# Biodiversity Assessment of the Fishes of Saba Bank Atoll, Netherlands Antilles

Two formal corrections to the article have been published:
* The species of the fish in figure 68 is given as Priacanthus arenatus; the species is actually Heteropriacanthus cruentatus.
* The species of the fish in figure 124 is given as Xyrichtys splendens; the species is actually Doratonotus megalepis.
​Biodiversity Assessment of the Fishes of Saba Bank Atoll, Netherlands Antilles
Jeffrey T. Williams; Kent E. Carpenter; James L. Van Tassell; Paul Hoetjes; Wes Toller; Peter Etnoyer; Michael Smith
* ↑ Fish Division, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Suitland, Maryland, United States of America
* ↑ Department of Biological Sciences, Old Dominion University, Norfolk, Virginia, United States of America
* ↑ Department of Ichthyology, American Museum of Natural History, New York, New York, United States of America
* ↑ Department of Environment and Nature, Ministry of Public Health and Social Development, Curaçao, Netherlands Antilles
* ↑ Saba Conservation Foundation, Fort Bay, Saba, Netherlands Antilles
* ↑ Harte Research Institute, Texas A&M University-Corpus Christi, Corpus Christi, Texas, United States of America
* ↑ Center for Applied Biodiversity Science, Conservation International, Arlington, Virginia, United States of America
Abstract
Biodiversity surveys were conducted on Saba Bank, Netherlands Antilles, to assess ichthyofaunal richness and to compare with published surveys of other Caribbean localities. The primary objective was to estimate the total species richness of the Saba Bank ichthyofauna. A variety of sampling techniques was utilized to survey the fish species of both the visually accessible megafauna and the camouflaged and small-sized species comprising the cryptic ichthyofauna. Based on results presented herein, the number of species known on Saba Bank is increased from 42 previously known species to 270 species. Expected species-accumulation curves demonstrate that the current estimate of species richness of fishes for Saba Bank under represents the actual richness, and our knowledge of the ichthyofauna has not plateaued. The total expected fish-species richness may be somewhere between 320 and 411 species. The Saba Bank ichthyofaunal assemblage is compared to fish assemblages found elsewhere in the Caribbean. Despite the absence of shallow or emergent shore habitats like mangroves, Saba Bank ranks as having the eighth highest ichthyofaunal richness of surveyed localities in the Greater Caribbean. Some degree of habitat heterogeneity was evident. Fore-reef, patch-reef, and lagoonal habitats were sampled. Fish assemblages were significantly different between habitats. Species richness was highest on the fore reef, but 11 species were found only at lagoonal sites. A comprehensive, annotated list of the fishes currently known to occur on Saba Bank, Netherland Antilles, is provided and color photographs of freshly collected specimens are presented for 165 of the listed species of Saba Bank fishes to facilitate identification and taxonomic comparison with similar taxa at other localities. Coloration of some species is shown for the first time. Preliminary analysis indicates that at least six undescribed new species were collected during the survey and these are indicated in the annotated list.
Citation: Williams JT, Carpenter KE, Van Tassell JL, Hoetjes P, Toller W, Etnoyer P, Smith M. (2010) Biodiversity Assessment of the Fishes of Saba Bank Atoll, Netherlands Antilles. PLoS ONE 5(5): e10676. https://doi.org/10.1371/journal.pone.0010676
Editor: Brian Gratwicke, Smithsonian's National Zoological Park, United States of America
Received: November 13, 2009; Accepted: January 21, 2010; Published: May 21, 2010
This is an open-access article distributed under the terms of the Creative Commons Public Domain declaration which stipulates that, once placed in the public domain, this work may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose.
Funding: The Department of Environment and Nature of the Netherlands Antilles and Conservation International provided funding for this project. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: Dr. Peter Etnoyer co-authored the Deep Sea News blog with PLoS One editor Dr. Craig McClain 2005–2009. Dr. McClain has recused himself from the Saba Bank Biodiversity Assessment volume for this reason.
* ↑ Current address: MSC Accreditation Program, Accreditation Services International GmbH, Bonn, Germany
* ↑ Current address: Center for Coastal Environmental Health and Biomolecular Research, National Oceanic and Atmospheric Administration, Charleston, South Carolina, United States of America
Introduction
Saba Bank is the largest atoll in the Atlantic Ocean Basin and one of the three largest atolls on earth $^1$. Located in the Dutch Windward Islands about 250 km east of Puerto Rico, it is a flat-topped seamount rising 1800 m from the surrounding sea floor. Except for the fact that it does not break the water surface, Saba Bank is a classic atoll consisting of a submerged mountain crowned at the summit with a ring of actively growing coral reefs $^2$. Saba Bank is relatively free of the problems that are degrading many Caribbean reef systems, and the few problems it faces include anchoring and abrasion by oil tankers maneuvering off the petroleum transshipment facilities on St. Eustatius, potential petroleum spillage and subsequent use of dispersants, general vessel passage in a zone of high maritime traffic, possible overfishing for certain species, and exploration for petroleum reserves (so far unsuccessful). Saba Bank's fisheries and dive operations are economically significant to the small community on Saba Island (about 1500 residents) that has direct responsibility for its management.
