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STATUS OF CORAL REEFS OF LITTLE CAYMAN AND GRAND CAYMAN, BRITISH WEST INDIES, IN 1999 (PART 2: FISHES)
BY
CHRISTY V. PATTENGILL-SEMMENS,1 and BRICE X. SEMMENS2
ABSTRACT
The fish assemblages at 33 sites around the islands of Grand Cayman and Little Cayman were assessed in June 1999 for the Atlantic and Gulf Rapid Reef Assessment initiative using belt transects and Roving Diver Technique surveys. A comprehensive species list, with 58 new records, was compiled for the Cayman Islands based on these data and survey data from the Reef Environmental Education Foundation database. In general, the reefs on Little Cayman appeared to support larger and more individual fishes than those of Grand Cayman. A multidimensional scaling ordination plot showed no clear island pattern but did reveal that the windward or leeward location of each site was an important factor affecting fish community composition. All but two sites followed a pattern of distinct windward and leeward clusters, and these clusters also correlated to macroalgal abundance. The relationship between macroalgal abundance and herbivore density was analyzed and significant correlations were found with surgeonfishes (Acanthuridae) and parrotfishes (Scaridae) using multiple regression.
INTRODUCTION
Fishes have the potential to provide sensitive indices of reef health. Certain predatory fish species dominate the top of coral reef food webs, hence their density reflects a vast number of human and natural disturbances from habitat alteration to direct exploitation (Ferreira et al., 1998). Similarly, the presence and abundance of herbivorous fishes affect algal composition and cover (Ogden and Lobel, 1978).
In response to concerns about the widespread deterioration of reef condition in the Caribbean basin, the Atlantic and Gulf Rapid Reef Assessment (AGRRA) initiative was designed to provide a regional perspective using a standardized methodology. The rapid assessment protocol is focused on three main components of the reef community: stony corals, fish, and algae. As part of this initiative, the reefs of Grand Cayman (GC) and Little Cayman (LC) were assessed in June 1999.
1 Reef Environmental Education Foundation, P.O. Box 246, Key Largo, FL 33037. Email: [email protected]
2 University of Washington, Dept. of Zoology, Box 351800, Seattle, WA 98195-1800.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 1 (actual page number in volume to be determined)
The Cayman Islands are a British Crown Colony located in the western Caribbean. The three islands lie between 19° 15' and 19° 45' N latitude and between 79° 44 ' and 81° 27' W longitude (Fig. 1). GC is the largest and most populous. LC lies approximately 145 km to its east-northeast and is about 10 km from Cayman Brac. The three islands are limestone, horst-and-graben structures associated with the Cayman Ridge (Jones, 1994). Freshwater is scarce and the islands lack rivers and streams. The fringing reefs that surround most of the islands contain shallow reef crests (rubble ramparts) as well as mid-shelf and shelf-edge fore reefs (Blanchon and Jones, 1997). These fringing reefs are particularly well-developed on the windward (eastern and southern) coasts of both islands. Other submerged benthic habitats include seagrass beds and mangrove fringes.
The level of human disturbance on GC is significantly greater than on LC, which is relatively remote and undeveloped. Anthropogenic impacts on GC reefs include habitat destruction from anchors and increased suspended sediment load from dredging and mangrove removal. Fishing pressure is considerably greater on GC than around LC. Five spawning aggregations of Nassau grouper (Epinephelus striatus) have been heavily harvested (during the 2002 spawning season, all but one had been depleted). Five hundred local residents are licensed to snorkel with spearguns. Fish pots (Antillean Z-traps) probably represent the biggest threat to the fish communities of both islands (personal observations).
An extensive marine park system was established in the Cayman Islands in 1986. Reefs in marine park and replenishment zone areas are protected from fish traps, spearguns, anchoring, and line fishing, although line fishing from shore and beyond the drop-off (shelf edge) is allowed. The Cayman Islands’ Department of the Environment maintains a system of 257 permanent mooring buoys throughout the three islands.
The benthos of the Cayman Islands has been well studied, including descriptions of the coral communities, reef status, and analysis of spatial patterns (Roberts, 1988; Logan, 1994; Roberts, 1994). In contrast, apart from descriptions of Nassau grouper spawning aggregations (Colin et al., 1987; Tucker et al., 1993), there are few scientific descriptions of its reef fishes. However, Burgess et al.’s (1994) taxonomic review of collection expeditions contained an annotated list of 381 species known to occur in the Cayman Islands, including the endemic y-lined blenny (Starksia y-lineata) described by Gilbert (1965).
Since 1994, fish sighting and relative abundance data have been collected around the Cayman Islands as part of the Reef Environmental Education Foundation (REEF) Fish Survey Project, an ongoing volunteer monitoring effort. REEF volunteers use the Roving Diver Technique (RDT) (Schmitt and Sullivan, 1996) and the survey data are maintained in a publicly-accessible database. By the end of 2001, the REEF database contained over 40,000 surveys from over 2,000 sites, including approximately 2,200 surveys from the Cayman Islands.
This paper describes the fish assemblages of the Cayman Islands using the 1999 AGRRA data for GC and LC, along with REEF data from the two islands collected between 1994 and 2001. An updated species list and comparisons between islands and among sites are provided. The relationship between herbivorous fishes and macroalgal abundance is also investigated.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 2 (actual page number in volume to be determined)
METHODS
In June 1999, AGRRA fish and benthos surveys were simultaneously conducted at 15 sites on GC and 18 sites on LC (Fig. 1, Table 1). Sites were chosen by a mixed representative/strategic strategy: 12 were on the windward sides of the islands and 21 were on their leeward sides (the southwest side of GC was underrepresented). Six sites on LC and three on GC were located within marine park or replenishment zone areas. The benthic component is reported by Manfrino et al. (this volume). To assess the fishes, the AGRRA protocol Version 2 was used (Appendix One). At each site, a team of three (occasionally two) divers conducted at least 10 2 m x 30 m belt transects. Counts of serranids (groupers) were restricted to species of Epinephelus and Mycteroperca; scarids (parrotfishes) and haemulids (grunts) less than 5 cm in length were not tallied. Each diver also conducted a 45-60 minute RDT survey at each site. All fieldwork was undertaken between 9:00 a.m. and 3:00 p.m. Field identifications were based on Humann (1994), Stokes (1980), and Robins et al. (1986).
The fish transect data were entered into a custom AGRRA Excel spreadsheet. REEF provided the RDT data in ASCII format. Using the transects as replicates, the average density (#/100 m2) and size (cm) of each species and family were calculated for each site. Analyses were done at the regional (GC versus LC) and site levels, incorporating reef location (windward, leeward) and benthic parameters when appropriate. The average density and size of each species and family were compared between regions using a t-test after testing the data for normality. Due to confounding factors such as differences in use (e.g., recreation, harvest) and hydrographic features, comparisons between protected (marine park) and unprotected sites were not attempted. The site data were used in a hierarchical cluster analysis using Pearson’s similarity index. The similarity matrix was generated using log-transformed density values for each species documented in at least three (10%) of the sites; the other 22 rare species were eliminated (per Grossman et al., 1982). A two-dimensional multidimensional scaling (MDS) ordination plot was also generated using the similarity matrix. The transect data were also used to investigate interactions between the fish assemblages and the benthic community. This preliminary investigation was focused on herbivore/algae interactions. A regression was calculated on the densities of parrotfish and surgeonfish against percent absolute macroalgal abundance in quadrats (hereafter macroalgal abundance) and height at each site. Other coral factors (percent live coral cover, average colony height, percent diseased colonies) and environmental (windward/leeward) parameters were also plotted against each fish family. All values were transformed prior to regression (transformations were log+1 for fish density and algal height and arcsine of the square root for proportions).
The RDT survey data provided species lists, frequency of occurrence, and relative abundance estimates. Percent sighting frequency (%SF) for each species was the percentage of all dives in which the species was recorded. An estimate of abundance was calculated as: abundance score = D x %SF, where the density score (D) for each species was a weighted average index based on the frequency of observations in different abundance categories. Density score was calculated as:
D= ((nSx1)+(nFx2)+(nMx3)+(nAx4)) / (nS + nF + nM + nA), where nS, nF, nM, and nA represented the number of times each abundance category (Single, Few, Many,
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 3 (actual page number in volume to be determined)
Figure 1. AGRRA survey sites in Grand Cayman and Little Cayman, Cayman Islands. See Table 1 for site codes.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 4 (actual page number in volume to be determined)
Abundant) was assigned for a given species. The RDT data were pooled and compared by island using the Wilcoxon Sign Rank test. Only species that were seen in at least 10% of the RDT AGRRA surveys were included in the analysis (103 species), reducing the effect of rare species (Grossman et al., 1982). SYSTAT 7.0 was used for all the analyses.
All expert-level REEF data from GC and LC, including the RDT data collected during the AGRRA expedition, were used to compile a species list of reef fishes for the Cayman Islands (REEF, 2001).
