final report prediction and verification of multi-species snapper...

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Final Report Prediction and Verification of Multi-Species Snapper Grouper Spawning Areas in the U.S. South Atlantic

Submitted to: Bob Mahood, South Atlantic Fisheries Management Council

Submitted by: William D. Heyman, LGL Ecological Research Associates

Contract #: SA-(14)-36

This Draft: 20 May 2015

Original date: 31 January 2015

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Project Goal: The overall goal of this project is to develop and test a process to identify/verify snapper grouper spawning locations in the South Atlantic using analyses of physical geomorphology and the knowledge of expert fishermen for prediction and various methods for verification. If successful in verifying the presence of even a single fish spawning area (FSA), this methodology could then be applied to predict and verify other locations in the SA region. Results can support efforts to manage snapper grouper fisheries through discreet time area closures (Special Management Zones or SMZs) via implementation of the proposed Amendment 36 to the Snapper Grouper Fishery Management Plan.

Work Completed: Heyman organized and completed two offshore research trips in the South Atlantic Region, one with Mark Marhefka, 17 – 20 July 2014 out of Charleston, South Carolina and one with Jack Cox, 11 - 13 July 2014 out of Wilmington, North Carolina. In addition Heyman met with many relevant persons in an effort to foster development of a collaborative research and monitoring program for spawning areas in the South Atlantic Region. An interim report was submitted to the SAFMC on 20 August, and an oral presentation was presented to the SAFMC Snapper Grouper Committee on 5 December 2014. This report serves to fulfill the obligations of the contract.

South Atlantic Snapper Grouper AMENDMENT 36

L-1 Appendix L. Final Report: Prediction & Validation of Spawning

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Figure Captions

Figure 1: Spawning season for selected species within the grouper/snapper complex in the U.S. South Atlantic, that may form large transient spawning aggregations and the references for this information. Grey boxes indicates spawning season; black illustrates peak spawning season (Farmer et al., in prep.).

Figure 2: Overview map of sampling locations. NC shelf area sampled 11 – 13 July with Jack Cox; Georgetown Hole Area sampled 17 – 20 July 2014 with Mark Marhefka. Numbers on the map indicate sampling locations and correspond to GPS coordinates in Tables 1 and 2.

Figure 3: Detailed map of sampling locations in relation to bathymetry (m) along the NC shelf edge as sampled with Jack Cox, 11 – 13 July 2014.

Figure 4: Female mutton snapper Lutjanus analis caught near the shelf edge in 46 m water depth on 12 July 2014 at GPS position 95 (on Figure 3). The late development stage gonads provides some indirect evidence that the fish will likely spawn relatively soon in the vicinity. Mutton snapper are known to spawn in July in Florida waters. This contributes to the limited evidence that mutton snapper spawn farther north.

Figure 5: Vermilion snapper Rhomboplites aurorubens with late development stage gonads caught along the NC Shelf edge by Jack Cox. All of the 11 vermilion snapper captured during the trip, including both male and female were spawning capable. This provides evidence for the timing of the spawning season, consistent with the known spawning season for this species in Figure 1. Full moon occurred on 12 July, the day this (and other similarly staged) fish were caught so Vermilion snapper spawning likely takes place sometime after full moon.

Figure 6: Male Almaco jack Seriola rivoliana caught in 80 m water depth is ripe and running. Males can be ripe and running through the entire reproductive season and are therefore not a robust indicator of spawning location or time. Female reproductive state only can be used as a reliable indicator of spawning time and location.

Figure 7: Female Greater Amberjack Seriola dumerili with late development gonads. Since the primary female reproductive season for this species is April and May tailing off in June this provides evidence that the season may be longer than previously documented.

Figure 8: Gag grouper Mycteroperca microlepis showing “black belly” coloration,

photographed on the NC shelf. Gilmore and Jones (1992) documented this non-ephemeral coloration for large, old male gag groupers. They are less common now due to fishing pressure that removes larger individuals. Gilmore and Jones suggest that these fish form haremic aggregations with a single black belly. Since large gag groupers stay at spawning aggregation sites throughout



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the year in the Gulf of Mexico (Coleman et al. 2011) the presence of this fish may provide evidence that this is a spawning site. This observation is not indicative of spawning time or seasonality.

Figure 9: Georgetown Hole locator map (top) showing all locations sampled during 2014

in relation to the coastline and shelf edge. Bottom map is a detailed view and shows only sites sampled during July 2014.

Figure 10: Yellowedge grouper Hyporthodus flavolimbatus collected in 202 m water depth at what appeared to be a pinnacle, east of Georgetown Hole (see location of GPS 136 on Figure 9). The gonad appears hydrated but this can only be confirmed when histology results are returned from MARMAP. If indeed hydrated, this provides direct evidence of spawning in the area within 24 h and potentially less.

Figure 11: American Red Snapper Lutjanus campechanus female showing late development gonads in 44 m water depth, more than 20 miles inshore from the shelf edge. This finding is consistent with the known spawning season for this species shown at the bottom of this figure and in Figure 1.

Figure 12: Scamp Mycteroperca phenax courtship coloration documented on 25 April 2014 in the Georgetown Hole area. The inset drawing (Gilmore and Jones 1992) shows courtship coloration that matches that in the photo above. This behavior and ephemeral coloration was documented multiple times in several locations (see also Figure 9) during April but not in July (this report) or February 2014 in the same area.

Figure 13: Locations where scamp courtship behavior was observed (magenta triangles). The locations where Warsaw grouper (black squares) and speckled hind (yellow pentagons) were landed as part of this study are also noted. The locations of all GoPro camera drops during the 2014 sampling year are shown with green stars.



