Fishery-Independent Monitoring of Coral Reef Fishes, Coral Reefs, and Macro-invertebrates in the Dry Tortugas

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1 Sustaining Dry Tortugas National Park Coral Reef Resources FINAL REPORT FY 2006 Page 1.1 Fishery-Independent Monitoring of Coral Reef Fishes, Coral Reefs, and Macro-invertebrates in the Dry Tortugas Jerald S. Ault, Steven G. Smith, James A. Bohnsack, Jiangang Luo, Nicholas A. Farmer, Douglas E. Harper and David B. McClellan FINAL REPORT February 2007

2 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.2 FINAL REPORT FY 2006 Fishery-Independent Monitoring of Coral Reef Fishes and Macro-invertebrates in the Dry Tortugas Jerald S. Ault, Steven G. Smith, James A. Bohnsack 1, Jiangang Luo, Douglas E. Harper 1 and David B. McClellan 1 University of Miami, Rosenstiel School of Marine and Atmospheric Science, 4600 Rickenbacker Causeway, Miami, FL 33149, (305) ph; jault@rsmas.miami.edu 1 NOAA Fisheries, Southeast Fisheries Science Center, 75 Virginia Beach Drive, Miami, FL 33149, (305) ph; jim.bohnsack@noaa.gov National Park Service Contract No. H500000B494-J NMFS Coral Reef Program NA17RJ1226 February 2007 Cover photo: Large school of French grunt (Haemuelon flavolineatum) and a single creole wrasse (Clepticus parrae) seen by RVC divers on Tortugas Bank during 2006 expedition.

3 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.3 FINAL REPORT FY 2006 EXECUTIVE SUMMARY In November 2006, the Florida governor and cabinet approved implementation of a management plan for a Research Natural Area (RNA) or no-take marine reserve in the Dry Tortugas National Park (DTNP) to become effective in January The Florida Fish and Wildlife Conservation Commission also concurred with the proposed National Park Service regulations related to marine fishing in the park. In 2001 no-take marine reserves (NTMRs) covering approximately 566 km 2 were established in Florida Keys National Marine Sanctuary waters near Dry Tortugas National Park. The park s new RNA, coupled with marine reserves in the Florida Keys National Marine Sanctuary, is designed to protect precious coral reefs, fishery, and cultural resources, and to ensure sustainability of intensely exploited regional reef fisheries resources benefiting the Tortugas, the Florida Keys and beyond. To evaluate monitor and evaluate baseline conditions of coral reef resources in and around DTNP, this program focused on two primary areas of research: (1) fishery-independent monitoring of coral reef fishes, coral reefs, and macroinvertebrates in DTNP; and, (2) a pilot acoustic telemetry tracking study of reef fishes to determine population flux rates in open areas and fully-protected marine reserves. The primary objective of fishery-independent monitoring was to conduct a synoptic visual census survey using SCUBA/nitrox to assess the resource status (occurrence, abundance, and spatial distribution), and MPA performance for the reef fish community in Dry Tortugas National Park. The goals of the 2006 Tortugas research expedition were: (1) to conduct a quantitative visual census assessment of coral reef fishery and habitat resources in the Tortugas region five years after implementation of the Tortugas Ecological Reserve (TER); (2) to sample all fish species and sizes in all representative coral reef habitats both inside and outside reserve areas; and, (3) to monitor trends in coral reef fish populations and the effectiveness of current management practices. Using a sampling design-based approach in 2006 we conducted a research cruise to the Dry Tortugas that resulted in 1,344 scientific dives in the region monitoring reef fish, benthic habitats, and spiny lobster, with 817 dives occurring the DTNP. We compared these data to a series of synoptic research cruises with over 4,000 research dives to survey reef fish populations and habitats in the Dry Tortugas before and three years after the NTMRs were implemented. We recorded the presence, abundance and size of 267 fish species from eight reef habitats in three management areas offering different levels of resource protection: the Tortugas North Ecological Reserve (a NTMR), Dry Tortugas National Park (recreational angling only), and southern Tortugas Bank (open to all fishing under regional regulations). Species richness and composition remained stable between , 2004, and 2006, within the overall survey domain. Greatest reef fish biodiversity was found in the more rugose habitats. We detected significant domain-wide increases in abundance for several exploited and non-exploited species, while no declines were detected. In the Tortugas Bank NTMR, we found significantly greater abundances and shifts in length composition structures towards a higher proportion of exploited phase animals in 2004 and 2006 compared to for some species (e.g., black grouper and red grouper). Consistent with predictions from marine reserve theory, we did not detect any declines for exploited species in the NTMR, while for non-target species we detected both increases and declines in population abundance in the NTMR for non-target species. The observed upsurge in exploited populations, however, may have also been influenced by other factors including past or recent fishery management actions that increased minimum sizes or reduced fishing mortality rates; the passage of recent

4 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.4 FINAL REPORT FY 2006 hurricanes; and, the occurrence of good recruitment year classes. Although still early in the recovery process, our results after three years are encouraging and suggest that NTMRs, in conjunction with traditional management, can potentially help build sustainable fisheries while protecting the Florida Keys coral reef ecosystem. The primary objective of the second research foci was to develop a pilot study using state-of-theart acoustic telemetry technology to track real-time movements of groupers in key habitats of DTNP to precisely estimate population flux rates in fully-protected areas free from exploitation. Rosenstiel School fisheries researchers tracked red and black grouper in the Dry Tortugas National Park to develop a better understanding of species movement and habitat require-ments, so they can help more efficiently design and assess future marine-protected areas. Through funding from the National Park Service and transportation support from Yankee Fleet Ferry Service, scientists were able to conduct this high-tech observation that involves surgically implanted transmitters for approximately a year. A pilot field study was designed that used acoustic telemetry technology to track continuously the movements and habitat use of red and black grouper in the Dry Tortugas National Park, the 46-square-nautical-mile marine reserve. The groupers were fitted with transmitters or pingers that emit unique acoustic codes underwater approximately every 20 seconds. Passive listening stations or receivers were placed in a submersed array that detected the transmitters. Receivers recorded an acoustic tag s presence when it was within range, usually 250-1,000 meters, depending on the oceanographic conditions. Because the tags each emit unique identification numbers and time stamps, individual receivers could potentially detect up to 4,000 different fish at any given time. An array of submersed hydroacoustic receivers were deployed in a representative suite of benthic habitats along the northwestern boundary of the newly-approved Research Natural Area (RNA) in Dry Tortugas National Park, with a corresponding smaller array in the adjacent waters of Tortugas North Ecological Reserve in Florida Keys National Marine Sanctuary to detect flux of tagged fish across reserve boundaries. Receiver signal detection range and resultant position fixes for acoustic tags were calibrated using a variety of methods. Submersed receivers function as passive acoustic listening stations, capable of archiving a unique ID code, date and time of detection for any tagged fish that passes within range. Groupers and snappers were surgically implanted with acoustic transmitters and their movements were passively monitored for four months. Fish were systematically tracked to obtain continuous, precise tracks of animal locations, including daily travels and frequently visited areas. The following points summarize our findings: (1) reef fish within Dry Tortugas National Park are capable of, and do, cross reserve boundaries; (2) red grouper demonstrated relatively high site; (3) red grouper seldom moved across large expanses of sand and all movements out of the RNA occurred across contiguous reef habitat; and, (5) red grouper are apparently crepuscular feeders as frequency of detections increased during dawn-dusk. The results of this pilot study have provided us with previously unavailable insights into grouper movements that are critical towards improving the experimental methodology to better address the goal of quantifying the rate of flux across reserve boundaries for these important species, a focus of our current research. Ultimately, these data will be used to parameterize a spatially-explicit models that provide insights into how flux of exploited species across reserve boundaries will impact NTMR effectiveness as a fisheries management tools and help ensure sustainable fisheries in the Dry Tortugas and Florida Keys ecosystems.