RESULTS
A total of 341 transects (142 - GC; 199 - LC) and 79 RDT surveys (32- GC and 47- LC) documenting 173 species were conducted at 33 reefs (Table 1). The RDT survey data were added to the existing REEF database. The total number of species recorded by REEF experts on the Cayman Islands between 1994 and 2001 was 276 (Appendix A, this paper). When compared with Burgess et al.’s (1994) ichthyofaunal list, the REEF survey data added 58 new species records for a total of 423 reef fishes documented on the Cayman Islands (five freshwater species, 10 deepwater (>300 m) species, and a misidentification (Stegastes mellis) listed by Burgess et al. (1994) were not included in this tally). The 25 most common species, according to %SF in the REEF database, are noted in Table 2.
Parrotfish (Scaridae) was the most abundant family recorded during the belt transects (Fig. 2). Average density of snapper (Lutjanidae) on LC was approximately twice that of GC reefs. Size frequency distributions of carnivores (select grouper genera and all snappers) and herbivores (parrotfishes ≥5 cm, surgeonfish (Acanthuridae), and the
0
5
10
15
20
25
30
Acanth
urida
eBali
stida
eCha
etodo
ntida
eHae
mulida
eLu
tjanid
aePo
macan
thida
eSc
arida
eSe
rranid
ae
Other
Mea
n D
ensi
ty (#
/100
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) Little CaymanGrand Cayman
0
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eBali
stida
eCha
etodo
ntida
eHae
mulida
eLu
tjanid
aePo
macan
thida
eSc
arida
eSe
rranid
ae
Other
Acanth
urida
eBali
stida
eCha
etodo
ntida
eHae
mulida
eLu
tjanid
aePo
macan
thida
eSc
arida
eSe
rranid
ae
Other
Mea
n D
ensi
ty (#
/100
m2
)M
ean
Den
sity
(#/1
00m
2) Little CaymanLittle Cayman
Grand CaymanGrand Cayman
Figure 2. Mean fish density (no. individuals/100m2 + sd) for AGRRA fishes in Grand Cayman and Little Cayman. Other = Bodianus rufus, Caranx ruber, Lachnolaimus maximus, Microspathodon chrysurus, Sphyraena barracuda.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 5 (actual page number in volume to be determined)
yellowtail damselfish Microspathodon chrysurus) are shown in Figure 3. Approximately 75% of the carnivores were less than 30 cm in length, and 85% of the herbivores were less than 20 cm in length. T-tests on these data showed that the average density and size for most species and families did not differ between islands. However, many species were reported in RDT surveys with greater than average abundance on the LC reefs (Wilcoxon Sign Rank p<0.0005). In particular, the sighting frequencies of six species of large groupers were considerably greater in LC (Table 3; Wilcoxon Sign Rank p<0.05). Exceptions included yellowtail snapper (Ocyurus chrysurus) and sergeant major (Abudefduf saxatilis), two species that become abundant when fed regularly by divers. Fish feeding is much more commonplace on GC reefs (Burgess et al., 1994; personal observations).
Site comparisons at the assemblage level showed no clear, intra-island groupings. However, two distinct clusters were obvious in the MDS plot (Fig. 4A) and, to a lesser extent, in the cluster diagram (Fig. 4B). The only environmental characteristics significantly related to fish density were reef location (windward/leeward) and macroalgal abundance. The windward (high-wave exposure) or leeward/protected windward (low wave-exposure) location of the sites was an important factor in the MDS cluster for all but two of the sites (LC02 and GC30). Leeward sites also had significantly higher macroalgal abundance than windward sites (45% versus 31%, respectively: F-test p<0.001; multiple R = 0.560).
Figure 3. Size frequency distribution of (A) carnivores (all Lutjanids, select Serranids) and (B) herbivores (Acanthurids, Scarids > 5 cm, Microspathodon chrysurus) in GC (Grand Cayman) and LC (Little Cayman). Total number of individuals counted (n) is given.
A Carnivores
Size (cm)
Prop
ortio
n (%
)
n=666LCGC n=292
0-5 6-10 11-20 21-30 31-40 >400
10
20
30
40
50
60Pr
opor
tion
(%)
0
10
20
30
40
50
60
0-5 6-10 11-20 21-30 31-40 >40
Size (cm)
B Herbivores
n=3884LCGC n=2825
A Carnivores
Size (cm)
Prop
ortio
n (%
)
n=666LCGC n=292
0-5 6-10 11-20 21-30 31-40 >400
10
20
30
40
50
60Pr
opor
tion
(%)
0
10
20
30
40
50
60
0-5 6-10 11-20 21-30 31-40 >40
Size (cm)
B Herbivores
n=3884LCGC n=2825
A Carnivores
Size (cm)
Prop
ortio
n (%
)
n=666LCGC n=292
0-5 6-10 11-20 21-30 31-40 >400
10
20
30
40
50
60
A Carnivores
Size (cm)
Prop
ortio
n (%
)
n=666LCGC n=292
n=666LC n=666LCGC n=292GC n=292
0-5 6-10 11-20 21-30 31-40 >400
10
20
30
40
50
60
0-5 6-10 11-20 21-30 31-40 >400-5 6-10 11-20 21-30 31-40 >400
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20
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50
60
0
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20
30
40
50
60
0
10
20
30
40
50
60Pr
opor
tion
(%)
0
10
20
30
40
50
60
0-5 6-10 11-20 21-30 31-40 >40
Size (cm)
B Herbivores
n=3884LCGC n=2825
Prop
ortio
n (%
)
0
10
20
30
40
50
60
0-5 6-10 11-20 21-30 31-40 >400
10
20
30
40
50
60
0
10
20
30
40
50
60
0
10
20
30
40
50
60
0-5 6-10 11-20 21-30 31-40 >400-5 6-10 11-20 21-30 31-40 >40
Size (cm)
B Herbivores
n=3884LCGC n=2825
n=3884LC n=3884LCGC n=2825GC n=2825
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 6 (actual page number in volume to be determined)
Surgeonfish density showed an inverse relationship with macroalgal abundance (p<0.01; r2 = 0.209), whereas parrotfish density was positively related to macroalgal abundance (p<0.01; r2 = 0.215) (Fig. 5). Adding macroalgal height to a multiple regression significantly improved the relationships with macroalgal abundance for parrotfish (p<0.001; r2 = 0.413) and surgeonfish (p=0.001; r2 = 0.367) densities. A strong inverse relationship between parrotfish and surgeonfish densities was also found (p<0.001; r2 = 0.384).
B Hierarchical cluster analysis A MDS ordination plot
Figure 4. (A) MDS ordination plot (left cluster is windward) and (B) hierarchical cluster analysis of AGRRA reef fish transect data in GC (Grand Cayman) and LC (Little Cayman).
0
5
10
15
20
25
30
35
40
0 10 20 30 40 50 60%Macro Algae Cover
Den
sity
(#/1
00m
2 )
Parrotfish SurgeonfishLinear (Surgeonfish)Linear (Parrotfish)
0
5
10
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20
25
30
35
40
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Den
sity
(#/1
00m
2 )
Parrotfish SurgeonfishLinear (Surgeonfish)Linear (Parrotfish)
ParrotfishParrotfish SurgeonfishSurgeonfishLinear (Surgeonfish)Linear (Surgeonfish)Linear (Parrotfish)Linear (Parrotfish)
Figure 5. Regression plot between mean parrotfish density (♦) and mean surgeonfish density (□) (no. individuals/100m2) and mean absolute macroalgal density, by site in the Cayman Islands.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 7 (actual page number in volume to be determined)
DISCUSSION
The reefs of the Cayman Islands support relatively diverse and abundant fish
assemblages. This richness is probably a result of several factors including high local habitat diversity, a significant (34%) area of coastal reserves, and a reef system that is generally in fair condition (Manfrino et al., this volume). However, significant differences were revealed between GC and LC, most likely a result of the greater anthropogenic impacts on GC reefs. Higher harvest pressure on GC was reflected in the lower density and size of large groupers, parrotfishes and snappers (Table 4) and lower sighting frequencies of large groupers (Table 3). Analyses of RDT data indicated that regardless of commercial importance, the average abundance of most fish species was higher on LC, hence other factors, such as coastal development and water pollution, may also adversely impact fish communities on GC.
The site-level transect density data correlated most strongly with relative wave exposure (Fig. 4A,B). Macroalgae were significantly less abundant overall on windward (high-wave exposure) sites than on leeward and protected windward (low-wave exposure) sites, where parrotfish were the most abundant fishes in the transects. It is clear, however, that macroalgal abundance does not by itself adequately explain site-level assemblage composition, given that LC sites had only slight differences between wave-exposed and non wave-exposed sites (Manfrino et al., this volume). The correlation between fish communities at sites with similar wave exposure highlights the effect of physical parameters on fish assemblage structure, and should be taken into consideration in future analyses of fish data for the Cayman Islands.
In a simple system, one might expect the presence and density of herbivorous species to be negatively correlated with algal abundance and height. In other words, a site with many herbivorous fish would have relatively low algal abundance due to grazing. Our analysis at the site level indicates that this expectation holds true for surgeonfish. However, the inverse is evident in parrotfish. This implies either or both of the following: 1) there is a direct or indirect interaction between parrotfish and surgeonfish, or, more generally 2) the dynamic spatial and temporal characteristics of reef fish confound simple relationships between resource availability and fish abundance. Recent work on stoplight parrotfish (Sparisoma viride) indicates that whereas there are few, if any, direct interactions between surgeonfish and parrotfish, the use of space on the reef by individual fish is complex (territorial behavior, depth partitioning based on social grouping), and varies as a function of social status and intraspecific interactions (van Rooij et al., 1996a; van Rooij et al., 1996b). Clearly, more research is needed to understand the use of space by reef fish if accurate conclusions are to be drawn from relationships between fish abundance and benthic conditions.