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Table Captions

Table 1: Sampling locations and landings in the Cubera Hole area with Jack Cox, July 2014.

Table 2: Video observations taken along NC shelf by GPS location and depth.

Table 3: Landings and sampling points at Georgetown Hole and environs with Mark Marhefka, July 2014

Table 4: Samples collected and being processed by MARMAP for histology and age.

Table 5: Video observations taken along SC shelf by GPS location and depth.



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Introduction

Evidence of Spawning Areas

Spawning aggregations are broadly defined herein based on Domeier et al. (1997) the most commonly cited definition as “a gathering of conspecific fish, for the purposes of spawning that consist of fish densities significantly higher than are found during the non-reproductive period”. Spawning can be documented in one of two ways. Direct evidence of spawning can be provided by video or photo documentation of actual spawning (gamete release) events which is generally accomplished using SCUBA in the tropics (Domeier et al., 1997; Colin et al., 2003; Heyman et al., 2004). Alternately, the reproductive phases of fishes (including spawning) can be documented through observations of gonadal tissues. Specifically the presence of female fishes with hydrated oocytes in their ovaries indicate imminent spawning while females with new post-ovulatory follicles ovaries indicate recent spawning (Brown-Peterson et al., 2011; see also methods section).

If spawning has been documented through histology, an aggregation is defined as a 3-fold increase in the density of the species at the same site during non-aggregation times or in other areas (Domeier et al., 1997). Since there exist a wide variety of types aggregations and the number of individuals and their densities vary among species and locations (Domeier et al., 1997; Lindeman et al., 2000), what constitutes a spawning aggregation is difficult to define, precisely. Documenting an aggregation is further confounded for species with low abundance (naturally or from exploitation) and in the relatively deep waters where they can occur for some species in the US South Atlantic (i.e. deeper than can be observed with SCUBA). Several indirect methods have therefore been used traditionally to verify the presence of spawning aggregations.

Indirect evidence of spawning aggregations include anecdotal evidence from fishermen, extraordinary high catch per unit effort at the site, particularly when at least some of the fishes caught at the site have large, late development stage gonads. In addition, several photos or video documentation of courtship coloration or behaviors, swollen or distended abdomens, recent bite marks, or a species density 3x the normal density at the site can also provide indirect evidence for the presence of spawning aggregations (Domeier et al., 1997; Colin et al., 2003; Heyman et al., 2004). Most of these techniques were developed in tropical waters and on shelf edges in 25 – 60 m water depth (Kobara et al., 2013). This study therefore required development of new techniques to verify spawning areas that are more appropriate for the subtropical waters of the US South Atlantic.

Spawning seasons for selected reef fishes in the US South Atlantic Region

The knowledge about when fish spawn in the region is somewhat limited for some species. At the beginning of this project, Dave Wyanski, Nick Farmer and the author compiled a table that illustrates the most generally accepted information about the spawning seasons of target species (Farmer et al., in prep; Figure 1). This table served as a guide in directing the timing of our sampling efforts.



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Methodology

This study continued testing of several methods for the prediction and verification of grouper snapper spawning areas in the South Atlantic. The development and test of these methods are leading to the Cooperative Research and Monitoring Program for US South Atlantic Spawning Aggregations (Heyman in prep). Three primary methods are being evaluated for their utility in verification of spawning areas and spawning aggregations:

1. Landings and catch per effort

2. Biological sampling (age, growth, reproductive status, fecundity)

3. Underwater visual and video observation and census

Monitoring techniques can be an extension of baseline data collected during prediction and verification but different in many ways. Since this study focuses only on prediction and verification, however, we have concentrated analysis efforts on those techniques that are likely to offer the most defendable direct or indirect evidence of spawning presence – biological observations and sampling and underwater video observations.

Landings and catch per effort

At each sampling location, commercial fishermen chose the spot and either anchored or hovered in the area, motor fishing. Landings were closely monitored on board. The time, date, location (GPS position), weather conditions, water depth, water temperature at fishing depth, and all catch (landings numbers of each species caught and discarded) and effort (gear type, number of hooks, lines, start and stop time for each line) data were recorded on either a field notebook or data sheet. Catch per effort data has not been analyzed in this report due to lack of time for analysis. The use of catch per effort measurements is part of developing a long-term cooperative monitoring methodology and not as important for verification purposes. For all samples collected in South Carolina, SCDNR sampled those relevant fish for age/growth/reproduction.

Biological Sampling

When possible, gonad condition for each fish landed from all key species was evaluated using macroscopic observation and photographs (either at sea or at the dock). When possible, (Trip 2) detailed histological studies of collected gonads were collected for analysis in the MARMAP lab. Histological data was processed at MARMAP and was not available at the time of the first draft of this document but was recently summarized (LGL Ecological Research Associates, 2015).

This method recognizes that females with hydrated oocytes are actively spawning (< 24 hours). Hydrated

Box 1: Go Pro video monitoring setup



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oocytes are clearly visible macroscopically and their presence can be confirmed using histology. Female fishes that have recently spawned possess new postovulatory follicles in their gonads, which can also be determined using histology (Brown-Peterson et al., 2011).

Underwater Video and Visual Census

We tested the use of GoPro cameras, deployed on commercial bandit reel rigs to document species composition and density as well as courtship coloration or spawning behavior. We have also evaluated the use of a bottom mounted, rotating camera system designed by Chris Koenig. Each video was captured and stored with relevant position and time information in a digital database. Video effort for each drop was calculated from the time the camera hit bottom and was adjusted to being upright until the time that the camera started its ascent back up to the boat. Total underwater video effort summed to 2.5 hours of footage between the two trips in July. Videos were examined and analyzed for bottom type, species presence, absence and abundance indexing, and most importantly for any signs of courtship or spawning behavior.