The known fish fauna of Saba Bank prior to our survey consisted of 42 fish species. Most of these species were taken during fishery bottom-trawl surveys on Saba Bank, including two M/V Oregon stations in 1958 and nine stations in 1959, and two trawl hauls taken in 1969 by the R/V Pillsbury. Although four of these trawls were taken on or near the top of Saba Bank, nine were on the deep outer slopes. The habitats sampled during these surveys were restricted to relatively soft-bottom habitats due to the exclusive use of trawling techniques. These trawl samples provided valuable records of fishes living on soft bottoms and on the outer slopes of Saba Bank.
​A biodiversity-assessment survey was carried out on Saba Bank during 2006 and 2007, with a major goal being to improve knowledge of the biodiversity on one of the world's most significant, though poorly known, coral-capped seamounts. In an effort to record as many fish species as possible in the short period of time available for the survey, we utilized a variety of fish sampling techniques. These techniques included visual surveys by divers, use of SCUBA to apply ichthyocide (a natural fish toxicant consisting of dried and powdered Derris root - assayed at 7.5% rotenone), hand-line fishing, by-catch from lobster and fish traps taken by local fishermen, and port sampling observations of fish landings. During the surveys, an attempt was made to obtain a photograph documenting the fresh colors of as many species as possible, a tissue sample of each species, and preserved specimen vouchers that have been archived in the National fish collection (USNM) of the National Museum of Natural History (NMNH), Smithsonian Institution (SI). Specimens representing six or more undescribed species and two rare gobies, Pycnomma roosevelti and Psilotris boehlkei, were collected.
The habitats surveyed on the Saba Bank “atoll” during this study are classified as: fore reef, patch reef, lagoonal and Small Bank. Fore-reef areas are located around the outer rim of the submerged atoll. Patch reef is an isolated portion of “reef” situated on the lagoonal (interior) side of the fore reef. Lagoonal is the central portion of the atoll interior to the fore-reefs around the atoll rim and may have a variety of bottom types. Small Bank is a small, independent “seamount-like” structure located off the northwest corner of Saba Bank with its shallowest depth at about 11 m. Depth categories were arbitrarily assigned as shallow (11–24 m), mid-depth (25–34 m), and deep (35–38 m).
The primary goals of the overall biodiversity survey were to provide data and analysis to support designation of Saba Bank as a marine protected area, to support the development of a management plan, and to contribute to a petition to the International Maritime Organization to designate appropriate parts of Saba Bank as a Particularly Sensitive Sea Area.
Results
Specimens collected during the RAP survey were preserved as vouchers and processed into the fish collection (available online at http://vertebrates.si.edu/fishes/fishes_collections.html) at the NMNH (as many species as possible were also tissue sampled ​and photographed in the field). Toller collected and preserved vouchers from fishery landings when possible and photographed specimens representing new records. Toller's vouchers were processed into the fish collection at the NMNH and are also available in the online database.
Some species were only taken during fishery bottom trawl surveys by the M/V Oregon (1958, 1959) and the R/V Pillsbury (1969) on Saba Bank. Species represented by voucher specimens in museum collections are included in our comprehensive species list.
There were significant differences in the fish assemblages when classified by depth - shallow, middle, and deep (ANOSIM Global test, R = 0.618, P = 0.006). Differences in species composition were most evident between mid-depth and deep sites (ANOSIM Pairwise test, R = 0.829, P = 0.008). Fore-reef sites were typically at mid to shallow depths (20 to 34 m) whereas the patch-reef sites were typically deep (35 to 38 m). There is some evidence of habitat heterogeneity and vertical zonation for fish assemblages on Saba Bank, but more sampling is necessary to discern whether habitat or depth best explains the differences among groups.
The horizontal axis in the MDS plot represents a gradient in species richness with patch-reef and lagoonal sites having lowest richness ( Figure 2). The vertical axis in the MDS plot illustrates differences related to depth with deepest station (Small Bank) higher in the vertical axis, and shallower sites lower in the vertical axis. The two-dimensional stress value was low in the MDS (stress = 0.1) indicating a slight chance of misrepresentation.
A principal components analysis (PCA) was used to understand which species assemblages were responsible for differences among stations. The first three components explained 43.9% of the variation. The first component (18.6% of variation explained) was dominated by ubiquitous species and most common species. Strongest loadings (negative) on the first principal component (PC1) included the four species found at all stations (Halichoeres garnoti, Serranus tigrinus, Stegastes partitus, and Thalassoma bifasciatum) and 13 species found at all but one to four of the stations. These 13 common species are rarely found at the small bank, patch-reef, and lagoonal stations and, therefore, PC1 also serves to define the fore-reef sites. For example, Acanthurus bahianus, Coryphopterus glaucofraenum, Coryphopterus dicrus, and Scarus taeniopterus were found at all fore-reef sites but rarely at the Small Bank, lagoonal and at least one of the patch-reef sites. The strongest positive loadings on PC1 are from 11 species only found at the lagoonal site. Principal components 2 and 3 correspond strongly to the horizontal or species richness component on the MDS with strongest positive ​loadings on species typically found at the low to medium species richness stations 3, 5, 6 and 8–12 (e.g. Cryptotomus roseus, Haemulon melanurum, Halichoeres bivittatus, Astrapogon puncticulatus, and Serranus baldwini). Strongest negative scores were on species that were frequently found at high to medium species richness stations 1, 2, 4, 7 and 8 (e.g. Hypoplectrus puella, Lythrypnus elasson, Prognathodes aculeatus, Neoniphon marianus, and Gramma loreto).