5 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.5 FINAL REPORT FY 2006 TABLE OF CONTENTS Executive Summary. 1.3 Section 1.0 Fishery-Independent Monitoring of Coral Reef Fishes, Coral Reefs, and Macroinvertebrates in the Dry Tortugas 1.0 Introduction: Background and Rationale Materials and Methods Study Area Survey Design and Operations Statistical Analysis Results Sampling Effort Preliminary Analysis of Change, Baseline to Next Steps in the Research Analysis Acknowledgments Literature Cited Section 2.0 Acoustic Telemetry Tracking of Reef Fish to Determine Population Flux Rates in Open Areas and Fully-Protected Marine Reserves 2.0 Introduction Methods Range-testing: broad scale Range-testing: fine scale Conventional anchor tagging Acoustic telemetry tagging Results Range-testing: broad scale Range-testing: fine scale Conventional anchor tagging Acoustic telemetry tagging Missing transmissions Movement patterns and habitat use Flux across RNA boundaries Home range Discussion Range-testing Acoustic telemetry Missing transmissions Movement patterns and habitat use Flux across RNA boundaries Home range Summary and Conclusions Relevant Literature. 2.17

6 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.6 FINAL REPORT FY INTRODUCTION: BACKGROUND AND RATIONALE The Dry Tortugas are located about 70 miles west of Key West at the southwestern end of the west Florida shelf (Figure 1.1). The Tortugas region is a unique tropical marine environment of national significance, renown for its luxuriant and productive coral reef ecosystem, diverse natural resources, broad recreational fishing opportunities, and spectacular scenic beauty (Schmidt et al. 1999; Culhane 2002; Ault et al. 2002; Brock and Culhane 2004). The Tortugas region supports multibillion dollar fishing and tourism industries in southern Florida, including economically-important fisheries for pink shrimp, spiny lobster, reef fish (snapper-groupers), kingfish and Spanish mackerel. In the Florida Keys, increased fishing pressure from rapid regional human population growth and environmental changes associated with coastal development have raised concerns about fisheries sustainability and persistence of the coral reef ecosystem (Porter and Porter 2001; Ault et al. 2005a; Pandolfi et al. 2005). Historically intense commercial and rising recreational fishing pressures have resulted in unsustainable rates of exploitation for seventy percent of the snapper-grouper complex (Ault et al. 1998, 2005b), which consists of over 50 species of mostly groupers and snappers, but also grunts, jacks, porgies, and hogfish. Over the last 40 years, the number of registered recreational vessels in southern Florida has grown by more than 500%. Sport fishing effort is expected to continue to grow in proportion to regional human populations which have doubled about every 20 years (Ault et al. 2005a). At present, the recreational fleet accounts for a substantial proportion of the total regional catches for some key exploited species (NOAA MRFSS Database; Florida FWC Trip Ticket Database; Coleman et al. 2004), and this trend will likely continue to increase. Reef fisheries in the Florida Keys ecosystem are complex and regulated by several entities including the Florida Fish and Wildlife Conservation Commission (FWC, ), the National Park Service (NPS, DOI, and the National Marine Fisheries Service in conjunction with the South Atlantic Fishery Management Council (SAFMC; NOAA, and the Gulf of Mexico Fishery Management Council (GMFMC, NOAA, In response to declining trends in reef fishery catches, a series of regional federal and state management regulations were imposed including recreational bag limits, minimum size limits, commercial quotas and trip limits, seasonal closures, gear restrictions, limited commercial entry, closed fisheries, species moratoria, game

7 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.7 FINAL REPORT FY 2006 fish status, and restrictions on sale and possession. These regulations were implemented to stabilize catches, protect spawning stock biomass, and reduce fishing mortality rates. In general, the history of regional regulations for reef fishes has been complex, and tended to be more restrictive over time. Nonetheless, despite the bevy of regulations imposed in the Florida Keys, recent fishery assessments indicate that, for example, black grouper spawning stock biomass was less than 10% of its historical size (Ault et al. 2005b). In recent years, new ecosystem-based management measures have been enacted in the Florida Keys, including the 1997 implementation of a network of 23 NTMRs by the Florida Keys National Marine Sanctuary (FKNMS, NOAA, These are relatively small (mean 2 km 2, range km 2 ), comprising 46 km 2 in total area (Department of Commerce 1996) and have varying levels of protection: four allow catch-and-release surface trolling and four can only be accessed by special permit. In July 2001, the Florida Keys network was expanded with the implementation of two NTMRs in the Dry Tortugas region covering about 566 km 2. The Tortugas region is believed to be extremely important for coral reefs and fisheries as a source of recruitment because of its upstream location in the Florida Current that facilitates advective dispersion and transport of eggs and larvae to the rest of the Keys (Lee and Williams 1999; Dahlgren and Sobel 2000; Lindeman et al. 2000; Ault et al. 2002; Yeung and Lee 2002; Domeier 2004). Following implementation of conventional management measures or implementation of spatial controls like NTMRs, rebuilding of reef fish population biomass and age-structures of the depleted resources is expected. In the longer-run, unrestricted growth of biomass within reserves should result in resource export through reserve boundaries to surrounding areas as either larval dispersal to proximal natal sites as well as the diffusive movements of fishable biomass (Bohnsack 1998; Roberts et al. 2001; Pauly et al. 2002; Russ 2002; Zeller and Russ 2004; Bohnsack et al. 2004). The rate at which these impacts on individual reef fish population s biomass will occur and could be detected depends greatly on the species life history, demographic characteristics, and survey precision. Given that snapper and grouper life spans are often measured in decades, the effects of management actions could take 20 years or more to reach their full potential (e.g., Beverton and Holt 1957). Using a sampling design-based approach we conducted a series of synoptic research cruises with over 4,000 research dives to survey reef fish populations and habitats in the Dry

8 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.8 FINAL REPORT FY 2006 Tortugas before and three years after the NTMRs were implemented (Ault et al. 2006). We recorded the presence, abundance and size of 267 fish species from eight reef habitats in three management areas offering different levels of resource protection: the Tortugas North Ecological Reserve (a NTMR), Dry Tortugas National Park (recreational angling only), and southern Tortugas Bank (open to all fishing under regional regulations). Species richness and composition remained stable between and 2004 within the overall survey domain. Greatest reef fish biodiversity was found in the more rugose habitats. We detected significant domain-wide increases in abundance for several exploited and non-exploited species, while no declines were detected. In the Tortugas Bank NTMR, we found significantly greater abundances and shifts in length composition structures towards a higher proportion of exploited phase animals in 2004 compared to for some species (e.g., black grouper and red grouper). Consistent with predictions from marine reserve theory, we did not detect any declines for exploited species in the NTMR, while for non-target species we detected both increases and declines in population abundance in the NTMR for non-target species. The observed upsurge in exploited populations, however, may have also been influenced by other factors including past or recent fishery management actions that increased minimum sizes or reduced fishing mortality rates; the passage of recent hurricanes; and, the occurrence of good recruitment year classes. Although still early in the recovery process, our results after three years of protection were encouraging and suggest that NTMRs, in conjunction with traditional management, can potentially help build sustainable fisheries while protecting the Florida Keys coral reef ecosystem. The goals of the 2006 Tortugas research expedition were: To conduct a quantitative visual census assessment of coral reef fishery and habitat resources in the Tortugas region five years after implementation of the Tortugas Ecological Reserve (TER). To sample all fish species and sizes in all representative coral reef habitats both inside and outside reserve areas. To monitor trends in coral reef fish populations and the effectiveness of current management practices. A potentially confounding factor in understanding the efficacy of management actions in the Tortugas region was the sharp increase in hurricane activity in the intervening time period

9 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.9 FINAL REPORT FY 2006 between the 2004 and 2006 surveys. As illustrated in Figure 1.2, one hurricane (Charley) and one tropical storm (Ivan) passed through the Dry Tortugas region in August-September 2004, and four hurricanes (Dennis, Katrina, Rita, and Wilma) impacted the region in July-October This report documents the scientific activities of the 2006 research expedition in the Tortugas region and presents preliminary findings on changes in the reef fish community five years after implementation of NTMRs. 1.1 MATERIALS AND METHODS Study Area The Florida Keys coral reef ecosystem extends 380 km from Miami to the Dry Tortugas (Figure 1.1). The Tortugas study area is located about 113 km west of Key West, and encompasses approximately 1686 km 2 in two principal areas: Dry Tortugas National Park (managed by Department of the Interior, DOI); and, Tortugas Bank (NOAA, FKNMS, Department of Commerce, DOC) (Figure 1.3) Survey Design and Operations We employed a stratified random diver visual survey to obtain fishery-independent data on the spatial distribution, abundance, size composition, and habitats of coral reef fishes in the Tortugas region (Bohnsack and Bannerot, 1986; Ault et al., 1998, 2002; Bohnsack et al., 1999). The principal survey domain encompassed coral reef habitats less than 33 m deep in Tortugas Bank and Dry Tortugas National Park (Figure 1.3). The domain was extended to depths of 42 m along the western edge of Tortugas Bank by a technical dive team equipped with tri-mix. The sampling domain was partitioned into habitat strata based on the degree of vertical relief (e.g., rugosity, complexity) and the degree of patchiness (e.g., amount of softbottom substrate interspersed among reef structures) of the hardbottom substrate (Franklin et al. 2003; Ault et al. 2006). This habitat-based stratification procedure was developed from the 1999 and 2000 baseline surveys, and was shown to be effective in partitioning the domain into areas of high, moderate, and low levels of mean fish density and associated variance for many principal reef species (Ault et al. 2002), thereby improving sampling efficiency and cost-effectiveness (Smith and Ault 1993; Ault et al. 1999, 2003). Management zones were incorporated as a second spatial stratification variable, designated as follows: Tortugas Bank NTMR -- closed to all types of fishing; Tortugas Bank Fished -- open to all types of commercial and recreational fishing under regional regulations; and, Dry Tortugas National Park -- open to only recreational hook-

