Marine Sanctuaries Conservation Series MSD-01-1

Distribution and Sighting Frequency of

Reef Fishes in the Florida Keys

National Marine Sanctuary

U.S. Department of Commerce

National Oceanic and Atmospheric Administration

National Ocean Service

Office of Ocean and Coastal Resource Management

Marine Sanctuaries Division

January 2001
About the Marine Sanctuaries Conservation Series

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SUGGESTED CITATION

Jeffrey, C.F.G., C. Pattengill-Semmens, S. Gittings, and M. E. Monaco. 2001.

Distribution and sighting frequency of reef fishes in the Florida Keys National Marine

Sanctuary. Marine Sanctuaries Conservation Series MSD-01-1. U.S. Department of

Commerce, National Oceanic and Atmospheric Administration, Marine Sanctuaries

Division, Silver Spring, MD. 51 pp.
DISTRIBUTION AND SIGHTING FREQUENCY OF REEF FISHES IN THE

FLORIDA KEYS NATIONAL MARINE SANCTUARY

Jeffrey, C.F.G1,4., C. Pattengill-Semmens2, 3, S. Gittings2, and M. E. Monaco1

1

National Center for Coastal Ocean Science/

Center for Coastal Monitoring and Assessment/

Biogeography Program

2

National Marine Sanctuaries Program

3

Reef Environmental Education Foundation

4 University of Georgia

January 2001
v

CONTACTS:

For more information on National Ocean Service (NOS) Biogeography Program:

Dr. Mark E. Monaco

NOAA/NOS/CCMA Biogeography Program

NSCI/1, SSMC-4, 9409

1305 East West Highway

Silver Spring, MD 20910

(301) 713-3028 ext. 160

Mark.Monaco@noaa.gov

Christopher F. Jeffrey

NOAA/NOS/CCMA Biogeography Program

NSCI/1, SSMC-4, 9222

1305 East West Highway

Silver Spring, MD 20910

(301) 713-3028 ext. 134

Chris.Jeffrey@noaa.gov

Web address: biogeo.nos.noaa.gov

For more information on Reef Environmental Foundation:

Dr. Christy Pattengill-Semmens

Reef Environmental Education Foundation

PO Box 246 Key Largo, FL 33037

(305) 451-0312

christy@reef.org

Laddie Akins

Reef Environmental Education Foundation

PO Box 246 Key Largo, FL 33037

(305) 451-0312

lad@reef.org

Web address: www.reef.org

For more information on NOS Marine Sanctuaries Division:

Dr. Steve Gittings

NOAA Sanctuaries and Reserves Division

N/ORM62, SSMC-4, 11645

1305 East West Highway

Silver Spring, MD 20910

 (301)713-3125 ext. 130

steve.gittings@noaa.gov

Web address: www.sanctuaries.nos.noaa.gov
vi

ABSTRACT

This study analyzed species richness, distribution, and sighting frequency of selected reef

fishes to describe species assemblage composition, abundance, and spatial distribution patterns

among sites and regions (Upper Keys, Middle Keys, Lower Keys, and Dry Tortugas) within the

Florida Keys National Marine Sanctuary (FKNMS) barrier reef ecosystem. Data were obtained

from the Reef Environmental Education Foundation (REEF) Fish Survey Project, a volunteer

fish-monitoring program. A total of 4,324 visual fish surveys conducted at 112 sites throughout

the FKNMS were used in these analyses. The data set contained sighting information on 341 fish

species comprising 68 families. Species richness was generally highest in the Upper Keys sites

(maximum was 220 species at Molasses Reef) and lowest in the Dry Tortugas sites. Encounter

rates differed among regions, with the Dry Tortugas having the highest rate, potentially a result

of differences in the evenness in fishes and the lower diversity of habitat types in the Dry

Tortugas region. Geographic coverage maps were developed for 29 frequently observed species.

Fourteen of these species showed significant regional variation in mean sighting frequency

(%SF). Six species had significantly lower mean %SF and eight species had significantly higher

mean %SF in the Dry Tortugas compared with other regions. Hierarchical clustering based on

species composition (presence-absence) and species % SF revealed interesting patterns of

similarities among sites that varied across spatial scales. Results presented here indicate that

phenomena affecting reef fish composition in the FKNMS operate at multiple spatial scales,

including a biogeographic scale that defines the character of the region as a whole, a reef scale

(~50-100 km) that include meso-scale physical oceanographic processes and regional variation in

reef structure and associated reef habitats, and a local scale that includes level of protection,

cross-shelf location and a suite of physical characteristics of a given reef. It is likely that at both

regional and local scales, species habitat requirements strongly influence the patterns revealed in

this study, and are particularly limiting for species that are less frequently observed in the Dry

Tortugas. The results of this report serve as a benchmark for the current status of the reef fishes

in the FKNMS. In addition, these data provide the basis for analyses on reserve effects and the

biogeographic coupling of benthic habitats and fish assemblages that are currently underway.

Key Words: Florida Keys National Marine Sanctuary, biogeography, distribution, reef

fish, volunteer data, Reef Environmental Education Foundation
vii

TABLE OF CONTENTS

Page

Abstract.................................................................................................................................. ii

Table of Contents................................................................................................................... iii

List of Tables......................................................................................................................... iv

List of Figures........................................................................................................................ v

Introduction........................................................................................................................... 1

Objectives.............................................................................................................................. 1

Project Overview................................................................................................................... 2

Methods................................................................................................................................. 3

Study Area: The Florida Keys................................................................................... 3

Field Data Collection................................................................................................. 3

Data Processing......................................................................................................... 3

Statistical Analysis and Development of Species Distribution Maps 4

Results................................................................................................................................... 5

Survey Effort............................................................................................................. 5

Patterns of Species Richness..................................................................................... 5

Regional Patterns in Species Distribution................................................................. 6

Regional Patterns in Species Composition................................................................ 7

Discussion............................................................................................................................. 8

Literature Cited...................................................................................................................... 10

Acknowledgements............................................................................................................... 11

Tables..................................................................................................................................... 12

Figures................................................................................................................................... 15

Appendix 1- List of sites in the FKNMS and Dry Tortugas surveyed by REEF................. 43

Appendix 2. Species list for the FKNMS and Dry Tortugas................................................. 46
viii

LIST OF TABLES

Page

Table 1. Most frequently observed species in the Florida Keys based on sighting

frequency (%SF). Data are from REEF (2000)........................................ 12

Table 2. The number of species per family ranked among the twenty most frequently

observed species in the Florida Keys and Dry Tortugas....................................... 12

Table 3. Most frequently observed species in the Dry Tortugas ranked by sighting

frequency.................................................................................... 13

Table 4. Results of Wilcoxon / Kruskal-Wallis non-parametric (Rank Sums) and Tukey

HSD tests............................................................................................................... 14
ix

LIST OF FIGURES

Figure 1. Map of the Florida Keys showing regional boundaries....................... 15

Figure 2. Classification scheme used in mapping fish abundance...................... 16

Figure 3. Frequency distribution and summary statistics for survey time........... 17

Figure 4. Distribution map of visual surveys done by REEF divers................... 18

Figure 5. Level of survey effort compared among expert and novice divers...... 19

Figure 6. Cumulative species-time curve for reef fishes in the FKNMS............ 20

Figure 7. Regional cumulative species richness and analysis of variance.......... 21

Figure 8. Proportion of species richness.............................................................. 22

Figure 9. Distribution map of species richness................................................... 23

Figure 10. Mean species richness per diver survey............................................. 24

Figure 11. Species and family richness among expert and novice divers............. 25

Figure 12. Spatial distribution and relative %SF of bluestriped grunt................. 26

Figure 13. Spatial distribution and relative %SF of yellowtail damselfish.......... 27

Figure 14. Spatial distribution and relative %SF of sergeant major..................... 28

Figure 15. Spatial distribution and relative %SF of porkfish.............................. 29

Figure 16. Spatial distribution and relative %SF of sharpnose puffer.................. 30

Figure 17. Spatial distribution and relative %SF of foureye butterflyfish............ 31

Figure 18. Spatial distribution and relative %SF of barred hamlet....................... 32

Figure 19. Spatial distribution and relative %SF of blue hamlet........................... 33

Figure 20. Spatial distribution and relative %SF of butter hamlet........................ 34

Figure 21. Spatial distribution and relative %SF of cocoa damselfish................. 35

Figure 22. Spatial distribution and relative %SF threespot damselfish................. 36

Figure 23. Spatial distribution and relative %SF of blue angelfish....................... 37
x

Figure 24. Spatial distribution and relative %SF of striped parrotfish.................. 38

Figure 25. Spatial distribution and relative %SF of neon goby............................. 39

Figure 26. Results of hierarchical clustering and correspondence analysis to

determine similarity among sites based on species composition......... 40

Figure 27. Results of hierarchical clustering and correspondence analysis to

determine similarity among sites based on species percent sighting

frequency (%SF).................................................................................. 41

Figure 28. Dendograms showing similarities in species composition and species

sighting frequency among regions........................................................ 42
1

INTRODUCTION

Reef fish communities are a major component of tropical and sub tropical fish faunas.

Fishes dominate the top of coral reef food webs and play an important role as herbivores, and

their presence and abundance reflect the overall condition of an area (Ogden and Lobel, 1978).

Reef fishes also support important commercial and recreational fisheries, and because these

fishes represent a large proportion of the biomass in coastal environments, intense fishing

pressure may have significant effects on ecosystem processes (Richards and Lindeman, 1987).

It is generally accepted that reef fishery resources of the Florida Keys are highly stressed,

and increased documentation of declining fisheries and marine environments has prompted the

use of no-take marine reserves to protect these important resources (Bohnsack and Ault, 1996;

NOAA, 1996; Allison et al., 1998; Ault et al., 1998). Additionally, the 1990 amendment of the

Magnuson Fishery Conservation and Management Act (H.R. 2061) has increased focus on

habitat protection within the United States coastal waters.

Implementing marine protected areas to enhance fishery resources (e.g. increase biomass)

requires an understanding of the complex interactions among several physical and biological

factors (e.g., ocean currents, habitat distribution, and reproductive behavior) that determine

broad-scale patterns of fish abundance and distribution. Specifically, resource managers need a

better understanding of the natural spatial and temporal variability exhibited by marine

populations as well as the ecological relationships among ecosystems, habitats, and the living

resources they contain. Reef fish populations and assemblages often vary greatly among habitat

patches at varying scales such as physiographic reef zones or reef types (Williams, 1991). Thus,

designing effective monitoring or resource management programs requires an understanding of a

population’s spatial and temporal patterns of distribution.

This report describes the reef fish assemblage composition, abundance, and spatial

distribution patterns within the Florida Keys National Marine Sanctuary (FKNMS) barrier reef

ecosystem. It is the first part of a larger project, and provides the basis for future analyses of

reserve effects and biogeographic coupling of benthic habitats and fish assemblages. Data were

obtained from the Reef Environmental Education Foundation (REEF) Fish Survey Project, a

volunteer fish-monitoring program (REEF 2000).

The specific objectives of this report are as follows:

1. Provide fish species composition of the FKNMS;

2. Describe the diversity and distribution of reef fishes in the FKNMS; and

3. Compare the species richness, distribution, and relative abundance of selected species

and families among sites and regions in the FKNMS.
2

PROJECT OVERVIEW

In 1999, the Biogeography Program of the National Oceanic and Atmospheric

Administration (NOAA) initiated a joint project with the NOAA’s Marine Sanctuaries Division

(MSD) and the Reef Environmental Education Foundation (REEF). The project’s goal was to

describe the distribution and abundance of reef fishes in the FKNMS, and to use that information

to evaluate fish-habitat interactions and the performance of management zones.

The Biogeography Program within the Center for Coastal Monitoring and Assessment

(CCMA) was established to develop a knowledge base of living marine resource distributions

and ecology throughout the Nation's estuarine and coastal environments and to provide resource

managers with an improved ecosystem basis for decision-making. The program integrates

information on the distribution and abundance of species, distribution of habitats, and defines the

strength of species-habitat associations using a suite of technologies including GIS and modeling

tools. The program’s goal is to provide resource managers with a variety of tools to successfully

manage living resources.