One of the crucial tasks that scientists face in implementing a “reef health scale” using AGRRA data is to determine exactly what indicators within the collected data track health. An additional challenge lies in assessing how to evaluate and analyze the broad and complementary set of information collected on fishes, stony corals, and algae. Results from this paper and others in this volume will provide valuable insight on these issues. Due to the inherently complex nature of coral reef communities, the manner in which AGRRA data will dictate a scale of reef condition is most certainly also complex.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 8 (actual page number in volume to be determined)
The negative relationship between surgeonfish and parrotfish at the site level is a good example of how community complexity may confound seemingly logical indicators of reef health such as herbivore biomass. Given our results, it is possible that Cayman Island reefs with similar herbivore biomass constituted by predominately different taxa may reflect dramatic differences in benthic conditions. The disparity between grouper abundance between the transect and RDT data and the dramatic increase in species reported in the Cayman Islands that resulted from the RDT surveys (18% based on the published list by Burgess et al., 1994) highlights the importance of using the two complementary visual fish-survey methods.
Because certain fish species dominate the top of coral reef food webs, a baseline of fish community composition and richness provides a useful tool for future assessment of reef health, given that a change in reef communities at lower trophic levels will most likely result in changes in the reef fish community composition (Choat, 1991; Jones et al., 1991). Additionally, because fish tend to be the most charismatic group of reef community members, changes in their community are most likely to be noticed and documented.
ACKNOWLEDGEMENTS
The authors wish to thank the Marine Education and Environmental Research Institute (MEERI) and Dr. C. Manfrino for coordinating this immense project. The data collection assistance of K. Hoshino, B. Hansrod, L. Whaylen, and P. Hillenbrand is also greatly appreciated. Funding support was provided by the Gilo Family Foundation, American Airlines, Island Air, Little Cayman Beach Resort, Southern Cross Club, the Reef Environmental Education Foundation, the Cayman Islands Department of Tourism and Tortuga Divers. The AGRRA Organizing Committee facilitated financial support as described in the Forward. The REEF volunteers who have contributed surveys from the Cayman Islands are also appreciated.
REFERENCES Blanchon P., and B. Jones 1997. Hurricane control on shelf-edge reef architecture around Grand Cayman.
Sedimentology 44:479-506. Burgess, G.H., S.H. Smith, and E.D. Lane 1994. Fishes of the Cayman Islands. Pp. 199-228. In: M. A. Brunt and J. E. Davies
(eds.), The Cayman Islands: Natural History and Biogeography. Dordrecht (Netherlands). Kluwer Academic Publishers. 576pp.
Choat, J.H. 1991. The biology of herbivorous fishes on coral reefs. Pp. 120-153. In: P. F. Sale
(ed.), The Ecology of Fishes on Coral Reefs. San Diego, California (USA). Academic Press, Inc. 754 pp.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 9 (actual page number in volume to be determined)
Colin, P.L., D.Y. Shapiro, and D. Weiler 1987 Aspects of the reproduction of two groupers, Epinephelus guttatus and E.
striatus in the West Indes. Bulletin of Marine Science 40: 220-230. Ferreira, C.E L., A.C. Peret, and R. Coutinho 1998. Seasonal grazing rates and food processing by tropical herbivorous fishes.
Journal of Fisheries Biology 53: Suppl. A 222-235. Gilbert, C.R.
1965. Starksia y-lineata, a new clinid fish from Grand Cayman Island, British West Indies. Notulae Naturae, Academy of Natural Sciences, Philadelphia 379.
6 pp. Grossman, G.D., P.B. Moyle, and J.J.O. Whitaker
1982. Stochasticity in structural and functional characteristics of an Indiana stream fish assemblage: a test of community theory. American Naturalist 120:
423-454. Humann, P. 1994. Reef Fish Identification (2nd ed.). Jacksonville, FL, New World Publications,
Inc. 396 pp. Jones, B.
1994. Geology of the Cayman Islands. Pp. 13-40. In: M. A. Brunt and J. E. Davies (eds.), The Cayman Islands: Natural History and Biogeography. Dordrecht (Netherlands). Kluwer Academic Publishers. 576pp.
Jones, G.P., D.J. Ferrell, and P.F. Sale 1991. Fish predation and its impact on the invertebrates of coral reefs and adjacent
sediments. Pp. 156-178. In: P.F. Sale (ed.). The Ecology of Fishes on Coral Reefs. San Diego, California (USA). Academic Press, Inc. 754 pp.
Logan, A. 1994. Reefs and lagoons of Cayman Brac and Little Cayman. Pp. 105-124. In:
M.A. Brunt and J.E. Davies (eds.), The Cayman Islands: Natural History and Biogeography. Dordrecht (Netherlands). Kluwer Academic Publishers. 576 pp.
Ogden, J.C., and P.S. Lobel 1978. The role of herbivorous fishes and urchins in coral reef communities.
Environmental Biology of Fishes 3:49-63. REEF 2001. Reef Environmental Education Foundation. World Wide Web electronic
publication. www.reef.org, date of download (31 December 2001). Roberts, H.H. 1988. Grand Cayman. Swamps and shallow marine substrates 1 and 2, Cayman
Islands 1: 25,000 (map). U.K.: Overseas Development Natural Resources Institute.
Roberts, H.H. 1994. Reefs and lagoons of Grand Cayman. Pp. 75-104. In: M. A. Brunt and J. E.
Davies (eds.), The Cayman Islands: Natural History and Biogeography. Dordrecht (Netherlands). Kluwer Academic Publishers. 576 pp.
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Robins, C.R., G.C. Ray, and J. Douglass 1986. Peterson Field Guides-Atlantic Coast Fishes. New York, NY. Houghton
Mifflin. 354 pp. Schmitt, E.F., and K.M. Sullivan 1996 Analysis of a volunteer method for collecting fish presence and abundance
data in the Florida Keys. Bulletin of Marine Science 59:404-416. Stokes, F.J. 1980 Handguide to the Coral Reef Fishes of the Caribbean and Adjacent Tropical
Waters Including Florida, Bermuda, and the Bahamas. New York, NY. Lippincott and Crowell, Publishers, 160 pp.
Tucker, J.W., P.G. Bush, and S.T. Slaybaugh 1993. Reproductive patterns of Cayman Islands Nassau grouper (Epinephelus
striatus) populations. Bulletin of Marine Science. 52:961-969. van Rooij, J.M., E.D. Jong, F. Vaandrager, and J.J. Videler 1996a. Resource and habitat sharing by stoplight parrotfish, Sparisoma viride, a
Caribbean reef herbivore. Environmental Biology of Fishes 47:81-91. van Rooij, J.M., F.J. Kroon, and J.J. Videler 1996b. The social and mating system of the herbivorous reef fish Sparisoma viride:
one-male versus multi-male groups. Environmental Biology of Fishes 47:353-378.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 11 (actual page number in volume to be determined)
Tabl
e 1.
Site
info
rmat
ion
for A
GR
RA
fish
surv
eys i
n Li
ttle
Cay
man
and
Gra
nd C
aym
an Is
land
s.
Nam
e1 Si
te
Prot
ectio
n2 Lo
catio
n3 R
eef T
ype4
Latit
ude °N
Lo
ngitu
de
°W
Surv
ey d
ate
Dep
th (m
) %
live
ston
y
cora
l cov
er
(mea
n ±
sd)5
30m
fish
tra
nsec
ts
(#)
RD
T su
rvey
s (#
)
Fish
spec
ies
(#)
Littl
e Ca
yman
Ji
gsaw
Puz
zle
LC02
op
en
lee
Hig
h S&
G
19 3
9.98
3'
80 0
6.39
0'
6-Ju
n-99
10
.5
27.1
± 9
.7
11
4 96
M
ixin
g B
owl
LC05
pa
rk
lee
She
lf ed
ge
19 4
1.09
6'
80 0
4.70
0'
8-Ju
n-99
12
.4
29.0
± 1
2.1
12
2 10
5 B
lack
Tip
Tun
nels
LC
06
repl
en
lee
S&G
19
42.
847'
79
57.
470'
9-
Jun-
99
12.4
15
.6 ±
7.4
12
2
74
Peng
uin'
s Lea
p LC
07
open
le
e H
ardp
an
19 4
2.55
1'
80 0
0.48
7'
9-Ju
n-99
16
.1
16.4
± 7
.9
12
2 64
M
eado
ws
LC08
pa
rk
lee
Patc
h
19 4
1.51
0'
80 0
4.13
0'
10-J
un-9
9 18
.4
37.1
± 1
1.6
14
3 88
N
ancy
's C
up o
f Tea
LC
09
park
le
e Sh
elf e
dge
19 4
1.63
9'
80 0
4.13
7'
10-J
un-9
9 12
.5
21.7
± 1
3.9
14
3 87
Jo
y's J
oy
LC10
pa
rk
lee
Shel
f edg
e 19
40.
690'
80
05.