10 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.10 FINAL REPORT FY 2006 and-line fishing (Figure 1.3). In the Tortugas region, areas open to fishing such as Tortugas Bank Fished, allow a variety of types of legal fishing activities under regional management and represent the lowest level of resource protection in the study area. Dry Tortugas National Park represents an intermediate level of resource protection by allowing only recreational angling. Commercial fishing has been prohibited since 1935 when it was established as a National Monument. Recreational lobster diving was prohibited in After conversion to Dry Tortugas National Park in 1992, protection increased with exclusion of headboats for recreational fishing in The Tortugas Bank NTMR, a no-take and no anchoring reserve, represents the highest level of resource protection. Prior to 1 July 2001, Tortugas Bank was open to fishing under GMFMC and Florida FWC regulations. We used a geographical information system (GIS) and digital spatial databases of benthic habitats, bathymetry, and management zone boundaries to facilitate spatial delineation of the survey domain, sampling strata, and sample units. The Tortugas sampling domain was overlain in a GIS with a grid of 200 by 200 m cells that represented the minimum mapping units for benthic habitat types (Figure 1.4). A two-stage stratified-random sampling design was employed in which the primary sample unit was the 200 by 200 m habitat grid cell and the second-stage unit was a 15 m diameter visual census circular plot (described below) (Figure 1.4). Stratum (h) sizes in terms of area (A h ) consisting of N h possible primary sampling units are given in Table 1.1. These values were updated from the 2004 survey with the incorporation of new, high-resolution bathymetry data for portions of the Tortugas region graciously provided by NOAA NOS, US Geological Survey, and the National Park Service. Allocation of the number of primary units to be sampled among strata was based on stratum area and variance of fish density for a representative suite of species (i.e., a Neyman allocation scheme; Cochran, 1977). Within a stratum, specific primary units to be sampled were randomly selected a priori with equal probability from the complete list of N h units using a discrete uniform distribution (Law and Kelton, 2000). To ensure replication, two pairs of second-stage sample units (i.e., diver visual census plots) were randomly positioned within each selected primary unit. Because of diving safety concerns and statistical concerns about sample autocorrelation, in our computations each second-stage unit estimate consisted of the arithmetic average of stationary plots from two individual divers (i.e., a buddy pair ). Thus, each primary sample unit location in Figure 1.3 denotes where at least four scientific divers were

11 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.11 FINAL REPORT FY 2006 deployed to conduct visual census samples (i.e., one pair of divers at each of two second-stage locations within a primary sampling unit). Highly trained and experienced divers collected biological data using Nitrox or Trimix SCUBA and the reef fish visual census (RVC) protocol, a standard, non-destructive, in situ visual monitoring method. In the RVC protocol, a stationary diver collects reef fish data while centered in a randomly selected 15 m diameter circular plot (Bohnsack and Bannerot, 1986; Bohnsack et al., 1999; Ault et al., 2002). First, all fish species observed within 7.5 m in an imaginary cylinder extending from the bottom to the limits of vertical visibility (usually the surface) are listed for 5 minutes. After the initial 5 min listing, data are then collected on the abundance, and minimum, mean, and maximum lengths for each species sighted. A ruler connected perpendicularly to the end of a meter stick is used as a reference to reduce apparent magnification errors in fish size estimates. For each plot, depth, bottom substrate composition, estimated benthic percentage cover, and vertical relief characteristics of the seafloor were recorded from the polar perspective of the centrally located observer. Digital photographs were taken at each station to assist with habitat classification and identification of uncommon fish species. The time required to record each sample averaged 15 to 20 min, depending on the habitat. The synoptic 2006 survey was conducted over a three-week period from June 5 to June 26, with 20 days of onsite sampling, using a 30 m live-aboard dive vessel equipped with 4 compressor banks of Nitrox (M/V Spree, Gulf Diving, Houston, TX) and additional tanks of helium for making Trimix. The onboard scientific crew, consisting of 24 persons on any given sampling day, was comprised of a fish census team, a benthic habitat team, and a spiny lobster Panilurus argus team, as well as two full-time divemasters to oversee the complex diving operations. Visual survey data were entered onboard into a digital database using a laptop-based data entry system that includes extensive error-checking and validation protocols. Although our primary objective was reef fish visual assessment, data were obtained to study possible synergistic effects of fish density, lobster density, habitat, and prey availability by conducting habitat and lobster assessments in conjunction with visual fish censuses. These data will assist development of statistical models to link fish, habitat, and food resources from a broad ecological perspective.

12 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.12 FINAL REPORT FY Statistical Analysis Our statistical analyses focused on changes between baseline years 1999 and 2000 (before) and 2006 (after), with comparison to changes between baseline and 2004 reported by Ault et al. (2006). Statistical analysis of change was evaluated using a community metric, species richness, and two population metrics: frequency of occurrence and abundance. Statistical estimation procedures followed Cochran (1977) for a two-stage stratified random sampling design. In these procedures, strata means and variances of a given metric are weighted by strata sizes, i.e., W = N / N, to obtain overall means and variances for either specific h h h h management zones, or for the entire Tortugas domain. Species richness was estimated on a primary sample unit basis (i.e., the number of unique species observed within a primary unit by the group of divers) to ensure a sufficient search area for obtaining reliable estimates. In this case, the statistical sample size was n, the number of sampled primary units. Both frequency of occurrence and abundance were estimated by species on a second-stage unit basis, the standard approach for two-stage designs (Cochran, 1977), where the number of second-stage units nm was the statistical sample size. Since benthic habitat classification, digital mapping, and development of the Tortugas survey design occurred concurrently with the baseline surveys of 1999 and 2000 (Ault et al., 2002), each population and community metric was estimated as a composite of the two baseline years to alleviate problems of misclassification of habitats and misallocation of samples among habitat strata. In this procedure, strata means and variance components were computed as two-year averages weighted by respective sample sizes in 1999 and Species chosen for detailed analyses reflected the range of population dynamic processes (growth and survivorship) for relatively abundant exploited and non-exploited components of the reef fish community. Statistical tests for differences among estimates of mean density, total abundance and mean proportion of samples for the sampling design configuration were conducted by inspection of confidence intervals (CI) using Bonferroni adjustments (Cochran, 1977). Detection of change was defined as the ability to discriminate between the 95% CI of mean responses between the two time periods. We used the Bonferoni CI t-test because it is more suited to sample design statistics and does not require homogenous variance between two distributions to test differences in the mean responses. The absolute ability to detect changes was thus determined by the precision of the survey estimates (e.g., standard error).

13 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.13 FINAL REPORT FY RESULTS Sampling Effort Forty-three scientists from various academic, state and federal government agencies and organizations participated in the field survey (Table 1.2). Basic scientific dive statistics are shown in Table 1.3. The fish, lobster, and habitat survey teams conducted a total of 1,344 scientific dives in the Tortugas region, with 817 dives in Dry Tortugas National Park and 527 dives in Tortugas Bank. A new addition for the 2006 survey was a fish team equipped with Trimix to survey reef habitats deeper than 33 m. This team conducted a total of 24 scientific dives on the western edge of Tortugas Bank in depths to 42 m. Table 1.4 shows statistical sample sizes in terms of primary (n) and second-stage (nm) sample units by year, habitat, and management zone. A total of 254 primary units were sampled by the reef fish survey team, just slightly less than the target allocation of n=270 even though diving operations were suspended for several days due to Tropical Storm Alberto. A subset of the 254 primary units were cosampled by the spiny lobster or benthic habitat survey teams along with the reef fish teams (Figure 1.3). A complete descriptive list of these sampling sites is provided in Table 1.5 for Dry Tortugas National Park and Table 1.6 for Tortugas Bank Preliminary Analysis of Change, Baseline to 2006 Fish species richness ranged from 23 to 67 species per primary sample unit (psu) and, in general, was correlated with habitat class (Figure 1.5). Greatest reef fish species diversity was found in high rugosity habitats. For the Tortugas sampling domain, we detected a significant increase in mean species richness (mean number of species per psu) from 37.1 in the baseline to 41.5 in 2006 (Table 1.7). Diversity of exploited species in the snapper-grouper complex was also highest in high rugosity habitats (Figure 1.6); however, we detected no change in mean richness for snapper-grouper species (Table 1,7). Domain-wide estimates of frequency of occurrence, or sighting frequency, showed a relatively stable fish community structure. Although there were minor changes in ranks between years, only five of the top 50 species for the 2006 survey were not among the top 50 species for the surveys (Table 1.8). The top 50 species in 2006 included 10 species from the exploited snapper-grouper complex. Domain-wide estimates of frequency of occurrence and abundance for representative species of principal families are given in Tables 1.9 and 1.10, respectively. Among species in