Mapping

All of the data collected with this methodology are tied to specific GPS positions and times. These points and the data from each location were entered into a GIS database for analysis and mapping purposes. Additional data including bathymetry and the locations of existing protected areas were included with the maps for reference. This report presents maps and descriptions of the sites and data collected in July 2014. These data include visual observations made with GoPro cameras, macroscopic observations of gonads, and species (and numbers) harvested by location. Underwater videos and CPUE data are being maintained for possible future analysis.

Training

Heyman collected the data along with Jack Cox in NC. Due to time and weather constraints, and to evaluate technician acquisition and use of the techniques Heyman trained Aaron Swersky who then collected the data along with Mark Marhefka at Georgetown Hole.

Results and Discussion of July 2014 Data

Trip 1: Captain Jack Cox, M/V Elizabeth; 11 - 13 July 2014

The initial goal of this trip was to evaluate/verify spawning areas within the “780 Bottom” an anecdotally reported multi-species spawning aggregation site (SAFMC 2012). Unfortunately, the currents were so strong that neither fishing nor filming would have been possible. We therefore explored an area farther south and west of the 780 Bottom along the shelf at a location locally called Cubera Hole and near the 366 Wreck (Figures 2, 3). We successfully deployed GoPro cameras on commercial fishing gear, caught fish and examined gonads. Because the fish were landed in NC, we did not bring the fish for MARMAP workup so do not have histology data from this trip.



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Landings

Fishing was monitored at a series of sites around Cubera Hole and the 366 Wreck on the NC shelf edge. A list of all of the sampling sites, showing latitude, longitude, time, depth, species and number of each caught at each site is included as Table 1. Effort data has not been analyzed herein, but has been recorded and stored.

Biological sampling

We recorded macroscopic observations and photographs of the gonads from all species of concern, when possible. Unfortunately, biological samples and measurements could not be taken because of the distance and logistical difficulty of getting fish to MARMAP. Most of these observations did not have any indication of imminent or recent spawning activity.

We recorded a female mutton snapper in late development stage with large swollen gonads (Figure 4). While this did not indicate spawning was imminent, it does provide evidence that spawning in this species occurs in the vicinity during this time. We recorded several vermilion snapper, both males and females, with late development stage gonads (Figure 5).

We recorded a male Almaco jack in ripe and running condition, releasing milt from the gonopore (Figure 6). If this were female, it would be highly indicative of spawning. However, males can be in spawning condition through most of the spawning season so this is not a robust indicator of spawning. We also observed two female Greater Amberjacks with late development gonads (Figure 7). The reported spawning season for Greater Amberjack ends in June according to Harris et al. (2007) but that study only had access to 17 female specimens north of 310 N. Observations presented herein suggest a more protracted spawning season for this species in the US South Atlantic.

Underwater video observation

Total video effort for this trip was 1.3 hours of usable footage for analysis and a full set of observations is included as Table 2. We recorded several gag grouper (Mycteroperca microlepis) exhibiting “black belly” coloration (Figure 8) as described by Gilmore and Jones (1992). This coloration is not an ephemeral coloration pattern associated with courtship, but rather the color of most large gag grouper, generally male (Gilmore and Jones 1992, J. Cox, pers. comm., M. Marhefka, pers. comm.) (but also see Collins et al., 1998). Male gag groupers are known to remain on shelf-edge reefs associated with spawning sites while females move inshore after spawning has occurred as described by Coleman et. al. (2011). We did not observe any other gag or courtship coloration during this trip.

Trip 2: Captain Mark Marhefka, M/V Amy Marie; 17 – 20 July 2014

Aaron Swersky conducted sampling on this trip after intensive three day training from Heyman. He successfully collected high quality data, indicating that the techniques can be used by trained technicians, lending to their applicability in a cooperative research program. The goal of the trip was to continue sampling in the area of Georgetown Hole



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(Figures 2, 9) during July, a month in which blueline tilefish, snowy grouper, Cubera snapper, red snapper, mutton snapper, and vermilion snapper are believed to be at or near their peak spawning period. Since the spawning season for red porgy and vermilion snapper is fairly well documented, we did not focus analyses on these species.

Landings and catch per effort

Fishing was monitored at a series of sites around Georgetown Hole, the elbow north of Georgetown hole, and on the transit in and out of Charleston. A list of all of the sampling sites, showing latitude, longitude, time, depth, species and number of each caught at each site is included as Table 3. Effort data has not been analyzed herein, but has been recorded and stored. Though vermilion snapper (158 individuals) and red porgy (141 individuals) dominated the catch, they were not sampled specifically for gonad condition.

Biological sampling

Several target species were harvested including Warsaw grouper, gag, scamp, snowy and yellowedge grouper, mutton snapper, American Red Snapper, and various tilefish. Specimens were collected and delivered to MARMAP personnel for full biological sample workups at the end of the trip (Table 4). The histological and age data were still being analyzed at the time the first draft of this report was completed.