575'
10
-Jun
-99
12.1
18
.9 ±
6.4
14
3
82
Paul
's A
ncho
r LC
12
park
le
e Sh
elf e
dge
19 4
1.66
1'
80 0
4.18
1'
13-J
un-9
9 12
.9
17.1
± 1
2.1
9 3
73
Roc
k B
otto
m W
all
LC13
op
en
lee
S&G
19
42.
057'
80
03.
421'
13
-Jun
-99
12.6
22
.6 ±
10.
3 10
2
70
LC
16
open
le
e S&
G
19 3
9.70
2'
80 0
6.72
8'
15-J
un-9
9 14
25
.7 ±
5.3
10
3
76
LC
17
open
le
e S&
G
19 4
2.47
0'
80 0
0.49
5'
15-J
un-9
9 8.
8 22
.7 ±
4.7
10
3
71
LC
18
repl
en
pro
win
d H
ardp
an
19 4
2.99
6'
79 5
8.92
1'
15-J
un-9
9 10
.8
14.9
± 5
.6
10
3 62
G
rund
y's G
arde
ns
LC01
pa
rk
win
d S&
G
19 3
9.42
1'
80 0
5.32
1'
6-Ju
n-99
9.
4 37
.7 ±
11.
7 10
3
80
Dis
neyl
and
LC03
re
plen
w
ind
S&G
19
49.
831'
80
01.
374'
7-
Jun-
99
9.9
25.4
± 6
.3
11
2 73
C
harle
s Bay
LC
04
open
w
ind
S&G
19
41.
628'
79
58.
459'
7-
Jun-
99
10.7
17
.3 ±
6.3
11
2
76
Mai
n C
hann
el E
ast
LC11
op
en
win
d Fr
ingi
ng
19 3
9.41
2'
80 0
4.36
8'
11-J
un-9
9 2.
8 15
.8 ±
5.5
9
1 30
Lu
cas's
Led
ge
LC14
re
plen
w
ind
S&G
19
40.
155'
80
02.
595'
14
-Jun
-99
13.6
24
.6 ±
5.9
10
3
68
LC
15
repl
en
win
d S&
G
19 4
0.62
8'
80 0
1.56
2'
14-J
un-9
9 10
.9
26.3
± 8
.7
10
3 65
G
rand
Cay
man
H
epp'
s Min
i Wal
l G
C28
pa
rk
lee
Patc
h/S&
G
19 2
3.12
6'
81 2
4.99
2'
20-J
un-9
9 11
.5
22.0
± 5
.6
5 1
35
Cem
eter
y R
eef
GC
32
park
le
e Pa
tch/
S&G
19
21.
917'
81
23.
726'
22
-Jun
-99
9 17
.3 ±
4.9
9
2 74
Su
nset
Hou
se
GC
33
park
le
e Pa
tch/
S&G
19
17.
172'
81
23.
463'
22
-Jun
-99
9.1
22.7
± 1
1.9
12
3 74
Is
abel
's R
eef
GC
22
open
pr
o w
ind
Hig
h S&
G
19 2
1.46
0'
81 0
8.14
5'
18-J
un-9
9 10
.6
24.3
± 7
.4
10
2 52
B
abyl
on
GC
23
repl
en
pro
win
d S&
G
19 2
1.20
0'
81 0
9.84
2'
18-J
un-9
9 9.
5 18
.2 ±
3.4
10
2
51
Del
ila's
Del
ight
G
C24
op
en
pro
win
d S&
G
19 2
1.51
8'
81 1
4.80
1'
19-J
un-9
9 7.
1 22
.8 ±
13.
1 10
2
56
Que
en's
Thro
ne
GC
25
repl
en
pro
win
d H
ardp
an
19 2
2.81
8'
81 1
7.49
3'
19-J
un-9
9 12
.1
14.7
± 3
.2
10
2 61
G
C26
re
plen
pr
o w
ind
S&G
19
21.
202'
81
11.
746'
19
-Jun
-99
13.9
12
.7 ±
4.1
10
2
49
Bea
r's P
aw
GC
27
repl
en
pro
win
d S&
G
19 2
3.85
4'
81 2
1.61
7'
20-J
un-9
9 10
.7
15.7
± 3
.4
5 1
46
GC
19
open
w
ind
Hig
h S&
G
19 1
9.05
8'
81 0
4.48
4'
17-J
un-9
9 6.
7 23
.1 ±
9.3
10
2
38
GC
20
open
w
ind
Hig
h S&
G
19 2
0.00
2'
81 0
4.59
6'
17-J
un-9
9 8.
9 17
.9 ±
4.6
10
2
49
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 12 (actual page number in volume to be determined)
Tabl
e 1,
Con
tinue
d
Nam
e Si
te1
Prot
ectio
n2 Lo
catio
n3 R
eef T
ype4
Latit
ude °N
Lo
ngitu
de °W
Su
rvey
dat
e D
epth
(m)
% li
ve st
ony
co
ral c
over
(m
ean ±
sd)5
30m
fish
tra
nsec
ts (#
) R
DT
surv
eys
(#)
Fish
spec
ies
(#)
Snap
per H
ole
GC
21
open
w
ind
Hig
h S&
G
19 2
0.63
4'
81 0
4.67
6'
18-J
un-9
9 10
.1
23.8
± 1
0.5
10
3 72
B
reak
ers
GC
30
open
w
ind
S&G
19
17.
507'
81
12.
069'
21
-Jun
-99
12.5
22
.4 ±
4.6
10
2
59
Play
ing
Fiel
ds
GC
31
open
w
ind
Hig
h S&
G
19 1
7.56
5'
81 0
6.31
8'
21-J
un-9
9 7.
5 17
.5 ±
4.7
10
3
71
1 Site
nam
e gi
ven
if it
corr
espo
nds t
o a
know
n C
aym
an Is
land
Dep
artm
ent o
f Env
ironm
ent b
uoy.
2 Pa
rk =
Mar
ine
Park
Are
a; o
pen
= no
pro
tect
ion;
repl
en =
Rep
leni
shm
ent Z
one
Are
a
3 lee
= le
ewar
d; w
ind
= w
indw
ard;
pro
win
d =
prot
ecte
d w
indw
ard
4 Ree
f typ
es fo
llow
Man
frin
o et
al.
(this
vol
ume)
, S&
G =
spur
and
gro
ove,
Hig
h S&
G =
hig
h pr
ofile
spur
and
gro
ove.
5 C
over
val
ues f
rom
ben
thic
AG
RR
A tr
anse
cts (
Man
frin
o et
al.,
this
vol
ume)
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 13 (actual page number in volume to be determined)
Tabl
e 2.
Tw
enty
-fiv
e m
ost f
requ
ently
sigh
ted
fish
spec
ies o
n th
e C
aym
an Is
land
s. D
ata
(Sig
htin
g Fr
eque
ncy
and
Den
sity
Sco
re)
wer
e co
mpi
led
from
the
REE
F da
taba
se, u
sing
exp
ert s
ight
ings
from
199
4 th
roug
h 20
01 (N
=670
RD
T Su
rvey
s).
Scie
ntifi
c na
me
C
omm
on n
ame
Sigh
ting
freq
uenc
y (%
) D
ensi
ty sc
ore1
Acan
thur
us c
oeru
leus
B
lue
Tang
98
2.
8 Th
alas
som
a bi
fasc
iatu
m
Blu
ehea
d 97
3.
3 C
hrom
is c
yane
a B
lue
Chr
omis
97
3.
8 St
egas
tes p
artit
us
Bic
olor
Dam
selfi
sh
96
3.6
Spar
isom
a vi
ride
St
oplig
ht P
arro
tfish
96
2.
7 C
aran
x ru
ber
Bar
Jack
95
2.
3 C
hrom
is m
ultil
inea
ta
Bro
wn
Chr
omis
95
3.
4 G
ram
ma
lore
to
Fairy
Bas
slet
94
3.
6 C
haet
odon
cap
istr
atus
Fo
urey
e B
utte
rfly
fish
94
2.1
Spar
isom
a au
rofr
enat
um
Red
band
Par
rotfi
sh
93
2.7
Hal
icho
eres
gar
noti
Yel
low
head
Wra
sse
92
2.7
Can
thig
aste
r ros
trat
a Sh
arpn
ose
Puff
er
92
2.1
Epin
ephe
lus c
ruen
tatu
s G
rays
by
92
2.1
Lutja
nus a
podu
s Sc
hool
mas
ter
91
2.4
Ocy
urus
chr
ysur
us
Yel
low
tail
Snap
per
89
2.5
Hae
mul
on fl
avol
inea
tum
Fr
ench
Gru
nt
89
2.3
Scar
us c
roic
ensi
s St
riped
Par
rotfi
sh
89
2.4
Scar
us ta
enio
pter
us
Prin
cess
Par
rotfi
sh
88
2.5
Cle
ptic
us p
arra
e C
reol
e W
rass
e 87
3.
7 M
elic
hthy
s nig
er
Bla
ck D
urgo
n 87
2.
8 M
ullo
idic
hthy
s mar
tinic
us
Yel
low
Goa
tfish
84
2.
4 H
olac
anth
us tr
icol
or
Roc
k B
eaut
y 84
1.
