Journal of Coastal Research 00 0 000–000 Coconut Creek, Month 0000 OccupancyPatternsofGulfSturgeon, oxyrinchus desotoi, Associated with Ship Island, Mississippi Page E. Vick†‡, Mark S. Peterson†, William T. Slack§, and Paul O. Grammer†

†Division of Coastal Sciences §U.S. Army Engineer Research and Development Center School of Ocean Science and Technology Waterways Experiment Station EE-A The University of Southern Mississippi Vicksburg, MS 39180, U.S.A. Ocean Springs, MS 39564, U.S.A.

ABSTRACT

Vick, P.E.; Peterson, M.S.; Slack, W.T., and Grammer, P.O., 0000. Occupancy patterns of Gulf , desotoi, associated with Ship Island, Mississippi. Journal of Coastal Research, 00(0), 000–000. Coconut Creek (Florida), ISSN 0749-0208.

In order to reduce wave energy associated with tropical storms and hurricanes in the western Mississippi Sound, the Mississippi Coastal Improvements Program will close a 5.6 km breach (i.e. Camille Cut) separating East and West Ship Island, Mississippi, and restore sediments to the southern shoreline of East Ship Island. As part of this program, federally designated critical habitat associated with Ship Island was monitored to establish baseline patterns of use prior to island restoration using a passive acoustic array during overwintering periods from 2011 to 2014. Gulf Sturgeon of both western and eastern populations occupied the passes, cuts, and ends of Ship Island over the study period, but no difference in occupancy was observed among four initially established zones. However, when comparing the new most eastern zone, Dog Keys Pass, to the four initial zones, occupancy was four times greater in this zone than any other monitored zone, indicative of foraging behavior associated with high abundance of benthic resources. Reconnecting East and West Ship Island through filling Camille Cut proper may increase Gulf Sturgeon occupancy in Dog Keys Pass and other areas near the ends and passes of the barrier islands. The loss of Camille Cut proper as critical habitat may be partially augmented by ‘‘new’’ nearshore habitat north and south of Camille Cut; however, it is uncertain whether filling in Camille Cut will alter the physical components of the habitat, causing a shift in benthic prey availability and, thus, quality of forage habitat.

ADDITIONAL INDEX WORDS: Critical habitat, overwintering, anadromous, habitat restoration, threatened species, acoustic telemetry.

INTRODUCTION work brought the channel to the current authorized dimen- The barrier islands of the Mississippi Sound have been sions of 122 m wide and 11.6 m deep (Byrnes et al., 2012). The heavily impacted by natural events but have partially Pascagoula Channel has been dredged from Horn Island Pass recovered due to westward littoral drift, which maintains and to the city of Pascagoula since 1880; currently, the channel is replenishes the barrier islands. Winter storms and hurricanes 137.2 m wide and 13.4 m deep (E. Godsey, U.S. Army Corps of have contributed to island narrowing, lateral movement, and Engineers [USACE] Mobile District, personal communication). breakup of the islands in Mississippi Sound (Morton, 2008), Channel dredging can limit sediment availability and compo- including Ship Island. Ship Island did not naturally recover sition and can limit down drift sediment transport in some from Hurricane Camille, which separated Ship Island into Mississippi Barrier Islands (Morton, 2008). However, Ship distinct east and west segments in 1969, and the island Island Pass is at the end of the littoral sand transport along the segments were further separated in 2005 by Hurricanes Cindy, barrier island shorelines, and, therefore, sand placement was Katrina, and Rita (Morton, 2008; USACE, 2016). not considered a net loss to the natural transport system from Anthropogenic activities have also significantly contributed channel dredging in the Gulfport Ship Channel (Byrnes et al., to habitat loss due to erosion and alteration of the barrier 2012; J. McDonald, USACE Mobile District, personal commu- islands, including dredging shipping channels in the tidal nication). inlets of the Mississippi Sound as well as storm events (Morton, Repeated storm events and dredging have contributed to 60% 2008). The Gulfport Ship Channel has been dredged from the land loss of Ship Island (Morton, 2008); thus, the USACE Ship Island Pass to the city of Gulfport since 1899; the original Mobile District was tasked by Congress in 2009 to restore the ship channel was 90 m wide and 5.7 m deep and has been Mississippi barrier islands (USACE, 2016). One of the widened several times (Morton, 2008). In 2012, new dredging components of the restoration, as developed by the USACE Mississippi Coastal Improvements Program (MsCIP) team, DOI: 10.2112/JCOASTRES-D-17-00027.1 received 8 February 2017; accepted in revision 12 August 2017; corrected proofs received was the restoration of Ship Island by filling in a 5.6 km wide 21 September 2017; published pre-print online XX Month XXXX. breach in the island known as Camille Cut and by restoring the ‡Current address: Independent Monitoring Program, southern shoreline of East Ship Island (see figure ES-1 in Fish and Wildlife Research Institute, Florida Fish and Wildlife USACE, 2016). It is anticipated that the restored barrier Conservation Commission (FWC), St. Petersburg, FL 33701, U.S.A.; [email protected] islands will increase coastal protection from wave energy ÓCoastal Education and Research Foundation, Inc. 2017 associated with tropical storms and hurricanes in the western

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:39 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 1 0 Vick et al.