Based on macroscopic observations in the field, a female Yellowedge grouper appeared to have hydrated gonads with clear and visible eggs (Figure 10). This fish was captured in over 200 m water depth at GPS #136 (see Figure 9) on 19 July 2014 at 7:10 AM local time. Hydrated female gonads indicate that spawning was imminent and probably occurred that day or the next, in relatively close proximity to the location of capture, approximately seven days after full moon in July. The site of capture appears like a pinnacle on existing bathymetric charts but detailed surveys using Marhefka’s WinPlot 3D sonar plotter illustrated that instead it was more of a hump than a pinnacle, yet still distinct from surrounding bottom. There is very little reproductive information available for this species. Sedberry et al. (2006) report the spawning season for species is between August and September but the sample size is relatively small (n=52). Spawning season for Yellowedge grouper extends begins at least as early as July, as shown by this specimen.

An American red snapper female was caught in 44 m (Figure 11) water depth on the shelf edge, about 20 miles inshore of the shelf edge on 17 July 2014 at GPS #142 (Figure 9). Macroscopic examination of the gonads reveals the fish as late development (spawning ready but not hydrated). This is not indicative of a spawning aggregation but does occur to be consistent with the reported spawning season.

Underwater video observation

Seventeen individual underwater videos (~1.2 hours usable video) were recorded and analyzed for the presence of courtship coloration, behavior or other indications of spawning (Table 5). Though a relatively large group (>10) scamp were seen in one location in July GPS #142, (Figure 9) no courtship coloration or behavior were observed



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at any of the observation locations during this time. This site will be re-examined during spawning season (April) in 2015.

In addition to the cameras deployed on commercial bandit reels (Box 1) we also tested the utility of a rotating GoPro mount developed by Chris Koenig (Box 2).

Data and Conclusions Based on All Data Collected During 2014

2014 Sampling

This report describes findings from two trips in the South Atlantic in July 2014. In addition, however, two additional trips were also conducted in the Georgetown Hole area using the same techniques. Though this report will not provide a comprehensive analysis of all data collected from the entire year, it does offer some highlights from the other trips as relevant to the prediction and verification of spawning areas and the value of the areas surveyed for possible SMZ for spawning protection and for Warsaw grouper and speckled hind.

April 2014 included many observations and underwater video recordings of scamp in courtship coloration, exhibiting what appeared to be courtship behavior. As described by Gilmore and Jones (1992) we observed “gray head” phase scamp flashing, rubbing, and nuzzling other individuals (e.g. Figure 12). In one instance, on 25 April 2014 at 8:48 AM local time, scamp courtship behavior was observed extensively within a small aggregation (15+ individuals) of scamp grouper. These observations took place at GPS # 84 where the bottom was highly rugose with high relief and provided ample habitat for groupers to hide. In addition, a large black-bellied gag patrolled the area. Large male gags in the Gulf of Mexico are known to remain at their shelf edge spawning aggregation sites throughout the year (Coleman et al. 2011). The combination of high numbers of scamp showing courtship behaviors and the presence of large (presumed male) gag grouper on the same site suggests that a spawning area may be located in the vicinity. This site in particular will be re-examined during 2015. These observations also suggest that spawning had not yet occurred on the dates of these observations, and forthcoming histological samples will confirm or refute this claim. Histological samples will help to elucidate the seasonal and lunar spawning period for this species. Congeneric Mycteroperca bonaci and M. venenosa spawn approximately 10 – 13 days after full moon in Belize (Heyman et al., 2004; Heyman and Kjerfve 2008). In the only study in the US South Atlantic that reports on lunar periodicity of spawning, Harris et al. (2002) showed that the proportion of scamp with spawning indicators is highest around both new and full moons.

Box 2: Koenig's camera rotator



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This study has also helped to identify habitat for Warsaw grouper and speckled hind by noting the location of all individuals captured (Figure 13). Though these data are only preliminary, it does provide evidence that these sites do harbor these species presently. In sum, these data contribute evidence to, but do not by themselves serve to confirm spawning in the vicinity.

Methodological observations

This study reports on techniques that were being developed, ad hoc in partnership with commercial fishermen in the South Atlantic. Some techniques have been adapted from other studies and used for the first time for this purpose in this region (e.g. biological sampling and CPUE). As far as we are aware, this study was the first ever to use GoPro deployments on commercial fishing gear to study spawning behavior in reef fishes. After a year of testing and various experiments and adjustments (e.g. camera angle, deployment depth, attachment techniques, baited versus non-baited, time of day, sea state, etc.) the technique has proved itself worthy of further exploration. We have found that the best results were obtained when the camera was deployed on an L-bar as close to the bottom as possible and set to aim horizontally, with no bait on the L-bar but a normal long leader and baited grouper hook. The L-bar and line provide a bit of a vane that stabilizes the camera from lateral movements (wagging) in the current. Though it restricts the field of view to what is behind the vessel, it provides a very still and clear image. The bait provides some attraction for fish, but not too close to the camera as to be disruptive. The blinking red light on the GoPro serves to attract feeding fish in many instances, which has caused damage to the housing. More durable housings would be desirable and GoPro has come out with a new “dive housing” that is rated for deeper water than their normal one. That said, this study has effectively gathered data in over 100 m water depth, though housings did seem to slowly be crushed and eventually flood in waters deeper than ~ 164 m. We will continue to experiment with Koenig’s circulating camera mount in 2015. Improvements will include better anchoring and fastening the retrieve line to the base (not the top) and allowed slack while the camera is recording. The constant movement of the camera seemed to blur (pixilate) the images in the limited tests that we did.

Data recording, storage and reporting

This has been a year of rapid learning. Beginning with simple field notes, no data sheets, database, or method for data organization, it has been a painful year of learning by doing. One year later, at the beginning of 2015, we have explored various data sheets and recording and storage methods and are nearing a workable system that will be tested in 2015.