9 St
egas
tes d
ienc
aeus
Lo
ngfin
Dam
selfi
sh
82
2.3
Epin
ephe
lus f
ulvu
s C
oney
81
2.
1 St
egas
tes p
lani
fron
s Th
rees
pot D
amse
lfish
80
2.
2 1 Se
e M
etho
ds fo
r def
initi
on o
f Den
sity
Sco
re.
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 14 (actual page number in volume to be determined)
Tabl
e 3.
Mea
n pe
rcen
t sig
htin
g fr
eque
ncy
of se
lect
gro
uper
s dur
ing
AG
RR
A ro
ving
div
er su
rvey
s in
LC a
nd G
C, C
aym
an Is
land
s.
Scie
ntifi
c na
me
Com
mon
nam
e Si
ghtin
g fr
eque
ncy
(%)
LC
GC
M
ycte
rope
rca
bona
ci
Bla
ck G
roup
er
7%
3%
Epin
ephe
lus s
tria
tus
Nas
sau
Gro
uper
69
%
9%
Epin
ephe
lus g
utta
tus
Red
Hin
d 44
%
21%
M
ycte
rope
rca
tigri
s Ti
ger G
roup
er
50%
12
%
Myc
tero
perc
a ve
neno
sa
Yel
low
fin G
roup
er
22%
0%
M
ycte
rope
rca
inte
rstit
ialis
Y
ello
wm
outh
Gro
uper
6%
3%
Ta
ble
4. D
ensi
ty a
nd le
ngth
(mea
n ±
stan
dard
dev
iatio
n) o
f AG
RR
A fi
shes
, and
mac
roal
gal i
ndex
val
ues f
or L
C a
nd G
C, C
aym
an
Isla
nds.
Isla
nd
H
erbi
vore
s
Car
nivo
res
Aca
nthu
ridae
Sc
arid
ae
(≥5
cm)
H
aem
ulid
ae
(≥5
cm)
Lutja
nida
e Se
rran
idae
1 M
acro
alga
l In
dex2
LC d
ensi
ty (#
/100
m2 )
10
.67 ±
7.22
21
.20 ±
7.56
4.59
± 5
.98
3.17
± 3
.61
2.38
± 0
.96
LC le
ngth
(cm
)
11.8
5 ±
2.08
15
.87 ±
2.67
19.4
5 ±
2.58
27
.78 ±
3.42
19
.11 ±
3.02
10
5
GC
den
sity
(#/1
00m
2 )
8.35
± 6
.66
22.1
3 ±
7.04
8.72
± 1
0.04
1.
86 ±
2.0
4 2.
00 ±
0.6
7 G
C le
ngth
(cm
)
12.0
2 ±
1.60
15
.33 ±
2.19
19.2
1 ±
1.36
28
.58 ±
7.29
18
.95 ±
1.92
32
1 Epin
ephe
lus s
pp. a
nd M
ycte
rope
rca
spp.
2 M
acro
alga
l ind
ex =
mac
roal
gal a
bund
ance
x c
anop
y he
ight
; dat
a fr
om M
anfr
ino
et a
l. (th
is v
olum
e).
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 15 (actual page number in volume to be determined)
App
endi
x A
. Cay
man
Isla
nds S
peci
es L
ist.
Dat
a co
mpi
led
from
the
REE
F da
taba
se, u
sing
exp
ert s
ight
ings
from
199
4 th
roug
h 20
01.
A to
tal o
f 670
exp
ert s
urve
ys (3
2- C
aym
an B
rac;
258
- Litt
le C
aym
an; 3
80- G
rand
Cay
man
) rep
orte
d 27
6 sp
ecie
s. F
or e
ach
spec
ies,
perc
ent s
ight
ing
freq
uenc
y (%
SF) a
nd d
ensi
ty sc
ore
(DEN
) are
giv
en.
Fifty
-eig
ht sp
ecie
s pre
viou
sly
unre
porte
d fr
om th
e C
aym
an
Isla
nds a
re li
sted
and
indi
cate
d by
an
aste
risk
(*).
Scie
ntifi
c N
ame
Com
mon
Nam
e SF
%
DEN
Sc
ient
ific
Nam
e C
omm
on N
ame
SF%
D
EN
Aca
nthu
rida
e Su
rgon
fishe
s
B
elon
idae
N
eedl
efis
hes
Acan
thur
us b
ahia
nus
Oce
an S
urge
onfis
h 84
%
2.3
*Pla
ybel
one
arga
lus
Kee
ltail
Nee
dlef
ish
0.1%
2.
0 Ac
anth
urus
chi
rurg
us
Doc
torf
ish
39%
2.
0 Ty
losu
rus c
roco
dilu
s H
ound
fish
3%
1.6
Acan
thur
us c
oeru
leus
B
lue
Tang
98
%
2.9
Ble
nniid
ae
Ble
nnie
s (C
ombt
ooth
)
A
pogo
nida
e C
ardi
nalfi
shes
En
tom
acro
dus n
igri
cans
Pe
arl B
lenn
y 0.
3%
1.0
*Apo
gon
affin
is
Big
toot
h C
ardi
nalfi
sh
0.1%
2.
0 O
phio
blen
nius
atla
ntic
us
Red
lip B
lenn
y 32
%
1.9
*Apo
gon
auro
linea
tus
Brid
le C
ardi
nalfi
sh
0.4%
1.
6 *P
arab
lenn
ius m
arm
oreu
s Se
awee
d B
lenn
y 1%
1.
3 Ap
ogon
bin
otat
us
Bar
red
Car
dina
lfish
17
%
2.0
Bot
hida
e Fl
ound
ers (
Lef
teye
)
Ap
ogon
lach
neri
W
hite
star
Car
dina
lfish
17
%
2.0
Both
us o
cella
tus
Eyed
Flo
unde
r 0.
3%
1.0
Apog
on m
acul
atus
Fl
amef
ish
14%
1.
7 Bo
thus
luna
tus
Peac
ock
Flou
nder
12
%
1.2
*Apo
gon
pilli
onat
us
Bro
adsa
ddle
Car
dina
lfish
SO
1
Cal
liony
mid
ae
Dra
gone
ts
Apog
on p
lani
fron
s Pa
le C
ardi
nalfi
sh
1%
1.8
Para
dipl
ogra
mm
us b
aird
i La
ncer
Dra
gone
t 3%
1.
3 Ap
ogon
pse
udom
acul
atus
Tw
ospo
t Car
dina
lfish
2%
1.
6 C
aran
gida
e Ja
cks
Apog
on q
uadr
isqu
amat
us
Saw
chee
k C
ardi
nalfi
sh
1%
1.8
Alec
tis c
iliar
is
Afr
ican
Pom
pano
SO
1
Apog
on to
wns
endi
B
elte
d C
ardi
nalfi
sh
26%
2.
2 C
aran
x ba
rtho
lom
aei
Yel
low
Jack
3%
1.
6 As
trap
ogon
pun
ctic
ulat
us
Bla
ckfin
Car
dina
lfish
0.
4%
2.0
Car
anx
crys
os
Blu
e R
unne
r 1%
1.
4 Ph
aeop
tyx
pigm
enta
ria
Dus
ky C
ardi
nalfi
sh
3%
2.3
Car
anx
hipp
os
Cre
valle
Jack
2%
1.
9 *P
haeo
ptyx
xen
us
Spon
ge C
ardi
nalfi
sh
11%
1.
6 C
aran
x la
tus
Hor
se-E
ye Ja
ck
33%
2.
3 A
ulos
tom
idae
T
rum
petf
ishe
s
C
aran
x lu
gubr
is
Bla
ck Ja
ck
8%
1.4
Aulo
stom
us m
acul
atus
Tr
umpe
tfish
69
%
1.7
Car
anx
rube
r B
ar Ja
ck
95%
2.
3 B
alis
tidae
L
eath
erja
cket
s
El
agat
is b
ipin
nula
ta
Rai
nbow
Run
ner
0.4%
1.
3 Al
uter
us sc
ript
us
Scra
wle
d Fi
lefis
h 9%
1.
1 Sc
ombe
rom
orus
rega
lis
Cer
o 18
%
1.3
Balis
tes v
etul
a Q
ueen
Trig
gerf
ish
34%
1.
3 Tr
achi
notu
s fal
catu
s Pe
rmit
3%
1.2
Can
ther
hine
s mac
roce
rus
Whi
tesp
otte
d Fi
lefis
h 22
%
1.4
Trac
hino
tus g
oode
i Pa
lom
eta
0.3%
2.
0 C
anth
erhi
nes p
ullu
s O
rang
espo
tted
File
fish
13%
1.
3 C
arch
arhi
nida
e Sh
arks
(Req
ueim
)
C
anth
ider
mis
suffl
amen
O
cean
Trig
gerf
ish
11%
1.
3 *C
arch
arhi
nus l
imba
tus
Bla
cktip
Sha
rk
0.3%
1.
5 M
elic
hthy
s nig
er
Bla
ck D
urgo
n 87
%
2.8
*Car
char
hinu
s per
ezi
Ree
f Sha
rk
2%
1.1
Mon
acan
thus
tuck
eri
Slen
der F
ilefis
h 8%
1.
4 C
haen
opsi
dae
Ble
nnie
s (T
ube)
Ac
anth
embl
emar
ia a
sper
a R
ough
head
Ble
nny
11%
1.