Mississippi Sound (USACE, 2016). Since waters around Ship Island are important overwintering habitat for Gulf Sturgeon (Acipenser oxyrinchus desotoi; Figure 1; Rogillio et al., 2007; Ross et al., 2009), understanding their use of this assumed foraging area before and after the Ship Island restoration is needed to best manage the population’s recovery and mitigate any impacts to the species. Gulf Sturgeon are listed as a threatened species under the Endangered Species Act (USFWS, 1991) and endangered in Mississippi (MMNS, 2014). Without safeguards placed on critical habitats (Manson and Hogarth, 2003), the conservation and recovery of Gulf Sturgeon would be impacted. Life history characteristics combined with anthropogenic and environmen- tal factors, such as habitat loss, dams/sills, coastal develop- ment, overfishing, and storm events (e.g., hurricanes; Ahrens and Pine, 2014; Flowers et al., 2009; Rudd et al., 2014) have played a role in declining population size among sturgeon species (Cooke, Paukert, and Hogan, 2012; Haxton, Sulak, and Figure 1. Gulf Sturgeon swimming near tagging location on the Pascagoula Hildebrand, 2016; IUCN, 2010; Nelson et al., 2013), including River, Mississippi. Photo courtesy of Michael Andres. (Color for this figure is available in the online version of this paper.) Gulf Sturgeon. In particular, the Pearl and Pascagoula river systems (i.e. western population) have the lowest Gulf Sturgeon population numbers (Ahrens and Pine, 2014; Morrow observed that eastern fish migrated much further west of et al., 1999; Rudd et al., 2014) among the eight core drainages Mobile Bay, with at least 23 adult Gulf Sturgeon traveling to (described in USFWS and NMFS, 2009). Therefore, preserving the Ship Island vicinity to overwinter, where presumed feeding spawning and foraging habitats protect threatened and occurred. endangered species from further decline (McCauley et al., Gulf Sturgeon from the eastern population segment over- 2015; Wilcove et al., 1998) and may aid in the population winter in habitats that are characterized as nearshore shallow recovery of anadromous Gulf Sturgeon (Ahrens and Pine, 2014; areas (2 to 4 m) with sandy bottoms (Fox, Hightower, and Havrylkoff, Peterson, and Slack, 2012; Peterson et al., 2013). Parauka, 2002), which is prevalent at the Mississippi barrier Overwintering occurs in the fall with fish emigrating from islands (Ross et al., 2009). In Ross et al. (2009), only western natal rivers to more saline waters in mid-to-late September population Gulf Sturgeon were detected using shallow, sandy- through mid-November (Havrylkoff, Peterson, and Slack, bottom areas surrounding the Mississippi barrier islands; no 2012; Heise et al., 2005; Peterson et al., 2016). Large subadult Gulf Sturgeon were detected in the middle of the Mississippi and adult Gulf Sturgeon are benthic cruisers and are found in Sound or at inshore, nonisland sites. Ross et al. (2009) suggests marine foraging areas during overwintering periods (Carr, that this lack of Gulf Sturgeon detections may be due to quick 1983; Huff, 1975; Mason and Clugston, 1993), which are offshore movement of the fish; however, the lack of detections characterized as shallow, sandy bottoms (Fox, Hightower, and may be due to the limitations of manual tracking techniques. In Parauka, 2002), with clear and well-oxygenated waters (Ross et contrast, Peterson et al. (2013) documented high occupancy al., 2009). Previous acoustic studies showed greater detection nearshore in the western distributary of the Pascagoula River events of Gulf Sturgeon in areas with higher feeding potential estuary compared to the urbanized eastern distributary, and (Brooks and Sulak, 2005; Fox, Hightower, and Parauka, 2002; these habitats had finer grain size characteristics compared to Harris, Parkyn, and Murie, 2005; Peterson et al., 2013). Mississippi barrier islands and eastern population segment Previous manual tracking, a much more laborious method, habitats (Brooks and Sulak, 2005; Fox, Hightower, and Para- detected Gulf Sturgeon tagged in the Pearl and Pascagoula uka, 2002; Harris, Parkyn, and Murie, 2005; Ross et al., 2009; rivers using the barrier island passes of the Mississippi Sound, Sulak et al., 2009). Therefore, the nearshore and pass habitats but fish from the rivers east of Mobile Bay (i.e. eastern associated with Mississippi barrier islands are more like the population) were not detected (Rogillio et al., 2007; Ross et al., nearshore areas of the eastern populations habitats (see 2009). While intermittent and limited, Gulf Sturgeon move- Peterson et al., 2013). Since Gulf Sturgeon are characterized ment has occurred between natal rivers within population as benthic cruisers, it is presumed that the fish are more segments (Dugo et al., 2004); previous passive tracking studies concentrated in the sandy nearshore areas of islands or in the rarely detected eastern population Gulf Sturgeon within the passes due to a high availability of prey items (Fox, Hightower, designated feeding habitat of the Pascagoula River estuary and Parauka, 2002; Parauka, Duncan, and Lang, 2011; (Havrylkoff, Peterson, and Slack, 2012; Parauka, Duncan, and Peterson et al., 2013; Ross et al., 2009). Herein, the occupancy Lang, 2011). A compilation of data for the Deepwater Horizon ‘‘patterns’’ of Gulf Sturgeon around Ship Island and associated injury assessment did note Gulf Sturgeon from as far east as passes and cuts were examined during these overwintering Escambia Bay (Escambia, Blackwater, and Yellow rivers) and periods (mid-September to mid-April) prior to MsCIP restora- the Choctawhatchee River spent a considerable amount of time tion using a passive acoustic telemetry array (see Appendix A west of Mobile Bay, especially near Dauphin Island and the for all zones and scenarios). The first objective of this study was mouth of the Pascagoula River (USFWS, 2015). This study to quantify baseline (prerestoration) occupancy index (OI)

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:39 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 2 Gulf Sturgeon Occupancy 0

‘‘patterns’’ (sensu Peterson et al., 2013) of Gulf Sturgeon for the Choctawhatchee; based on core river individuals, no associated with waters around Ship Island (Scenario 1). The clear spatial structure of occupancy was observed within any second objective was to evaluate if the OI is sensitive to changes zone or during any monitoring period (Vick, 2016). in receiver number and area of coverage by the acoustic array (Scenario 2). Ship Island Acoustic Array Each Vemco VR2W receiver was attached to a 152.4 cm METHODS length 3 3.81 cm diameter conduit aluminum pipe, which was secured to a CC-3 Mooring buoy (Polyform U.S.) with stainless Gulf Sturgeon were captured and acoustically tagged steel (SS) hardware, and anodes were attached to the top and annually in the Pearl and Pascagoula rivers, from early bottom of the aluminum pipe for corrosion control. The October to late November. In both systems, Gulf Sturgeon receivers were attached to the bottom of the pipe so that the were captured with anchored multifilament (60.9 3 3.0 m, 20.3 cm bar mesh or 45.7 3 3.0 m, 12.7 cm bar mesh) and receivers could pivot with the movement of the buoy (Peterson monofilament (71.0 m 3 2.4 m, 5.1 cm bar mesh) gill nets set et al., 2013, 2016; Sulak et al., 2009). Attached to the pipe was a parallel and perpendicular to the flow (Peterson et al., 2016; bridle composed of 9.53 mm SS cable and SS hardware; the Rogillio et al., 2007). Nets were checked every 2 hours, and Gulf bridle was connected to a SS cable (9.53 mm) that was anchored Sturgeon were weighed (nearest 0.1 kg), measured for fork and to a 68 kg concrete block. Cable length at each buoy mooring total length (FL and TL, mm), and assessed for external tags was three times greater than water depth to provide adequate and internal passive integrated transponder (PIT) tags. New scope. The detection radius of each Vemco VR2W receiver was captures were tagged with T-bar and PIT tags as described by assumed to be 300 m, based on range tests conducted in the Heise et al. (2004). Subadult fish (891–1250 mm FL) were nearshore sandy bottom habitats off Pensacola, Florida tagged externally at the base of the dorsal fin (Havrylkoff, (Robydek and Nunley, 2011). Detections of the acoustic tags Peterson, and Slack, 2012; Sulak et al., 2009) with uniquely were recorded and stored in the receiver until downloaded, coded low-powered acoustic tags (either Model V9-2 L or V13-1 which occurred monthly. Each detection was linked to an L; 69 kHz; 90 s mean random delay; Vemco, Halifax, Nova individual Gulf Sturgeon by the unique code of the Vemco Scotia). Adult fish (.1251 mm FL) were tagged internally acoustic tag. Using the NOAA Gulf Sturgeon database (https:// (Baremore and Rosati, 2013; Moser et al., 2000; USFWS, 1991) grunt.sefsc.noaa.gov/gsp/index.jsp), the river of origin, original with high-powered uniquely coded and coated (clear platinum measurements (FL, TL, and weight), date tagged, and owners silicone elastomer) V16-6H acoustic tags (69 kHz; 90 s mean of each transmitter (i.e. fish) were identified based on unique random delay). Betadyne-impregnated petroleum jelly was transmitter numbers downloaded from the VR2W receivers. used on all tagging wounds. Both the Pearl and Pascagoula Transient Gulf Sturgeon identities, measurements, and acous- rivers empty into the Mississippi Sound, where the tagged tic transmitter specifications were confirmed or provided by the large subadult and adult Gulf Sturgeon were detected tag owners. overwintering on the offshore Ship Island acoustic array from Occupancy Data mid-November to mid-April. Detailed descriptions of both core Data from the Ship Island receiver array was organized so rivers and specific tagging areas can be found in Partyka and that individual OI values could be calculated to estimate Peterson (2008), Peterson et al. (2007, 2016), and Rogillio et al. occupancy patterns for Scenarios 1 and 2 (e.g., configuration of (2001, 2007). receivers within zones over the monitoring periods). Using the Annually, the acoustic receivers (Vemco VR2W; 69 kHz, compiled detection data of both western and eastern population Halifax, Nova Scotia) of the Ship Island array were deployed in segments, variability in Gulf Sturgeon OI patterns was early-to-mid September and removed in mid-to-late June analyzed by zone within and among monitoring periods for (hereafter called monitoring periods) to avoid hurricane all fish detected within the array (Peterson et al., 2013, 2016). season; therefore, no detections were recorded from mid-June To examine OI patterns as the array area and number of to mid-September. Initially, 21 receivers were deployed in four receivers increased over the course of this study, OI patterns of zones (1 to 4) between the west end of West Ship Island through Scenarios 1 and 2 were directly compared to establish whether Camille Cut to the east end of East Ship Island in fall 2011 the OI was sensitive to these changes and therefore whether (Figure 2A; Appendix A). With the addition of eight receivers in the OI would be useful in future acoustic studies. 2012, Dog Keys Pass, the area between East Ship Island and Horn Island, was added to the passive acoustic array as zone 5 Telemetry Data Organization (Figure 2B). While the exact position of each receiver changed Individual Gulf Sturgeon were considered to be present slightly between monitoring periods, the number and relative within the array if the fish was detected at least two times on a position of each receiver within the array remained constant. single receiver on the same date (Peterson et al., 2016). Conservative estimates of distance (in kilometers) from the Individual Gulf Sturgeon detected during multiple monitoring closest receiver in any zone to the closest receiver in the next periods were treated as new individuals annually when zone (not including land) were determined with ArcMap and compiling detection data and calculating OI values over the ranged from 0.54 to 3.24 km (x¯ 6 1 standard deviation [SD]; course of the study as annual movement patterns to offshore 2.30 6 1.90). Furthermore, the conservative distance (in barrier islands were considered to be independent behaviors. kilometers) between the nearest Ship Island receiver and an Prior to data analysis, detections from each monitoring period eastern or western core river receiver (emigration location) were sorted chronologically and then by transmitter. Following ranged from 30.57 6 9.89 for the Pascagoula to 261.08 6 23.60 data organization, the time between successive detections for