An annotated list of the fishes of Saba Bank is provided below. In the list, we include the family, genus and species, author and English common name (as common names are not standardized internationally, we strove to apply the most widely used English common name based on FishBase (http://www.fishbase.org) listings). The use of “cf” before a species name indicates that the specimen photographed is similar to that species, but probably represents an undescribed species. Voucher specimens are archived at the National Museum of Natural History (USNM) and the Florida Museum of Natural History (UF) and each species with vouchers is annotated with the museum's acronym where the specimens are housed. The basis of each species record is indicated by: I – ichthyocide station, F – caught by a local fisherman and photographed, T – bottom trawl, O – visual sighting during Toller survey, V – visual sighting during RAP survey at roving and rotenone station. Lengths of specimens are recorded in mm for either standard length (SL), total length (TL), or fork length (FL). Photographs showing the color pattern of freshly collected specimens are included for as many of the species as possible. Images illustrating observed sexual and developmental (juvenile to adult) variability in color pattern are included where possible.
Ginglymostomatidae—nurse sharks
Ginglymostoma cirratum (Bonnaterre, 1788)—nurse shark; OV; Figure 3
Squalidae—dogfish sharks
Squalus cubensis Howell Rivero, 1936—Cuban dogfish; F; Figure 4
Carcharhinidae—requiem sharks
Carcharhinus perezii (Poey, 1876)—reef shark; F
Galeocerdo cuvier (Péron & Lesueur, 1822)—tiger shark; F,O
Etmopteridae—lantern sharks
Etmopterus bullisi Bigelow & Schroeder, 1957—lined lantern shark; USNM, T
Dasyatidae—whiptail stingrays
Dasyatis americana Hildebrand & Schroeder, 1928—southern stingray; V
Muraenidae—morays
Anarchias similis (Lea, 1913)—pygmy moray; USNM, I
Enchelycore carychroa (Böhlke & Böhlke, 1976)—chestnut moray; USNM, I, F;Figure 5
Enchelycore nigricans (Bonnaterre, 1788)—viper moray; USNM, I
Gymnothorax conspersus Poey, 1867—saddled moray; USNM, F; Figure 6
Gymnothorax maderensis(Johnson, 1862)—sharktooth moray; USNM, F; Figure 7
Gymnothorax miliaris (Kaup, 1856)—goldentail moray; USNM, I, O; Figure 8
Gymnothorax moringa (Cuvier, 1829)—spotted moray; USNM, I, O, V; Figure 9
Gymnothorax polygonius Poey, 1876—polygon moray; USNM, F; Figure 10
Gymnothorax vicinus (Castelnau, 1855)—purplemouth moray; USNM, I, O; Figure 11
Monopenchelys acuta (Parr, 1930)—redface moray; USNM, I; Figure 12
Uropterygius macularius (Lesueur, 1825)—marbled moray; USNM, I
Ophichthidae—snake eels
Ahlia egmontis (Jordan, 1884)—key worm eel; USNM, I; Figures 13, 14
Aprognathodon platyventris Böhlke, 1967—stripe eel; USNM, I; Figures 15, 16
Myrichthys breviceps (Richardson, 1848) —sharptail eel; O
Myrichthys ocellatus (Lesueur, 1825)—goldspotted eel; USNM, I; Figures 17, 18
Congridae—conger eels
Bathycongrus thysanochilus (Reid, 1934)—conger eel; USNM, T
Conger esculentus Poey, 1866—grey conger; USNM, F
Chlopsidae—false morays
Kaupichthys hyoproroides (Strömman, 1896)—false moray; USNM, I; Figure 19
Kaupichthys nuchalis Böhlke, 1967—collared eel; USNM, I; Figure 20
Moringuidae—spaghetti eels
Moringua edwardsi (Jordan & Bollman, 1889)—spaghetti eel; USNM, I; Figure 21
Synodontidae—lizardfishes
Saurida brasiliensis Norman 1935—largescale lizardfish; UF, T
Saurida normani Longley 1935—shortjaw lizardfish; UF, T
Synodus intermedius (Spix & Agassiz, 1829)—sand diver; USNM, UF, F, O; Figure 22
Synodus poeyi Jordan, 1887—offshore lizardfish; UF, T
Synodus saurus (Linnaeus, 1758)—Atlantic lizardfish; UF, I, V
Synodus synodus (Linnaeus, 1758)—red lizardfish; UF, USNM, I;Figure 23
Trachinocephalus myops (Forster, 1801)—snakefish; V
Ophidiidae—cusk-eels
Brotula barbata (Bloch & Schneider, 1801)—bearded brotula; USNM, O; Figure 24
Neobythites ocellatus Günther 1887—ocellate cusk-eel; USNM, T
Neobythites unicolor Nielsen & Retzer 1994—unicolor cusk-eel; USNM, T
Ophidion antipholus Lea & Robins, 2003—longnose cusk-eel; UF, T
Otophidium omostigma (Jordan & Gilbert, 1882)—polka-dot cusk-eel; UF, T
Parophidion schmidti (Woods & Kanazawa, 1951)—dusky cusk-eel; USNM, I; Figure 25
Petrotyx sanguineus (Meek & Hildebrand, 1928)—redfin brotula; USNM, I; Figure 26
Bythitidae—viviparous brotulas
Ogilbia sabaji Moller, Schwarzhans & Nielsen, 2005—Sabaj coralbrotula; USNM, I; Figure 27
Antennariidae—frogfishes
Antennarius pauciradiatus Schultz, 1957 —dwarf frogfish; USNM, I; Figures 28, 29
Antennarius multiocellatus (Valenciennes, 1837) —longlure frogfish; USNM, I
A single juvenile specimen was collected in 2007. Although adults of this species have a very long first spine, our juvenile specimen has the first dorsal spine about the same length as the second. Böhlke and Chaplin $^3$ mention this allometric growth pattern in which the first spine is short in young specimens, but increases in length with growth. The juvenile exhibits the typical adult color pattern.