5
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 16 (actual page number in volume to be determined)
App
endi
x A
, Con
tinue
d Sc
ient
ific
Nam
e C
omm
on N
ame
SF%
D
EN
Scie
ntifi
c N
ame
Com
mon
Nam
e SF
%
DEN
C
haen
opsi
dae
(con
t.)
Ble
nnie
s (T
ube)
G
erre
idae
(con
t.)
Moj
arra
*A
cant
hem
blem
aria
cha
plin
i Pa
pillo
se B
lenn
y 0.
1%
1.0
*Euc
inos
tom
us g
ula
Silv
er Je
nny
0.1%
2.
0 Ac
anth
embl
emar
ia m
aria
Se
cret
ary
Ble
nny
16%
1.
6 Eu
cino
stom
us jo
nesi
Sl
ende
r Moj
arra
0.
1%
2.0
Embl
emar
ia p
andi
onis
Sa
ilfin
Ble
nny
14%
1.
7 G
erre
s cin
ereu
s Y
ello
wfin
Moj
arra
11
%
1.9
Embl
emar
iops
is sp
. D
arkh
ead
Ble
nny
4%
1.2
Gob
ieso
cida
e C
lingf
ishe
s
Lu
caya
blen
nius
zing
aro
Arr
ow B
lenn
y 19
%
1.5
Arco
s rub
igin
osus
R
ed C
lingf
ish
1%
1.4
Cha
eton
dont
idae
B
utte
rfly
fishe
s
G
obie
sox
punc
tula
tus
Stip
pled
Clin
gfis
h 0.
1%
1.0
Cha
etod
on a
cule
atus
Lo
ngsn
out B
utte
rfly
fish
26%
1.
5 G
obiid
ae
Gob
ies
Cha
etod
on c
apis
trat
us
Four
eye
But
terf
lyfis
h 94
%
2.1
Cor
ypho
pter
us
Mas
ked/
Gla
ss G
oby
82%
3.
7 C
haet
odon
oce
llatu
s Sp
otfin
But
terf
lyfis
h 38
%
1.7
p
erso
natu
s/hy
alin
us
C
haet
odon
sede
ntar
ius
Ree
f But
terf
lyfis
h 1%
1.
1 C
oryp
hopt
erus
dic
rus
Col
on G
oby
3%
1.2
Cha
etod
on st
riat
us
Ban
ded
But
terf
lyfis
h 73
%
1.8
Cor
ypho
pter
us e
idol
on
Palli
d G
oby
20%
1.
6 C
irrh
itida
e H
awkf
ishe
s
C
oryp
hopt
erus
gla
ucof
raen
um
Brid
led
Gob
y 63
%
2.3
Ambl
ycir
rhitu
s pin
os
Red
spot
ted
Haw
kfis
h 29
%
1.3
Cor
ypho
pter
us li
pern
es
Pepp
erm
int G
oby
46%
1.
9 C
ongr
idae
E
els (
Con
ger)
G
nath
olep
is th
omps
oni
Gol
dspo
t Gob
y 69
%
2.4
Het
eroc
onge
r hal
is
Bro
wn
Gar
den
Eel
22%
3.
5 G
obio
nellu
s sae
pepa
llens
D
ash
Gob
y 1%
1.
4 D
acty
lopt
erid
ae
Flyi
ng G
urna
rds
*Gob
ioso
ma
chan
cei
Shor
tstri
pe G
oby
3%
1.7
Dac
tylo
pter
us v
olita
ns
Flyi
ng G
urna
rd
0.3%
1.
5 G
obio
som
a di
leps
is
Ora
nges
ided
Gob
y 28
%
1.8
Das
yatid
ae
Ray
s (St
ing)
G
obio
som
a ev
elyn
ae
Shar
knos
e G
oby
20%
1.
9 D
asya
tis a
mer
ican
a So
uthe
rn S
tingr
ay
23%
1.
3 G
obio
som
a ge
nie
Cle
anin
g G
oby
49%
2.
4 E
chen
eidi
dae
Rem
oras
G
obio
som
a ho
rsti
Yel
low
line
Gob
y 45
%
1.9
Eche
neis
nau
crat
es
Shar
ksuc
ker
5%
1.1
Gob
ioso
ma
loui
sae
Spot
light
Gob
y 6%
1.
7 E
lopi
dae
Tar
pon
Gob
ioso
ma
mul
tifas
ciat
um
Gre
enba
nded
Gob
y 6%
2.
0 M
egal
ops a
tlant
icus
Ta
rpon
22
%
2.2
Gob
ioso
ma
palle
ns
Sem
isca
led
Gob
y 4%
1.
3 E
phip
pida
e Sp
adef
ishe
s
*G
obio
som
a sa
ucru
m
Leop
ard
Gob
y 0.
3%
1.5
*Cha
etod
ipte
rus f
aber
A
tlant
ic S
pade
fish
1%
1.2
*Gob
ioso
ma
xant
hipr
ora
Yel
low
prow
Gob
y 0.
1%
2.0
Exo
coet
idae
Fl
ying
fishe
s/H
alfb
eeks
*I
oglo
ssus
hel
enae
H
over
ing
Gob
y 2%
1.
4 *H
emir
amph
us b
alao
B
alao
0.
1%
3.0
*Mic
rogo
bius
car
ri
Sem
inol
e G
oby
0.1%
1.
0 *H
irun
dich
thys
spec
ulig
er
Mirr
orw
ing
Flyi
ngfis
h SO
1
*Nes
long
us
Ora
nges
potte
d G
oby
0.3%
1.
5 Fi
stul
ariid
ae
Cor
netf
ishe
s
Pr
iole
pis h
ipol
iti
Rus
ty G
oby
3%
1.2
Fist
ular
ia ta
baca
ria
Blu
espo
tted
Cor
netfi
sh
1%
1.5
Riso
r rub
er
Tusk
ed G
oby
1%
1.5
Ger
reid
ae
Moj
arra
G
ram
mat
idae
B
assl
ets
*Euc
inos
tom
us m
elan
opte
rus
Flag
fin M
ojar
ra
0.3%
2.
0 *G
ram
ma
linki
Y
ello
wch
eek
Bas
slet
0.
1%
1.0
*Euc
inos
tom
us le
froy
i M
ottle
d M
ojar
ra
1%
2.2
Gra
mm
a lo
reto
Fa
iry B
assl
et
94%
3.
6
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 17 (actual page number in volume to be determined)
App
endi
x A
, Con
tinue
d Sc
ient
ific
Nam
e C
omm
on N
ame
SF%
D
EN
Scie
ntifi
c N
ame
Com
mon
Nam
e SF
%
DEN
G
ram
mat
idae
(con
t.)
Bas
slet
s
L
abri
dae
(con
t.)
Wra
sses
s
G
ram
ma
mel
acar
a B
lack
cap
Bas
slet
34
%
3.4
Dor
aton
otus
meg
alep
is
Dw
arf W
rass
e 1%
1.
0 Li
pogr
amm
a tr
iline
atum
Th
reel
ine
Bas
slet
2%
1.
4 H
alic
hoer
es b
ivitt
atus
Sl
ippe
ry D
ick
42%
2.
2 H
aem
ulid
ae
Gru
nts
*Hal
icho
eres
cya
noce
phal
us
Yel
low
chee
k W
rass
e 2%
1.
1 An
isot
rem
us su
rina
men
sis
Bla
ck M
arga
te
3%
1.5
Hal
icho
eres
gar
noti
Yel
low
head
Wra
sse
93%
2.
8 An
isot
rem
us v
irgi
nicu
s Po
rkfis
h 1%
1.
3 H
alic
hoer
es m
acul
ipin
na
Clo
wn
Wra
sse
43%
2.
0 H
aem
ulon
alb
um
Whi
te M
arga
te
17%
1.
5 *H
alic
hoer
es p
ictu
s R
ainb
ow W
rass
e 16
%
2.0
Hae
mul
on a
urol
inea
tum
To
mta
te
6%
2.3
*Hal
icho
eres
poe
yi
Bla
ckea
r Wra
sse
0.1%
2.
0 H
aem
ulon
car
bona
rium
C
aesa
r Gru
nt
26%
2.
3 H
alic
hoer
es ra
diat
us
Pudd
ingw
ife
19%
1.
4 H
aem
ulon
chr
ysar
gyre
um
Smal
lmou
th G
runt
1%
2.
5 *H
emip
tero
notu
s mar
tinic
ensi
s R
osy
Raz
orfis
h 19
%
2.2
Hae
mul
on fl
avol
inea
tum
Fr
ench
Gru
nt
89%
2.
3 H
emip
tero
notu
s spl
ende
ns
Gre
en R
azor
fish
28%
1.
9 *H
aem
ulon
mac
rost
omum
Sp
anis
h G
runt
3%
1.
7 La
chno
laim
us m
axim
us
Hog
fish
44%
1.
3 *H
aem
ulon
mel
anur
um
Cot
tonw
ick
1%
1.7
Thal
asso
ma
bifa
scia
tum
B
lueh
ead
97%
3.