14 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.14 FINAL REPORT FY 2006 the exploited snapper-grouper complex, we detected an increase in domain-wide percent occurrence between and 2006 for mutton snapper, and we detected declines in red grouper, gray snapper, and hogfish. Concomitantly, we detected an increase in domain-wide abundance for mutton snapper and decreases in abundance for red grouper and hogfish. Notably, estimates of abundance in 2006 for black grouper and yellowtail snapper were similar to the estimates for , in contrast to the positive changes observed between and Changes in abundance within management zones between and 2006 were also observed for some snapper-grouper species (Table 1.11). Declines for red grouper and hogfish were detected in Tortugas Bank Fished, whereas mutton snapper increased in Dry Tortugas National Park. Again, positive changes in abundance from baseline observed in 2004 within Tortugas Bank NTMR for red grouper, black grouper, and mutton snapper were not detected in Similarly, positive abundance changes observed in 2004 in Dry Tortugas National Park for black grouper and yellowtail snapper were not detected in Among species not targeted by exploitation, we detected domain-wide increases in occurrence between baseline and 2006 for 4 species and decreases for 3 species (Table 1.9). Likewise, we detected domain-wide increases in abundance for 3 species and decreases for 4 species (Table 1.10). These changes are in marked contrast to our findings for the 2004 survey in which no domain-wide declines in abundance were observed for non-target species. Within spatial management zones, we detected both increases and decreases in abundance between baseline and 2006 for a number of non-target species irrespective of zone (Table 1.11). Moreover, in many cases the changes observed in 2006 differed from those observed in the 2004 survey. For black grouper, the shift observed in domain-wide length composition between and 2004 towards a higher proportion of exploited-phase animals seems to have progressed even further in 2006 (Figure 1.7). In contrast, length composition of red grouper in 2006 shows only a modest increase in the proportion of exploited-phase animals from , and the shift toward larger animals is less pronounced than observed in 2004 (Figure 1.8). Temporal changes in length composition within management zones are documented in Figures 1.9 to 1.11 for black grouper and in Figures 1.12 to 1.14 for red grouper. For both species, the length composition data for 2006 suggest that the proportion of exploited-phase animals is directly

15 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.15 FINAL REPORT FY 2006 related to the level of resource protection, with highest proportions of exploited-phase animals in the no-take reserve and lowest proportions in the area open to both commercial and recreational fishing. 1.3 NEXT STEPS IN THE RESEARCH ANALYSIS Our preliminary findings indicate that changes in abundance metrics for a suite of exploited and non-target fish species observed between baseline surveys in the Tortugas region in and the 2006 survey were in many cases very different from the changes observed between baseline and We suspect that the increased hurricane activity in the intervening period between the 2004 and 2006 surveys was a contributing factor to the discrepancies in the 2004 and 2006 results. Our next research focus and challenge will be to develop, refine, and apply analysis methods for understanding the relative contributions of NTMRs, various traditional fishery management actions, community interactions, and environmental factors and events such as hurricanes in terms of impacts on fish population abundance and size metrics, and how these impacts in turn influence long-term resource sustainability. 1.4 Acknowledgments We thank Captain Frank Wasson and the crew of the M/V Spree for their expert seamanship, dive support, and their special friendship. We thank mission coordinator Jay Styron and the expert staff from the National Undersea Research Center including Doug Kesling and Tom Potts for their exceptional mission coordination and dive support in carrying out the 2006 reef fish census survey. We sincerely appreciate the enthusiasm and professionalism of all 43 participating scientists from a host of University, State and Federal institutions and for their assistance in meeting the rigor and demands of this challenging endeavor. This study was supported by funding from the National Park Service through the Cooperative Ecosystems Study Unit, Contract No. H500000B494-J , NOAA Fisheries Coral Reef Program (NA17RJ1226), and personnel support from the Florida Fish and Wildlife Conservation Commission. We thank Tom Schmidt, Bob Howard, Bob Johnson and Bob Zepp of Everglades/Dry Tortugas National Park; Brian Keller, Billy Causey and Cheva Heck of the Florida Keys National Marine Sanctuary; Alex Chester, Peter Thompson and Nancy Thompson of the National Marine Fisheries Service; Rose Mann of the Pew Institute of Ocean Sciences; and Annie Reisewitz and Ivy Kupec of the University of Miami.

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19 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.19 FINAL REPORT FY 2006 Porter, J.W., and K.G. Porter. (eds.) The Everglades, Florida Bay, and coral reefs of the Florida Keys. CRC Press, Boca Raton p. Roberts, C.M Ecological advice for the global fisheries crisis. Trends in Ecology and Evolution 12: Roberts, C.M., and N.V.C. Polunin Are marine reserves effective in the management of reef fisheries? Reviews in Fish Biology and Fisheries 1: Roberts, C.M., Bohnsack, J.A., Gell, F., Hawkins, J.P., and R. Goodridge Effects of marine reserves on adjacent fisheries. Science 294: Russ, G.R Yet another review of marine reserves as reef fisheries management tools. In: Sale, P.F., (Ed.), Coral reef fishes: dynamics and diversity in a complex ecosystem. Academic Press, San Diego, pp Russ, G.R., Alcala, A.C., Maypa, A.P., Calumpong, H.P., and A.T. White Marine reserves benefit local fisheries. Ecological Applications 14(2): Sale, P.F., Cowen, R.K., Danilowicz, B.S., Jones, G.P., Kritzer, J.P., Lindeman, K.C., Planes, S., Polunin, N.V.C., Russ, G.R., Sadovy, Y.J., and R.S. Steneck Critical science gaps impede use of no-take fishery reserves. Trends in Ecology and Evolution 20: Smith, S.G. and J.S. Ault Statistical sampling design analysis of the Puerto Rico shallow-water reef fish monitoring survey. NOAA Tech. Memo. NMFS-SEFSC-331: Steele, J., and P. Hoagland Are fisheries sustainable? Fisheries Research 64: 1-3. Stefansson, G. and A. A. Rosenberg Combining control measures for more effective management of fisheries under uncertainty; quotas, effort limitation and protected areas. Phil. Trans. Royal Soc Ser. B. 360: Stewart-Oaten, A., Murdoch, W.W., and K.R. Parker Environmental impact assessment: pseudoreplication in time? Ecology 67: Underwood, A.J Experiments in ecology and management: their logics, functions and interpretations. Australian Journal of Ecology 15: Underwood, A.J The mechanics of spatially replicated sampling programmes to detect environmental impacts in a variable world. Australian Journal of Ecology 18: US Commission on Ocean Policy An Ocean Blueprint for the 21 st Century. Final Report. Washington, DC. 676 p. Willis, T.J., Millar, R.B., Babcock, R.C., and N. Tolimieri Burdens of evidence and the benefits of marine reserves: putting Descartes before des horse? Environmental Conservation 30: Yeung C., and T.N. Lee Larval transport and retention of the spiny lobster, Panulirus argus, in the coastal zone of the Florida Keys, USA. Fisheries Oceanography 11(5): Zeller, D., and G.R. Russ Are fisheries sustainable? A counterpoint to Steele and Hoagland. Fisheries Research 67: Additional References Ault, J.S., and Ehrhardt, N.M Correction to the Beverton and Holt Z-estimator for truncated catch length-frequency distributions. ICLARM Fishbyte 9(1): Ault, J.S., Bohnsack, J.A. and G.A. Meester A retrospective ( ) multispecies assessment of coral reef fish stocks in the Florida Keys. Fishery Bulletin 96(3):