Chaunacidae—sea toads
Chaunax suttkusi Caruso, 1989—Suttkus sea toad; USNM, T
Ogcocephalidae—batfishes
All of the batfish records from Saba Bank are based on trawl collections with vouchered museum specimens.
Dibranchus atlanticus Peters, 1876—Atlantic batfish; UF, T
Halieutichthys aculeatus (Mitchill, 1818)—pancake batfish; UF, T
Ogcocephalus pumilus Bradbury, 1980—dwarf batfish; USNM, T
Exocoetidae—flyingfishes
Cypselurus comatus (Mitchill, 1815)—clearwing flyingfish; USNM
This specimen was probably captured at the surface using a dip net.
Syngnathidae—pipefishes
Anarchopterus tectus (Dawson, 1978) —insular pipefish; USNM, I
Bryx randalli (Herald, 1965)—ocellated pipefish; USNM, I; Figure 30
Micrognathus crinitus (Jenyns, 1842) —banded pipefish; USNM, I; Figures 31, 32
Aulostomidae—trumpetfishes
Aulostomus maculatus Valenciennes, 1837—trumpetfish; USNM, I, O, V
Holocentridae—squirrelfishes
Holocentrus adscensionis (Osbeck, 1765)—squirrelfish; USNM, I, O, V; Figure 33
Holocentrus rufus (Walbaum, 1792)—longspine squirrelfish; USNM, I, O, V; Figure 34
Myripristis jacobus Cuvier, 1829—blackbar soldierfish; USNM, I, O; Figure 35
Neoniphon marianus (Cuvier, 1829)—longjaw squirrelfish; USNM, I; Figure 36
Plectrypops retrospinis (Guichenot, 1853)—cardinal soldierfish; USNM, I; Figure 37
Sargocentron coruscum (Poey, 1860)—reef squirrelfish; USNM, I, O; Figures 38, 39
Scorpaenidae—scorpionfishes
Scorpaena albifimbria Evermann & Marsh, 1900—coral scorpionfish; USNM, I; Figure 40
Scorpaena bergii Evermann & Marsh, 1900—goosehead scorpionfish; USNM, I
Scorpaena grandicornis Cuvier, 1829—plumed scorpionfish; USNM, I; Figure 41
Scorpaena inermis Cuvier, 1829 —mushroom scorpionfish; USNM, I, T; Figures 42, 43
Two color morphs were collected at Saba Bank, a red morph (Figure 42) and a yellow morph (Figure 43).
Scorpaena plumieri Bloch, 1789—spotted scorpionfish; USNM, I
Scorpaenodes caribbaeus Meek & Hildebrand, 1928—reef scorpionfish; USNM, I; Figure 44
Triglidae—searobins
All of the searobin records from Saba Bank are based on trawl collections with vouchered museum specimens.