3 H
aem
ulon
par
ra
Sailo
rs C
hoic
e 18
%
2.2
Lab
riso
mid
ae
Ble
nnie
s (Sc
aly)
H
aem
ulon
plu
mie
ri
Whi
te G
runt
69
%
1.8
Acan
them
blem
aria
spin
osa
Spin
yhea
d B
lenn
y 4%
1.
4 H
aem
ulon
sciu
rus
Blu
estri
ped
Gru
nt
65%
2.
0 C
haen
opsi
s lim
baug
hi
Yel
low
face
Pik
eble
nny
1%
1.0
*Hae
mul
on st
riat
um
Strip
ed G
runt
0.
3%
2.0
*Hem
iem
blem
aria
sim
ulus
W
rass
e B
lenn
y 1%
1.
2 H
oloc
entr
idae
Sq
uirr
elfis
hes
*Lab
riso
mus
fila
men
tosu
s Q
uillf
in B
lenn
y 0%
2.
0 H
oloc
entr
us a
dsce
nsio
nis
Squi
rrel
fish
57%
1.
9 La
bris
omus
gob
io
Pale
head
Ble
nny
0.1%
1.
0 H
oloc
entr
us c
orus
cum
R
eef S
quirr
elfis
h 3%
1.
7 *L
abri
som
us k
alis
hera
e D
owny
Ble
nny
0.1%
1.
0 H
oloc
entr
us m
aria
nus
Long
jaw
Squ
irrel
fish
69%
2.
0 M
alac
octe
nus a
urol
inea
tus
Gol
dlin
e B
lenn
y 2%
1.
7 H
oloc
entr
us ru
fus
Long
spin
e Sq
uirr
elfis
h 68
%
2.0
Mal
acoc
tenu
s boe
hlke
i D
iam
ond
Ble
nny
25%
1.
5 H
oloc
entr
us v
exill
ariu
s D
usky
Squ
irrel
fish
19%
1.
9 M
alac
octe
nus m
acro
pus
Ros
y B
lenn
y 7%
1.
5 M
yrip
rist
is ja
cobu
s B
lack
bar S
oldi
erfis
h 54
%
2.0
Mal
acoc
tenu
s tri
angu
latu
s Sa
ddle
d B
lenn
y 75
%
2.3
Plec
tryp
ops r
etro
spin
is
Car
dina
l Sol
dier
fish
2%
1.2
Mal
acoc
tenu
s ver
sico
lor
Bar
fin B
lenn
y 0.
1%
1.0
Iner
miid
ae
Bon
netm
outh
s
St
arks
ia n
anod
es
Dw
arf B
lenn
y 0.
1%
2.0
*Em
mel
icht
hyop
s atla
ntic
us
Bon
netm
outh
0.
4%
3.0
Lut
jani
dae
Snap
pers
*I
nerm
ia v
ittat
a B
oga
17%
3.
0 Ap
silu
s den
tatu
s B
lack
Sna
pper
0.
6%
1.2
Kyp
hosi
dae
Chu
bs
Lutja
nus a
nalis
M
utto
n Sn
appe
r 59
%
1.5
Kyp
hosu
s sec
tatr
ix/in
ciso
r B
erm
uda/
Yel
low
Chu
b 78
%
2.4
Lutja
nus a
podu
s Sc
hool
mas
ter
91%
2.
4 L
abri
dae
Wra
sses
s
Lu
tjanu
s buc
cane
lla
Bla
ckfin
Sna
pper
17
%
2.0
Bodi
anus
pul
chel
lus
Spot
fin H
ogfis
h 1%
1.
5 Lu
tjanu
s cya
nopt
erus
C
uber
a Sn
appe
r 3%
1.
2 Bo
dian
us ru
fus
Span
ish
Hog
fish
74%
1.
9 Lu
tjanu
s gri
seus
G
ray
Snap
per
4%
1.9
Cle
ptic
us p
arra
e C
reol
e W
rass
e 87
%
3.7
Lutja
nus j
ocu
Dog
Sna
pper
10
%
1.3
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 18 (actual page number in volume to be determined)
App
endi
x A
, Con
tinue
d Sc
ient
ific
Nam
e C
omm
on N
ame
SF%
D
EN
Scie
ntifi
c N
ame
Com
mon
Nam
e SF
%
DEN
L
utja
nida
e (c
ont.)
Sn
appe
rs
Pem
pher
idae
Sw
eepe
rs
Lutja
nus m
ahog
oni
Mah
ogan
y Sn
appe
r 69
%
2.2
Pem
pher
is sc
hom
burg
ki
Gla
ssy
Swee
per
2%
2.2
Lutja
nus s
ynag
ris
Lane
Sna
pper
2%
1.
9 Po
mac
anth
idae
A
ngel
fishe
s
O
cyur
us c
hrys
urus
Y
ello
wta
il Sn
appe
r 89
%
2.5
Cen
trop
yge
argi
C
heru
bfis
h 1%
1.
2 M
atac
anth
idae
T
ilefis
hes
*Hol
acan
thus
ber
mud
ensi
s B
lue
Ang
elfis
h 0.
1%
1.0
Mal
acan
thus
plu
mie
ri
Sand
Tile
fish
74%
2.
0 H
olac
anth
us c
iliar
is
Que
en A
ngel
fish
35%
1.
4 M
obul
idae
R
ay (M
anta
s)
Hol
acan
thus
tric
olor
R
ock
Bea
uty
84%
1.
9 M
anta
bir
ostr
is
Man
ta
0.3%
1.
0 Po
mac
anth
us a
rcua
tus
Gra
y A
ngel
fish
39%
1.
4 M
ullid
ae
Goa
tfis
hes
Pom
acan
thus
par
u Fr
ench
Ang
elfis
h 51
%
1.3
Mul
loid
icht
hys m
artin
icus
Y
ello
w G
oatfi
sh
84%
2.
4 Po
mac
entr
idae
D
amse
lfish
es
Pseu
dupe
neus
mac
ulat
us
Spot
ted
Goa
tfish
46
%
1.7
Abud
efdu
f sax
atili
s Se
rgea
nt M
ajor
49
%
2.5
Mur
anid
ae
Eel
s (M
oray
)
Ab
udef
duf t
auru
s N
ight
Ser
gean
t 0.
4%
2.0
Echi
dna
cate
nata
C
hain
Mor
ay
0.1%
1.
0 C
hrom
is c
yane
a B
lue
Chr
omis
97
%
3.8
Ench
elyc
ore
cary
chro
a C
hest
nut M
oray
0.
1%
1.0
*Chr
omis
inso
lata
Su
nshi
nefis
h 18
%
2.2
Gym
noth
orax
fune
bris
G
reen
Mor
ay
5%
1.0
Chr
omis
mul
tilin
eata
B
row
n C
hrom
is
95%
3.
4 G
ymno
thor
ax m
iliar
is
Gol
dent
ail M
oray
4%
1.
0 M
icro
spat
hodo
n ch
rysu
rus
Yel
low
tail
Dam
selfi
sh
57%
2.
0 G
ymno
thor
ax m
orin
ga
Spot
ted
Mor
ay
3%
1.2
Steg
aste
s die
ncae
us
Long
fin D
amse
lfish
82
%
2.3
Gym
noth
orax
vic
inus
Pu
rple
mou
th M
oray
0.
3%
2.0
Steg
aste
s fus
cus
Dus
ky D
amse
lfish
18
%
1.8
Myl
ioba
tidae
R
ays (
Eag
le)
Steg
aste
s leu
cost
ictu
s B
eaug
rego
ry
35%
1.
8 Ae
toba
tus n
arin
ari
Spot
ted
Eagl
e R
ay
9%
1.1
Steg
aste
s par
titus
B
icol
or D
amse
lfish
96
%
3.6
Ogc
ocep
halid
ae
Bat
fishe
s
St
egas
tes p
lani
fron
s Th
rees
pot D
amse
lfish
80
%
2.2
Ogc
ocep
halu
s nas
utus
Sh
ortn
ose
Bat
fish
SO1
St
egas
tes v
aria
bilis
C
ocoa
Dam
selfi
sh
17%
1.
5 O
phic
hthi
dae
Eel
s (Sn
ake)
Pr
iaca
nthi
dae
Big
eyes
M
yric
hthy
s bre
vice
ps
Shar
ptai
l Eel
0.
3%
1.0
Pria
cant
hus a
rena
tus
Big
eye
0.4%
1.
0 O
pist
ogna
thid
ae
Jaw
fishe
s
Pr
iaca
nthu
s cru
enta
tus
Gla
ssey
e Sn
appe
r 9%
1.
3 *O
pist
ogna
thus
aur
ifron
s Y
ello
whe
ad Ja
wfis
h 42
%
2.1
Rhi
ncod
ontid
ae
Shar
ks (C
arpe
t)
*Opi
stog
nath
us m
acro
gnat
hus
Ban
ded
Jaw
fish
0.3%
1.
5 G
ingl
ymos
tom
a ci
rrat
um
Nur
se S
hark
5%
1.
0 *O
pist
ogna
thus
whi
tehu
rsti
Dus
ky Ja
wfis
h 0.
3%
1.5
Scar
idae
Pa
rrot
fishe
s
O
stra
cion
tidae
B
oxfis
hes
Cry
ptot
omus
rose
us
Blu
elip
Par
rotfi
sh
6%
1.9
Lact
ophr
ys b
icau
dalis
Sp
otte
d Tr
unkf
ish
30%
1.