Coordination and support from MARMAP have been superb. They generously provided sample workup for free this year. This will not be possible in coming years given the increasing workload. Also, MARMAP cannot necessarily process samples collected in other locations and that are not landed in Charleston (e.g. our first trip with Jack Cox). For these reasons, we are exploring other methods/systems/partners to ensure that these data are collected and archived with sufficient oversight and quality control that the samples are useable for stock assessments.



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In addition, we successfully trained two technicians in these techniques, Aaron Swersky from South Carolina and Kyle McCain from LGL in Texas. These two initial trainees will work with the system again in 2015 to help standardize and systematize the techniques – making it possible to streamline training and to train additional observers and thus increase our sampling capacity in space and time.

Cooperative Research

Great thanks are due to Capt. Mark Marhefka and Capt. Jack Cox and their crews and mates. Without them, none of this would have been possible. They have all helped to think through, implement and refine data collection techniques. They are helping to build a system that is cost effective (i.e. can be accomplished with observers aboard commercial vessels instead of research vessels) and efficient (in that data can be collected without disturbing and disrupting (too much) commercial fishing activity. These and other captains have been incredibly forthcoming with their extensive historical geographic knowledge of fishing areas and times. The best way to find spawning areas has always been and will likely always be from the observations of local fishermen in the area. Their technical abilities in the design and construction of various sampling methods has been invaluable, e.g. the GoPro observations on commercial fishing gear. It is an important goal of this research that will indeed assist managers and grouper snapper fishermen and in the region to locate, monitor, and protect spawning areas such that they can contribute to the resilience and sustainable management of these resources.

Summary and Next Steps

This report serves as a final report to the SAFMC for contract SA-(14)-36. It offers an update on the prediction and verification of reef fish spawning areas in the US South Atlantic, based on two trips funded by this project conducted in July 2014. No evidence of spawning was recorded during our trips using video but histological results were still pending at the first drafting of this report. In addition to the two July trips, which form the focus of this report. We also completed two additional trips in the region, February and April 2014. Some observations from the full set of data have been included in this report. The methods employed herein, appear to come together as valuable and workable system for verification. The techniques described herein may prove useful for verifying the presence of spawning sites throughout the South Atlantic region. We will be testing the same techniques both in the US South Atlantic and the US Gulf of Mexico regions during 2015. The objectives for 2015 are to:

1. Collect additional data at Georgetown Hole and 780 Bottom.

2. Hone the system and techniques and develop a clear training manual and monitoring protocol, that can be used far more broadly in years to come.

3. Develop the database and the institutional housing to store, QA/QC, and serve the data for management and research purposes.



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Schobernd, C.M. and Sedberry, G.R., 2009. Shelf-edge and upper-slope reef fish assemblages in the South Atlantic Bight: Habitat characteristics, spatial variation, and reproductive behavior. Bulletin of Marine Science 84, 67–92.

SEDAR 2011. SEDAR 25 - South Atlantic Black Sea Bass Stock Assessment Report. SEDAR, North Charleston SC. 146 pp. available online at: http://www.sefsc.noaa.gov/sedar/Sedar_Workshops.jsp?WorkshopNum=25

SEDAR. 2013. SEDAR 36 – South Atlantic Snowy Grouper Stock Assessment Report. SEDAR, North Charleston SC. 146 pp. available online at: http://www.sefsc.noaa.gov/sedar/Sedar_Workshops.jsp?WorkshopNum=36

Sedberry, G. R., O. Pashuk, D. M. Wyanski, J. A. Stephen, and P. Weinbach. 2006. Spawning locations for Atlantic reef fishes off the southeastern U.S. Proceedings of the Gulf and Caribbean Fisheries Institute 57:464–514.

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Wyanski, D.M., White, D.B., Barans, C.A., 2000. Growth, population age structure, and aspects of the reproductive biology of snowy grouper, Epinephelus niveatus, off North Carolina and South Carolina. Fishery Bulletin 98, 199–218.

Ziskin, G.L., Harris, P.J., Wyanski, D.M., Reichert, M.J.M., 2011. Indications of continued overexploitation of speckled hind along the Atlantic coast of the southeastern United States. Transactions of the American Fisheries Society 140, 384–398.



16

Figure 1: Spawning season for selected species within the grouper/snapper complex in the U.S. South Atlantic, that may form large transient spawning aggregations and the references for this information. Grey boxes indicates spawning season; black illustrates peak spawning season (Farmer et al., in prep.)



17

Figure 2: Overview map of sampling locations. NC shelf area sampled 11 – 13 July with Jack Cox; Georgetown Hole Area sampled 17 – 20 July 2014 with Mark Marhefka. Numbers on the map indicate sampling locations and correspond to GPS coordinates in Tables 1- 4.



18

Figure 3: Detailed map of sampling locations in relation to bathymetry (m) along the NC shelf edge as sampled with Jack Cox, 11 – 13 July 2014.



19

Figure 4: Female mutton snapper Lutjanus analis caught near the shelf edge in 46 m water depth on 12 July 2014 at GPS position 95 (on Figure 3). The late development stage gonads provides some indirect evidence that the fish will likely spawn relatively soon in the vicinity. Mutton snapper are known to spawn in July in Florida waters. This contributes to the limited evidence that mutton snapper spawn farther north.