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:39 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 3 0 Vick et al.

Figure 2. Ship Island acoustic array for (A) Scenario 1 and (B) Scenario 2. Note: Scenario 1 includes zones 1 to 4 for monitoring periods 1 to 3 (2011–2014). Scenario 2 included zones 1 to 5 for monitoring periods 2 to 3 (2012–2014).

an individual tag was calculated during each monitoring period detections for each individual Gulf Sturgeon within a partic-

(Peterson et al., 2013, 2016). Duplicate detections and ular zone (xi) to obtain effort-adjusted detections (EADs, simultaneous detections were removed from the database prior Equation [2]; Peterson et al., 2013). These EADs were then to analysis. Duplicate detections are detections made on a normalized (Z scores) for each fish by zone and monitoring single receiver from the same transmitter with no time period (Equation [3]) using a global mean EAD (x¯ g) and global between successive detections, whereas simultaneous detec- standard deviation EAD (SDg), which were calculated from the tions are detections made on two or more receivers when the total number of detections recorded for each individual Gulf time between successive detections is less than the minimum Sturgeon on the entire Ship Island acoustic array for a given tag interval minus 10 seconds (Peterson et al., 2013, 2016). If monitoring period (Peterson et al., 2013). Afterward, these OI there were several simultaneous detections in a row, the values were scaled by adding the absolute value of the lowest receiver assignment of the retained detection was determined OI (Z score) to each OI value, making the lowest scaled OI value by the last valid detection within the detection record (i.e.the zero (Peterson et al., 2013). This OI was used to compare last receiver to record a valid single detection before the group occupancy among zones and monitoring periods. A high OI of simultaneous detections). If there were no valid records prior indicates that a particular zone has the greatest mean number to the simultaneous group, then the first detection of that group was considered a valid detection and the remaining simulta- of EADs out of all zones for that monitoring period (Peterson et neous detections were removed (Peterson et al., 2016). al., 2013). Occupancy Equations: Occupancy Index Calculations Following data organization and false detection removal, the No: of receivers in zone w ¼ 1 ð1Þ OI values were generated using several calculations. First, an total No: of receivers effort-adjusted weighting value (w) was calculated for each zone (Equation [1]) and applied to the number of acoustic tag effort-adjusted detections ¼ w 3 xi ð2Þ

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:39 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 4 Gulf Sturgeon Occupancy 0