In November 1990, the United States Congress passed the Florida Keys National Marine

Sanctuary and Protection Act (HR 5909) that designated the FKNMS. The Act authorized

NOAA to develop and implement a comprehensive management plan to manage and protect

Sanctuary resources for the enjoyment of present and future generations. The FKNMS comprises

approximately 9,500 km2

 of coastal and oceanic water and submerged lands organized into five

management areas: Wildlife Management Areas (WMA), Ecological Reserves (ER), Sanctuary

Preservation Areas (SPA), Existing Management Areas (EMA), and Special Use/Research Only

Areas (SA/RO). On July 1, 1997, the FKNMS Management Plan, which included a large-scale

marine zoning plan became effective and closed 23 areas (eighteen SPAs, four SAs, and one ER)

to all extractive use (NOAA, 1996). These zones aim to protect the biological diversity and

integrity of the marine environment in the Keys. In addition to providing areas that are limited to

non-extractive recreation, these no-take zones are intended to act as replenishment zones where

the total abundance of fishes, their average size, and their overall egg production may increase.

In the year 2002, State and Federal managers will reevaluate the use of zones as a management

tool. Therefore, the FKNMS and NOAA's South Florida Restoration Fund are supporting a widescale, three-tiered monitoring program to evaluate the effects of the zones on biodiversity and

human activities. Monitoring projects include research on coral, algae, fish, lobster, and human

values, and focus on three levels: ecosystem, human/ecosystem interface, and volunteer

monitoring of ecosystem health.

In 1997, REEF’s Advanced Assessment Team (AAT), made up of highly trained and

active volunteers, was contracted to annually monitor reef fish populations at 31 sites within the

FKNMS as part of the Sanctuary’s zone monitoring program. REEF is a nonprofit organization

founded in 1990 that educates the public about marine resources and enables divers and

snorkelers to participate in long-term monitoring. REEF accomplishes this through its Fish

Survey Project, which was developed by REEF with support from The Nature Conservancy

(TNC) and guidance from the National Marine Fisheries Service (NMFS) Southeast Fisheries
3

Science Center. The Fish Survey Project is now in place in the tropical western Atlantic, the west

coast of the US and Canada, and in the tropical eastern Pacific (Gulf of California to the

Galapagos Islands). Participants conduct standardized surveys as part of their regular diving

activities, and the data collected are input into REEF’s database. Data summaries are available

on the REEF Website (http://www.reef.org) and data files are available to the research,

conservation, and management communities. REEF has been monitoring fishes in the Florida

Keys since 1993, and to date its members have completed more than 6,387 fish surveys in

Florida. This extensive data set has become an important source of information on reef fishes for

the FKNMS.

METHODS

Study area: the Florida Keys

The Florida Keys comprise an island archipelago that extends 320 km southwest from

Soldier Key in Biscayne Bay to the Dry Tortugas (Figure 1). To the north and west, the Keys are

bounded by Biscayne Bay and the Gulf of Mexico; to the east and south, they are bounded the

Straits of Florida. Submerged aquatic habitats include extensive seagrass beds and an extensive

coral reef tract that extends 8 km off-shore toward the Atlantic Ocean. For this study, the

FKNMS was divided into four regions reflecting geomorphological differences (FMRI, 1998).

The regions were the Upper Keys (Key Largo to Upper Matecumbe Key), Middle Keys (Upper

Matecumbe Key to Pigeon Key), Lower Keys (Little Duck Key to Marquesas Key), and the Dry

Tortugas (Figure 1). Hereafter, the term “Florida Keys” will be used to collectively describe the

Upper, Middle and Lower Keys excluding the Dry Tortugas region.

Field data collection

Data on fish presence and relative abundance were collected between Feb 7, 1994 and

August 9, 1999, by REEF volunteers using a visual survey method called the Roving Diver

Technique (RDT; Schmitt and Sullivan, 1996). The RDT involves divers swimming freely about

a dive site (within a 100-m radius of the starting point) and recording every fish species that can

be positively identified. The survey begins as soon as the diver enters the water. At the

conclusion of each survey, the diver assigns each recorded species to one of four log10 abundance

categories [single (1); few (2-10); many (11-100); and abundant (>100)] based on the

approximate number of individuals seen. Survey location, survey time, depth, temperature, and

other environmental data pertinent to the survey are also noted. All data are recorded on REEF

survey sheets printed on underwater paper and are transferred to standardized data scan sheets,

which are returned to REEF and optically scanned into a database (REEF, 2000).

Data processing

RDT survey data files obtained from REEF were imported into JMP statistical software

(Version 3.2.2, SAS Institute Inc.) for processing and analysis. Each survey was assigned a

unique identification number and was used as a replicate within survey sites. Site (point sample)

locations were identified by unique geographic zone codes and by latitude and longitude.
4

Five hundred and eighteen REEF divers conducted 4,431 surveys at 119 sites in the

Florida Keys and Dry Tortugas from July 1993 to August 1999 (Appendix 1). Recorded survey

time varied significantly, ranging from 10 minutes to 245 minutes (4.75 hr), and were normally

distributed around a mean of 59.4 min ± 0.2 min (Figure 3). Approximately 96% (4,331) of the

surveys ranged between 30 and 100 minutes (Figure 3). Because species richness data may be

influenced by observation time, surveys shorter than 30 minutes or greater than 100 minutes

were considered outliers, and data from these surveys were not used in statistical analyses.

Additionally, sites with fewer than three replicate surveys were excluded, resulting in 4,324

surveys from 112 sites being used for analysis of species richness and sighting frequency.

Species richness (R), defined as the total number of species documented, was calculated

for each survey, site, and region. Three parameters, percent sighting frequency (%SF), density

score (D), and abundance score (A), were calculated for each species by site and by region (after

Schmitt and Sullivan 1996). Percent sighting frequency was the percentage of all survey dives in

which the particular species or family was recorded. The density score for each species, a

weighted average index based on the frequency of observations in different abundance

categories, was calculated as:

D = [(nS x 1)+(nF x 2) + (nM x 3) + (nA x 4)] / (nS + nF + nM + nA),

where nS, nF, nM, and nA represent the number of times each abundance category was assigned

for a given species. An abundance score (where A = D x %SF) was calculated to account for

zero observations.

Statistical analysis and development of species distribution maps

A series of N-way ANOVA or Kruskal-Wallis non-parametric Rank Sums tests was

conducted with JMP statistical software (Ver. 3.2.6, SAS Institute, 1999) to determine the

important factors that may be influencing two parameters of reef fishes in the Florida Keys:

species richness and sighting frequency. Raw data and estimated variables were checked for

normality to determine whether parametric or non-parametric statistical procedures should be

used. Surveys were considered replicates, whereas location, region and other factors (e.g., diver

experience and habitat types) were considered treatment effects. Statistical analyses (including

comparisons of means among treatments) were done with α = 0.05 to test for significant

differences.

Distribution maps for the most frequently observed species were created using four equal

%SF quartiles (Figure 2). Data were imported into Arc View GIS software (ver. 3.1.1, ESRI Inc.,

1999) and geographic coverages were created with base maps of the FKNMS region.

Similarity in species assemblage composition among sites and regions was determined by

hierarchical clustering (Ward’s minimum variance) and correspondence analysis. Hierarchical

clustering was used to group sites based on species composition (presence-absence) and species

%SF such that sites that were most similar clustered more closely than sites that were more

dissimilar. The clustering analysis included only sites with at least five surveys and all species
5

were included. Resulting clusters were plotted as dendograms so that regional patterns of

assemblage composition could be detected more easily. Correspondence analysis was used to

determine if any significant associations existed between the resulting clusters and the four

geographic regions of the FKNMS.

RESULTS

Survey effort

Survey effort varied substantially among regions and between diver types (Figures 4 and

5). The Upper Keys sites were most intensely surveyed (2,595 surveys) and Dry Tortugas sites

were the least surveyed (411 surveys; Figure 4). REEF volunteer divers are classified into two

groups, novice and expert. Expert divers, those with at least 35 surveys and a score of 90% or

greater on the REEF Advanced Exam, conducted about half as many surveys as novice divers in

all regions. The pattern of decreasing survey effort from the Upper Keys to the Dry Tortugas was

similar for both diver types (Figure 5).

Patterns of species richness

The Florida Keys and Dry Tortugas REEF data set contained sighting information on 341

fish species comprising 68 families (Appendix 2). The Molasses Reef site in the Upper Keys

(25.009° N, 87.3737° W) had 220 species, the highest number of species observed per location.

Molasses Reef also had the greatest number of surveys (277 surveys; approximately 261hr.).

To determine the relationship between species richness and survey effort, the cumulative

number of species was plotted against cumulative survey time for each location (Figure 6).

Cumulative species richness at survey sites varied strongly with total survey time (R2

 = 0.82, P <

0.0001; Figure 6). The cumulative number of species observed at survey sites increased loglinearly with increasing cumulative survey time, such that log-linear model accounted for 82% of

the observed variation in cumulative species richness. Approximately 75% of the species

richness was observed after 50 hr., and 90% of the species richness was observed after 130 hr. of

survey time (Figure 6).

Cumulative species richness also varied strongly with cumulative survey time within and

among regions (Figure 7). Within regions, observed species richness increased log-linearly with

cumulative survey time. Observed species richness was strongly correlated with cumulative

survey time at sites in the Upper, Middle, and Lower Keys, with the model accounting for 81-94

% of the observed variation in species richness in those regions (see R2

 values in Figure 7).

Species richness was not as strongly correlated with survey time in the Dry Tortugas (Figure 7).

The species encounter rate also varied among regions such that the encounter rate relative to the

total number of species present was faster in the Dry Tortugas than in other regions of the

Florida Keys (Figure 8). Approximately 95% of the species seen in the Dry Tortugas were

observed within 50 hr compared with 85% for the Middle and Lower Keys and 75% for the

Upper Keys within the same period.
6

Significant patterns were observed when mean species richness was compared among

regions. Chi-square comparisons of mean richness among regions showed significantly fewer

species in the Dry Tortugas compared with other regions, P < 0.0001 (Figure 9). The number of

species observed per survey were similar among the Upper, Middle, and Lower Keys but was

significantly lower in the Dry Tortugas, P < 0.0001 (Figure 10). Patterns of species richness were

similar between novice and expert divers using the RDT method. Family richness did not vary

significantly among regions or diver types (Figure 11).

Regional patterns in species distribution

Geographic coverages were developed for the twenty most frequently observed species in

the Florida Keys and the Dry Tortugas (Table 1). Blue tang (Acanthurus coeruleus) was the most

frequently seen fish and had a sighting frequency of 91%. Other frequently observed species

included the stoplight parrotfish (Sparisoma viride), yellowtail snapper (Ocyurus chrysurus),

sergeant major (Abudefduf saxatilis), bluehead (Thalassoma bifasciatum), and the french grunt

(Haemulon flavolineatum). Grunts (Haemulidae) and damselfishes (Pomacentridae) were more

highly represented among the top twenty species than other reef fish families (Table 2). No

groupers (Serranidae) were ranked among the top twenty most frequently observed species.

Several of the most frequently observed species in the Dry Tortugas did not rank among the

twenty most frequently observed species overall (Table 3); distribution maps were developed for

those nine species.

A series of non-parametric analyses and Tukey HSD tests showed significant regional

variation in the mean % SF of fourteen of the 29 species whose distributions were mapped

(Table 4). Six species had significantly lower mean % SF, and eight species had significantly

higher mean % SF in the Dry Tortugas compared with other regions (Tukey HSD, P < 0.05;

Table 4). Bluestriped grunt (Haemulon sciurus) was significantly less frequent at Dry Tortugas

sites (38.0 % ± 8.2) compared with the Lower (81.2 % ± 3.2), Middle (87.1% ± 4.3), and Upper

Keys (86.1 % ± 2.6) but differences among the Lower, Middle, and Lower Keys were not

significant (Tukey HSD, P < 0.05; Figure 12). Yellowtail damselfish (Microspathodon

chrysurus), sergeant major (A. saxatilis), and porkfish (Anisotremus virginicus) had distribution

patterns similar to that of the bluestriped grunts (Figures 13-15), all having significantly greater

%SF in the Upper, Middle, and Lower Keys than in the Dry Tortugas.