20

Figure 5: Vermilion snapper Rhomboplites aurorubens female with late development stage gonads caught along the NC Shelf edge by Jack Cox. All of the 11 vermilion snapper captured during the trip, including both male and female were spawning capable. This provides evidence for the timing of the spawning season, consistent with the known spawning season for this species shown in Figure 1 and below the photos in this figure. Full moon occurred on 12 July, the day this (and other similarly staged) fish were caught so Vermilion snapper spawning likely takes place sometime after full moon.

Vermilion snapper

NC Shelf Edge 33.xxx N 76.xxx W (GPS 106)

Date caught: 12 July 2014 5:40 PM local time 164 m water depth

Female, Late Development

Captain: Jack Cox Vessel: F/V Elizabeth



21

Figure 6: Male Almaco jack Seriola rivoliana caught in 80 m water depth is ripe and running. Males can be ripe and running through the entire reproductive season and are therefore not a robust indicator of spawning location or time. Female reproductive state only can be used as a reliable indicator of spawning time and location.



22

Figure 7: Female Greater Amberjack Seriola dumerili with late development gonads.

Since the primary female reproductive season for this species is April and May tailing off in June this provides evidence that the season may be longer than previously documented.



23

Figure 8: Gag grouper Mycteroperca microlepis showing “black belly” coloration,

photographed on the NC shelf. Gilmore and Jones (1992) documented this non-ephemeral coloration for large, old male gag groupers. They are less common now due to fishing pressure that removes larger individuals. Gilmore and Jones suggest that these fish form haremic aggregations with a single black belly. Since large gag groupers stay at spawning aggregation sites throughout the year in the Gulf of Mexico (Coleman et al. 2011) the presence of this fish may provide evidence that this is a spawning site in the South Atlantic. This observation is not indicative of spawning time or seasonality.



24

Figure 9: Georgetown Hole locator map (top) showing all locations sampled during 2014

in relation to the coastline and shelf edge. Bottom map is a detailed view and shows only sites sampled during July 2014.



25

Figure 10: Yellowedge grouper Hyporthodus flavolimbatus collected in 202 m water depth at what appeared to be a pinnacle, east of Georgetown Hole (see location of GPS 136 on Figure 9). The gonad appears hydrated based on visual examination. This was confirmed subsequently when histology results were returned from SCDNR MARMAP (Table 1; LGL 2015) providing direct evidence of spawning in the area within < 24 h.



26

Figure 11: American Red Snapper Lutjanus campechanus female showing late development gonads in 44 m water depth, more than 20 miles inshore from the shelf edge. This finding is consistent with the known spawning season for this species shown at the bottom of this figure and in Figure 1.



27

Figure 12: Scamp Mycteroperca phenax courtship coloration documented on 25 April 2014 in the Georgetown Hole area. The inset drawing (Gilmore and Jones 1992) shows courtship coloration that matches that in the photo above. This behavior and ephemeral coloration was documented multiple times in several locations (see also Figure 9) during April but not in July (this report) or February 2014 in the same area.

Plate 8: Scamp

Georgetown Hole 32.xxx N 78.xxx W (GPS 88)

Date filmed: 25 April 2014 10:40 AM local time Depth: 42 m

Courtship coloration and behavior

Captain: Mark Marhefka Vessel: F/V Amy Marie

Still photo extracted from GoPro Hero 3+ video (file GoPro 173.mp4)



28

Figure 13: Locations where scamp courtship behavior was observed (magenta triangles). The locations where Warsaw grouper (black squares) and speckled hind (yellow pentagons) were landed as part of this study are also noted. The locations of all GoPro camera drops during the 2014 sampling year are shown with green stars.



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Table 1: Sampling locations and landings in the Cubera Hole area with Jack Cox, July 2014.



30

Table 2: Video observations taken along NC shelf by site number and depth (fathoms). Site%# Local%Time Depth%(Fathoms) File%Number Film%Observa;ons%=%1 Video%effort%(min)

6:20 23 GOPRO471.MP4/

course/sand,/elongate/small/roll/</1m/ver@cal,/with/rela@vely/dense/growth/but/less/than;/lots/of/fish.//Scamp/(black),/amberjacks,/big/school/unident./Maybe/tomtate;/grey/trigger,/black/sea/bass,/lesser/amberjack,

0:59

6:20 23 GOPRO473.MP4boQom/with/1m/roll/described/above./>8/scamp/most/in/very/dark/phase./~4/large/amberjacks.//Grey/trigger

1:00

96 7:27 23 GOPRO176.MP4boQom/smooth/U/course/sand,/small/patches/of/algae,/1U2'/hardboQom/ridges,/very/few/fish,/gray/triggers

4:53

97 8:15 23 GOPRO177.MP4BoQom/as/above:/red/porgy,/queen/trigger,/gray/trigger,/blue/tang,/amberjack,/vermillion/snapper,/black/belly/gag/4:50/U/(good/for/s@ll/photo/grab)

5:00

8:30 24 GOPRO178.MP4fine/U/course/sand,/scaQered/boQom/growth,/red/porgy,/queen/trigger,/black/belly/gag/(/2:51)

0:41

8:30 24 GOPRO179.MP4 grey/@gger,/black/bellied/gag,/porcupine/fish 1:20

10:29 40 GOPRO180.MP4

boQom/largely/hardboQom/with/short/growth/but/high/areal/coverage/algae,/greater/amberjack,/lesser/amberjack,/french/angelfish/(courtship/colora@on?),/2/scamp/with/courtship/behavior/but/not/colora@on/at/5:33/one/the/scamp/caught/(good/s@ll/grab/at/7

4:17

10:29 40 GOPRO184.MP4boQom/as/above:/four/jacks/circling/hook/then/one/took/it.//Big/Almaco