20 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.20 FINAL REPORT FY 2006 Ault, J.S., Diaz, G.A., Smith, S.G., Luo, J. and J.E. Serafy An efficient sampling survey design to estimate pink shrimp population abundance in Biscayne Bay, Florida. North American Journal of Fisheries Management 19(3): Ault, J.S., Meester, G.A., Luo, J., Smith, S.G., and K.C. Lindeman. 2000b. Natural resources affected environment. In Dry Tortugas National Park General Management Plan, National Park Service. Denver, CO. 250 p. Ault, J.S., Smith, S.G., Meester, G.A., Luo, J., and J.A. Bohnsack Site characterization for Biscayne National Park: assessment of fisheries resources and habitats. NOAA Technical Memorandum NMFS-SEFSC p. Ault, J.S., Smith, S.G., Luo, J., Meester, G.A., Bohnsack, J.A., and S.L. Miller Baseline multispecies coral reef fish stock assessment for the Dry Tortugas. NOAA Tech. Memo. NMFS-SEFSC-487. Ault, J.S., Smith, S.G., Diaz, G.A., and E.C. Franklin. 2003a. Florida hogfish fishery stock assessment. Report to the Florida Fish and Wildlife Conservation Commission. 67 pp + Figures and Tables. Ault, J.S., S.G. Smith, E.C. Franklin, J. Luo and J.A. Bohnsack. 2003b. Sampling design analysis for coral reef fish stock assessment in Dry Tortugas National Park. Final Report, National Park Service Contract No. H Ault, J.S., Bohnsack, J.A., and S.G. Smith. 2005a. Towards sustainable multispecies fisheries in the Florida USA coral reef ecosystem. Bulletin of Marine Science 76(2), in press Ault, J.S., Smith, S.G., and Bohnsack, J.A. 2005b. Evaluation of average length as an indicator of exploitation status for the Florida coral reef fish community. ICES Journal of Marine Science, 61: Bohnsack, J.A., and J.S. Ault Management strategies to conserve marine biodiversity. Oceanography 9(1): Bohnsack, J.A. and S.P. Bannerot A stationary visual census technique for quantitatively assessing community structure of coral reef fishes. U.S. Dept. Commer., NOAA Tech. Report NMFS 41, 15 p. Bohnsack, J.A., D.B. McClellan, D.E. Harper, G.S. Davenport, G.J. Konoval, A.M. Eklund, J.P. Contillo, S.K. Bolden, P.C. Fischel, G.S. Sandorff, J.C. Javech, M.W. White, M.H. Pickett, M.W. Hulsbeck, J.L. Tobias, J.S. Ault, G.A. Meester, S.G. Smith, J. Luo Baseline data for evaluating reef fish populations in the Florida Keys, NOAA Technical Memorandum NMFS-SEFSC-427. Bohnsack, J.A., Ault, J.S., and B. Causey Why have no-take marine protected areas? American Fisheries Society Symposium 42: Bohnsack, J.A., D.E. Harper, and D.B. McClellan Fisheries trends from Monroe County, Florida. Bull. Mar. Sci. 54(3): Brock, R.J., and B.F. Culhane The no-take research natural area of Dry Tortugas National Park (Florida): wishful thinking or responsible planning? American Fisheries Society Symposium 42: Claro, R., Lindeman. K.C., and Parenti, L.R Ecology of the Marine Fishes of Cuba. Smithsonian Institution Press, Washington, D.C., 253 p. Cochran, W.G Sampling techniques. 3rd edition. Wiley, New York. Culhane, B A new era for marine resource protection at Dry Tortugas and the Florida Keys. Pages in J. Selleck, ed. Natural Resource Year in Review U.S. Department of the Interior, National Park Service, Lakewood, CO.

21 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.21 FINAL REPORT FY 2006 Ehrhardt, N.M., and Ault, J.S Analysis of two length-based mortality models applied to bounded catch length frequencies. Transactions of the American Fisheries Society 121(1): Franklin, E.C., Ault, J.S., Smith, S.G., Luo, J., Meester, G.A., Diaz, G.A., Chiappone, M., Swanson, D.W., Miller, S.L., and J.A. Bohnsack Benthic habitat mapping in the Tortugas region, Florida. Marine Geodesy 26(1-2): Meester, G.A., J.S. Ault, and J.A. Bohnsack Visual censussing and the extraction of average length as a biological indicator of stock health. Naturalista sicil. Vol. XXIII (Suppl.): Meester, G.A., Ault, J.S., Smith, S.G., and A. Mehrotra An integrated simulation modeling and operations research approach to spatial management decision making. Sarsia 86: Meester, G.A., Mehrotra, A., Ault, J.S., and E.K. Baker Designing marine reserves for fishery management. Management Science 50(8): Methot, R.D Synthesis model: an adaptive framework for analysis of diverse stock assessment data. International North Pacific Fisheries Commission Bulletin, 50: Quinn, T.J., and R.B. Deriso Quantitative fish dynamics. Oxford University Press, 542 pp. Prager, M.H A suite of extensions to a non-equilibrium surplus-production model. Fishery Bulletin, 92: Schmidt, T.W., Ault, J.S., and J.A. Bohnsack Site characterization for the Dry Tortugas region: fisheries and essential habitats. Florida Keys National Marine Sanctuary and National Park Service. NOAA Technical Memorandum NMFS-SEFSC p. Shyu, M-L., Ranasingha, C., Ault, J.S., Smith, S., Harper, D., Wong, S., and J.A. Bohnsack Caribbean-wide Reef Fish Visual Census Oracle Database Management System Design and Development, Technical Report Draft, in Preparation. Smith, S.G. and J.S. Ault Statistical sampling design analysis of the Puerto Rico shallow-water reef fish monitoring survey. NOAA Technical Memorandum NMFS- SEFSC-331:1-36.

22 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.22 FINAL REPORT FY 2006 Table 1.1. (A) Habitat strata (h) characteristics and sizes in terms of primary sampling units (N h ) and area (A h ) for the Dry Tortugas sampling domain. (B) Habitat strata sizes for three management zones within the Dry Tortugas sampling domain; dashes denote habitats not found in a given management zone. (A) Reef Habitat Classification Habitat Code Degree of Patchiness Degree of Domain-wide Area Vertical Relief N h A h (km 2 ) Low-relief hardbottom LRHB Low Low 4, Low-relief spur & groove LRSG Moderate Low Patchy hardbottom in sand PHBS High Low Medium profile reef MDPR Low Moderate Rocky outcrops RKOC Moderate-High Moderate Reef terrace RFTC Low High High-relief spur & groove HRSG Moderate High Pinnacle reef RFPN High High Total 8, (B) Habitat Tortugas Bank Fished Tortugas Bank NTMR Dry Tortugas National Park Code N h A h (km 2 ) N h A h (km 2 ) N h A h (km 2 ) LRHB 1, , , LRSG PHBS MDPR RKOC RFTC HRSG RFPN Total 1, , ,

23 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.23 FINAL REPORT FY 2006 Table 1.2. Onboard scientific personnel for the 2006 Tortugas survey. Name Organization Scientific Role Ault, Dr. Jerald UM-RSMAS Principal Investigator/Fish Diver Brandt, Marilyn UM-RSMAS Fish-Tech Diver Farmer, Nick UM-RSMAS Fish Diver Feeley, Mike UM-RSMAS Fish-Tech Diver Fiechter, Jerome UM-RSMAS Fish Diver Gomez, Rick UM-RSMAS Fish-Tech Diver Kleisner, Kristin UM-RSMAS Fish Diver Larkin, Mike UM-RSMAS Fish Diver Luo, Dr. Jiangang UM-RSMAS Chief Videographer/Photographer McCrea, Ashley UM-RSMAS Fish Diver Smith, Dr. Steve UM-RSMAS Logistics Coordinator/Fish Diver Swanson, Dione UM-RSMAS Coral Diver Zurcher, Natalia UM-RSMAS Logistics Specialist/Fish-Tech Diver Baertlein, Neil NOAA Fisheries Fish Diver Balchowsky, Heather NOAA Fisheries Fish Diver Bohnsack, Dr. Jim NOAA Fisheries Co-Principal Investigator/Fish Diver Contillo, Joe NOAA Fisheries Fish Diver Davenport, Guy NOAA Fisheries Fish Diver Harman, Leah NOAA Fisheries Fish Diver Harper, Doug NOAA Fisheries Data Manager/Fish Diver Jackson, Tom NOAA Fisheries Fish Diver Javech, Jack NOAA Fisheries Fish Diver Judge, Mike NOAA Fisheries Fish Diver Kellison, Dr. Todd NOAA Fisheries Fish Diver McClellan, Dave NOAA Fisheries Logistics Specialist/Fish Diver Waara, Robert NPS Coral Diver Caldow, Chris NOAA NOS Fish Diver Woody, Kimberly NOAA NOS Fish Diver Franklin, Erik NOAA NOS Fish Diver Bertelsen, Dr. Rod FWRI Lobster Diver Braynard, Michelle FWRI Lobster Diver Cox, Carollyn FWRI Lobster Diver Eaken, Dave FWRI Lobster Diver Hawtof, David FWRI Lobster Diver Lewis, Cindy FWRI Lobster Diver Maxwell, Kerry FWRI Lobster Diver Sympson, Bill FWRI Lobster Diver Walsh, Aaron FWRI Lobster Diver Chiappone, Mark NURC/UNCW Coral Diver Kesling, Doug NURC/UNCW Coral Diver Potts, Tom NURC/UNCW Divemaster Rutten, Leann NURC/UNCW Divemaster Styron, Jay NURC/UNCW Divemaster