Mean sharpnose puffer (Canthigaster rostrata) % SF was less variable among regions

(Figure 16). Comparisons showed that the mean % SF of the Dry Tortugas region (48.5% ± 7.9)

was significantly lower than the mean of the Upper Keys (71.8% ± 3.4) but was not different

from the means of the Lower and Middle Keys (58.0% ± 6.0 and 56% ± 5.1). Results also

showed that the Lower and Middle Keys were not significantly different from the Upper Keys

(Tukey HSD, P > 0.05).

The mean % SF of foureye butterflyfish (Chaetodon capistratus) was lowest in the Dry

Tortugas region (55.1 % ± 7.7) and highest in the Lower and Middle Keys (81.4 % ± 5.5 and

85.6% ± 5.4 SF); this difference was significant (Tukey HSD, P < 0.05; Figure 17). Mean % SF

in the Upper Keys was not significantly different from that of other regions (73.5% ± 4.4 ).
7

Eight species were consistently more frequently observed in the Dry Tortugas than other

regions of the Keys (Table 4). Barred hamlets (Hypoplectrus puella) occurred almost twice as

often at Dry Tortugas sites than at sites in other regions, and were not observed at 47% and 20%

of sampled sites in the Middle and Upper Keys, respectively (Figure 18). The distribution of blue

hamlets (Hypoplectrus gemma) was similar. In the Dry Tortugas, the mean % SF of blue hamlets

was 59.7% (±5.9), whereas it ranged from 25.4% ± 6.3 to 30.0% ± 3.9 in the rest of the Keys

(Figure 19). Blue hamlets were not observed in 16-20% of the Florida Keys sites compared with

8% of sampled sites in the Dry Tortugas (Figure 19). These differences were significant (q* =

2.609, P < 0.05, Tukey HSD).

Butter hamlets (Hypoplectrus unicolor) were observed in 78% of sampled sites in the

Middle, Lower Keys and the Dry Tortugas region compared with 67% in Upper Keys sites.

Butter hamlets had a mean % SF of 70.5% ± 5.5 in the Dry Tortugas region, which was

significantly different from that of the Upper Keys (55.6% ± 3.8), the Middle Keys (40.5% ±

6.2), and Lower Keys (38.6% ± 6.0), (q* = 2.609, P < 0.05, Tukey HSD; Figure 20). The mean

% SF of butter hamlets in the Upper Keys was significantly different from that of the Lower and

Middle Keys but differences between Middle and Lower Keys were not significant (P > 0.05).

The cocoa damselfish (Stegastes variabilis) had significantly higher mean % SF in the

Dry Tortugas region (69.8 ± 5.0%) than the rest of the Florida Keys (50.1% ± 5.5 to 53.1% ±

5.6; P = 0.0046; Figure 21). Differences among mean % SF means for the Upper, Middle, and

Lower Keys were small and not significant (Figure 21). Threespot damelfish (Stegastes

planifrons) distribution (Figure 22) was different from that seen in the cocoa damselfish in that

the Dry Tortugas and the Upper Keys had significantly higher mean % SF (70.9% ± 5.8 and

62.7% ± 4.1) than the Middle Keys (43.2% ± 6.5). Other pairwise comparisons of threespot

damselfish mean % SF among regions were not significant (P > 0.05).

Other species with significant variation among regions included the blue angelfish

(Holacanthus bermudensis), striped parrotfish (Scarus croicensis), and the neon goby

(Gobiosoma oceanops). Blue angelfish were sighted significantly more frequently in the Dry

Tortugas (66.7% ± 5.2) than in the Lower (30.6% ± 5.7), Middle (36.6% ± 5.8), and Upper Keys

(22.9% ± 3.6) (Figure 23). Differences in blue angelfish mean % SF among the Lower, Middle,

and Upper Keys were not significant (P > 0.05). Striped parrotfish %SF was highest in the Dry

Tortugas (70.9% ± 4.8) and decreased eastward through the Florida Keys (Figure 24). Significant

differences in striped parrotfish mean %SF occurred between the Dry Tortugas region and the

Upper Keys only (P < 0.05). Neon goby %SF was high in the Dry Tortugas and Lower Keys

(66.7% ± 5.5 and 64.1% ± 6.0) but significantly lower in the Middle and Upper Keys (39.6% ±

6.1 and 43.7% ± 3.8; P < 0.05; Figure 25).

Patterns in species composition

Hierarchical clustering based on species composition (presence-absence) and species

%SF revealed interesting patterns of similarities among sites that varied across spatial scales

(Figures 26-28). Clustering based on species presence-absence data revealed 6 main site clusters

within the Florida Keys and Dry Tortugas (Figure 26). Similar patterns were observed in
8

clustering based on species sighting frequency data. For most clusters (e.g., Cluster 1), sites

within regions tended to cluster together more closely than with sites from other regions. For

example, sites in the Dry Tortugas clustered together and separately from other sites, as did

several Upper Key sites (Figures 26 and 27). However, several sub-clusters did not necessarily

follow geographical location, suggesting that much of the variation in species composition and

sighting frequency cannot be explained solely by regional differences or geographic variation.

These sub-clusters may have resulted from cross-shelf location of sites. For example, inshore

patch reef sites (e.g., Hens and Chickens and Cheeca Rocks) tended to cluster together, even

though they were located in different regions. Similarly, offshore bank reef sites tended to cluster

together (Figure 26). Correspondence analysis revealed highly significant associations between

regions and cluster groups (Figure 28, P < 0.0001).

DISCUSSION

Critical to the design of effective monitoring programs is a preliminary understanding of

the populations of interest. Estimates of parameters such as species encounter rates and levels of

sample variance each provide researchers with information that can be used to determine the

effort required to test specific hypotheses or to detect changes of varying magnitude. This study

is intended, in part, to provide such background, and utilizes a database generated by a large,

volunteer-based survey effort to accomplish this. In addition, the study extends the effort to

ascertain regional patterns of diversity and abundance.

The species/effort curves (Figure 7) suggested that encounter rate was inversely

proportional to richness for each of the four regions, with the Tortugas region having the highest

rate. Higher encounter rates would generally be expected in populations with higher evenness,

that is, in populations with individuals apportioned more equally among species. This may be the

case for the Tortugas. However, it is not clear whether the lower richness estimates for the

Tortugas are real or due to reduced survey effort. If they are real, the likely explanations relate to

the limited geographic extent of the area relative to the rest of the Florida Keys, as well as the

more limited variety and extent of habitats in the Dry Tortugas region. The other regions of the

Florida Keys contain considerably more areas of seagrass and mangrove, both important nursery

and early life habitats for reef fishes.

The sighting frequency data confirmed the uniqueness of the Tortugas reefs relative to

other regions of the Florida Keys. Among the 14 species (of the 29 most frequently observed in

the data set) with significant differences between regions, eight were more frequently observed

in the Tortugas (barred, blue and butter hamlets, cocoa and threespot damselfish, blue angelfish,

neon goby, and striped parrotfish) than in most or all the other regions. The six others were less

frequently encountered in the Tortugas (bluestriped grunt, yellowtail damselfish, sergeant major,

porkfish, foureye butterflyfish, and sharpnose puffer).

Beyond the regional differences revealed in the study, site differences were identified in

the cluster analysis. Most sites clustered according to region, supporting other analyses, but
9

others did not. Probable reasons for this vary, and are likely to be related to reef size and

structure, proximity between sites, distance from shore, stochastic events, and other factors.

Thus, it is clear that phenomena affecting reef fish composition in the FKNMS operate at

multiple spatial scales. Beyond the biogeographic scale that defines the character of the region as

a whole (the reefs contain tropical species of the Caribbean Province), processes operating on a

scale of the order of ~50-100 km account for differences between the Tortugas and the rest of the

Florida Keys. These are likely to include meso-scale physical oceanographic processes such as

those recently identified by Lee et al. (1992, 1994), which may localize recruitment, to an extent,

within several areas of the Keys. Regional variation in reef structure may also contribute to

differences at this scale. Upper Keys reefs tend to be more complex than those in the Middle and

Lower Keys. The influence of Florida Bay may operate at similar scales, as passes are not

common in the Upper Keys, but are numerous in the Middle and Lower Keys. Finally,

submerged aquatic vegetation, habitat utilized by species during portions of their life, is

abundant in the Upper Keys, less so in the Middle and Lower Keys, and is rare by comparison in

the Tortugas.

It is likely that at both regional and local scales, habitat requirements strongly influence

the patterns revealed in this study, and are particularly limiting for species that are less frequently

observed in the Tortugas. This is because of constraints imposed by the reduced variety and

extent of habitats in that region. The importance of specific characteristics of habitats in

controlling fish populations indicates the need to investigate sub-regional scale phenomena, such

as the proximity of reefs to seagrass beds and to shore-associated habitats (e.g. mangroves).

Also, the extent of reef habitats themselves at each sample site should be evaluated against reef

fish community composition, as this factor is also likely to limit community development.

Currently, we are using the REEF data set to analyze these relationships. Ongoing analyses

include testing hypotheses of non-uniform fish distribution among benthic habitats and

management zones, and analyzing spatial trends and correlations between fish community

structure and benthic habitat parameters (e.g., diversity and reef proximity to submerged aquatic

vegetation) within the Florida Keys National Marine Sanctuary.

The results of this report serve as a benchmark for the current status of the reef fishes in

the FKNMS. These data provide the basis for analyses on reserve effects and the biogeographic

coupling of benthic habitats and fish assemblages, and ultimately, may be useful in developing

future management zones.
10

LITERATURE CITED

Allison, G. W., J. Lubchenco, and M. H. Carr. 1998. Marine reserves are necessary but not

sufficient for marine conservation. Ecol. App. 8 (1): S79-S92.

Ault J.S., J.A. Bohnsack, and G.A. Meester. 1998. A retrospective (1979-1996) multispecies

assessment of coral reef fish stocks in the Florida Keys. Fish. Bull. 96: 395-414.

Bohnsack J.A. and J.S. Ault. 1996. Management strategies to conserve marine biodiversity.

Oceanography 9: 73-82.

FMRI (Florida Marine Research Institute). 1998. Benthic Habitats of the Florida Keys. FMRI

Technical Report TR-4. 53 pp.

Lee T. N., C. Rooth, E. Williams, M. McGowan, A. F. Szmant, and M. E. Clarke. 1992.

Influence of Florida Current, gyres and wind-driven circulation on transport of larvae and

recruitment in the Florida Keys coral reefs. Cont. Shelf Res. 12: 971-1002.

Lee T. N., M. E. Clarke, E. Williams, A.F. Szmant, and T. Berger. 1994. Evolution of the

Tortugas Gyre and its influence on recruitment in the Florida Keys. in J. M. Prospero and

C.C. Harwell, eds. 1992. Symposium on Florida Keys Regional Ecosystem, Miami, FL

(USA), Nov. 1992.

NOAA (National Oceanic and Atmospheric Administration). 1996. Florida Keys National

Marine Sanctuary Final Management Plan/Environmental Impact Statement, Volume 1:

Management Plan. 319 pp.

Ogden, J. C. and P. S. Lobel. 1978. The role of herbivorous fishes and urchins in coral reef

communities. Env. Biol. Fish. 3: 49-63.

REEF (Reef Environmental Education Foundation). 2000. World Wide Web electronic

publication. www.reef.org, 1 May 2000.

Richards, W. J. and K. C. Lindeman 1987. Recruitment dynamics of reef fishes: planktonic

processes, settlement and demersal ecologies and fishery analysis. Bull. Mar. Sci. 41 (2):

392-410.

Schmitt, E. F. and K. M. Sullivan. 1996. Analysis of a volunteer method for collecting fish

presence and abundance data in the Florida Keys. Bull. Mar. Sci. 59: 404-416.