0:39

10:29 40 GOPRO185.MP4amberjack,/solitary/grouper/(sp?),/very/few/fish,/amberjack,/lesser/amberjack,/caught/amberjack

6:18

10:29 40 GOPRO186.MP4shallow/behind/boat,/schooly/mahi/bi@ng/hooks/and/being/caught,/sargassum/in/the/water

6:08

102 11:45 64 GOPRO187.MP4boQom/fairly/fine/sand/and/smooth/boQom/with/liQle/algae/or/hard/boQom;/amberjacks,/very/few/fish

1:50

14:00 74 GOPRO188.MP4 loads/of/red/porgy,/caught/snowy 0:47

14:00 74 GOPRO189.MP4boQom/course/sand/with/shell,/no/vegeta@on/or/structure,/very/flat,/loads/of/red/porgy;/caught/snowy

1:41

14:00 74 GOPRO190,/GOP010190

boQom/of/course/sand,/but/smooth/without/ripples,/red/porgy,/amberjack,/lots/of/red/porgy/figh@ng/over/bait,/rig/was/brought/up,/rebaited/and/reUdeployed;/two/part/video/first/15/min,/second/less/than/two.

13:54

14:00 74 GOPRO191.MP4boQom/same/as/148.//loads/of/red/porgy,/some/black/sea/bass;/caught/snowy

1:36

14:00 74 GOPRO198.MP4 very/few/fish/visible./ 2:17

15:42 79 GOPRO210.MP4BoQom/with/rela@vely/fine/sand,/showing/meandering/ridges;/Amberjack,/grouper/(sp?),/very/few/fish

6:46

15:42 79 GOPRO211.MP4

BoQom/with/rela@vely/fine/sand,/showing/meandering/ridges;/a/2m/high/elongate/hillock/with/many/ledges/providing/good/habitat/and/lots/of/fish.//Several/species/there,/but/tough/to/ID,/AJ,/scamp,/snowy,/others./Caught/snowy

3:20

15:42 79 GOPRO212.MP4Very/dark,/tough/to/ID/fish;//school/(15)/of/amberjacks,/large/shark/sp?,/several/large/groupers/sp?

7:50

105 17:11 82 GOPRO215.MP4 To/dark/to/ID/fish 5:28

95

98

101

103

104



31

Table 3: Landings and sampling locations in Georgetown Hole and environs with Mark Marhefka, July 2014

Site%Number Species%(common%name) Species%(la4n%name) Species%(MARMAP%Code) #%CaughtBlack&Sea&Bass Centropris*s+striata A177 1Gag&Grouper Mycteroperca+microlepis A189 1Gray&Trigger Balistes+capriscus A428 5Red&Porgy Pagrus+pagrus A272 14Vermillion&Snapper Rhomboplites+aurorubens A252 28Pinfish Lagodon+rhomboides A271 4Red&Porgy Pagrus+pagrus A272 1Vermillion&Snapper Rhomboplites+aurorubens A252 4Gray&Trigger Balistes+capriscus A428 3Vermillion&Snapper Rhomboplites+aurorubens A252 4Gray&Trigger Balistes+capriscus A428 3Red&Porgy Pagrus+pagrus A272 1Vermillion&Snapper Rhomboplites+aurorubens A252 32Red&Porgy Pagrus+pagrus A272 3Vermillion&Snapper Rhomboplites+aurorubens A252 52Goldface&Elefish Caulola*lus+chrysops B464 1Red&Porgy Pagrus+pagrus A272 1Scamp Mycteroperca+phenax A190 1Greater&Amberjack Seriola+dumerili A230 1Jolthead&porgy Calamus+bajonado A265 1Warsaw&grouper Epinephelus+nigritus A881 1

124 Greater&Amberjack Seriola+dumerili A230 1Blueline&Tilefish Caulola*lus+microps B438 1Red&Porgy Pagrus+pagrus A272 1Scamp Mycteroperca+phenax A190 2Almaco&Jack Seriola+rivoliana A232 1Gray&Trigger Balistes+capriscus A428 1Lesser&Amberjack Seriola+fasciata A231 1Red&Porgy Pagrus+pagrus A272 1Scamp Mycteroperca+phenax A190 2Snowy&grouper Epinephelus+niveatus A185 4Red&Porgy Pagrus+pagrus A272 3Scamp Mycteroperca+phenax A190 3Snowy&grouper Epinephelus+niveatus A185 4Red&Porgy Pagrus+pagrus A272 18Snowy&grouper Epinephelus+niveatus A185 3Red&Porgy Pagrus+pagrus A272 10Warsaw&grouper Epinephelus+nigritus A881 1Blueline&Tilefish Caulola*lus+microps B438 1Red&Porgy Pagrus+pagrus A272 8Snowy&grouper Epinephelus+niveatus A185 1Warsaw&grouper Epinephelus+nigritus A881 1

132 Red&Porgy Pagrus+pagrus A272 12Gray&Trigger Balistes+capriscus A428 1Greater&Amberjack Seriola+dumerili A230 1Saucereye&Porgy Calamus+calamus A336 1Snowy&grouper Epinephelus+niveatus A185 1

135 Red&Porgy Pagrus+pagrus A272 8Blueline&Tilefish Caulola*lus+microps B438 3Goldface&Elefish Caulola*lus+chrysops B464 1Snowy&grouper Epinephelus+niveatus A185 5Yellowedge&Grouper Epinephelus+flavolimbatus A880 1Gag&Grouper Mycteroperca+microlepis A189 2Gray&Trigger Balistes+capriscus A428 1Red&Porgy Pagrus+pagrus A272 2Scamp Mycteroperca+phenax A190 2Vermillion&Snapper Rhomboplites+aurorubens A252 3