24 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.24 FINAL REPORT FY 2006 Table 1.3. Sampling effort by reef fish, benthic habitat, and spiny lobster teams in the Dry Tortugas region for Survey Team Number of Scientific Dives Dry Tortugas Tortugas National Park Bank Combined Areas Reef Fish-Nitrox Reef Fish-Trimix Benthic Habitat Spiny Lobster All Teams ,344 Table 1.4. Reef fish survey sample sizes in terms of primary (n) and second-stage (nm) units by habitat class and management zone for Habitat codes are defined in Table 1; dashes denote habitats not found in a given management zone. Habitat Tortugas Bank Fished Tortugas Bank NTMR Dry Tortugas National Park Domain-wide Code n nm n nm n nm n nm LRHB LRSG PHBS MDPR RKOC RFTC HRSG RFPN Total

25 Sustaining Dry Tortugas National Park Coral Reef Resources Page 1.25 FINAL REPORT FY 2006 Table 1.5. Primary sampling unit (200 by 200 m grid cell) site list for the summer 2006 survey in Dry Tortugas National Park denoting the type of sampling (Survey Team), location, spatial management zone, and habitat classification (see Table 1.1 for habitat codes). Primary Unit ID Survey Team Latitude Longitude Spatial Zone Habitat 001U Coral-Fish outside RNA PHBS 002U Fish outside RNA LRHB 003U Lobster-Fish outside RNA LRSG 004U Lobster-Fish outside RNA RKOC 005U Fish outside RNA MDPR 006U Lobster-Fish outside RNA LRHB 007U Lobster-Fish outside RNA MDPR 008U Fish outside RNA LRHB 009U Lobster-Fish outside RNA LRHB 010U Coral-Fish outside RNA LRHB 011U Lobster-Fish outside RNA PHBS 012U Fish outside RNA LRHB 013U Lobster-Fish outside RNA LRSG 014U Lobster-Fish outside RNA LRHB 015U Coral-Fish outside RNA LRHB 016U Coral-Fish outside RNA MDPR 017H Fish outside RNA LRHB 017U Fish outside RNA MDPR 018U Lobster-Fish outside RNA MDPR 019U Lobster-Fish outside RNA MDPR 020H Fish outside RNA LRHB 020U Fish outside RNA Sand 021U Lobster-Fish outside RNA HRSG 022U Lobster-Fish outside RNA LRSG 023U Coral-Fish outside RNA LRHB 024U Fish outside RNA RKOC 025U Lobster-Fish outside RNA RFPN 026U Lobster-Fish outside RNA HRSG 027U Fish outside RNA RKOC 028U Lobster-Fish outside RNA PHBS 029U Coral-Fish outside RNA MDPR 030U Lobster-Fish outside RNA RKOC 031U Fish outside RNA MDPR 032U Lobster-Fish outside RNA LRHB 033U Lobster-Fish outside RNA LRHB 034U Coral-Fish outside RNA MDPR 035U Fish RNA RFTC 036U Lobster-Fish RNA RFTC 037U Lobster-Fish RNA RFTC 038U Fish outside RNA RFTC 039U Lobster-Fish outside RNA RFTC 040U Lobster-Fish outside RNA RFTC 041U Coral-Fish outside RNA MDPR 042U Fish outside RNA RFTC

26 Positive Signs for Florida s Coral Reef Fisheries Page 1.26 Table 1.5, cont. Primary Unit ID Survey Team Latitude Longitude Spatial Zone Habitat 043U Lobster-Fish RNA MDPR 044U Lobster-Fish RNA LRHB 045U Fish outside RNA LRHB 046U Coral-Fish outside RNA MDPR 047U Lobster-Fish outside RNA LRHB 048U Lobster-Fish outside RNA LRHB 064U Coral-Fish RNA RFPN 065U Fish RNA RKOC 066U Lobster-Fish RNA RKOC 067U Lobster-Fish RNA RFPN 068U Fish RNA MDPR 069U Lobster-Fish RNA LRHB 070U Lobster-Fish RNA LRHB 071U Coral-Fish RNA LRHB 072U Fish RNA LRHB 073U Coral-Fish RNA LRHB 074U Fish RNA LRHB 100U Coral-Fish outside RNA LRHB 101U Fish outside RNA LRHB 102U Lobster-Fish outside RNA LRHB 103U Lobster-Fish outside RNA LRHB 104U Fish outside RNA LRHB 105U Coral-Fish outside RNA LRHB 106U Lobster-Fish outside RNA LRHB 107U Lobster-Fish outside RNA LRHB 165H Coral-Fish RNA LRHB 165U Coral-Fish RNA MDPR 166U Lobster-Fish RNA RKOC 167H Fish RNA RKOC 167U Fish RNA LRHB 168U Lobster-Fish RNA RKOC 169U Coral-Fish RNA HRSG 170U Fish outside RNA HRSG 171U Lobster-Fish RNA LRSG 172U Lobster-Fish outside RNA RKOC 173U Coral-Fish RNA MDPR 174H Fish RNA MDPR 174U Fish RNA LRHB 175U Lobster-Fish RNA LRHB 176U Lobster-Fish RNA LRHB 177U Fish RNA MDPR 178U Lobster-Fish RNA MDPR 179U Lobster-Fish RNA MDPR 180U Fish RNA LRHB 181U Coral-Fish RNA LRHB 182U Lobster-Fish RNA LRHB 183U Lobster-Fish RNA LRHB

27 Positive Signs for Florida s Coral Reef Fisheries Page 1.27 Table 1.5, cont. Primary Unit ID Survey Team Latitude Longitude Spatial Zone Habitat 184U Fish RNA RKOC 185U Lobster-Fish RNA LRHB 186U Coral-Fish RNA LRHB 187U Lobster-Fish RNA LRHB 188U Fish RNA LRHB 189U Lobster-Fish RNA RKOC 190U Lobster-Fish RNA RKOC 191U Coral-Fish RNA RFTC 192U Fish RNA RFPN 193U Lobster-Fish RNA RFTC 194U Lobster-Fish RNA RFTC 195U Fish RNA MDPR 196U Lobster-Fish RNA RFTC 197U Coral-Fish RNA LRHB 198U Lobster-Fish RNA LRHB 199U Fish RNA LRHB 200U Lobster-Fish RNA RKOC 201U Lobster-Fish RNA RFPN 202U Coral-Fish RNA PHBS 203U Fish RNA LRSG 204U Lobster-Fish RNA LRSG 205U Coral-Fish RNA LRHB 206U Lobster-Fish RNA PHBS 207U Coral-Fish outside RNA LRHB 208U Fish outside RNA LRHB 209U Lobster-Fish outside RNA LRHB 210U Lobster-Fish outside RNA LRHB 211U Fish outside RNA LRHB 212U Lobster-Fish outside RNA LRHB 213U Lobster-Fish RNA PHBS 214U Coral-Fish RNA MDPR 215U Fish RNA RKOC 216U Lobster-Fish RNA RKOC 217U Lobster-Fish RNA RKOC 218U Fish RNA RKOC 219U Lobster-Fish RNA RKOC 220U Lobster-Fish RNA RKOC 221U Coral-Fish RNA RFPN 222U Fish RNA LRHB 223U Lobster-Fish RNA RKOC 224U Lobster-Fish RNA RKOC 225U Coral-Fish outside RNA RKOC 226U Fish outside RNA LRSG 227U Lobster-Fish outside RNA LRSG 228U Lobster-Fish outside RNA LRSG 229U Coral-Fish outside RNA LRHB 230U Fish RNA RFPN

28 Positive Signs for Florida s Coral Reef Fisheries Page 1.28 Table 1.5, cont. Primary Unit ID Survey Team Latitude Longitude Spatial Zone Habitat 231H Lobster-Fish outside RNA Sand 231U Fish outside RNA LRHB 232U Lobster-Fish outside RNA RKOC 233H Fish outside RNA Sand-Seagrass 233U Fish outside RNA RKOC 234U Coral-Fish RNA RKOC 235U Lobster-Fish RNA MDPR 236U Lobster-Fish RNA RKOC 237U Fish RNA RKOC 238U Lobster-Fish RNA LRHB 239U Lobster-Fish RNA RKOC 240U Coral-Fish RNA RKOC 241U Fish RNA PHBS 242U Lobster-Fish RNA LRHB 243U Lobster-Fish RNA RKOC 244U Coral-Fish outside RNA RFTC 245U Fish outside RNA MDPR 246U Lobster-Fish outside RNA LRHB 247U Lobster-Fish outside RNA LRHB