Williams, D. McB. 1991. Patterns and processes in the distribution of coral reef fishes. pp 437-

473 in P. F. Sale, ed. The Ecology of Fishes on Coral Reefs. Academic Press, Ca. 754 pp.
11

ACKNOWLEDGEMENTS

This project was made possible through a NOAA Cooperative Agreement with the

University of Georgia (NOAA Award No.: NA07OA0066). Tracy Gill (NOAA) and Mac

Rawson (Georgia College Sea Grant Program, University of Georgia) provided invaluable

administrative support for this project. Brian Keller (FKNMS), Paula Souik (NOAA), and David

Moe Nelson (NOAA) provided constructive comments on the manuscript. The support of

NOAA’s Biogeography Team (Tim Battista, Ken Buja, Chris Caldow, John Christensen,

Michael Coyne, Tracy Gill, Matt Kendall, and David Moe Nelson) is appreciated. We give

sincere thanks to REEF (Laddie Akins, Leslie Whaylen, and the volunteers) whose data formed

the basis of this project.
12

TABLES

Table 1. Most frequently observed species in the Florida Keys based on sighting frequency

(%SF). Data are from REEF (2000).

Table 2. The number of species per family ranked among the twenty most

frequently observed species in the Florida Keys and Dry Tortugas.

Rank Common Name Scientific Name Family SF (%)

1 Blue tang Acanthurus coeruleus Acanthuridae 91.1

2 Stoplight parrotfish Sparisoma viride Scaridae 87.7

3 Yellowtail snapper Ocyurus chrysurus Lutjanidae 84.1

4 Sergeant major Abudefduf saxatilis Pomacentridae 83.4

5 Bluehead Thalassoma bifasciatum Labridae 83.4

6 French grunt Haemulon flavolineatum Haemulidae 82.4

7 Bicolor damselfish Stegastes partitus Pomacentridae 81.6

8 Ocean surgeonfish Acanthurus bahianus Acanthuridae 77.5

9 Bluestriped grunt Haemulon sciurus Haemulidae 77.3

10 Yellowtail damselfish Microspathodon chrysurus Pomacentridae 74.8

11 Porkfish Anisotremus virginicus Haemulidae 74.2

12 Foureye butterflyfish Chaetodon capistratus Chaetodontidae 73.9

13 White grunt Haemulon plumieri Haemulidae 73.9

14 Redband parrotfish Sparisoma aurofrenatum Scaridae 71.1

15 Spotfin butterflyfish Chaetodon ocellatus Chaetodontidae 70.5

16 Yellowhead wrasse Halichoeres garnoti Labridae 70.1

17 Great barracuda Sphyraena barracuda Sphyraenidae 68.5

18 Gray angelfish Pomacanthus arcuatus Pomacanthidae 68.0

19 Bar jack (Skipjack) Caranx ruber Carangidae 66.4

20 Sharpnose puffer Canthigaster rostrata Tetraodontidae 62.4

Family # of species 

Haemulidae Grunts 4

Pomacentridae Damselfishes 3

Scaridae Parrotfishes 2

Labridae Wrasses 2

Chaetodontidae Butterflyfish 2

Acanthuridae Surgeonfishes 2

Tetradontidae Puffers 1

Sphyraenidae Barracuda 1

Pomacanthidae Angelfishes 1

Lutjanidae Snappers 1

Carangidae Jacks 1
13

Table 3. Twenty most common species found in the Dry Tortugas sites ranked by sighting

frequency (%SF). Asterisks denote species that were not ranked among the twenty most

frequently observed species in the Florida Keys.

Rank Common name Species Family SF (%)

1 Blue tang Acanthurus coeruleus Acanthuridae 93.4

2 Bluehead Thalassoma bifasciatum Labridae 89.5

3 Stoplight parrotfish Sparisoma viride Scaridae 83.9

4 Gray angelfish Pomacanthus arcuatus Pomacanthidae 83.7

5 Yellowtail snapper Ocyurus chrysurus Lutjanidae 83.2

6 White grunt Haemulon plumieri Haemulidae 78.8

7 Cocoa damselfish* Stegastes variabilis Pomacentridae 78.1

8 Spotfin butterflyfish Chaetodon ocellatus Chaetodontidae 77.1

9 Threespot damselfish* Stegastes planifrons Pomacentridae 75.9

10 Neon goby* Gobiosoma oceanops Gobiidae 74.9

11 Redband parrotfish Sparisoma aurofrenatum Scaridae 69.6

12 Butter hamlet* Hypoplectrus unicolor Serranidae 68.6

13 Blue angelfish* Holacanthus bermudensis Pomacanthidae 68.4

14 French grunt Haemulon flavolineatum Haemulidae 64.0

15 Striped parrotfish* Scarus croicensis Scaridae 63.8

16 Bicolor damselfish Stegastes partitus Pomacentridae 63.5

17 Blue hamlet* Hypoplectrus gemma Serranidae 63.0

18 Yellowhead wrasse Halichoeres garnoti Labridae 61.6

19 Slippery dick* Halichoeres bivittatus Labridae 59.9

20 Barred hamlet* Hypoplectrus puella Serranidae 57.4
HF Highly frequent F Frequent C Common UC Uncommon

Table 4. Results of Wilcoxon / Kruskal-Wallis non-parametric (Rank Sums) and Tukey HSD tests of no sdifference in mean percent sighting frequency among regions, for twenty-nine frequently sighted species Keys and the Dry Tortugas (alpha = 0.05, df = 3). Asterisks indicate species ranked in the Dry Tortugas bthe twenty most observed species in the Florida Keys. Frequency classes are based on equal quantiles detthe distribution of the mean sighting frequencies (n = 112 site means). DT = Dry Tortugas, UK = Upper KMiddle Keys, LK = Lower Keys. A positive (+) sign indicates significant results.

Family Chi-Sq P(Chi-Sq) Sig.

DT 

Bluestriped grunt Haemulon sciurus Haemulidae 21.00 0.00 + C 

Yellowtail damselfish Microspathodon chrysurus Pomacentridae 20.52 0.00 + C 

Sergeant major Abudefduf saxatilis Pomacentridae 18.42 0.00 + C 

Porkfish Anisotremus virginicus Haemulidae 14.85 0.00 + C 

Sharpnose puffer Canthigaster rostrata Tetradontidae 11.29 0.01 + C 

Foureye butterflyfish Chaetodon capistratus Chaetodontidae 7.86 0.05 + F 

Barred hamlet* Hypoplectrus puella Serranidae 23.57 0.00 + C 

Blue angelfish* Holacanthus bermudensis Pomacanthidae 29.65 0.00 + F 

Neon goby* Gobiosoma oceanops Gobiidae 19.41 0.00 + F 

Butter hamlet* Hypoplectrus unicolor Serranidae 18.72 0.00 + F 

Blue hamlet* Hypoplectrus gemma Serranidae 15.66 0.00 + F 

Cocoa damselfish* Stegastes variabilis Pomacentridae 13.03 0.00 + F 

Threespot damselfish* Stegastes planifrons Pomacentridae 11.49 0.01 + F 

Striped parrotfish* Scarus croicensis Scaridae 9.48 0.02 + F 

Redband parrotfish Sparisoma aurofrenatum Scaridae 7.68 0.05 F 

Bar jack Caranx ruber Carangidae 7.35 0.06 F 

Ocean surgeonfish Acanthurus bahianus Acanthuridae 6.77 0.08 F 

French grunt Haemulon flavolineatum Haemulidae 6.50 0.09 F 

Bluehead Thalassoma bifasciatum Labridae 6.34 0.10 HF 

Gray angelfish Pomacanthus arcuatus Pomacanthidae 6.12 0.11 HF 

Yellowhead wrasse Halichoeres garnoti Labridae 5.05 0.17 F 

Spotfin butterflyfish Chaetodon ocellatus Chaetodontidae 4.87 0.18 HF 

Yellowtail snapper Ocyurus chrysurus Lutjanidae 4.63 0.20 HF 

Bicolor damselfish Stegastes partitus Pomacentridae 4.40 0.22 F 

Great barracuda Sphyraena barracuda Sphyraenidae 3.05 0.38 F 

Blue tang Acanthurus coeruleus Acanthuridae 1.68 0.64 HF 

White grunt Haemulon plumieri Haemulidae 1.28 0.73 HF 

Stoplight parrotfish Sparisoma viride Scaridae 1.17 0.76 HF 

Slippery dick* Halichoeres bivittatus Labridae 1.31 0.73 F 

Species Ave

14
Middle

Keys

0 100 KM

 Key 



Upper Matecumbe Ke



Marquesas



Pigeon Key



Little Duck Key

Florida Keys National Marine Sa

DryTortugas

Lower Keys Upper

Keys

Figure 1. Map of the Florida Keys showing geographic boundaries among four regions.

15
Uncommon

Highly Frequent

Common

Frequent

0.4

10

20

30

40

50

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80

90

100

Sighting Frequency (%)

Figure 2. Classification scheme used in mapping fish abundance and

summary statistics for fish data obtained from the Reef Environmental

Education Foundation (REEF, 2000).

Frequency

16
Figure 3. The frequency distribution and summary statistics for time spent by

REEF divers while surveying fishes in the Florida Keys. Only data from

surveys with time ranging between 30 –100 minutes (in shaded box) were

included for analysis.

0

100

200

Survey time (minutes)

Maximum

Median

Minimum

100.0

99.5

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Quantiles % Minutes

245.00

100.84

75.00

67.00

60.00

50.00

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0.00

Mean

Std Dev

Std Error Mean

Upper 95% Mean

Lower 95% Mean

Number of surveys

59.384

14.578

0.219

59.813

58.955

4431

10.0 41.00

Frequency

17
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Dry Tortugas, 377 (348)Lower Keys, 739 (753)Middle Keys, 560 (595)Upper Keys, 2603 (2569)

Surveys

5 - 1314 - 3132 - 6364 - 277

Sanctuary boundary

Total surveys, # (hr.)

5

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5

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Figure 4. Distribution map of visual surveys done by Reef Environmental Education Foundation (Rvolunteers using the Roving Diver Technique (RDT).

18
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UK MK LK DT UK MK LK DT

#

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Expert

Surveys (no.) Total Hours

Figure 5. Level of survey effort compared among expert and novice divers

stratified by region (UK = Upper Keys, MK = Middle Keys, LK = Lower

Keys, DT = Dry Tortugas).
Figure 6. Cumulative species-time curve for reef fishes observed in the Florida Keys 

with summary of fit and analysis of variance results for the fitted curve.

Source DF Sum of Squares Mean square F Ratio Prob> F

Model 1 156350.56 156351 487.1304 <.0001

Error 110 35305.87 321

Total 111 191656.43

Summary of Fit

R2 

0.82

Root Mean Square Error 

17.91542

Mean

125.8214

N

112

SE

3.92637

0

50

100

150

200

250

0 50 100 150 200 250

Cumulative species = 36.8739 + 29.8516 Ln (Cumulative hr.)

Cumulative hr.

Cummulative Species (#)

0.2

0.4

0.6

0.8

1.0

Proportion of richness

0

20
Figure 7. Regional cumulative species richness and analysis of variance 

results for fitted curves for sites among four regions of the Florida Keys.

Region Source DF Type 1 SS F R2 Prob>F

Dry Tortugas Model 1 11916.702 26.832 0.549 0.0001

Error 22 9770.631

Total 23 21687.333

Lower Keys Model 1 27195.207 297.804 0.943 0.0001

Error 18 1643.743

Total 19 28838.950

Middle Keys Model 1 20412.881 191.293 0.918 0.0001

Error 17 1814.067

 Total 18 2226.947

Upper Keys Model 1 66422.296 205.955 0.814 0.0001

Error 47 15157.949

Total 48 81580.245

0

50

100

150

200

0 50 100 150 200 250

Cummulative hours

Dry Tortugas

Lower Keys

Middle Keys

Upper Keys

Cumulative species

21
Figure 8. Proportion of species richness for regions of the Florida Keys.

Data are predicted estimates from fitted curves in Figure 6.

0.2

0.4

0.6

0.8

1.0

0 50 100 150 200 250

Cumulative hours

Proportion of species

Dry Tortugas

Lower Keys

Middle Keys

Upper Keys

22
Figure 9. Distribution map of species richness for four regions of the Florida Keys. The bar grap

regional differences in the mean number of species and tests for significant differences among re(Kruskal-Wallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means wit

symbols (+ or -) were not significantly different.