130

134

136

138

129

126

127

128

120

114

116

117

118

119

123

125



32

Table 3 (continued): Landings and sampling points at Georgetown Hole and environs with Mark Marhefka, July 2014

Site%Number Species%(common%name) Species%(la4n%name) Species%(MARMAP%Code) #%CaughtGray%Trigger Balistes(capriscus A428 2Vermillion%Snapper Rhomboplites(aurorubens A252 3

140 Gray%Trigger Balistes(capriscus A428 2Mahi%Mahi Coryphaena(hippurus A240 1Remora%sp. Remora(sp. A944 1

141 American%Red%Snapper Lutjanus(campechanus A244 2Gag%Grouper Mycteroperca(microlepis A189 3Gray%Trigger Balistes(capriscus A428 2Jolthead%porgy Calamus(bajonado A265 1MuEon%Snapper Lutjanus(analis A241 1Vermillion%Snapper Rhomboplites(aurorubens A252 7

142 American%Red%Snapper Lutjanus(campechanus A244 1Gray%Trigger Balistes(capriscus A428 1Remora%sp. Remora(sp. A944 1Scamp Mycteroperca(phenax A190 2

143 Black%Sea%Bass Centropris;s(striata A177 6144 American%Red%Snapper Lutjanus(campechanus A244 1

Black%Sea%Bass Centropris;s(striata A177 4Gray%Trigger Balistes(capriscus A428 2Red%Porgy Pagrus(pagrus A272 58Scamp Mycteroperca(phenax A190 1Vermillion%Snapper Rhomboplites(aurorubens A252 25White%Grunt Haemulon(plumierii A260 3

139



33

Table 4: Samples collected and being processed by MARMAP for histology and age during July 2014.



34

Table 5: Video observations taken along SC shelf in the Georgetown Hole area with Mark Marhefka by location (site number) and depth (fathoms)

Site%# Local%Time Depth%(Fathoms) File%Number Film%Observa;ons%=%1 Video%effort%(min)

114 12:15 15.5 Trig+GoPro003

course+to+sand+botom+with+ripples+and+sparse+algae;+a+1+@+rocky+ledge+emerges+from+sand;+water+green+top+to+boDom,+lots+of+fish+at+this+site+mostly+vermillion+snapper,+red+porgy,+gray+trigger,+a+small+shark+(sp?),+several+(8)+scamp,+other+larger+grouper

9:24

116 15:18 20.3 Trig+GoPro004boDom+course+to+fine+sand,+no+boDom+vegetaNon+visible,+water+blue+in+the+surface+but+very+green+and+turbid+at+the+boDom,+high+fish+density+=+vermillion,+red+porgy,+white+grunt,+gray+trigger,

5:53

117 15:48 21.5 Trig+GoPro005boDom+course+and+fine+sand+some+gravel+and+small+rock+in+clumps+that+form+patch+reefswith+sponge+and+gorgonian+growth,+water+blue+near+the+surface,+green+near+the+botom:+gray+trigger,+black+sea+bass,+vermilliion,+lesser+amberjack,+porcupine/balloon+fish,

5:59

118 17:51 20.6 Trig+GoPro006almost+no+structure+on+boDom,+some+wave+paDerns+in+sand,+grey+trigger,+red+porgy,+white+grunt,+jack+(amber+or+almaco),+scamp

6:15

120 7:48 27.5 Trig+GoPro007BoDom+showing+many+humps+and+vegetaNon.+Triggerfish+(sps?),+Eel+(sp?),Grouper+(sp.),+2+lionfish(2:05U6:40)+chasing+grouper.+

8:57

121 8:14 27 Trig+GoPro008water+blue+top+to+boDom,+boDom+shows+liDle+relief+but+a+grass+like+field+of+growth,+low+fish+density,+red+porgy,+sand+Nlefish

5:07

123 8:55 26.5 Trig+GoPro009BoDom+densely+covered+in+gorgonians+and+algae+on+what+appears+to+be+a+1+U+2m+roll.++Queen+trigger,+wrasses,+tang,+jack+(amber?),+red+porgy,+porgy+spp.?++Low+fish+density

4:22

124 9:45 46 Trig+GoPro011

boDom+covered+with+rocky+clumps+over+hard+packed+sand+and+encrusted+in+somehwat+dense+algae,+rolling+undulaNng+boDom+with+possible+dropoff+edge+but+too+far+to+see.++Amberjacks+(10),+scamp+(8),+shark+sp?+And+lesser+amberjack.++Medium+to+low+number+of+fi

6:07

128 12:09 67 Trig+GoPro012

very+dark+and+grainy.++BoDom+course+sediment,+very+sparse+bits+of+colonized+rock+but+mostly+flat.++Low+fish+density;+Red+porgy,+shark+U+blackNp?,+scamp,+other+grouper+U+gag?+Snowy?+Nice+shark+while+camera+is+going+up

5:40

129 14:20 64 Trig+GoPro013,+GoPro1013very+dark+and+grainy,+Red+porgy,+Nice+shark+shot+3:10;+nice+sNngray+shot,+8:20

8:53

139 14:50 23.5 Gag+GoPro005 Coarse+sand,+rock+patches.+Shark+sp.,+ 3:15

142 17:31 21.7 Gag+GoPro007boDom+course+sand+with+wave+induced+ripples;+big+school+of+scamps+(10?)+3:01+U+3:08;+red+porgy

1:47



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