1 Sc
arus
coe
lest
inus
M
idni
ght P
arro
tfish
8%
1.
2 La
ctop
hrys
pol
ygon
ia
Hon
eyco
mb
Cow
fish
26%
1.
2 Sc
arus
coe
rule
us
Blu
e Pa
rrot
fish
2%
1.5
Lact
ophr
ys q
uadr
icor
nis
Scra
wle
d C
owfis
h 1%
1.
6 Sc
arus
cro
icen
sis
Strip
ed P
arro
tfish
89
%
2.4
Lact
ophr
ys tr
igon
us
Trun
kfis
h 0.
4%
1.3
Scar
us g
uaca
mai
a R
ainb
ow P
arro
tfish
15
%
1.3
Lact
ophr
ys tr
ique
ter
Smoo
th T
runk
fish
23%
1.
2 Sc
arus
taen
iopt
erus
Pr
ince
ss P
arro
tfish
88
%
2.5
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 19 (actual page number in volume to be determined)
App
endi
x A
, Con
tinue
d Sc
ient
ific
Nam
e C
omm
on N
ame
SF%
D
EN
Scie
ntifi
c N
ame
Com
mon
Nam
e SF
%
DEN
Sc
arid
ae (c
ont.)
Pa
rrot
fishe
s
Se
rran
idae
(con
t.)
Sea
Bas
ses
Scar
us v
etul
a Q
ueen
Par
rotfi
sh
59%
1.
9 *L
iopr
opom
a ca
rmab
i C
andy
Bas
s 1%
1.
0 *S
pari
som
a at
omar
ium
G
reen
blot
ch P
arro
tfish
41
%
2.2
Liop
ropo
ma
mow
bray
i C
ave
Bas
s 3%
1.
3 Sp
aris
oma
auro
fren
atum
R
edba
nd P
arro
tfish
93
%
2.7
Liop
ropo
ma
rubr
e Pe
pper
min
t Bas
s 19
%
1.4
Spar
isom
a ch
ryso
pter
um
Red
tail
Parr
otfis
h 62
%
2.0
Myc
tero
perc
a bo
naci
B
lack
Gro
uper
11
%
1.2
Spar
isom
a ra
dian
s B
uckt
ooth
Par
rotfi
sh
2%
2.2
Myc
tero
perc
a in
ters
titia
lis
Yel
low
mou
th G
roup
er
6%
1.1
Spar
isom
a ru
brip
inne
Y
ello
wta
il Pa
rrot
fish
46%
2.
0 *M
ycte
rope
rca
phen
ax
Scam
p 1%
1.
2 Sp
aris
oma
viri
de
Stop
light
Par
rotfi
sh
96%
2.
7 M
ycte
rope
rca
tigri
s Ti
ger G
roup
er
54%
1.
5 Sc
iaen
idae
D
rum
s
M
ycte
rope
rca
vene
nosa
Y
ello
wfin
Gro
uper
18
%
1.3
Eque
tus a
cum
inat
us
Hig
hhat
1%
1.
5 *P
aran
thia
s fur
cife
r C
reol
e-fis
h 1%
2.
0 Eq
uetu
s lan
ceol
atus
Ja
ckni
fe F
ish
0.1%
1.
0 Ry
ptic
us sa
pona
ceus
G
reat
er S
oapf
ish
18%
1.
2 Eq
uetu
s pun
ctat
us
Spot
ted
Dru
m
15%
1.
2 *S
erra
nus b
aldw
ini
Lant
ern
Bas
s 11
%
1.5
Odo
ntos
cion
den
tex
Ree
f Cro
aker
0.
4%
1.6
Serr
anus
taba
cari
us
Toba
ccof
ish
53%
1.
8 Sc
orpi
onid
ae
Scor
pion
fishe
s
Se
rran
us ti
grin
us
Har
lequ
in B
ass
75%
1.
9 Sc
orpa
ena
plum
ieri
Sp
otte
d Sc
orpi
onfis
h 6%
1.
1 *S
erra
nus t
ortu
garu
m
Cha
lk B
ass
7%
1.7
Scor
paen
odes
car
ibba
eus
Ree
f Sco
rpio
nfis
h 1%
1.
0 Sp
arid
ae
Porg
ies
Serr
anid
ae
Sea
Bas
ses
Cal
amus
baj
onad
o Jo
lthea
d Po
rgy
15%
1.
2 Ep
inep
helu
s ads
cens
ioni
s R
ock
Hin
d 2%
1.
1 C
alam
us c
alam
us
Sauc
erey
e Po
rgy
27%
1.
3 Ep
inep
helu
s cru
enta
tus
Gra
ysby
92
%
2.1
*Cal
amus
pen
natu
la
Plum
a 1%
1.
1 Ep
inep
helu
s ful
vus
Con
ey
81%
2.
1 Sp
hyra
enid
ae
Bar
racu
das
Epin
ephe
lus g
utta
tus
Red
Hin
d 27
%
1.3
Sphy
raen
a ba
rrac
uda
Gre
at B
arra
cuda
49
%
1.5
Epin
ephe
lus i
taja
ra
Gol
iath
Gro
uper
0.
6%
1.2
Sphy
raen
a pi
cudi
lla
Sout
hern
Sen
net
1%
3.5
*Epi
neph
elus
mor
io
Red
Gro
uper
0.
3%
1.5
Sphy
rnid
ae
Shar
ks (H
amm
erhe
ad)
Epin
ephe
lus s
tria
tus
Nas
sau
Gro
uper
50
%
1.5
*Sph
yrna
lew
ini
Scal
lope
d H
amm
erhe
ad
SO1
H
ypop
lect
rus a
berr
ans
Yel
low
belly
Ham
let
4%
1.2
Syng
nath
idae
Pi
pefis
hes/
Seah
orse
s
*H
ypop
lect
rus c
hlor
urus
Y
ello
wta
il H
amle
t 0.
3%
1.0
*Ace
ntro
nura
den
driti
ca
Pipe
hors
e 0.
4%
1.6
Hyp
ople
ctru
s gum
mig
utta
G
olde
n H
amle
t 0.
4%
1.0
*Cos
moc
ampu
s alb
iros
tris
W
hite
nose
Pip
efis
h 0.
1%
1.0
Hyp
ople
ctru
s gut
tava
rius
Sh
y H
amle
t 21
%
1.3
*Hip
poca
mpu
s ere
ctus
Li
ned
Seah
orse
0.
4%
1.0
Hyp
ople
ctru
s ind
igo
Indi
go H
amle
t 9%
1.
3 Sy
nodo
tidae
L
izza
rdfis
hes
Hyp
ople
ctru
s nig
rica
ns
Bla
ck H
amle
t 14
%
1.4
Syno
dus i
nter
med
ius
Sand
Div
er
5%
1.1
Hyp
ople
ctru
s pue
lla
Bar
red
Ham
let
64%
1.
8 *S
ynod
us sa
urus
B
lues
tripe
d Li
zard
fish
0.3%
1.
0 *H
ypop
lect
rus s
p.
Mas
ked
Ham
let
2%
1.0
Syno
dus s
ynod
us
Red
Liz
ardf
ish
0.4%
1.
0 *H
ypop
lect
rus s
p.
Tan
Ham
let
2%
1.0
Tet
rado
ntid
ae
Puff
ers
Hyp
ople
ctru
s uni
colo
r B
utte
r Ham
let
19%
1.
3 C
anth
igas
ter r
ostr
ata
Shar
pnos
e Pu
ffer
92
%
2.1
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 20 (actual page number in volume to be determined)
App
endi
x A
, Con
tinue
d Sc
ient
ific
Nam
e C
omm
on N
ame
SF%
D
EN
Scie
ntifi
c N
ame
Com
mon
Nam
e SF
%
DEN
T
etra
dont
idae
(con
t.)
Puff
ers
Tri
pter
ygiid
ae
Ble
nnie
s (T
ripp
lefin
)
*C
hilo
myc
teru
s ant
enna
tus
Brid
led
Bur
rfis
h 0.
3%
1.0
Enne
anec
tes a
ltive
lis
Lofty
Trip
lefin
2%
1.
0 D
iodo
n ho
loca
nthu
s B
allo
onfis
h 4%
1.
0 En
nean
ecte
s atr
orus
B
lack
edge
Trip
lefin
0.
1%
1.0
*Dio
don
hyst
rix
Porc
upin
efis
h 13
%
1.1
Enne
anec
tes b
oehl
kei
Rou
ghhe
ad T
riple
fin
2%
1.4
Spho
eroi
des s
peng
leri
B
andt
ail P
uffe
r 5%
1.
2 En
nean
ecte
s pec
tora
lis
Red
eye
Trip
lefin
3%
1.
1 T
orpe
dini
dae
Ray
s (T
orpe
do E
lect
ric)
U
rolo
phid
ae
Ray
s (R
ound
)
*N
arci
ne b
rasi
liens
is
Less
er E
lect
ric R
ay
0.1%
2.
0 U
rolo
phus
jam
aice
nsis
Y
ello
w S
tingr
ay
6%
1.2
Pattengill-Semmens and Semmens, Atol Research Bulletin, 2002 p. 21 (actual page number in volume to be determined)