Bellator egretta (Goode & Bean, 1896)—streamer searobin; USNM, T
Bellator militaris (Goode & Bean, 1896)—horned searobin; UF, T
Prionotus ophryas Jordan & Swain, 1885—bandtail searobin; UF, T
Symphysanodontidae—slopefishes
Symphysanodon berryi Anderson, 1970—slope bass; USNM, F; Figure 45
Serranidae—sea basses
Alphestes afer (Bloch, 1793)—mutton hamlet; F, O; Figure 46
Cephalopholis cruentata (Lacepède, 1802)—graysby; USNM, I, F, O, V; Figure 47
Cephalopholis fulva (Linnaeus, 1758)—coney; UF,'USNM, I, F, O, V'; Figure 48
Diplectrum bivittatum (Valenciennes, 1828) —dwarf sand perch; UF, T
Epinephelus flavolimbatus Poey, 1865—yellowedge grouper; F; Figure 49
Epinephelus guttatus (Linnaeus, 1758)—red hind; UF, USNM, F, I, O, T, V; Figure 50
Epinephelus morio (Valenciennes, 1828)—red grouper; F
Epinephelus niveatus Valenciennes, 1828—snowy grouper; USNM, F; Figure 51
Epinephelus striatus (Bloch, 1792)—Nassau grouper; F
Hypoplectrus chlorurus (Cuvier, 1828) —yellowtail hamlet; O
Hypoplectrus nigricans (Poey, 1852)—black hamlet; USNM, I, O, V; Figure 52
Hypoplectrus puella (Cuvier, 1828)—barred hamlet; O, V
Liopropoma rubre Poey, 1861—peppermint basslet; USNM, I; Figure 53
Mycteroperca interstitialis (Poey, 1860)—yellowmouth grouper; F; Figure 54
Mycteroperca tigris (Valenciennes, 1833)—tiger grouper; V
Mycteroperca venenosa (Linnaeus, 1758)—yellowfin grouper; F, OBS
Paranthias furcifer (Valenciennes, 1828)—Atlantic creolefish; UF, USNM, I, O, V; Figure 55
Pseudogramma gregoryi (Breder, 1927)—reef bass; USNM, I; Figure 56
Rypticus bistripinus (Mitchill, 1818) —freckled soapfish; USNM, I; Figure 57
Rypticus saponaceus (Bloch & Schneider, 1801)—greater soapfish; USNM, I; Figure 58
Rypticus new species; USNM, I
This new species of soapfish is very similar in appearance to Rypticus subbifrenatus. The new species is being described by C Baldwin and DG Smith (pers. comm.).
Rypticus subbifrenatus Gill, 1861—spotted soapfish; USNM, I; Figure 59
Schultzea beta (Hildebrand, 1940)—school bass; UF, USNM, I
Serranus baldwini (Evermann & Marsh, 1899)—lantern bass; UF, USNM, I, O, V; Figure 60
Serranus maytagi Robins & Starck, 1961 —maytag bass; UF
Serranus notospilus Longley, 1935—saddle bass; USNM, F; Figure 61
Serranus tabacarius (Cuvier, 1829)—tobaccofish; UF, O, V
Serranus tigrinus (Bloch, 1790)—harlequin bass; USNM, I, O, V; Figure 62
Serranus tortugarum Longley, 1935—chalk bass; UF, USNM, I, O; Figure 63
Grammatidae—basslets
Gramma loreto Poey, 1868—fairy basslet; USNM, I, O, V; Figure 64
Opistognathidae—jawfishes
Opistognathus aurifrons (Jordan & Thompson, 1905)—yellowhead jawfish; USNM, I, O, V; Figure 65
Opistognathus whitehursti (Longley, 1927)—dusky jawfish; USNM, I; Figures 66, 67
Priacanthidae—bigeyes
Heteropriacanthus cruentatus (Lacepède, 1801)—glasseye snapper; O
Priacanthus arenatus Cuvier, 1829—bigeye; USNM, I; Figure 68
Apogonidae—cardinalfishes
Apogon aurolineatus (Mowbray in Breder, 1927)—barred cardinalfish; USNM, I
Apogon binotatus (Poey, 1867)—barred cardinalfish; USNM, I
Apogon maculatus (Poey, 1860)—flamefish; USNM, I; Figure 69
Apogon pillionatus Böhlke and Randall, 1968 —broadsaddle cardinalfish; UF, USNM, I
Apogon cf quadrisquamatus Longley, 1934 —sawcheek cardinalfish; USNM, I; Figure 70
The specimens identified here as A.cf quadrisquamatus have been found to represent a new undescribed species closely related to A. quadrisquamatus (C. Baldwin & D.G. Smith, pers. comm. 2009).
Apogon robinsi Böhlke & Randall, 1968 —roughlip cardinalfish; USNM, I; Figure 71
Apogon townsendi (Breder, 1927)—belted cardinalfish; USNM, I; Figure 72
Astrapogon puncticulatus (Poey, 1867) —blackfin cardinalfish; USNM, I; Figure 73
Phaeoptyx conklini (Silvester, 1916)—freckled cardinalfish; USNM, I; Figure 74
Phaeoptyx pigmentaria (Poey, 1860)—dusky cardinalfish; USNM, I
Malacanthidae—tilefishes
Caulolatilus cyanops Poey, 1866—blackline tilefish; USNM, F; Figure 75
Malacanthus plumieri (Bloch, 1786)—sand tilefish; UF, O, V
Coryphaenidae-dolphinfishes
Coryphaena hippurus Linnaeus, 1758—common dolphinfish; F
Rachycentridae-cobias
Rachycentron canadum (Linnaeus, 1766)—cobia
Our record for this species is based on an underwater video recently filmed by Yap Films Inc (Toronto, Canada) at the shipwreck on Saba Bank. The video clearly shows a cobia swimming along the side of the shipwreck.