Âà Z score ¼ðw 3 xiÞx¯g =SDg ð3Þ

Data Processing and Analysis A two-way analysis of variance (ANOVA) was used to compare the calculated OI values by zone (n ¼ 4), monitoring period (n¼3), and the interaction term for Scenario 1 as well as zone (n¼5), monitoring period (n¼2), and the interaction term for Scenario 2. If normality and homogeneity of variance assumptions were not met for these data sets, then ANOVA was considered robust to assumption violations (Field, 2013; Underwood, 1997) and robust enough to run the effort- adjusted, normalized, and scaled OI values because the data were balanced and large (see Appendix B). If a significant main effect was found with no interaction effect, a Sidak (homoge- nous variances) or Games-Howell (heterogeneous variances) post hoc test was used to statistically separate mean OI zone values for each main effect. Additionally, linear regression was Figure 3. Occupancy index (6SEM) by zone and monitoring period for used to examine the strength and predictability of the Scenario 1 (2011–2014). Note: See Figure 2 and Appendix A for descriptions relationship between the Gulf Sturgeon OI values and EADs of zones. by zone for Scenarios 1 and 2. All data calculations were completed with Excel, and all analysis with IBM SPSS (version detected (Figure 2A) with 25 fish from the western population 23; IBM, Armonk, New York, U.S.A.). (1040–1700 mm FL) and 19 from the eastern population (1290– Occupancy patterns of Gulf Sturgeon surrounding Ship 1880 mm FL; Appendix C). Twelve fish were detected during Island were calculated using two different scenarios. The two multiple monitoring periods, and, because returning Gulf calculations tested the flexibility and robustness of the OI by Sturgeon were characterized as unique individuals, the total accounting for potential increases or decreases in total number of detected individuals for Scenario 1 was 59 (Appendix detections within the deployment monitoring period relative C). Nine individuals were detected in the study area over two to increasing the area (number of receivers and zones) of the monitoring periods, and three were detected over three acoustic array field. For Scenario 1, OI values (Peterson et al., monitoring periods. During monitoring periods 2 to 3 in zones 2013) were calculated for zones 1 through 4 as they were 1 to 5 (Scenario 2), 45 Gulf Sturgeon were detected (Figure 2B); consistently deployed for monitoring periods 1 through 3 22 fish were from the western population (1040–1700 mm FL), (2011–2014, Figure 2A; Appendix A). With Scenario 2, OI and 23 fish were from the eastern population (1290–1950 mm values were calculated for zones 1 through 5 (Figure 2B; FL; Appendix D). In the second scenario, 10 fish were detected Appendix A) for monitoring periods 2 and 3 (2012–2014) to in the study area during both monitoring periods, and, because analyze the larger geographic area covered by the array and returning Gulf Sturgeon were characterized as unique indi- the greater number of tag detections from the increased viduals, a total of 55 individuals were detected. Of the 10 number of receivers and tagged Gulf Sturgeon in the system. returning fish, three were from the western population and Finally, OI patterns of Scenario 1 were compared to those seven were from the eastern population. For both Scenarios 1 patterns of Scenario 2 for zones 1 through 4 to test the utility of and 2, the majority of the eastern population fish were from the the OI as geographic area, receiver number, and zones within Escambia Bay region (Escambia, Blackwater, and Yellow the array increased. If there was no difference in the ‘‘pattern’’ rivers), with a few from the Choctawhatchee River. Gulf by zones as the array expanded, then the OI that generated the Sturgeon from the Pearl and Pascagoula drainages (western occupancy ‘‘pattern’’ was considered flexible to increases in the population) were equally represented. In Scenario 1, western data recorded within the acoustic array field’s new geography population fish contributed 57% of EADs, while eastern and increased number of receivers and zones. population fish contributed 43% of EADs (Appendix C); however, in Scenario 2 the opposite was true, with western RESULTS population fish contributing 47% of detections and eastern From 2011 to 2014, Gulf Sturgeon from both western and population fish contributing 53% of detections on the array eastern populations were detected on the Ship Island array (Appendix D). during all monitoring periods. Gulf Sturgeon from the Pearl, Pascagoula, Escambia, Blackwater, Yellow, and Chocta- Occupancy Index In the analysis of Scenario 1, there was no significant whatchee rivers were found occupying the habitat around Ship interaction effect (monitoring period3zone, F ¼1.0001, p¼ Island during overwintering months. 6,224 0.425) and no significant difference among monitoring periods

Gulf Sturgeon on the Ship Island Array and among zones (F2,224¼1.186, p¼0.307 and F3,224¼0.808, p¼ Overwintering adult and subadult Gulf Sturgeon were 0.491; Figure 3). Standard error of the mean (SEM) bars were detected during all three monitoring periods (2011–2014) on plotted on OI values, and the wide SEM bars indicated that the Ship Island array. During monitoring periods 1 to 3 in mean OI value by zone varied more in zones 2 and 3 than zones zones 1 to 4 of the array (Scenario 1), 44 Gulf Sturgeon were 1 and 4. The OI values of individuals by zone ranged from 0.0

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:39 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 5 0 Vick et al.

Figure 4. Occupancy index (6SEM) by zone and monitoring period for Scenario 2 (2012–2014). Note: See Figure 2 and Appendix A for descriptions of zones. The horizontal black line illustrates no significant differences (Sidak post hoc test, p 0.05) among zones 1 to 4, but all four zones are significantly different than zone 5.

mean occupancy (0 EADs) to a mean occupancy of about 0.4 Figure 5. Occupancy index directly compared to effort-adjusted detections (600 EADs; Figure 3). for (A) Scenario 1 (2011–2014) and (B) Scenario 2 (2012–2014). Note: (A) The In the analysis of Scenario 2, there was no significant data points represent the x¯ occupancy and x¯ effort-adjusted detections of interaction effect of zone and monitoring period (F4,265 ¼ 0.402, scenario 1 for zones 1 to 4 after monitoring period was pooled. (B) The data p ¼ 0.807), and there was no significant difference among points represent the x¯ occupancy and x¯ effort-adjusted detections of scenario 2 for zones 1 to 5 after monitoring period was pooled. All error bars represent monitoring periods (F 0.002, p 0.969). However, there 1,265 ¼ ¼ 61 SEM for occupancy values. The horizontal black lines represent no was a significant difference among zones (F4,265 ¼ 9.704, p , significant differences (Sidak post hoc tests, p 0.05) among zones 1 to 4 0.0001), with Sidak post hoc tests indicating zone 5 (Dog Keys (panels A and B) but all are significantly different than zone 5 (panel B). Pass) was significantly greater than all other zones (p , 0.0001 for all zones); zones 1 through 4 were not significantly different conservation of Gulf Sturgeon. The Gulf Sturgeon OI values in from each other (Figure 4). Wide SEM bars indicated marked Dog Keys Pass (zone 5) were, on average, nearly four times variability in mean OI values by zone ranging from 0.0 (0 greater than OI values compared to Gulf Sturgeon using any EADs) to 0.5 (2100 EADs; Figure 4). other island pass or cut (zones 1 to 4) of Ship Island, which Occupancy Index Strength and Testing the Robustness implies foraging behavior associated with high abundance of of the Index benthic resources (Fox, Hightower, and Parauka, 2002). Since there were no significant interaction effects or Furthermore, the OI spatial patterns observed did not appear monitoring period effects on the OI values of Scenarios 1 and to be related to the emigration pathways of either eastern or 2, the data were pooled by monitoring period and analyzed by western individuals (see Vick, 2016), since these individuals zone for each scenario. Linear regression indicated that the indiscriminately entered waters monitored by the Ship Island relationship between OI and EADs was strongly supported and acoustic array at various locations throughout the study period. predictive in both Scenario 1 (r2 ¼ 0.814, p , 0.0001) and With the alteration of Camille Cut proper through reconnect- Scenario 2 (r2 ¼ 1.0, p , 0.0001). The wide SEM bars again ing East and West Ship Island, moderate to high OI patterns of indicated marked variability in mean OI values and, thus, Gulf Sturgeon north and south of Camille Cut may decrease. EADs among Gulf Sturgeon (Figure 5A,B). For Scenario 1, an Although Camille Cut will be filled, the ‘‘new’’ nearshore OI value of 0.2 was, on average, about 400 EADs (Figure 5A), habitat north and south of Camille Cut will be available for and for Scenario 2, an OI value of 0.1 was about 500 EADs, on east–west migration and foraging habitat; it is, however, average, while an OI value 0.45 was about 2000 EADs, on uncertain whether filling in Camille Cut will alter the physical average, as observed in zone 5 (Dog Keys Pass; Figure 5B). components of the habitat, causing a shift in benthic prey availability. DISCUSSION Others have found that Gulf Sturgeon use a mosaic of Large subadult and adult Gulf Sturgeon from eastern and habitats comprised of metrics including depth, sediment grain western population segments use the habitat around Ship size, and prey availability (density or biomass; see Brooks and Island during the overwintering period presumably to forage, Sulak, 2005; Fox, Hightower, and Parauka, 2002; Harris, and this critical habitat must be protected for the recovery and Parkyn, and Murie, 2005; Peterson et al., 2013). Few data are

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:40 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 6 Gulf Sturgeon Occupancy 0