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Species richness 5 16 - 97

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5 208 - 234

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Mean richness

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(# Species)

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23
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n = 2,595 n = 580 n = 738 n = 411

χ2 = 128.32; df = 3; P < 0.0001

UK = Upper Keys

MK = Middle Keys

LK = Lower Keys

DT = Dry Tortugas

Figure 10. Mean species richness per diver survey for four regions of the

Florida Keys. Differences between means are significant. Means were

tested with a Wilcoxon / Kruskal-Wallis one-way Chi-Squared (Rank

Sums) test. The table shows the results of pairwise comparisons among

regions using Tukey-Kramer HSD test (α = 0.05; q* = 2.57003). A ‘+’

indicates significant differences among paired means; a ‘-’ shows no

significant differences among paired means.

24
#

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UK MK LK DT

Species

Novice

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Total

0

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#

Family

Figure 11. Species and family richness among expert and novice divers in the Florida

Keys.

25
Figure 12. Spatial distribution and relative %SF of bluestriped grunt among regions of the Florida K

scatter plot shows regional differences in mean %SF and tests for significant differences among mea(Kruskal-Wallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with th

symbols (+ or -) were not significantly different.

-

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26
Figure 13. Spatial distribution and relative %SF of yellowtail damselfish among regions of the Florid

The scatterplot shows regional differences in mean %SF and tests for significant differences among m(Kruskal-Wallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the 

symbols (+ or -) were not significantly different.

χ220.5203

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27
Figure 14. Spatial distribution and relative %SF of sergeant major among regions of the Florida Ke

scatterplot shows regional differences in mean %SF and tests for significant differences among mea(Kruskal-Wallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with th

symbols (+ or -) were not significantly different.

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% SF

28
Figure 15. Spatial distribution and relative %SF of porkfish among regions of the Florida Keys. The 

shows regional differences in mean %SF and tests for significant differences among means (Kruskal-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the same symbols (+ or 

significantly different.

5

5 5 5

5

5

7

7 55

5

7

5

7

5

5

5

75 5

5

5

5

5

5

5

5

5

5

5

55

5

5

5

5

55

5

55

5

5

5

5

5

5

5

5

5555

5

5

5

5

5

5

5

5

5

5

5

5

5

5

55

7

5

5

5

5

5

5

5

5

5555

5

5

5

55

5

55

555

5

7

55

5

5

5

N

Dry TortugasLower KeysMiddle KeysUpper KeysFKNMS boun

5 Highly Frequ

5 Frequent

5 Common

5 Uncommon o

7 Not observed

0 100 KM

20406080

100

DT LK MK UK

-

+ + +

χ2

14.8548

DF

3

P0.0019

%SF

Porkfish %SF(Anisotremus virg

29
Figure 16. Spatial distribution and relative %SF of sharpnose puffer among regions of the Florida Ke

scatterplot shows regional differences in mean %SF and tests for significant differences among mean(Kruskal-Wallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the 

symbols (+ or -) were not significantly different.

5

5

5

7

5

5

5

5 7

5

7

5

5

5

7

5

5

75

5

7

5

7

5

5

55

5

5

55

5

5

5

5

55

7

55

5

5

5

5

5

5

5

5

555

5

5

5

5

5

5

5

5

5

5

5

5

555

5

5

5

5

5

5

5

55

5

5

5

5

5

5

5

7

5

5

N

Dry TortugLower KeyMiddle KeUpper KeyFKNMS bo

5 Highly Fre

5 Frequent

5 Common

5 Uncommo

7 Not observ

0 100 KM

Sharpnose puf(Canthigaster r

20406080

100

DT LK MK UK

-

+

- +

-

+

χ2

11.2894

DF

3

P0.0103

%SF

30
Figure 17. Spatial distribution and relative %SF of foureye butterflyfish among regions of the Florida Ke

scatterplot shows regional differences in mean %SF and tests for significant differences among means (KWallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the same symbo

were not significantly different.

7

5

5

5

5

5

5

5 57

5

5

5

5

7

5

5

55 5

7

5

7

5

5

55

5

5

55

5

5

5

5

55

5

55

5

5

5

5

5

5

5

5

5555

5

5

5

5

5

5

7

5

5

55

5

5

5

5

5

5

5

5

5

55

5

5

N

Dry TortLower KMiddle KUpper KFKNMS 

5 Highly F

5 Frequen

5 Commo

5 Uncomm

7 Not obs

0 100 KM

Foureye butterf(Chaetodon ca

20406080

100

DT LK MK UK

-

+ +

+

-

χ2

7.8559

DF

3

P0.0491

%SF

31
Figure 18. Spatial distribution and relative %SF of barred hamlet among regions of the Florida Key

scatterplot shows regional differences in mean %SF and tests for significant differences among mea(Kruskal-Wallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with th

symbols (+ or -) were not significantly different.

5

5

5 5

7

5

5

5

55

5

5

5

5

7

5

5

5

75 5

7

5

5

5

5 5

5

5

5

5

75

5 5

5

5 55

5

55 5

5

5

7

5

5

7

7

55757

7

7 5

7

7 5

5

5

5

7

5

5

5

5

55

5

7

7

7

5

5

55

5

5555

5

5

5

55

7

57

7

55555

7

7 5

5

5

5

7

N

Dry TortugasLower KeysMiddle KeysUpper KeysFKNMS boun

5 Highly Frequ

5 Frequent

5 Common

5 Uncommon o

7 Not observed

0 100 KM

Barred hamlet %S(Hypoplectrus pue

20406080

100

DT LK MK UK

+

-

-

-

χ223.5729

DF

3

P0.0001

%SF

32
Figure 19. Spatial distribution and relative %SF of blue hamlet among regions of the Florida Keys. The

scatterplot shows regional differences in mean %SF and tests for significant differences among means (KWallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the same symbo

were not significantly different.

5

5

5

5

5

5

5

555

5

5

5

5

7

5

5

5

75

5

5

5

5

7

5 5

5

5

5

5

77

5 5

5

5

55

7

55 5 5

5

7

5

5

5

5

555

5

5

7

5 5 5

5

7

5

5

5

5

5

75

5

55

5

7

5

7

5

5

5

5 5

5

555 5

5

5

5

55

5

75

5

755555

7

7

55

5

5

7

N

Dry TortugasLower KeysMiddle KeysUpper KeysFKNMS boundary

5 Highly Frequent

5 Frequent

5 Common

5 Uncommon or Ra

7 Not observed

0 100 KM

Blue hamlet %SF(Hypoplectrus gemma

20406080

100

DT LK MK UK

+

- - -

χ2

15.664

DF

3

P0.0013

%SF

33
Figure 20. Spatial distribution and relative %SF of butter hamlet among regions of the Florida Keys. 

scatterplot shows regional differences in mean %SF and tests for significant differences among mean(Kruskal-Wallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the 

symbols (+, -, or *) were not significantly different.

5

5

5

5

5

5

5

5

5

5

5

7

5

7

5

5

5

5 5

5

5

5

5

7

5

5

5

5

5

5

55

5 5

5

5

55

7

55

5 5

5

7

5

5

5

7

55

555

5

5

5

5

5

5

5

5

5

5

5

5

5

55

5

55

5

7

5

5

5

5

5

5

5

55555

5

5

5

55

5

55

5

5555

5

7

55

5

5

7

N

Dry TortugasLower KeysMiddle KeysUpper KeysFKNMS boundar

5 Highly Frequent

5 Frequent

5 Common

5 Uncommon or R

7 Not observed

0 100 KM

Butter hamlet %SF(Hypoplectrus unicolor

20406080

100

DT LK MK UK

+

- -

*

χ2

18.7218

DF

3

P0.0003

%SF

34
Figure 21. Spatial distribution and relative %SF of cocoa damselfish among regions of the Florida Keys. 

scatterplot shows regional differences in mean %SF and tests for significant differences among means (KWallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the same symbol

were not significantly different.

5

5

5

5

5

5

5

55

5

5

5

5

5

7

5

5

5

5 5

5

5

5

5

7

5 5

5

5

5

5

55 5 5

5

5

55

7

55

5

5

5

5

5

5

5

5

555

5

5 55 5

5

5

5

5

5

5

5

55

5

5

5

55

5

7

5

5

5

5

5

5

5

55

55

5

5

5

5

55

5

55

5

5557

55

5

5 55

5

5

5

N

Dry TortugasLower KeysMiddle KeysUpper KeysFKNMS boundary

5 Highly Frequent

5 Frequent

5 Common

5 Uncommon or Rare

7 Not observed

0 100 KM

Cocoa damselfish %SF(Stegastes variabilis)

20406080

100

DT LK MK UK

+

- - -

χ2

13.0258

DF

3

P0.0046

%SF

35
Figure 22. Spatial distribution and relative %SF threespot damselfish among regions of the Florida 

scatterplot shows regional differences in mean %SF and tests for significant differences among meWallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the same s

were not significantly different.

5

5

5

5

5

5

5

555

5

5

7

5

7

5

5

75

5

5

5

5

5

5

5

5

5

5

5

5 5 5

5

5

55

7

55

5

5

5

7

5

5

5

5

5557

7

5

5

5 5 5

5

7

5

5

5

5

55

5

5

5

5

5

5

5

55

5

5

5

5

55

5

7

5

5

N

Dry TortugLower KeyMiddle KeUpper KeyFKNMS bo

5 Highly Fre

5 Frequent

5 Common

5 Uncommo

7 Not observ

0 100 KM

Threespot dams(Stegastes plani

20406080

100

DT LK MK UK

-

- +

-

χ2

11.4876

DF

3

P0.0094

%SF

36
Figure 23. Spatial distribution and relative %SF of blue angelfish among regions of the Florida Keys. 

scatterplot shows regional differences in mean %SF and tests for significant differences among meanWallis Chi-Squared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the same sym

were not significantly different.

5

5

5

5

5

5

555

5

5

5

5

5

7

5

5

5

55

5

5

5

5

7

5 5

5

5

5

5

55

5 5

5

5 55

5

55 5

5

5

7

5

5

5

5

557

55 5

5

75 5 5

5

5

5

5

5

5

55

75

5

55

5

5

5

5

5

5

55

5

555 5

5

5

7 55

5

55

5

7

55555

7

7

75

5

5

5

N

Dry TortugasLower KeysMiddle KeysUpper KeysFKNMS bound

5 Highly Frequen

5 Frequent

5 Common

5 Uncommon or 

7 Not observed

0 100 KM

Blue angelfish %SF(Holacanthus bermud

20406080

100

DT LK MK UK

-

+ +

+

χ2

29.6475

DF

3

P0.0001

%SF

37
5

5

5 5

5

5 55 55 5 5

5

5

5

5

5

5

5 5 5

5

5

5

7

5 5

55

5

5

55

5 5

5 5 55

5

55 5

5

5

5

5

5

5

5

5555 5

5

5

5 5 5

5

5

5

5

5

5

5

5

5

55

5

7

5

5

5

5

55 5

5555

5

5

5

55

5

55

5

55555

5

5 555

55

N

Dry TortugasLower KeysMiddle KeysUpper KeysFKNMS bound

5 Highly Freque

5 Frequent

5 Common

5 Uncommon or 

7 Not observed

0 100 KM

Figure 24. Spatial distribution and relative %SF of striped parrotfish among regions of the Florida Kscatterplot shows regional differences in mean %SF and tests for significant differences among meanWallis Chi-Squared [χ2] test, α = 0.05; Tukey-Kramer HSD test, α = 0.05). Means with the same sym

were not significantly different.

Striped parrotfish %S(Sparisoma viride)

20406080

100

DT LK MK UK

-

-

+

-

+

+

χ2

9.4833

DF

3

P0.0235

%SF

38
Figure 25. Spatial distribution and relative %SF of neon goby among regions of the Florida Keys. Th

shows regional differences in mean %SF and tests for significant differences among means (KruskalSquared [

χ2] test, 

α = 0.05; Tukey-Kramer HSD test, 

α = 0.05). Means with the same symbols (+ or 

significantly different.

5

5

5

5

5

5

5

55

5

5

5

7

7

7

5

5

5

5 5

5

5

5

5

5

5

5

5

5

5

5

55

5 5

5

5

55

5

55

5

5

5

7

5

5

5

5 5575 5

5

55 5

5

5

7

5

5

5

5

5

75

5

55

5

7

5

5

5

5

5

5

5

5

55 5

5

5

5

55

5

55

5

555

5

7

55

5

5

5

N

Dry TortugasLower KeysMiddle KeysUpper KeysFKNMS bound

5 Highly Freque

5 Frequent

5 Common

5 Uncommon o

7 Not observed

0 100 KM

Neon goby %SF(Gobiosoma oceano

20406080

100

DT LK MK UK

-

+

-

+

χ2

19.4089

DF

3

P0.0002

%SF

39
Figure 26. Correspondence analysis plot of associations among site clusters

and regions of the FKNMS. Site clusters were identified from hierarchical

clustering (Ward’s minimum variance technique) of species composition (i.e.

the presence or absence of a species at a given site). The table shows Chisquare (χ2

) and - log-likelihood tests of significant correlation among cluster

groups and regions.