Carangidae—jacks
Alectis ciliaris (Bloch, 1787)—African pompano; O; Figure 76
Caranx bartholomaei Cuvier, 1833—yellow jack; O, V
Caranx crysos (Mitchill, 1815) —blue runner; O
Caranx latus Agassiz, 1831—horse-eye jack; O, V
Caranx lugubris Poey, 1860—black jack; O, V
Caranx ruber (Bloch, 1793)—bar jack; UF, F, O, V; Figure 77
Decapterus macarellus (Cuvier, 1833)—mackerel scad; O; Figure 78
Elagatis bipinnulata (Quoy & Gaimard, 1825)—rainbow runner; F, OBS
Selar crumenophthalmus (Bloch, 1793)—bigeye scad; F; Figure 79
Seriola dumerili (Risso, 1810)—greater amberjack; F; Figure 80
Seriola rivoliana Valenciennes, 1833—almaco jack; F; Figure 80
Lutjanidae—snappers
Apsilus dentatus Guichenot, 1853—black snapper; USNM, F; Figure 81
Etelis oculatus (Valenciennes, 1828) —queen snapper; F
Lutjanus apodus (Walbaum, 1792)—schoolmaster; O, VIS
Lutjanus buccanella (Cuvier, 1828)—blackfin snapper; USNM, F, O; Figure 82
Lutjanus cyanopterus (Cuvier, 1828)—cubera snapper; USNM, I
Lutjanus mahogoni (Cuvier, 1828) —mahogany snapper; O
Lutjanus purpureus (Poey, 1866)—Caribbean red snapper; USNM, F; Figure 83
Lutjanus synagris (Linnaeus, 1758)—lane snapper; USNM, F; Figure 84
Lutjanus vivanus (Cuvier, 1828)—silk snapper; USNM, F; Figure 85
Ocyurus chrysurus (Bloch, 1791)—yellowtail snapper; UF, O, V
Pristipomoides aquilonaris (Goode & Bean, 1896)—wenchman; USNM, F; Figure 86
Rhomboplites aurorubens (Cuvier, 1829) —vermilion snapper; USNM, F; Figure 87
Lobotidae—tripletails
Lobotes surinamensis (Bloch, 1790)—Atlantic tripletail; F; Figure 88
Haemulidae—grunts
Haemulon album Cuvier, 1830—margate; F; Figure 89
Haemulon aurolineatum Cuvier, 1830—tomtate; F, O; Figure 90
Haemulon carbonarium Poey, 1860—caesar grunt; F; Figure 91
Haemulon flavolineatum (Desmarest, 1823)—French grunt; O, V
Haemulon melanurum (Linnaeus, 1758)—cottonwick; UF, USNM, F, I, O, V; Figure 92
Haemulon plumierii (Lacepède, 1801)—white grunt; USNM, F, I, O, V; Figure 93
Haemulon striatum (Linnaeus, 1758)—striped grunt; F; Figure 94
Inermiidae—bonnetmouths
Inermia vittata Poey, 1860—boga; O
Sparidae—porgies
Calamus calamus (Valenciennes, 1830)—saucereye porgy; V
Sciaenidae—drums and croakers
Equetus punctatus (Bloch & Schneider, 1801)—spotted drum; USNM, I; Figure 95
Pareques acuminatus (Bloch & Schneider, 1801)—high-hat; USNM, I; Figure 96
Mullidae—goatfishes
Mulloidichthys martinicus (Cuvier, 1829)—yellow goatfish; USNM, F, I, O; Figure 97
Pseudupeneus maculatus (Bloch, 1793)—spotted goatfish; USNM, I, O, V; Figure 98
Chaetodontidae—butterflyfishes
Chaetodon capistratus Linnaeus, 1758—foureye butterflyfish; USNM, I, O, V; Figure 99
Chaetodon ocellatus Bloch, 1787—spotfin butterflyfish; USNM, F, O, V
Chaetodon sedentarius Poey, 1860—reef butterflyfish; UF, O, V
Chaetodon striatus Linnaeus, 1758—banded butterflyfish; USNM, I, O, V; Figure 100
Prognathodes aculeatus (Poey, 1860)—longsnout butterflyfish; USNM, I, O, V; Figure 101
Pomacanthidae—angelfishes
Centropyge argi Woods & Kanazawa, 1951—cherubfish; USNM, I, O, V; Figure 102
Holacanthus ciliaris (Linnaeus, 1758)—queen angelfish; USNM, F, I, O, V
Holacanthus tricolor (Bloch, 1795)—rock beauty; USNM, I, O, V; Figures 103, 104
Pomacanthus arcuatus (Linnaeus, 1758)—gray angelfish; V
Pomacanthus paru (Bloch, 1787)—French angelfish; O, V
Kyphosidae—sea chub
Kyphosus incisor (Cuvier, 1831)—yellow chub; V
Kyphosus sectatrix (Linnaeus, 1766)—Bermuda chub; O, V
Cirrhitidae—hawkfishes
Amblycirrhitus pinos (Mowbray, 1927)—redspotted hawkfish; USNM, I, O; Figure 105
Pomacentridae-damselfishes
Chromis cyanea (Poey, 1860)—blue chromis; USNM, I, O, V; Figure 106
Chromis multilineata (Guichenot, 1853)—brown chromis; USNM, I, O, V; Figure 107
Microspathodon chrysurus (Cuvier, 1830)—yellowtail damselfish; O, V
Stegastes adustus (Troschel, 1865) —dusky damselfish; O
Stegastes leucostictus (Müller & Troschel, 1848) —beaugregory; O
Stegastes partitus (Poey, 1868)—bicolor damselfish; USNM, I, O, V; Figures 108, 109
This species has a variable color pattern throughout its range. Two color morphs were found at Saba Bank. Both morphs have a yellow pectoral fin and a reduced yellowish white area covering the caudal peduncle. One morph has a black caudal fin (Figure 108) and the other morph has a pale yellowish caudal fin with a dusky brown area in the middle of the upper and lower lobes (Figure 109).