Figure 6. Ship Island bathymetry in meters relative to receiver zones (n ¼ 5). Note: West Ship (zone 1) is highly disturbed due to the maintained Gulfport Ship Channel. Camille Cut (zones 2 and 3) and East Ship (zone 4) consists of shallow sandy bottoms, while Dog Keys Pass (zone 5) has both sand bars, shallow sandy bottoms, and naturally deep channels (NOAA and USGS, 2014). Bathymetry (m) values based on island elevation (positive) and water depth (negative).

available regarding prey availability on the Mississippi barrier making it an equally suitable habitat of greater area for Gulf islands (although see Ross et al., 2009), but bathymetry of the Sturgeon than Camille Cut. passes and cuts of the barrier islands and general sediment The need to rearrange the acoustic array configuration types (Morton, 2008) displays the variation in these important during this study provided an opportunity to test the flexibility habitat metrics among Gulf Sturgeon overwintering habitats, and, thus, utility of the OI methodology (Peterson et al., 2013), especially on Ship Island. For example, West Ship (zone 1) as the geographic coverage of study and number of receivers consists of a shallow, sandy area adjacent to the 11.6 m deep, and zones increased. The Gulf Sturgeon OI values exhibited a dredge-maintained Gulfport Ship Channel (Morton, 2008; strong relationship with the EADs, although actual EADs Figure 6). Camille Cut (zones 2 and 3) and East Ship (zone 4) varied by the OI values between the two scenarios. The overall consist of shallow, relatively smooth sandy bottoms, while Dog occupancy pattern of increased EADs and increased OI values Keys Pass (zone 5) is composed of several sandbars that are was predictable, and the pattern and spatial relationships of separated by naturally deep channels (Figure 6; NOAA and both were strongly supported. Both OI values and the EADs were much higher for Scenario 2 than for Scenario 1 due to the USGS, 2014). As previously noted, Gulf Sturgeon have been geographic expansion of the array with the addition of Dog found foraging over areas with shallow, sandy substrate, Keys Pass (zone 5) in Scenario 2. This expansion also included making the island ends and the open pass of Camille Cut ideal an increase in the number of receivers within the array, which feeding habitat (Ross et al., 2009), and these areas, although led to an increase in the number of tag detections. This occupied, did not differ in mean Gulf Sturgeon OI values from approach extends the earlier use of the OI method (Peterson et other nearby areas. Gulf Sturgeon will likely use Dog Keys al., 2013, 2016), which was developed in a fixed geographic Pass for foraging, since it appears to provide a more diverse coverage array over time; therefore, the OI method will be habitat mosaic. While Dog Keys Pass is further away from the useful in future telemetry studies as it appears to be a flexible maintained Gulfport Ship Channel, those zones closer to the approach when examining habitat use under real-world Gulfport Ship Channel may have a modified sediment adaptive management scenarios, as was the case in this study. composition compared to this more eastern zone due to navigation related disturbances (Brooks and Sulak, 2005; CONCLUSION Gutperlet et al., 2015; Parauka, Duncan, and Lang, 2011). These data provide resource managers a detailed quantita- Gulf Sturgeon have been found in areas of less disturbance tive baseline on Gulf Sturgeon use of waters associated with where food resources are stable and high (Peterson et al., 2013; nearshore areas of Ship Island that will be useful in any Ross et al., 2009); the high Gulf Sturgeon OI values within Dog mitigation requirements associated with the completion of the Keys Pass, as well as other barrier islands (e.g., Horn and Cat island restoration project. The loss of Camille Cut proper may islands), may indicate a greater availability of prey items, have limited negative impacts to Gulf Sturgeon since, over the

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:40 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 7 0 Vick et al.

