-276 T-262 T-261 T-260 T-259 T-275 T-277 Tag name T-274 T-258 T-248 T-257 T-255 T-256 T-252 T-249 T-254 T-253 T-250 T-251 261 247 267 251 242 249 245 271 269 265 262 259 270 252 255 266 257 246 263 243 248 268 260 256 258 264 253 254 244 Tag count 250 TABLE A1 (Continued) TABLE 318 | LUO et al. 7.92 7.93 9.40 9.61 1.26 4.70 4.48 1.16 0.75 0.27 0.20 6.57 2.58 2.70 8.70 8.00 14.28 13.30 13.58 15.44 28.70 Speed Speed (km/day) 4.8 37.6 37.6 59.1 77.8 13.3 15.6 43 28.7 58.3 30.1 451.2 339.4 229.9 594 214.2 494 923.2 666.7 Distance Distance (km) 1,127.9 1,066.4 4 9 1 8 1 24 75 13 26 57 32 39 15 29 68 34 141 111 167 104 258 Days at liberty NO NO NO NO YES NO NO NO NO NO NO NO Tag recover NO YES NO NO NO NO NO NO NO −89.525 −95.678 −96.702 −96.818 −96.599 −96.672 −96.237 −96.636 −96.521 −81.167 −85.305 −92.829 −82.604 −83.943 −83.930 −84.910 −84.342 −80.878 −80.052 −88.087 −88.711 Lon (dd) 26.862 29.714 29.947 29.107 28.178 29.878 25.427 28.889 17.598 27.989 28.184 30.039 Reporting Lat-Lon Reporting 24.820 26.887 28.170 28.136 28.379 Lat (dd) 28.212 18.738 20.047 18.780 06/19/18 07/20/18 10/04/16 07/14/17 Reporting Reporting date 09/16/16 09/13/14 11/15/14 06/17/16 09/14/15 09/26/14 09/12/16 07/26/14 01/20/16 05/24/16 12/01/16 09/19/16 10/16/13 09/17/14 03/02/15 11/27/14 01/09/15 12/10/18 SPOT5 SPOT5 SPOT5 Expected Expected date pop-off SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 SPOT5 09/01/16 12/01/16 SPOT5 SPOT5 SPOT5 03/01/15 SPOT5 03/01/15 −96.365 −96.365 −96.365 −96.365 −96.365 −96.365 −96.365 −96.365 −96.365 −84.503 −84.503 −84.503 −84.503 −84.503 −84.503 −84.503 −84.503 −84.503 −84.503 −88.056 −80.739 Lon (dd) 17.630 29.871 29.871 29.871 29.871 29.871 29.871 29.871 29.871 29.871 29.871 24.916 28.339 28.339 28.339 28.339 28.339 28.339 28.339 28.339 28.339 Release Lat-Lon Release Lat (dd) 47.7 57.3 59.1 59.1 59.1 49.9 49.9 77.2 99.0 51.0 41.3 52.2 45.7 52.6 62.5 54.5 54.5 66.0 66.0 80.1 64.2 Fish, wt (kg) Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Apalachicola, FL Apalachicola, Long Cay, Belize Apalachicola, FL Apalachicola, Port O'Connor, TX O'Connor, Port Florida FL Bay, Release location Release Port O'Connor, TX O'Connor, Port Apalachicola, FL Apalachicola, FL Apalachicola, FL Apalachicola, Port O'Connor, TX O'Connor, Port Port O'Connor, TX O'Connor, Port Port O'Connor, TX O'Connor, Port FL Apalachicola, Apalachicola, FL Apalachicola, Port O'Connor, TX O'Connor, Port Port O'Connor, TX O'Connor, Port Apalachicola, FL Apalachicola, Port O'Connor, TX O'Connor, Port Port O'Connor, TX O'Connor, Port 05/11/18 06/16/18 06/22/16 08/21/15 06/12/17 09/11/16 04/25/16 Tagging Tagging date 09/12/16 07/18/14 09/01/14 06/02/16 09/11/16 09/17/14 10/15/13 07/13/14 05/07/15 08/22/14 10/12/14 06/17/16 09/20/14 09/20/14 T-304 T-302 T-296 T-297 T-300 T-299 T-281 Tag name T-294 T-287 T-290 T-293 T-292 T-288 T-289 T-291 T-298 T-278 T-282 T-279 T-286 T-284 274 276 275 273 278 281 287 272 279 291 282 285 289 292 277 286 283 290 288 Tag count 284 280 PAT tagsNote: are PAT indicated by the expected pop-off date and SPOT tags are indicated by SPOT5 in the same column. Reporting date and locationlocation, refer to or the the point last tag at which location transmitted a PAT of SPOT tag transmission its first while still attached to the fish. Abbreviations: Pop-up Archival PAT, Transmitting; Smart SPOT, Position and Temperature. TABLE A1 (Continued) TABLE