Stegastes planifrons (Cuvier, 1830)—threespot damselfish; USNM, I, O; Figure 110
Labridae—wrasses
Bodianus rufus (Linnaeus, 1758)—Spanish hogfish; O, V
Clepticus parrae (Bloch & Schneider, 1801)—creole wrasse; USNM, I, O, V; Figure 111
Doratonotus megalepis Günther, 1862—dwarf wrasse; USNM, I, V; Figure 112
Halichoeres bivittatus (Bloch, 1791)—slippery dick; USNM, I, O, V; Figures 113, 114
Halichoeres cyanocephalus (Bloch, 1791)—yellowcheek wrasse; O, V
Halichoeres garnoti (Valenciennes, 1839)—yellowhead wrasse; UF, USNM, I, O, V; Figures 115, 116, 117
These figures show portions of the transitional color phases as individuals transform from juveniles (Figure 115) into initial phase females (Figure 116) and finally into terminal phase males (Figure 117).
Halichoeres maculipinna (Müller & Troschel, 1848)—clown wrasse; USNM, I, O
Halichoeres pictus (Poey, 1860)—rainbow wrasse; USNM, I, O; Figures 118, 119
Halichoeres poeyi (Steindachner, 1867)—blackear wrasse; UF, USNM, I, O, V; Figure 120
Halichoeres radiatus (Linnaeus, 1758)—puddingwife; O, V
Lachnolaimus maximus (Walbaum, 1792)—hogfish; USNM
Thalassoma bifasciatum (Bloch, 1791)—bluehead; USNM, I, O, V; Figures 121, 122
Xyrichtys splendens Castelnau, 1855—green razorfish; USNM, I, O; Figures 123, 124, 125, 126
Scaridae—parrotfishes
Cryptotomus roseus Cope, 1871—bluelip parrotfish; UF, USNM, I, O; Figure 127
Scarus coelestinus Valenciennes, 1840—midnight parrotfish; V
Scarus guacamaia Cuvier, 1829—rainbow parrotfish; F; Figure 128
Scarus iseri (Bloch, 1789)—striped parrotfish; USNM, O, V
Scarus taeniopterus Desmarest, 1831—princess parrotfish; USNM, I, O, V; Figure 129, 130
Scarus vetula Bloch & Schneider, 1801—queen parrotfish; O
Sparisoma atomarium (Poey, 1861)—greenblotch parrotfish; USNM, I, O; Figure 131
Sparisoma aurofrenatum (Valenciennes, 1840)—redband parrotfish; USNM, I, O, V; Figures 132, 133, 134
Sparisoma chrysopterum (Bloch & Schneider, 1801)—redtail parrotfish; USNM, I, O; Figure 135
Sparisoma radians (Valenciennes, 1840)—bucktooth parrotfish; USNM, I, O, V; Figure 136
Sparisoma viride (Bonnaterre, 1788)—stoplight parrotfish; USNM, I, O, V; Figure 137
Tripterygiidae—triplefins
Enneanectes altivelis Rosenblatt, 1960 —lofty triplefin; USNM, I; Figure 138
Enneanectes atrorus Rosenblatt, 1960—redeye triplefin; USNM, I
Enneanectes jordani (Evermann and Marsh, 1899) —mimic triplefin; USNM, I
Dactyloscopidae-sand stargazers
Dactyloscopus tridigitatus Gill, 1859—sand stargazer; USNM, I; Figures 139, 140
Gillellus uranidea Böhlke, 1968—warteye stargazer; USNM, I; Figure 141
Platygillellus rubrocinctus (Longley, 1934)—saddle stargazer; USNM, I; Figures 142, 143, 144
Blenniidae—combtooth blennies
Parablennius marmoreus (Poey, 1876)—seaweed blenny; USNM, I
Labrisomidae—scaly blennies
Labrisomus gobio (Valenciennes, 1836)—palehead blenny; USNM, I; Figure 145
Labrisomus haitiensis Beebe &Tee-Van, 1918—longfin blenny; USNM, I; Figure 146
Malacoctenus boehlkei Springer, 1959—diamond blenny; USNM, I; Figure 147
Paraclinus grandicomis (Rosen, 1911)—horned blenny; UF, USNM, I; Figure 148
Starksia atlantica Longley, 1934—smootheye blenny; USNM, I; Figures 149, 150
The Saba Bank population may be a distinct species in the species complex currently referred to as Starksia atlantica. This complex requires additional taxonomic study.