last 160–170 years (Morton, 2008), that area was periodically Field, A., 2013. Discovering Statistics using IBM SPSS Statistics, 4th land and unavailable to Gulf Sturgeon. Other more extensive edition. London: Sage Publications, 915p. Flowers, H.J.; Pine, W.E.; Dutterer, A.C.; Johnson, K.G.; Ziewitz, and permanent foraging habitats are available in nearby J.W.; Allen, M.S., and Parauka, F.M., 2009. Spawning site regions and islands. Having a better understanding of OI selection and potential implications of modified flow regimes on ‘‘patterns’’ and habitat use of Gulf Sturgeon aids in habitat viability of Gulf Sturgeon populations. Transactions of the preservation and science-based restoration, which is required American Fisheries Society, 138(6), 1266–1284. for the conservation and recovery of threatened and endan- Fox, D.A.; Hightower, J.E., and Parauka, F.M., 2002. Estuarine and nearshore marine habitat use of Gulf Sturgeon from the Chocta- gered species. Given the baseline data obtained in this study, whatchee river system, Florida. American Fisheries Society there should be a vast array of appropriate quality habitat Symposium Series, 28, 111–126. available for Gulf Sturgeon foraging on any Mississippi barrier Gutperlet, R.; Capperucci, R.M.; Bartholoma,¨ A., and Kroncke,¨ I., island given their current footprint. 2015. Benthic biodiversity changes in response to dredging activities during the construction of a deep-water port. Marine Biodiversity, 45(4), 819–839. ACKNOWLEDGMENTS Harris, J.E.; Parkyn, D.C., and Murie, D.J., 2005. Distribution of Gulf We would like to thank J.-M. Havrylkoff, P. Mickle, J. of Mexico sturgeon in relation to benthic invertebrate prey Green, T. Moncrief, M. Andres, A. Fogg, S. Ashworth, E. resources and environmental parameters in the Suwannee River Satterfield, C. Matten, M. Lowe, C. Griffin, B. Lewis, S. estuary, Florida. Transactions of the American Fisheries Society, George, A. Katzenmeyer, J. Collins, and J. Killgore for field 134(4), 975–990. Havrylkoff, J.-M.; Peterson, M.S., and Slack, W.T., 2012. Assessment and logistical assistance during the course of the study. We of the seasonal usage of the lower Pascagoula River estuary by Gulf thank G. Constant, K. Kimmel, C. Doolittle, and A. Baer, Sturgeon (Acipenser oxyrinchus desotoi). Journal of Applied USFWS Baton Rouge Conservation Office, for providing Ichthyology, 28(5), 681–686. tagging data and detection histories for some of the Pearl Haxton, T.J.; Sulak, K., and Hildebrand, L., 2016. Status of scientific knowledge of North American sturgeon. Journal of Applied River fish. Thank you to A. Kaeser, J. VanVrancken, F. Ichthyology, 32 (S1), 5–10. Parauka, A. Robydek, K. Herrington, S. Bolden, D. Fox, N. Heise, R.J.; Slack, W.T.; Ross, S.T., and Dugo, M.A., 2004. Spawning Willett, and R. Nelson for providing us with telemetry data. and associated movement patterns of Gulf Sturgeon in the The NOAA Gulf Sturgeon database provided data essential to Pascagoula River drainage, Mississippi. Transactions of the this study. This project was funded by a subcontract from American Fisheries Society, 133(1), 221–230. Heise, R.J.; Slack, W.T.; Ross, S.T., and Dugo, M.A., 2005. Gulf USACE-Mobile District (W912HZ-12-C-0045), and we appre- sturgeon summer habitat use and fall migration in the Pascagoula ciate the consistent and professional oversight of this project River, Mississippi, USA. Journal of Applied Ichthyology, 21(6), by S. Rees, J. McDonald, and E. Godsey, including excellent 461–468. comments on an earlier draft of this manuscript. We had an Huff, J.A., 1975. Life history of Gulf of Mexico Sturgeon, Acipenser annual Mississippi Museum of Natural Science state permit oxyrhynchus desotoi, in Suwannee River, Florida. Florida Marine Research Publications, 16, 1–32. to tag Gulf Sturgeon and a National Park Service federal IUCN (International Union for Conservation of Nature), 2010. permit to deploy acoustic receivers on Ship Island during this Sturgeon more critically endangered than any other group of study. This research was conducted under the University of species. Marine Pollution Bulletin, 60, 640–641. Southern Mississippi’s Institute of Care and Use Manson, C. and Hogarth, W.T., 2003. Endangered and threatened Committee 07081501, 09091702, 11092209, and 14091801. wildlife and plants; designation of critical habitat for the Gulf Sturgeon, final rule. Department of the Interior, Fish and Wildlife Service, 50 CFR Part 17, Department of Commerce, National LITERATURE CITED Oceanic and Atmospheric Administration, 50 CFR Part 226. Ahrens, R.N.M. and Pine, W.E., 2014. Informing recovery goals based Federal Register, 68(53), 13370–13495. on historical population size and extant habitat: A case study of the Mason, W.T. and Clugston, J.P., 1993. Foods of the Gulf sturgeon in Gulf Sturgeon. Marine and Coastal Fisheries: Dynamics, Manage- the Suwannee River, Florida. Transactions of the American ment, and Ecosystem Science, 6, 274–286. Fisheries Society, 122(3), 378–385. Baremore, I.E. and Rosati, J.D., 2013. Gulf Sturgeon Standardized McCauley, D.J.; Pinsky, M.L.; Palumbi, S.R.; Estes, J.A.; Joyce, F.H., Abundance and Mortality Study: Year Two Report. Silver Spring, and Warner, R.R., 2015. Marine defaunation: Animal loss in the Maryland: NOAA, NOAA Technical Memorandum NMFS-SEFSC- global ocean. Science, 347(6219), 247–254. 642, 25p. MMNS (Mississippi Museum of Natural Science), 2014. Endangered Brooks, R.A. and Sulak, K.J., 2005. Quantitative assessment of Species of Mississippi. Jackson, Mississippi: Mississippi Depart- benthic food resources for juvenile Gulf sturgeon, Acipenser ment of Wildlife, Fisheries and Parks, Museum of Natural Science, oxyrinchus desotoi, in the Suwannee River estuary, Florida, USA. 199p. Estuaries, 28(5), 767–775. Morrow, J.V.; Kirk, J.P., Killgore, K.J., and Rogillio, H.E., 1999. Byrnes, M.R.; Rosati, J.D.; Griffee, S.F., and Berlinghoff, J.L., 2012. Recommended enhancements to the Gulf Sturgeon recovery and Littoral Sediment Budget for the Mississippi Sound Barrier management plan based on Pearl River studies. North American Islands. Mobile, Alabama: Mississippi Coastal Improvements Journal of Fisheries Management, 19, 1117–1121. Program (MsCIP), ERDC/CHL TR-12-9, 106p plus 4 appendices. Morton, R.A., 2008. Historical changes in the Mississippi-Alabama Carr, S.H., 1983. All the way down upon the Suwannee River. barrier-island chain and the roles of extreme storms, sea level, and Audubon, 85, 78–101. human activities. Journal of Coastal Research, 24(6), 1587–1600. Cooke, S.J.; Paukert, C., and Hogan, Z., 2012. Endangered river fish: Moser, M.L.; Bain, M.; Collins, M.R.; Haley, N.; Kynard, B.; O’Herron Factors hindering conservation and restoration. Endangered II, J.C.; Rogers, G., and Squiers, T.S., 2000. A Protocol for Use of Species Research, 17(2), 179–191. Shortnose and Atlantic . Silver Spring, Maryland: Dugo, M.A.; Kreiser, B.R.; Ross, S.T.; Slack, W.T.; Heise, R.J., and NOAA, NOAA Technical Memorandum NMFS-OPR-18, 18p. Bowen, B.R., 2004. Conservation and management implications of Nelson, T.C.; Doukakis, P.; Lindley, S.T.; Schreier, A.D.; Hightower, fine-scale genetic structure of Gulf sturgeon in the Pascagoula J.E.; Hildebrand, L.R.; Whitlock, R.E., and Webb, M.A.H., 2013. River, Mississippi. Journal of Applied Ichthyology, 20(4), 243–251. Research tools to investigate movements, migrations, and life

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:41 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 8 Gulf Sturgeon Occupancy 0