Region Cluster

-2.0

-1.5

-1.0

-0.5

0.0

0.5

1.0

1.5

2.0

DT

LK

MK UK 1

2

3

4

5

6

-2.0 -1.5 -1.0 -0.5 .0 .5 1.0 1.5 2.0

c2

c1

Tests

Source

Model

Error

C Total

Total

DF

15

 77

 92

 97

-LogLikelihood

 42.372

 143.880

 186.252

R2(U)

0.2280

Test

Likelihood Ratio

Pearson

(χ2)

 84.745

 86.988

P(χ2)

 <.0001

 <.0001

40
Figure 27. Correspondence analysis plot of associations among site clusters

and regions of the FKNMS. Site clusters were identified from hierarchical

clustering (Ward’s minimum variance technique) of species frequency

(%SF). The table shows Chi-square (χ2) and - log-likelihood tests of

significant correlation among cluster groups and regions.

41

Tests

Source

Model

Error

C Total

Total

DF

15

 77

 92

 97

-LogLikelihood

 25.411

 103.203

 128.615

R2(U)

0.20

Test

Likelihood Ratio

Pearson

(χ2)

 50.824

 46.488

P(χ2

)

 <.0001

 <.0001

-1.0

-0.5

0.0

0.5

1.0 DT

LKMK

UK

1

2

3

4

5 6

-1.0 -0.5 .0 .5 1.0

c2

c1

Region Cluster
42

DT

LK

MK

UK

B. % SF

DT

LK

MK

UK

A. Species composition

Figure 28. Dendograms from hierarchical clustering (Figures 26

and 27) showing similarities in (A) species composition and (B)

species sighting frequency (%SF) among regions of the Florida

Keys. Symbols ( or +) indicates regions which were most

similar and clustered together.
43

Appendix 1. List of sites in the Florida Keys and Dry Tortugas surveyed by the Reef

Environmental Education Foundation (REEF) volunteers between 1993 and 1999.

Location Lat. (o N) Long. (o W) Surveys (no.) Hours

(tot.)

Species

(tot.)

Upper Keys

Carysfort Reef 25.2200 80.2123 63 60.2 209

Carysfort Deep Ledge 25.2200 80.2112 26 21.5 175

Carysfort Reef Johnny’s Spot 25.2135 80.2168 49 52.5 200

South Carysfort Reef 25.2105 80.2172 65 66.6 202

Watson’s Reef 25.1860 80.2425 3 3.0 132

Toadfish Flats (Hawks Ch.) 25.1792 80.3403 15 13.5 37

N. Carysfort - Fishbowl 25.1707 80.3680 12 11.7 120

Triple North (off Elbow) 25.1525 80.2673 7 7.1 102

Elpis Grounding Site 25.1483 80.2525 8 8.4 116

South-South Ledges E1 25.1473 80.2610 89 96.2 225

Civil War Wreck (Elbow) 25.1473 80.2577 8 6.3 99

City of Washington-E7/8 25.1460 80.2558 84 81.9 219

Anchor Chain E6 25.1450 80.2563 141 145.7 225

Mike’s Wreck E4/5 25.1447 80.2568 87 83.3 216

The Fingers E3 25.1423 80.2577 46 44.7 204

Train Wheel E2 25.1420 80.2578 44 43.6 186

South Ledges E9 25.1403 80.2590 94 98.5 219

The Elbow 25.1388 80.2610 56 53.6 213

Horseshoe Reef 25.1387 80.3050 31 30.6 205

Spanish Anchor (Elbow) 25.1382 80.2600 4 4.0 127

NN Dry Rocks 25.1363 80.2903 84 80.5 215

Pecks Place / Cap Happy’s 25.1343 80.2638 37 40.1 210

Spikes Ridge (off Elbow) 25.1333 80.2585 19 19.9 126

Minnow Caves/North Dry Rocks 25.1307 80.2943 64 64.1 213

Key Largo Dry Rocks 25.1225 80.2975 177 175.7 230

Little Grecian 25.1190 80.3002 48 52.6 201

Cannon Patch/Garret’s Reef 25.1118 80.3417 22 21.3 104

Grecian Rocks 25.1098 80.3042 123 119.6 222

Dixie Ledge 25.0773 80.3110 16 11.1 131

Benwood Wreck 25.0527 80.3337 114 110.9 226

White Banks 25.0417 80.3700 29 27.2 91

French Reef 25.0353 80.3473 123 121.2 227

Molasses Reef 25.0090 80.3737 277 261.0 234

Wellwood Grounding Site 25.0083 80.3750 11 11.4 125

The Pillars 24.9922 80.4085 3 3.1 87

Duane 24.9880 80.3805 7 3.6 87

Pickles Reef 24.9862 80.4157 52 46.5 209

Horseshoe (Near Conch) 24.9567 80.4570 2 1.4 82

Conch Reef 24.9518 80.4595 106 102.5 219

Mutton Snapper Reef 24.9435 80.4953 26 27.3 195

Hens and Chickens 24.9317 80.5483 72 75.3 210

Davis Reef 24.9220 80.5060 123 132.9 222

Pleasure Reef 24.9135 80.5158 11 10.7 114

Crocker Ridges 24.9032 80.5302 15 14.7 176

Crockers Wall 24.9002 80.5313 47 34.6 193

Pocket, The 24.8982 80.5363 14 13.5 149

Fish Bowl 24.8933 80.5527 15 16.1 107

Aquarium Reef 24.8912 80.5555 20 22.3 171

Hammerhead Reef 24.8888 80.5468 6 5.5 152
44

Appendix 1 continued

Location Lat. (o N) Long. (o W) Surveys (no.) Hours

(tot.)

Species

(tot.)

Middle Keys

Cheeca Rocks 24.9045 80.6155 28 30.5 110

Wreck of the Eagle 24.8695 80.5702 12 7.0 97

Alligator Reef 24.8512 80.6202 108 129.9 220

Tennessee Reef 24.7617 80.7550 57 67.6 195

Porkfish 24.7002 80.8938 5 5.3 104

Rusty’s 24.6953 80.9058 4 4.3 89

Donut Reef 24.6918 80.9478 2 1.9 92

Coffins Patch 24.6767 80.9750 63 68.0 200

Horseshoe 24.6612 80.9942 14 14.5 163

Samantha’s Ledge 24.6592 81.0040 66 71.3 210

Joanie’s Reef 24.6563 81.0095 36 38.2 208

Joanie’s Rock 24.6560 81.0098 4 4.3 95

Pot Holes 24.6517 81.0247 4 4.2 89

Hermans Behind 24.6510 81.0290 7 4.4 90

Herman’s Hole 24.6505 81.0313 16 16.7 167

Boom Ledge 24.6353 81.0793 2 1.7 71

Lucille’s Reef 24.6348 81.0415 4 4.2 86

Delta Shoals 24.6327 81.0900 18 17.2 115

Sombrero Reef 24.6283 81.1050 130 124.5 223

Lower Keys

The Alexander 24.6232 81.9822 3 2.5 50

Newfound Open/Captain’s Coral 24.6215 81.3805 12 8.3 79

Newfound Harbor Spa 24.6138 81.3953 12 9.2 87

Cottrel (Stingray) 24.6137 81.9213 2 2.0 36

No Name Reef 24.5965 81.2140 12 12.5 126

Nine Foot Stake 24.5683 81.5517 32 33.0 134

Looe Key - Research 24.5667 81.3933 32 31.6 183

Looe Key - East 24.5450 81.4083 48 52.4 204

Widow Fingers 24.5117 81.6172 38 40.5 190

Pelican Shoals 24.5020 81.6230 39 41.2 207

Middle Sambo 24.4952 81.6965 50 53.8 200

Eastern Sambo 24.4848 81.6648 63 69.4 213

Western Sambo 24.4730 81.7143 114 116.1 224

Research Site #1 24.4612 82.2047 18 21.8 155

Eastern Dry Rocks 24.4583 81.8407 57 55.1 203

Sand Key 24.4508 81.8778 100 102.8 223

Rock Key 24.4490 81.8563 51 50.5 210

Western Dry Rocks 24.4443 81.9305 39 35.5 150

Lost Reef 24.4433 81.9325 7 5.4 85

Trinity Cove 24.4338 81.9330 9 6.9 126
45

Appendix 1 continued

Location Lat. (o N) Long. (o W) Surveys (no.) Hours

(tot.)

Species

(tot.)

Dry Tortugas

The Wall (before Carysfort) 25.1693 80.2663 2 2.2 87

Shark’s Reef 25.1475 80.2927 20 20.7 175

Sherwood Forest 24.7115 83.0468 8 5.1 80

Robins Hood 24.7072 83.0475 7 4.0 64

Squid Row 24.7030 82.8593 10 8.5 60

Pulaski 24.6955 82.7713 31 29.4 172

Big Johnson 24.6843 82.8832 21 22.7 126

Oklahoma 24.6840 83.0505 7 6.8 81

Texas Rock 24.6817 82.8847 54 51.7 177

Cessies Peak (aka Bird In Hand) 24.6782 83.0375 6 5.0 84

Wreck Reef (Tortugas Banks) 24.6765 83.0242 13 9.9 113

Juanita’s Reef 24.6672 82.8920 37 38.0 149

The Gap 24.6660 80.9718 4 2.9 123

Blenny flats 24.6553 82.7877 16 17.7 86

G-Spot (near Pinnacles) 24.6538 83.0333 13 11.2 120

Oasis 24.6442 82.9295 18 16.8 97

Loggerhead Nursery 24.6385 82.9320 20 23.4 106

SW Loggerhead 24.6318 82.9362 2 2.0 16

Garlic Gardens (near Bird Key) 24.6217 82.9005 6 7.2 73

Windjammer Site 24.6212 82.9430 7 8.0 105

Bird Key 24.6128 82.8713 72 63.2 142

Simon’s Hump 24.5077 82.8775 18 11.7 105

Riley’s hump 24.4937 83.1218 13 7.9 94

Cuda Reef/Marquesas Rock 24.4593 82.2245 6 7.1 113
46

Appendix 2. Species list for the Flordia Keys and Dry Tortugas.