29 Positive Signs for Florida s Coral Reef Fisheries Page 1.29 Table 1.6. Primary sampling unit (200 by 200 m grid cell) site list for the summer 2006 survey in Tortugas Bank denoting the type of sampling (Survey Team), location, spatial management zone, and habitat classification (see Table for habitat codes). Primary Unit ID Survey Team Latitude Longitude Spatial Zone Habitat 049U Coral-Fish ER MDPR 050U Fish ER RKOC 051U Lobster-Fish ER MDPR 052U Lobster-Fish ER RKOC 053U Fish ER MDPR 054U Lobster-Fish ER RFTC 055U Lobster-Fish ER RKOC 056U Fish ER LRHB 057U Coral-Fish ER RKOC 058U Lobster-Fish ER MDPR 059U Fish ER Sand 060U Fish ER LRHB 061U Lobster-Fish ER RFPN 062U Coral-Fish ER RKOC 063U Lobster-Fish ER RKOC 075U Lobster-Fish ER RKOC 076U Lobster-Fish ER RKOC 077U Fish ER LRHB 078U Lobster-Fish ER RKOC 079U Coral-Fish ER RKOC 080H Lobster-Fish ER RKOC 080U Lobster-Fish ER LRHB 081U Fish ER LRHB 082U Coral-Fish ER RFTC 083U Fish ER RFTC 084U Lobster-Fish ER PHBS 085H Lobster-Fish ER LRHB 085U Lobster-Fish ER RKOC 086U Fish ER LRSG 087U Coral-Fish ER LRHB 088U Lobster-Fish ER MDPR 089U Lobster-Fish ER MDPR 090U Fish ER RKOC 091U Lobster-Fish ER MDPR 092U Lobster-Fish ER RKOC 093U Fish ER LRHB 094H Lobster-Fish ER RKOC 094U Lobster-Fish ER LRHB 095U Coral-Fish ER LRHB 096U Lobster-Fish outside ER LRHB 097U Fish ER RFPN 098U Lobster-Fish outside ER MDPR 099U Lobster-Fish outside ER RKOC 108U Lobster-Fish ER RFTC

30 Positive Signs for Florida s Coral Reef Fisheries Page 1.30 Table 1.6, cont. Primary Unit ID Survey Team Latitude Longitude Spatial Zone Habitat 109U Coral-Fish ER MDPR 110U Fish ER PHBS 111U Lobster-Fish ER RFTC 112U Lobster-Fish ER RFTC 113U Fish ER RFTC 114U Lobster-Fish ER RFTC 115U Lobster-Fish ER MDPR 116U Coral-Fish ER MDPR 117U Fish ER RFTC 118U Lobster-Fish ER RFTC 119U Lobster-Fish ER MDPR 120U Lobster-Fish ER RFTC 121U Lobster-Fish ER RFTC 122U Coral-Fish ER RFTC 123U Fish ER RFTC 124U Lobster-Fish ER RFTC 125U Lobster-Fish ER RFPN 126U Fish ER LRHB 127U Lobster-Fish ER LRHB 128U Lobster-Fish ER MDPR 129U Coral-Fish ER MDPR 130U Fish ER RFTC 131U Lobster-Fish ER PHBS 132U Lobster-Fish ER RKOC 133U Lobster-Fish ER RFPN 134U Lobster-Fish outside ER MDPR 135U Coral-Fish outside ER MDPR 136U Fish outside ER RFTC 137U Lobster-Fish outside ER LRHB 138U Lobster-Fish outside ER LRHB 139U Fish outside ER LRHB 140U Lobster-Fish outside ER PHBS 141U Lobster-Fish outside ER LRHB 142U Coral-Fish outside ER MDPR 143U Fish outside ER MDPR 144U Lobster-Fish outside ER LRHB 145U Lobster-Fish outside ER LRHB 146U Lobster-Fish outside ER RKOC 147U Coral-Fish outside ER LRHB 148U Fish outside ER LRHB 149U Lobster-Fish outside ER MDPR 150U Lobster-Fish outside ER LRHB 151U Fish outside ER LRHB 152U Lobster-Fish outside ER LRHB 153U Coral-Fish outside ER LRHB 154H Lobster-Fish outside ER RKOC

31 Positive Signs for Florida s Coral Reef Fisheries Page 1.31 Table 1.6, cont. Primary Unit ID Survey Team Latitude Longitude Spatial Zone Habitat 154U Lobster-Fish outside ER LRHB 155U Fish outside ER LRHB 156U Lobster-Fish outside ER LRHB 157U Lobster-Fish outside ER LRHB 158U Fish outside ER LRHB 159U Coral-Fish outside ER RKOC 160U Lobster-Fish outside ER LRHB 161U Lobster-Fish outside ER LRHB 162U Fish outside ER LRHB 163U Lobster-Fish outside ER LRHB 164U Lobster-Fish outside ER PHBS

32 Positive Signs for Florida s Coral Reef Fisheries Page 1.32 Table 1.7. Domain-wide estimates of mean species richness and associated standard errors (SE) for two taxa groups for baseline years and the 2004 and 2006 surveys. Statistical significant change from baseline: ns, not significant; *, p<0.05; **, p<0.01; ***, p< Mean Species Richness (SE) Taxa Change 2006 Change All Reef Fishes 37.1 (0.7) 38.1 (0.5) ns 41.5 (0.6) *** Exploited Snappers & Groupers 7.8 (0.2) 7.8 (0.2) ns 8.0 (0.2) ns

33 Positive Signs for Florida s Coral Reef Fisheries Page 1.33 Table 1.8. Ranked top 50 reef fish species in terms of percent occurrence for 2006 compared to 2004 and Common names in bold denote species in the exploited snapper-grouper complex. Occurrence Rank Common Name Scientific Name Family bluehead Thalassoma bifasciatum Labridae cocoa damselfish Stegastes variabilis Pomacentridae striped parrotfish Scarus iseri Scaridae yellowhead wrasse Halichoeres garnoti Labridae slippery dick Halichoeres bivittatus Labridae bridled goby Coryphopterus glaucofraenum Gobiidae saddled blenny Malacoctenus triangulatus Labrisomidae white grunt Haemulon plumierii Haemulidae redband parrotfish Sparisoma aurofrenatum Scaridae ocean surgeon Acanthurus bahianus Acanthuridae saucereye porgy Calamus calamus Sparidae spotted goatfish Pseudupeneus maculatus Mullidae bicolor damselfish Stegastes partitus Pomacentridae blue tang Acanthurus coeruleus Acanthuridae yellowtail snapper Ocyurus chrysurus Lutjanidae tomtate Haemulon aurolineatum Haemulidae seaweed blenny Parablennius marmoreus Blenniidae greenblotch parrotfish Sparisoma atomarium Scaridae yellowhead jawfish Opistognathus aurifrons Opistognathidae butter hamlet Hypoplectrus unicolor Serranidae red grouper Epinephelus morio Serranidae masked goby Coryphopterus personatus Gobiidae purple reeffish Chromis scotti Pomacentridae stoplight parrotfish Sparisoma viride Scaridae neon goby Elacatinus oceanops Gobiidae blue angelfish Holacanthus bermudensis Pomacanthidae spotfin butterflyfish Chaetodon ocellatus Chaetodontidae gray angelfish Pomacanthus arcuatus Pomacanthidae clown wrasse Halichoeres maculipinna Labridae hogfish Lachnolaimus maximus Labridae barred hamlet Hypoplectrus puella Serranidae beaugregory Stegastes leucostictus Pomacentridae blue hamlet Hypoplectrus gemma Serranidae reef butterflyfish Chaetodon sedentarius Chaetodontidae foureye butterflyfish Chaetodon capistratus Chaetodontidae blue dartfish Ptereleotris calliura Ptereleotridae sharpnose puffer Canthigaster rostrata Tetraodontidae doctorfish Acanthurus chirurgus Acanthuridae porkfish Anisotremus virginicus Haemulidae mutton snapper Lutjanus analis Lutjanidae threespot damselfish Stegastes planifrons Pomacentridae graysby Cephalopholis cruentatus Serranidae yellowtail reeffish Chromis enchrysura Pomacentridae harlequin bass Serranus tigrinus Serranidae bar jack Caranx ruber Carangidae Spanish hogfish Bodianus rufus Labridae queen angelfish Holacanthus ciliaris Pomacanthidae cero Scomberomorus regalis Scombridae black grouper Mycteroperca bonaci Serranidae bluelip parrotfish Cryptotomus roseus Scaridae