history of sturgeons (Acipenseridae), with an emphasis on marine- habitat use of Gulf sturgeon (Acipenser oxyrinchus desotoi) in the oriented populations. PLoS ONE, 8(8), 1–22. north-central Gulf of Mexico. Estuaries and Coasts, 32(2), 360–374. NOAA and USGS (National Oceanic and Atmospheric Association Rudd, M.B.; Ahrens, R.N.M.; Pine, W.E., and Bolden, S.K., 2014. and U.S. Geological Survey), 2014. CoNED Project: 2014 USGS Empirical, spatially explicit natural mortality and movement rate Topobathymetric Model of the Northern Gulf of Mexico (1888 to estimates for the threatened Gulf sturgeon (Acipenser oxyrinchus 2013). http://www.coast.noaa.gov/dataviewer/#/lidar/. desotoi). Canadian Journal of Fisheries and Aquatic Science, 71(9), Parauka, F.M.; Duncan, M.S., and Lang, P.A., 2011. Winter coastal 1407–1417. movement of Gulf of Mexico sturgeon through northwest Florida Sulak, K.J.; Randall, M.T.; Edwards, R.E.; Summers, T.M.; Luke, and southeast Alabama. Journal of Applied Ichthyology, 27(2), K.E.; Smith, W.T.; Norem A.D.; Harden, W.M.; Lukens, R.H.; 343–350. Parauka, F.; Bolden, S., and Lehnert, R., 2009. Defining winter Partyka, M.L. and Peterson, M.S., 2008. Habitat quality and salt- trophic habitat of juvenile Gulf sturgeon in the Suwannee and marsh species assemblages along an anthropogenic estuarine Apalachicola river-mouth estuaries, acoustic telemetry investiga- landscape. Journal of Coastal Research, 24(6), 1570–1581. tions. Journal of Applied Ichthyology, 25(5), 505–515. Peterson, M.S.; Havrylkoff, J.-M.; Grammer, P.O.; Mickle, P.F., and Underwood, A.J., 1997. Experiments in ecology: Their logical design Slack, W.T., 2016. Consistent spatio-temporal estuarine habitat and interpretation using analysis of variance. Cambridge, U.K.: use of a western population of the Gulf Sturgeon. Transaction of Cambridge University Press, 504p. the American Fisheries Society, 145(1), 27–43. USACE (U.S. Army Corps of Engineers), 2016. Mississippi Coastal Peterson, M.S.; Havrylkoff, J-M.; Grammer, P.O.; Mickle, P.F.; Slack, Improvements Program (MsCIP) Comprehensive Barrier Island Restoration Hancock, Harrison and Jackson Counties, Mississippi. W.T., and Yeager, K.M., 2013. Macrobenthic prey and physical Final Supplemental Environmental Impact Statement. Mobile, habitat characteristics in a western Gulf sturgeon population: Alabama: U.S. Army Corps of Engineers-Mobile District, 400p. Differential estuarine habitat use patterns. Endangered Species USFWS (U.S. Fish and Wildlife Service), 1991. Endangered and Research, 22(2), 159–174. threatened wildlife and plants; determination of a threatened Peterson, M.S.; Weber, M.R.; Partyka, M.L., and Ross, S.T., 2007. status for the Gulf Sturgeon. Federal Register, 56(189), 49653– Integrating in situ quantitative geographic information tools and 49568. size-specific, laboratory-based growth zones in a dynamic river- USFWS, 1993. Standard operating procedures for sturgeon. Panama mouth estuary. Aquatic Conservation: Marine and Freshwater City, Florida: USFWS, Panama City Field Office, 24p. Ecosystems, 17(6), 602–618. USFWS, 2015. Exposure and Injuries to Threatened Gulf Sturgeon Robydek, A. and Nunley, M., 2011. Determining marine migration (Acipenser oxyrinchus desotoi) as a Result of the Deepwater patterns and behavior of the Gulf Sturgeon in the Gulf Sturgeon Horizon Oil Spill. Fairhope, Alabama: USFWS. Draft Preliminary critical habitat of the Gulf testing and training range and Santa Technical Report, 48p. Rosa Island complex. Thirteenth annual Gulf Sturgeon workshop. USFWS and NMFS (U.S. Fish and Wildlife Service and National Niceville, Florida: Northwest Florida State College Conference Marine Fisheries Service), 2009. Gulf Sturgeon (Acipenser oxy- Center, November 16–18. rinchus desotoi), 5-Year Review: Summary and Evaluation. Rogillio, H.E.; Rabalais, E.A.; Forester, J.S.; Doolittle, C.N.; Granger, Panama City, Florida: USFWS, Southeast Region, Panama City W.J., and Kirk Jr., P.P., 2001. Status, Movement and Habitat Use Ecological Services Field Office, and St. Petersburg, Florida: of Gulf Sturgeon in the Lake Pontchartrain Basin, Louisiana— National Marine Fisheries Service, Southeast Region, Office of 2001. Baton Rouge, Louisiana: Final Report submitted to the Protected Resources, 49 p. National Fish and Wildlife Foundation, U.S. Fish and Wildlife Vick, P.E., 2016. Gulf Sturgeon (Acipenser oxyrinchus desotoi) Pre- Service, and Louisiana Department of Wildlife and Fisheries, 24p. Restoration Occupancy Patterns on Ship Island, Mississippi Sound Rogillio, H.E.; Ruth, R.T.; Behrens, E.H.; Doolittle, C.N.; Granger, with an Evaluation of Designated Critical Habitat Use by Eastern W.J., and Kirk, J.P., 2007. Gulf Sturgeon movements in the Pearl and Western Population Segments. Hattiesburg, Mississippi: The River drainage and the Mississippi Sound. North American University of Southern Mississippi, Master’s thesis, 84p. Journal of Fisheries Management, 27(1), 89–95. Wilcove, D.S.; Rothstein, D.; Dubow, J.; Phillips, A., and Losos, E., Ross, S.T.; Slack, W.T.; Heise, R.J.; Dugo, M.A.; Rogillio, H.; Bowen, 1998. Quantitative threats to imperiled species in the United B.R.; Mickle, P., and Heard, R.W., 2009. Estuarine and coastal States. Bioscience, 45(8), 607–615.

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:41 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 9 0 Vick et al.

Appendix A. Ship Island acoustic array from 2011 to 2014.

Monitoring No. of Period Dates Receivers Zones 1 2011–2012 21 1, 2, 3, 4 2 2012–2013 29 1, 2, 3, 4, 5 3 2013–2014 29 1, 2, 3, 4, 5

Zone 1 had four receivers, zone 2 had nine receivers, zone 3 had four receivers, zone 4 had four receivers, zone 5 had eight receivers. See Figure 1 for location of zones.

Appendix B. Summary of effort-adjusted detections (EADs) by scenario, monitoring period (MP), and zone.

EADs by Zone Total EADs Mean EADs Scenario MP Transmitters 12345by MP by MP 1 1 14 1594.3 2062.5 2394.5 4085.2 10,136.5 724.0 2 17 1613.1 7555.7 4080.2 5375.5 18,624.5 1095.6 3 28 4641.0 6594.9 23,084.4 2159.8 36,480.0 1302.9 Total EADs by Zone 7848.4 16,213.1 29,559.2 11,620.4 Mean EADs by Zone 133.0 274.8 501.0 197.0 2 2 21 1699.8 9287.5 4365.7 5755.2 43,230.2 64,338.5 3063.7 3 34 4929.4 7907.6 24,511.7 2296.1 62,666.4 102,311.2 3009.2 Total EADs by Zone 6630.1 17,197.2 28,880.4 8055.3 105,901.6 Mean EADs by Zone 120.5 312.7 525.1 146.5 1925.5

Transmitters refers to the total number of Gulf Sturgeon detected during a MP. Mean EADs by MP are the Total EADs by MP/Transmitters and Mean EADs by Zone are Total EADs by Zone/Transmitters.

Appendix C. Scenario 1 (zones 1 to 4, monitoring periods 1 to 3) Gulf Sturgeon detections from 2011 to 2014.

Transmitter Population River FL (mm) Wt (kg) MP1 EADs MP2 EADs MP3 EADs Total EADs A69-1303-45716 Eastern Blackwater 1470 22.4 0 229 0 229 A69-1303-45734 Eastern Blackwater 1520 26.9 0 0 315 315 A69-1303-45768 Eastern Blackwater 1580 NA 0 0 2 2 A69-1303-46420 Eastern Blackwater 1580 35.65 0 0 1 1 A69-1303-46423* Eastern Blackwater 1880 56 45 0 244 289 A69-1303-46432 Eastern Blackwater 1600 37.7 0 0 73 73 A69-1303-61034* Eastern Blackwater 1660 31.8 1567 0 20 1587 A69-1303-61037 Eastern Blackwater 1370 23.8 0 0 158 158 A69-1303-61040 Eastern Blackwater 1640 38.6 0 5631 0 5631 A69-1303-61041* Eastern Blackwater 1460 23.8 15 23 0 38 A69-9001-30534* Eastern Blackwater 1290 16.9 2 372 0 373 A69-9001-30542 Eastern Blackwater 1550 33.45 0 0 11 11 A69-1303-45862 Eastern Choctawhatchee 1860 54 0 1334 0 1334 A69-1303-46183 Eastern Choctawhatchee 1370 17.5 0 0 2481 2481 A69-9001-29907 Eastern Choctawhatchee 1710 44.5 0 0 58 58 A69-1303-45751 Eastern Escambia 1480 22 2548 0 0 2548 A69-1303-61008 Eastern Escambia 1778 53.6 0 47 0 47 A69-9001-30598 Eastern Escambia 1370 22.1 0 0 113 113 A69-9001-30554** Eastern Yellow 1730 48.35 1490 4070 7044 12,605 A69-1303-45053 Western Pascagoula 1040 6.55 0 420 0 420 A69-1303-46208* Western Pascagoula 1380 19.5 27 61 0 88 A69-1303-46210 Western Pascagoula 1472 26.7 0 0 52 52 A69-1303-46215* Western Pascagoula 1470 23.6 1029 36 0 1065 A69-1303-46567 Western Pascagoula 1020 7.98 179 0 0 179 A69-1601-31790 Western Pascagoula 1235 16.3 0 0 8 8 A69-9001-29896 Western Pascagoula 1528 27.2 0 0 15 15 A69-9001-29899* Western Pascagoula 1316 17.66 0 238 147 386 A69-9001-29902 Western Pascagoula 1350 21.77 0 0 1570 1570 A69-9001-29904 Western Pascagoula 1422 24.67 0 0 79 79 A69-9001-30587 Western Pascagoula 1398 24.9 0 0 1772 1772 A69-9001-30589* Western Pascagoula 1461 29 40 0 182 223 A69-1303-45711 Western Pearl 1360 23.8 0 0 5351 5351 A69-1303-45717** Western Pearl 1625 47.7 151 714 221 1086 A69-1303-45720* Western Pearl 1700 43.6 1368 276 0 1644 A69-1303-45721 Western Pearl 1480 16.5 0 33 0 33 A69-1303-45731 Western Pearl 1600 31.8 0 0 233 233