Family Common family Species Common species

Acanthuridae Surgeonfish Acanthurus coeruleus Blue Tang

Acanthuridae Surgeonfish Acanthurus chirurgus Doctorfish

Acanthuridae Surgeonfish Acanthurus bahianus Ocean Surgeonfish

Antennariidae Frogfish Antennarius multiocellatus Longlure Frogfish

Apogonidae Cardinalfish Apogon binotatus Barred Cardinalfish

Apogonidae Cardinalfish Apogon townsendi Belted Cardinalfish

Apogonidae Cardinalfish Apogon affinis Bigtooth Cardinalfish

Apogonidae Cardinalfish Apogon aurolineatus Bridle Cardinalfish

Apogonidae Cardinalfish Apogon maculatus Flamefish

Apogonidae Cardinalfish Apogon planifrons Pale Cardinalfish

Apogonidae Cardinalfish Apogon robinsi Roughlip Cardinalfish

Apogonidae Cardinalfish Apogon quadrisquamatus Sawcheek Cardinalfish

Apogonidae Cardinalfish Apogon pseudomaculatus Twospot Cardinalfish

Apogonidae Cardinalfish Apogon lachneri Whitestar Cardinalfish

Apogonidae Cardinalfish Astrapogon puncticulatus Blackfin Cardinalfish

Apogonidae Cardinalfish Astrapogon stellatus Conchfish

Apogonidae Cardinalfish Phaeoptyx pigmentaria Dusky Cardinalfish

Apogonidae Cardinalfish Phaeoptyx xenus Sponge Cardinalfish

Atherinidae/ Clupeidae/

Engraulidae

Silversides, Herrings,

Anchovies

Aulostomidae Trumpetfish Aulostomus maculatus Trumpetfish

Balistidae Leatherjacket Aluterus schoepfi Orange Filefish

Balistidae Leatherjacket Aluterus scriptus Scrawled Filefish

Balistidae Leatherjacket Balistes capriscus Gray Triggerfish

Balistidae Leatherjacket Balistes vetula Queen Triggerfish

Balistidae Leatherjacket Cantherhines pullus Orangespotted Filefish

Balistidae Leatherjacket Cantherhines macrocerus Whitespotted Filefish

Balistidae Leatherjacket Canthidermis sufflamen Ocean Triggerfish

Balistidae Leatherjacket Canthidermis maculata Rough Triggerfish

Balistidae Leatherjacket Melichthys niger Black Durgon

Balistidae Leatherjacket Monacanthus ciliatus Fringed Filefish

Balistidae Leatherjacket Monacanthus hispidus Planehead Filefish

Balistidae Leatherjacket Monacanthus setifer Pygmy Filefish

Balistidae Leatherjacket Monacanthus tuckeri Slender Filefish

Batrachoididae Toadfish Opsanus tau Oyster Toadfish

Belonidae Needlefish Playbelone argalus Keeltail Needlefish

Belonidae Needlefish Strongylura marina Atlantic Needlefish

Belonidae Needlefish Strongylura notata Redfin Needlefish

Belonidae Needlefish Tylosurus crocodilus Houndfish

Blenniidae Blenny Hypleurochilus bermudensis Barred Blenny

Blenniidae Blenny Ophioblennius atlanticus Redlip Blenny

Blenniidae Blenny Parablennius marmoreus Seaweed Blenny

Blenniidae Blenny Scartella cristata Molly Miller

Bothidae Flounder Bothus ocellatus Eyed Flounder

Bothidae Flounder Bothus lunatus Peacock Flounder

Bothidae Flounder Paralichthys albigutta Gulf Flounder

Bothidae Flounder Syacium micrurum Channel Flounder

Callionymidae Dragonet Diplogrammus pauciradiatus Spotted Dragonet

Callionymidae Dragonet Paradiplogrammus bairdi Lancer (coral) Dragonet

Carangidae Jack Alectis ciliaris African Pompano (Threadfin/threadfish)

Carangidae Jack Caranx ruber Bar Jack (Skipjack)

Carangidae Jack Caranx lugubris Black Jack

Carangidae Jack Caranx crysos Blue Runner (Hard-tailed Jack)

Carangidae Jack Caranx hippos Crevalle Jack

Carangidae Jack Caranx latus Horse-Eye Jack

Carangidae Jack Caranx bartholomaei Yellow Jack

Carangidae Jack Decapterus macarellus Mackerel Scad

Carangidae Jack Elagatis bipinnulata Rainbow Runner

Carangidae Jack Seriola rivoliana Almaco Jack

Carangidae Jack Seriola dumerili Greater Amberjack

Carangidae Jack Trachinotus goodei Palometa (Gaftopsail Pompano/Longfin

Pompano)

Carangidae Jack Trachinotus falcatus Permit (Round Pompano)

Carcharhinidae Requiem Shark Carcharhinus limbatus Blacktip Shark
47

Appendix 2 continued

Family Common family Species Common species

Carcharhinidae Requiem Shark Carcharhinus perezi Reef Shark

Centropomidae Snook Centropomus undecimalis Common Snook

Chaetodontidae Butterflyfish Chaetodon striatus Banded Butterflyfish

Chaetodontidae Butterflyfish Chaetodon capistratus Foureye Butterflyfish

Chaetodontidae Butterflyfish Chaetodon aculeatus Longsnout Butterflyfish

Chaetodontidae Butterflyfish Chaetodon sedentarius Reef Butterflyfish

Chaetodontidae Butterflyfish Chaetodon ocellatus Spotfin Butterflyfish

Cirrhitidae Hawkfish Amblycirrhitus pinos Redspotted Hawkfish

Clinidae Blenny Acanthemblemaria chaplini Papillose Blenny

Clinidae Blenny Acanthemblemaria aspera Roughhead Blenny

Clinidae Blenny Acanthemblemaria maria Secretary Blenny

Clinidae Blenny Acanthemblemaria spinosa Spinyhead Blenny

Clinidae Blenny Chaenopsis limbaughi Yellowface Pikeblenny

Clinidae Blenny Coralliozetus bahamensis Blackhead Blenny

Clinidae Blenny Emblemaria pandionis Sailfin Blenny

Clinidae Blenny Hemiemblemaria simulus Wrasse Blenny

Congridae Conger Heteroconger halis Brown Garden Eel

Coryphanidae Dolphin Coryphaena hippurus Dolphin (Mahi-Mahi)

Dasyatidae Stingray Dasyatis americana Southern Stingray

Echeneidae Remora Echeneis naucrates Sharksucker

Echeneidae Remora Echeneis neucratoides Whitefin Sharksucker

Echeneidae Remora Remora remora Remora

Elopidae Tarpon Megalops atlanticus Tarpon

Ephippidae Spadefish Chaetodipterus faber Atlantic Spadefish

Exocoetidae Flyingfish/Halfbeak Hemiramphus brasiliensis Ballyhoo

Exocoetidae Flyingfish/Halfbeak Hirundichthys speculiger Mirrorwing Flyingfish

Fistulariidae Cornetfish Fistularia tabacaria Bluespotted Cornetfish

Gerreidae Mojarra Eucinostomus jonesi Slender Mojarra

Gerreidae Mojarra Gerres cinereus Yellowfin Mojarra

Gobiidae Goby Bollmannia boqueronensis White-eye Goby

Gobiidae Goby Coryphopterus glaucofraenum Bridled Goby

Gobiidae Goby Coryphopterus dicrus Colon Goby

Gobiidae Goby Coryphopterus personatus/hyalinus Masked Goby/Glass Goby

Gobiidae Goby Coryphopterus eidolon Pallid Goby

Gobiidae Goby Coryphopterus lipernes Peppermint Goby

Gobiidae Goby Coryphopterus punctipectophorus Spotted Goby

Gobiidae Goby Gnatholepis thompsoni Goldspot Goby

Gobiidae Goby Gobionellus saepepallens Dash Goby

Gobiidae Goby Gobionellus stigmalophius Spotfin Goby

Gobiidae Goby Gobiosoma illecebrosum Barsnout Goby

Gobiidae Goby Gobiosoma prochilos Broadstripe Goby

Gobiidae Goby Gobiosoma genie Cleaning Goby

Gobiidae Goby Gobiosoma saucrum Leopard Goby

Gobiidae Goby Gobiosoma oceanops Neon Goby

Gobiidae Goby Gobiosoma dilepsis Orangesided Goby

Gobiidae Goby Gobiosoma grosvenori Rockcut Goby

Gobiidae Goby Gobiosoma evelynae Sharknose Goby

Gobiidae Goby Gobiosoma louisae Spotlight Goby

Gobiidae Goby Gobiosoma macrodon Tiger Goby

Gobiidae Goby Gobiosoma horsti Yellowline Goby

Gobiidae Goby Gobiosoma randalli Yellownose Goby

Gobiidae Goby Gobiosoma xanthiprora Yellowprow Goby

Gobiidae Goby Ioglossus calliuris Blue Goby

Gobiidae Goby Ioglossus helenae Hovering Goby

Gobiidae Goby Lophogobius cyprinoides Crested Goby

Gobiidae Goby Microgobius microlepis Banner Goby

Gobiidae Goby Microgobius carri Seminole Goby

Gobiidae Goby Nes longus Orangespotted Goby

Gobiidae Goby Priolepis hipoliti Rusty Goby

Gobiidae Goby Risor ruber Tusked Goby

Grammatidae Basslet Gramma melacara Blackcap Basslet

Grammatidae Basslet Gramma loreto Fairy Basslet (Royal gramma)

Haemulidae Grunt Anisotremus surinamensis Black Margate

Haemulidae Grunt Anisotremus virginicus Porkfish
48

Appendix 2 continued

Family Common family Species Common species

Haemulidae Grunt Haemulon bonariense Black Grunt

Haemulidae Grunt Haemulon sciurus Bluestriped Grunt

Haemulidae Grunt Haemulon carbonarium Caesar Grunt

Haemulidae Grunt Haemulon melanurum Cottonwick

Haemulidae Grunt Haemulon flavolineatum French Grunt

Haemulidae Grunt Haemulon parra Sailors Choice

Haemulidae Grunt Haemulon chrysargyreum Smallmouth Grunt

Haemulidae Grunt Haemulon macrostomum Spanish Grunt

Haemulidae Grunt Haemulon striatum Striped Grunt

Haemulidae Grunt Haemulon aurolineatum Tomtate

Haemulidae Grunt Haemulon plumieri White Grunt

Haemulidae Grunt Haemulon album White Margate

Haemulidae Grunt Orthopristis chrysoptera Pigfish

Holocentridae Squirrelfish Holocentrus vexillarius Dusky Squirrelfish

Holocentridae Squirrelfish Holocentrus marianus Longjaw Squirrelfish

Holocentridae Squirrelfish Holocentrus rufus Longspine Squirrelfish

Holocentridae Squirrelfish Holocentrus coruscum Reef Squirrelfish

Holocentridae Squirrelfish Holocentrus adscensionis Squirrelfish

Holocentridae Squirrelfish Myripristis jacobus Blackbar Soldierfish

Holocentridae Squirrelfish Plectrypops retrospinis Cardinal Soldierfish

Inermiidae Bonnetmouth Emmelichthyops atlanticus Bonnetmouth

Inermiidae Bonnetmouth Inermia vittata Boga

Kyphosidae Chub Kyphosus sectatrix/incisor Bermuda Chub/Yellow Chub

Labridae Wrasse Bodianus rufus Spanish Hogfish

Labridae Wrasse Bodianus pulchellus Spotfin Hogfish

Labridae Wrasse Clepticus parrae Creole Wrasse

Labridae Wrasse Doratonotus megalepis Dwarf Wrasse

Labridae Wrasse Halichoeres poeyi Blackear Wrasse

Labridae Wrasse Halichoeres maculipinna Clown Wrasse

Labridae Wrasse Halichoeres radiatus Puddingwife

Labridae Wrasse Halichoeres pictus Rainbow (painted) Wrasse

Labridae Wrasse Halichoeres bivittatus Slippery Dick

Labridae Wrasse Halichoeres cyanocephalus Yellowcheek Wrasse

Labridae Wrasse Halichoeres garnoti Yellowhead Wrasse

Labridae Wrasse Hemipteronotus splendens Green Razorfish

Labridae Wrasse Hemipteronotus novacula Pearly Razorfish

Labridae Wrasse Hemipteronotus martinicensis Rosy Razorfish

Labridae Wrasse Lachnolaimus maximus Hogfish

Labridae Wrasse Thalassoma bifasciatum Bluehead

Lutjanidae Snapper Lutjanus buccanella Blackfin Snapper

Lutjanidae Snapper Lutjanus cyanopterus Cubera Snapper

Lutjanidae Snapper Lutjanus jocu Dog Snapper

Lutjanidae Snapper Lutjanus griseus Gray (mangrove) Snapper

Lutjanidae Snapper Lutjanus synagris Lane Snapper

Lutjanidae Snapper Lutjanus mahogoni Mahogany Snapper

Lutjanidae Snapper Lutjanus analis Mutton Snapper

Lutjanidae Snapper Lutjanus apodus Schoolmaster

Lutjanidae Snapper Ocyurus chrysurus Yellowtail Snapper

Malacanthidae Tilefish Malacanthus plumieri Sand Tilefish

Mugilidae Mullet Mugil cephalus Striped Mullet

Mullidae Goatfish Mulloidichthys martinicus Yellow Goatfish

Mullidae Goatfish Mullus auratus Red Goatfish

Mullidae Goatfish Pseudupeneus maculatus Spotted Goatfish

Muraenidae Moray Echidna catenata Chain Moray

Muraenidae Moray Enchelycore carychroa Chestnut Moray

Muraenidae Moray Enchelycore nigricans Viper Moray

Muraenidae Moray Gymnothorax miliaris Goldentail Moray

Muraenidae Moray Gymnothorax funebris Green Moray

Muraenidae Moray Gymnothorax vicinus Purplemouth Moray

Muraenidae Moray Gymnothorax moringa Spotted Moray

Myliobatidae Eagle Ray Aetobatus narinari Spotted Eagle Ray

Ogcocephalidae Batfish Ogcocephalus radiatus Polka-dot Batfish

Ophichthidae Snake Eel Myrichthys ocellatus Goldspotted Eel

Opistognathidae Jawfish Opistognathus macrognathus Banded Jawfish
49

Appendix 2 continued

Family Common family Species Common species

Opistognathidae Jawfish Opistognathus whitehursti Dusky Jawfish

Opistognathidae Jawfish Opistognathus aurifrons Yellowhead Jawfish

Ostraciidae Boxfish Lactophrys trigonus BuffaloTrunkfish

Ostraciidae Boxfish Lactophrys polygonia Honeycomb Cowfish

Ostraciidae Boxfish Lactophrys quadricornis Scrawled Cowfish

Ostraciidae Boxfish Lactophrys triqueter Smooth Trunkfish

Ostraciidae Boxfish Lactophrys bicaudalis Spotted Trunkfish

Pempheridae Sweeper Pempheris schomburgki Glassy Sweeper (Copper

Sweeper/Hatchetfish)