34 Positive Signs for Florida s Coral Reef Fisheries Page 1.34 Table 1.9. Domain-wide estimates of percent occurrence for representative exploited and nontarget fish species for baseline years and the 2004 and 2006 surveys. Statistical significant change from baseline: ns, not significant; *, p<0.05; **, p<0.01; ***, p< Percent Occurrence (SE) Taxa Change 2006 Change Snapper-Grouper Complex Groupers (Serranidae) Goliath grouper (Epinephelus itajara) 0.5 (0.4) 1.3 (0.5) ns 1.1 (2.2) ns Red grouper (E. morio) 67.0 (3.3) 62.8 (3.1) ns 58.5 (3.2) * Nassau grouper (E. striatus) 1.0 (0.6) 0.3 (0.2) ns 0.9 (3.3) ns Black grouper (Mycteroperca bonaci) 19.5 (2.5) 28.8 (2.4) ** 15.9 (3.3) ns Snappers (Lutjanidae) Mutton snapper (Lutjanus analis) 14.8 (2.4) 25.8 (3.0) *** 24.8 (2.9) *** Gray snapper (L. griseus) 17.3 (2.5) 12.2 (1.5) * 10.4 (1.1) ** Yellowtail snapper (Ocyurus chrysurus) 74.7 (3.2) 68.1 (3.1) * 72.9 (3.4) ns Wrasses (Labridae) Hogfish (Lachnolaimus maximus) 52.8 (3.5) 42.6 (3.0) ** 44.6 (3.0) * Grunts (Haemulidae) White grunt (Haemulon plumieri) 82.0 (2.7) 71.5 (2.7) *** 80.2 (1.4) ns Bluestriped grunt (H. sciurus) 6.4 (1.7) 7.7 (1.2) ns 6.5 (2.9) ns Non-Target Fishes Surgeonfishes (Acanthuridae) Ocean surgeon (Acanthurus bahianus) 54.9 (3.3) 60.3 (2.7) ns 79.2 (2.6) *** Blue tang (A. coeruleus) 76.4 (3.1) 80.9 (2.2) ns 73.4 (2.9) ns Butterflyfishes (Chaetodontidae) Foureye butterflyfish (Chaetodon capistratus) 34.0 (3.3) 42.3 (2.8) * 33.7 (3.3) ns Spotfin butterflyfish (C. ocellatus) 56.4 (3.4) 49.9 (3.0) ns 47.7 (3.5) * Goatfishes (Mullidae) Spotted goatfish (Psuedupeneus maculatus) 50.7 (3.6) 71.7 (2.2) *** 76.9 (2.7) *** Angelfishes (Pomacanthidae) Blue angelfish (Holocanthus bermudensis) 57.9 (3.2) 55.9 (2.7) ns 49.8 (0.7) * Gray angelfish (Pomacanthus arcuatus) 45.5 (3.3) 43.9 (2.8) ns 45.7 (1.6) ns Damselfishes (Pomacentridae) Purple reeffish (Chromis scotti) 37.2 (3.4) 62.2 (3.1) *** 52.2 (2.8) *** Bicolor damselfish (Stegastes partitus) 72.7 (2.9) 72.6 (2.3) ns 73.9 (3.1) ns Cocoa damselfish (S. variabilis) 87.7 (2.3) 90.0 (2.0) ns 97.1 (3.5) *** Parrotfishes (Scaridae) Striped parrotfish (Scarus iseri) 88.4 (2.4) 94.3 (1.3) * 91.4 (0.3) ns Redband parrotfish (Sparisoma aurofrenatum) 80.8 (2.9) 86.9 (1.9) * 79.8 (1.8) ns Stoplight parrotfish (Sparisoma viride) 59.3 (3.5) 64.5 (3.3) ns 51.8 (0.4) *

35 Positive Signs for Florida s Coral Reef Fisheries Page 1.35 Table Domain-wide estimates of abundance (and associated coefficient of variation CV) for representative exploited and nontarget fish species for baseline years and the 2004 and 2006 surveys, and changes from baseline. Statistical significant change from baseline years: ns, not significant; *, p<0.05; **, p<0.01; ***, p< CV Abundance CV Abundance (%) (millions) (%) Change (millions) Abundance (millions) CV (%) Change Taxa Snapper-Grouper Complex Red grouper % ns % *** Black grouper % *** % ns Mutton snapper % *** % ** Gray snapper % ns % ns Yellowtail snapper % * % ns Hogfish % ns % ** White grunt % ns % ns Bluestriped grunt % ns % ns Non-Target Fishes Ocean surgeon % ns % ** Blue tang % *** % ns Foureye butterflyfish % ns % ns Spotfin butterflyfish % ns % ** Spotted goatfish % *** % *** Blue angelfish % ns % *** Gray angelfish % ns % ns Purple reeffish % *** % ns Bicolor damselfish % ** % *** Cocoa damselfish % ns % *** Striped parrotfish % * % ns Redband parrotfish % ns % ns Stoplight parrotfish % *** % *

36 Positive Signs for Florida s Coral Reef Fisheries Page 1.36 Table Population abundance changes from baseline years for the 2004 and 2006 surveys within management zones in the Dry Tortugas region. Statistical significant change from baseline years: ns, not significant; *, p<0.05; **, p<0.01; ***, p< Tortugas Bank, Fished Tortugas Bank, NTMR Dry Tortugas National Park Snapper-Grouper Complex Red grouper -43% * -48% ** +38% * -21% ns -9% ns -16% ns Black grouper +84% ns -69% ns +120% * -35% ns +128 *** +42% ns Mutton snapper -45% ns +23% ns +303% ** -20% ns +142% *** +94% ** Gray snapper -96% ns -96% ns -51% ns -54% ns +270% ns -55% ns Yellowtail snapper -19% ns -15% ns +367% ns +78% ns +132% *** -22% ns Hogfish -27% ns -78% *** +6% ns -13% ns -25% ns -16% ns White grunt +7% ns -18% ns +24% ns +531% ns +2% ns -1% ns Bluestriped grunt +50% ns -69% ns +13% ns +121 ns +242% ns -2% ns Non-Target Fishes Ocean surgeon +2% ns +29% ns +75% ** +51% ** -9% ns +47% * Blue tang +13% ns -22% ns +28% ns -19% ns +99% *** +25% ns Foureye butterflyfish +86% * +5% ns -18% ns -12% ns +32% ns -25% ns Spotfin butterflyfish +35% ns +13% ns -31% * -12% ns 0% ns -40% *** Spotted goatfish +133% ** +64% * +326% *** +273% *** +175% *** +135% *** Blue angelfish -18% ns +4% ns -20% ns -41% *** +31% * -32% * Gray angelfish -24% ns -10% ns +58% ns -5.8% ns +120% ns +31% * Purple reeffish +31% ns +49% ns +42% ns +11% ns +263% *** +94% ** Bicolor damselfish +6% ns -41% * +73% ** -40% ** +17% ns -29% ns Cocoa damselfish -28% ns +27% ns -21% ns +76% *** +6% ns +46% *** Striped parrotfish +51% ns +14% ns +127% * +39% ** +9% ns +7.3% ns Redband parrotfish +121% *** +1% ns +26% ns -7% ns +56% ns -30% ns Stoplight parrotfish +9% ns +30% ns +26% ns -41% * +84% *** -24% ns

37 Positive Signs for Florida s Coral Reef Fisheries Page 1.37 Figure 1.1. The Florida Keys coastal marine ecosystem. The coral reef tract runs offshore from Key Biscayne 380 km southwest to the Dry Tortugas. The Florida Keys National Marine Sanctuary boundary and Biscayne National Park and Dry Tortugas National Park are shown.

38 Positive Signs for Florida s Coral Reef Fisheries Page 1.38 Figure 1.2. Tracks of hurricanes impacting the Tortugas region in 2004 and 2005.

39 Positive Signs for Florida s Coral Reef Fisheries Page 1.39 Tortugas Bank NTMR Tortugas Bank Fished Dry Tortugas National Park FKNMS Figure 1.3. Map of the Dry Tortugas region showing bathymetry, spatial management boundaries, and primary units sampled by survey team (COR=coral-fish, FSH=fish, LOB=lobster-fish) during the 2006 expedition. Bathymetry is shaded from red (shallow, 0-3m) to blue (deep, >50m).

40 Positive Signs for Florida s Coral Reef Fisheries Page 1.40 Figure 1.4. Graphical illustration of the two-stage stratified random sampling design used in the Tortugas 2006 survey: (A) benthic habitats are mapped at 200 x 200 m grids termed primary sampling units (psu), shown for Dry Tortugas National Park; (B) within a psu, paired teams of divers are randomly placed in second-stage units where they concurrently sample for reef fishes (2 stationary RVC cylinders, solid dots), reef habitats (4 transects, pink lines), and/or lobsters (2 transects, red lines).

41 Positive Signs for Florida s Coral Reef Fisheries Page 1.41 Figure 1.5. Reef fish diversity as measured by species richness (number of species per 200 x 200 m primary unit) from the 2006 Tortugas monitoring expedition.

42 Positive Signs for Florida s Coral Reef Fisheries Page 1.42 Figure 1.6. Diversity of the exploited snapper-grouper complex as measured by species richness (number of species per 200 x 200 m primary unit) from the 2006 Tortugas monitoring expedition.

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