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:41 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 10 Gulf Sturgeon Occupancy 0

Appendix C. Continued.

Transmitter Population River FL (mm) Wt (kg) MP1 EADs MP2 EADs MP3 EADs Total EADs A69-1303-45737 Western Pearl 1510 26.4 0 0 13 13 A69-1303-45746** Western Pearl 1320 29.9 504 5007 2164 7675 A69-1303-45748 Western Pearl 1590 NA 0 0 420 420 A69-1303-45752 Western Pearl 1276 14.3 0 0 264 264 A69-1303-45753 Western Pearl 1370 NA 0 0 13,469 13,469 A69-1303-45765 Western Pearl 1210 26.4 0 19 0 19 A69-1303-45767 Western Pearl 1480 24.2 0 113 0 113 A69-1303-62661 Western Pearl 1670 51.3 1172 0 0 1172 Total 10,137 18,624 36,480 65,241

Note: FL refers to fork length, Wt refers to weight, EADs refers to effort-adjusted detections, and MP refers to monitoring period. Transmitters marked with an asterisk indicate a fish detected in two monitoring periods, while transmitters marked with two asterisks refer to a fish detected in three monitoring periods. For total analyzed individuals for Scenario 1, transmitters with an asterisk were counted twice and transmitters with two asterisks were counted three times.

Appendix D. Scenario 2 (zones 1 to 5, monitoring periods 2 and 3) Gulf Sturgeon detections from 2012 to 2014.

Transmitter Population River FL (mm) Wt (kg) MP2 EADs MP3 EADs Total EADs A69-1303-45716 Eastern Blackwater 1470 22.4 244 0 244 A69-1303-45734 Eastern Blackwater 1420 22.4 0 780 780 A69-1303-45768 Eastern Blackwater 1580 NA 0 2 2 A69-1303-46420 Eastern Blackwater 1530 NA 0 210 210 A69-1303-46423* Eastern Blackwater 1880 56 4382 7145 11,527 A69-1303-46432 Eastern Blackwater 1600 37.7 0 9037 9037 A69-1303-46434* Eastern Blackwater 1630 33.1 436 13 448 A69-1303-61034* Eastern Blackwater 1660 31.8 501 55 556 A69-1303-61037 Eastern Blackwater 1370 23.8 0 1634 1634 A69-1303-61040* Eastern Blackwater 1640 38.6 8110 1433 9543 A69-1303-61041* Eastern Blackwater 1460 23.8 5462 13,424 18,886 A69-9001-30534 Eastern Blackwater 1290 16.9 5593 0 5593 A69-9001-30539 Eastern Blackwater 1390 22.3 0 38 38 A69-9001-30542 Eastern Blackwater 1550 33.45 0 1028 1028 A69-1303-45862 Eastern Choctawhatchee 1860 54 1772 0 1772 A69-1303-46183 Eastern Choctawhatchee 1330 20 0 2886 2886 A69-9001-29907 Eastern Choctawhatchee 1710 44.5 0 622 622 A69-1303-45721 Eastern Escambia 1480 22 35 0 35 A69-1303-45751 Eastern Escambia 1480 22 0 1 1 A69-1303-61008 Eastern Escambia 1778 53.6 136 0 136 A69-9001-30598 Eastern Escambia 1370 22.1 0 260 260 A69-9001-30554* Eastern Yellow 1730 48.35 11,314 10,429 21,743 A69-9001-30564* Eastern Yellow 1950 54.7 1190 642 1833 A69-1303-45053 Western Pascagoula 1040 6.55 717 0 717 A69-1303-46208 Western Pascagoula 1380 19.5 69 0 69 A69-1303-46210 Western Pascagoula 1472 26.7 0 55 55 A69-1303-46215 Western Pascagoula 1470 23.6 60 0 60 A69-1601-31790 Western Pascagoula 1235 16.3 0 31 31 A69-9001-29896 Western Pascagoula 1528 27.2 0 1141 1141 A69-9001-29899* Western Pascagoula 1316 17.66 2428 2655 5084 A69-9001-29902 Western Pascagoula 1350 21.77 0 2788 2788 A69-9001-29904 Western Pascagoula 1422 24.67 0 832 832 A69-9001-30587 Western Pascagoula 1398 24.9 0 3392 3392 A69-9001-30589 Western Pascagoula 1461 29 0 2199 2199 A69-1303-45711 Western Pearl 1360 23.8 0 7090 7090 A69-1303-45717* Western Pearl 1625 47.7 802 605 1407 A69-1303-45720 Western Pearl 1700 43.6 15,063 0 15,063 A69-1303-45731 Western Pearl 1600 31.8 0 1628 1628 A69-1303-45737 Western Pearl 1510 26.4 0 14 14 A69-1303-45746* Western Pearl 1320 29.9 5890 8502 14,392 A69-1303-45748 Western Pearl 1590 NA 0 6582 6582 A69-1303-45752 Western Pearl 1276 14.3 0 635 635 A69-1303-45753 Western Pearl 1370 NA 0 14,523 14,523 A69-1303-45765 Western Pearl 1210 26.4 20 0 20 A69-1303-45767 Western Pearl 1320 29.9 113 0 113 Total 64,338 102,311 166,650

Note: FL refers to fork length, Wt refers to weight, EADs refers to effort-adjusted detections, and MP refers to monitoring period. Transmitters marked with an asterisk indicate a fish detected in two monitoring periods. For total analyzed individuals for Scenario 2, transmitters with an asterisk were counted twice.

Journal of Coastal Research, Vol. 00, No. 0, 0000

//titan/Production/c/coas/live_jobs/coas-34/coas-34-06/coas-34-06-12/layouts/coas-34-06-12.3d 23 October 2017 5:41 pm Allen Press, Inc. # JCOASTRES-D-17-00027.1 Page 11