Pomacanthidae Angelfish Centropyge argi Cherubfish

Pomacanthidae Angelfish Holacanthus bermudensis Blue Angelfish

Pomacanthidae Angelfish Holacanthus ciliaris Queen Angelfish

Pomacanthidae Angelfish Holacanthus tricolor Rock Beauty

Pomacanthidae Angelfish Holacanthus sp. (Hybrid) Townsend Angelfish

Pomacanthidae Angelfish Pomacanthus paru French Angelfish

Pomacanthidae Angelfish Pomacanthus arcuatus Gray Angelfish

Pomacentridae Damselfish Abudefduf taurus Night Sergeant

Pomacentridae Damselfish Abudefduf saxatilis Sergeant Major

Pomacentridae Damselfish Chromis cyanea Blue Chromis

Pomacentridae Damselfish Chromis multilineata Brown Chromis

Pomacentridae Damselfish Chromis scotti Purple Reeffish

Pomacentridae Damselfish Chromis insolata Sunshinefish

Pomacentridae Damselfish Chromis enchrysura Yellowtail Reeffish

Pomacentridae Damselfish Microspathodon chrysurus Yellowtail Damselfish

Pomacentridae Damselfish Stegastes leucostictus Beaugregory

Pomacentridae Damselfish Stegastes partitus Bicolor Damselfish

Pomacentridae Damselfish Stegastes variabilis Cocoa Damselfish

Pomacentridae Damselfish Stegastes fuscus Dusky Damselfish

Pomacentridae Damselfish Stegastes diencaeus Longfin Damselfish

Pomacentridae Damselfish Stegastes planifrons Threespot Damselfish

Priacanthidae Bigeye Priacanthus arenatus Bigeye

Priacanthidae Bigeye Priacanthus cruentatus Glasseye Snapper

Rachycentridae Cobia Rachycentron canadum Cobia

Rhincodontidae Carpet Shark Ginglymostoma cirratum Nurse Shark

Rhinobatidae Guitarfish Rhinobatos lentiginosus Atlantic Guitarfish

Rhinobatidae Guitarfish Rhinobatos percellens Southern Guitarfish

Scaridae Parrotfish Cryptotomus roseus Bluelip Parrotfish (Rosy Parrotfish/Slender

Parrotfish)

Scaridae Parrotfish Nicholsina usta Emerald Parrotfish

Scaridae Parrotfish Scarus coeruleus Blue Parrotfish

Scaridae Parrotfish Scarus coelestinus Midnight Parrotfish

Scaridae Parrotfish Scarus taeniopterus Princess Parrotfish

Scaridae Parrotfish Scarus vetula Queen Parrotfish

Scaridae Parrotfish Scarus guacamaia Rainbow Parrotfish

Scaridae Parrotfish Scarus croicensis Striped Parrotfish

Scaridae Parrotfish Sparisoma radians Bucktooth Parrotfish

Scaridae Parrotfish Sparisoma atomarium Greenblotch Parrotfish

Scaridae Parrotfish Sparisoma aurofrenatum Redband Parrotfish

Scaridae Parrotfish Sparisoma rubripinne Redfin (yellowtail) Parrotfish

Scaridae Parrotfish Sparisoma chrysopterum Redtail Parrotfish

Scaridae Parrotfish Sparisoma viride Stoplight Parrotfish

Sciaenidae Drum Bairdiella sanctaeluciae Striped Croaker

Sciaenidae Drum Equetus umbrosus Cubbyu

Sciaenidae Drum Equetus acuminatus Highhat

Sciaenidae Drum Equetus lanceolatus Jackknife-Fish

Sciaenidae Drum Equetus punctatus Spotted Drum

Sciaenidae Drum Odontoscion dentex Reef Croaker

Scombridae Mackerel Acanthocybium solandri Wahoo

Scombridae Mackerel Scomberomorus regalis Cero

Scombridae Mackerel Scomberomorus maculatus Spanish Mackerel

Scorpaenidae Scorpionfish Scorpaena plumieri Spotted Scorpionfish

Scorpaenidae Scorpionfish Scorpaenodes caribbaeus Reef Scorpionfish

Serranidae Seabass Alphestes afer Mutton Hamlet

Serranidae Seabass Diplectrum bivittatum Dwarf Sand Perch
50

Appendix 2 continued

Family Common family Species Common species

Serranidae Seabass Diplectrum formosum Sand Perch

Serranidae Seabass Epinephelus fulvus Coney

Serranidae Seabass Epinephelus cruentatus Graysby (Kitty Mitchell)

Serranidae Seabass Epinephelus itajara Jewfish

Serranidae Seabass Epinephelus striatus Nassau Grouper

Serranidae Seabass Epinephelus morio Red Grouper

Serranidae Seabass Epinephelus guttatus Red Hind (Speckled Hind/Strawberry

Grouper)

Serranidae Seabass Epinephelus adscensionis Rock Hind

Serranidae Seabass Epinephelus nigritus Warsaw Grouper

Serranidae Seabass Hypoplectrus puella Barred Hamlet

Serranidae Seabass Hypoplectrus nigricans Black Hamlet

Serranidae Seabass Hypoplectrus gemma Blue Hamlet

Serranidae Seabass Hypoplectrus unicolor Butter Hamlet

Serranidae Seabass Hypoplectrus gummigutta Golden Hamlet

Serranidae Seabass Hypoplectrus (Hybrid) Hybrid Hamlet

Serranidae Seabass Hypoplectrus indigo Indigo Hamlet

Serranidae Seabass Hypoplectrus sp. Masked Hamlet

Serranidae Seabass Hypoplectrus guttavarius Shy Hamlet

Serranidae Seabass Hypoplectrus sp. Tan Hamlet

Serranidae Seabass Hypoplectrus aberrans Yellowbelly Hamlet

Serranidae Seabass Hypoplectrus chlorurus Yellowtail Hamlet

Serranidae Seabass Liopropoma carmabi Candy Bass

Serranidae Seabass Liopropoma rubre Peppermint Bass

Serranidae Seabass Mycteroperca bonaci Black Grouper

Serranidae Seabass Mycteroperca rubra Comb Grouper

Serranidae Seabass Mycteroperca microlepis Gag

Serranidae Seabass Mycteroperca phenax Scamp (Salmon Rockfish)

Serranidae Seabass Mycteroperca tigris Tiger Grouper

Serranidae Seabass Mycteroperca venenosa Yellowfin Grouper

Serranidae Seabass Mycteroperca interstitialis Yellowmouth Grouper

Serranidae Seabass Paranthias furcifer Creole-fish

Serranidae Seabass Rypticus bistrispinus Freckled Soapfish

Serranidae Seabass Rypticus saponaceus Greater Soapfish

Serranidae Seabass Rypticus subbifrenatus Spotted Soapfish

Serranidae Seabass Rypticus maculatus Whitespotted Soapfish

Serranidae Seabass Serranus subligarius Belted Sandfish (Belted Sand Bass)

Serranidae Seabass Serranus tortugarum Chalk Bass

Serranidae Seabass Serranus tigrinus Harlequin Bass

Serranidae Seabass Serranus baldwini Lantern Bass

Serranidae Seabass Serranus annularis Orangeback Bass

Serranidae Seabass Serranus tabacarius Tobaccofish

Sparidae Porgy Archosargus rhomboidalis Sea Bream

Sparidae Porgy Archosargus probatocephalus Sheepshead

Sparidae Porgy Calamus bajonado Jolthead Porgy

Sparidae Porgy Calamus nodosus Knobbed Porgy

Sparidae Porgy Calamus proridens Littlehead Porgy

Sparidae Porgy Calamus pennatula Pluma

Sparidae Porgy Calamus calamus Saucereye Porgy

Sparidae Porgy Calamus penna Sheepshead Porgy

Sparidae Porgy Diplodus argenteus Silver Porgy

Sparidae Porgy Diplodus holbrooki Spottail Pinfish

Sparidae Porgy Lagodon rhomboides Pinfish

Sphyraenidae Barracuda Sphyraena barracuda Great Barracuda

Sphyraenidae Barracuda Sphyraena picudilla Southern Sennet

Sphyrnidae Hammerhead Shark Sphyrna tiburo Bonnethead

Sphyrnidae Hammerhead Shark Sphyrna mokarran Great Hammerhead

Sphyrnidae Hammerhead Shark Sphyrna lewini Scalloped Hammerhead

Syngnathidae Pipefish & Seahorse Cosmocampus elucens Shortfin Pipefish

Syngnathidae Pipefish & Seahorse Micrognathus ensenadae Harlequin Pipefish

Synodontidae Lizardfish Synodus saurus Bluestriped Lizardfish

Synodontidae Lizardfish Synodus foetens Inshore Lizardfish

Synodontidae Lizardfish Synodus synodus Red Lizardfish (Rockspear)

Synodontidae Lizardfish Synodus intermedius Sand Diver
51

Appendix 2 continued

Family Common family Species Common species

Tetraodontidae Puffer Canthigaster rostrata Sharpnose Puffer

Tetraodontidae Puffer Chilomycterus antennatus Bridled Burrfish

Tetraodontidae Puffer Chilomycterus schoepfi Striped Burrfish

Tetraodontidae Puffer Chilomycterus antillarum Web Burrfish

Tetraodontidae Puffer Diodon holocanthus Balloonfish (Spiny Puffer)

Tetraodontidae Puffer Diodon hystrix Porcupinefish (Spotted Spiny Puffer)

Tetraodontidae Puffer Sphoeroides spengleri Bandtail Puffer

Tetraodontidae Puffer Sphoeroides greeleyi Caribbean Puffer

Tetraodontidae Puffer Sphoeroides testudineus Checkered Puffer

Tetraodontidae Puffer Sphoeroides nephelus Southern Puffer

Torpedinidae Electric Ray Narcine brasiliensis Lesser Electric Ray

Tripterygiidae Triplefins Enneanectes altivelis Lofty Triplefin

Tripterygiidae Triplefins Enneanectes pectoralis Redeye Triplefin

Urolophidae Round Stingrays Urolophus jamaicensis Yellow Stingray

Stromateidae Butterfishes Nomeus gronovii Man-of-war Fish

Carapidae Carapuds Carapus bermudensis Pearlfish

Labrisomidae Labrisomids Labrisomus kalisherae Downy Blenny

Labrisomidae Labrisomids Labrisomus nuchipinnis Hairy Blenny

Labrisomidae Labrisomids Labrisomus gobio Palehead Blenny

Labrisomidae Labrisomids Labrisomus bucciferus Puffcheek Blenny

Labrisomidae Labrisomids Malacoctenus versicolor Barfin Blenny

Labrisomidae Labrisomids Malacoctenus boehlkei Diamond Blenny

Labrisomidae Labrisomids Malacoctenus macropus Rosy Blenny

Labrisomidae Labrisomids Malacoctenus triangulatus Saddled Blenny

Labrisomidae Labrisomids Starksia hassi Ringed Blenny

Istiophoridae Billfishes Makaira nigricans Blue Marlin