Starksia cf lepicoelia Böhlke & Springer, 1961—blackcheek blenny; USNM, I; Figures 151, 152
Females (Figure 151) lack the black spot on the cheek that is characteristic of mature males (Figure 152). The Saba Bank population may be a distinct species in the species complex currently referred to as Starksia lepicoelia. This complex requires additional taxonomic study.
Starksia melasma Williams & Mounts, 2003—black spot blenny; USNM, I
Starksia nanodes Böhlke & Springer, 1961—dwarf blenny; USNM, I; Figure 153
The Saba Bank population may be a distinct species in the species complex currently referred to as Starksia nanodes. This complex requires additional taxonomic study.
Chaenopsidae—tube blennies
Acanthemblemaria aspera (Longley, 1927)—roughhead blenny; USNM, I; Figures 154, 155, 156
Three different color patterns were observed at Saba Bank: juvenile/female (Figure 154), adult male (Figure 155), and a distinctive yellow, probably female, color morph (Figure 156). The adult male and the yellow morph were taken together at the same collecting station.
Emblemaria pandionis Evermann & Marsh, 1900—sailfin blenny; USNM, I; Figure 157
Emblemariopsis cf signifer (Ginsburg, 1942)—flagfin blenny; USNM, I; Figure 158
The signifer species complex ranges from Brazil throughout the Caribbean and includes a number of undescribed species in the Caribbean region. Additional taxonomic study is required to resolve the taxa.
Gobiesocidae—clingfishes
Acyrtus artius Briggs, 1955—papillate clingfish; USNM, I; Figures 159, 160, 161
Callionymidae—dragonets
Paradiplogrammus bairdi Jordan, 1888—lancer dragonet; USNM, I, O; Figures 162, 163
Gobiidae-gobies
Coryphopterus dicrus Böhlke & Robins, 1960—colon goby; USNM, I; Figure 164
Coryphopterus eidolon Böhlke & Robins, 1960—pallid goby; USNM, I; Figure 165
Coryphopterus glaucofraenum Gill, 1863—bridled goby; USNM, I, V; Figure 166
Coryphopterus personatus (Jordan & Thompson, 1905)—masked goby; USNM, I; Figure 167
Coryphopterus thrix Böhlke & Robins, 1960—bartail goby; USNM, I; Figure 168
Elacatinus chancei (Beebe & Hollister 1933)—shortstripe goby; USNM, I; Figure 169
Elacatinus evelynae (Böhlke & Robins, 1968)—sharknose goby; USNM, I; Figure 170
Elacatinus genie (Böhlke & Robins, 1968)—cleaner goby; USNM, I
Evermannichthys metzelaari Hubbs, 1923 —roughtail goby; USNM, I; Figure 171
Gnatholepis thompsoni Jordan, 1904—goldspot goby; USNM, I, V; Figure 172
Lythrypnus elasson Böhlke & Robins, 1960—dwarf goby; USNM, I; Figure 173
Lythrypnus minimus Garzón & Acero P., 1988 —pygmy goby; USNM, I; Figure 174
Lythrypnus nesiotes Böhlke & Robins, 1960—island goby; USNM, I; Figure 175
Priolepis hipoliti (Metzelaar, 1922)—rusty goby; USNM, I; Figures 176, 177
Psilotris batrachodes Böhlke, 1963—toadfish goby; USNM, I; Figure 178
Psilotris boehlkei Greenfield, 1993—yellowspot goby; I; Figure 179
The fresh colors of P. boehlkei are presented (Figure 179) for the first time. This very rare species was previously known from only five specimens taken at St. Barthelemey in 1965. The single specimen we collected extends the known distribution of P. boehlkei to Saba Bank. We have named this species the yellowspot goby in reference to the yellow spots on the head and body.
Pycnomma roosevelti Ginsburg, 1939 —Roosevelt's goby; USNM, I; Figure 180
Although P. roosevelti has previously been taken from a several scattered localities around the Caribbean (Isla Providencia, Guadeloupe, Belize and Puerto Rico), there are fewer than 10 specimens known and its fresh colors (Figure 180) have not been published previously.
Risor ruber (Rosén, 1911) —tusked goby; USNM, I; Figure 181
Acanthuridae—surgeonfishes
Acanthurus bahianus Castelnau, 1855—ocean surgeon; USNM, I, O, V; Figure 182
Acanthurus chirurgus (Bloch, 1787)—doctorfish; USNM, I, O, V; Figure 183
Acanthurus coeruleus Bloch & Schneider, 1801—blue tang; USNM, I, O, V; Figure 184
Sphyraenidae—barracudas
Sphyraena barracuda (Walbaum, 1792)—great barracuda; O, V
Scombridae—mackerels
Acanthocybium solandri (Cuvier, 1832)—wahoo; F
Euthynnus alletteratus (Rafinesque, 1810)—little tunny; USNM; Figure 185
Scomberomorus regalis (Bloch, 1793)—cero; USNM
Thunnus atlanticus (Lesson 1831)—blackfin tuna; F; Figure 186
Caproidae—boarfishes
Antigonia capros Lowe, 1843—deepbody boarfish; USNM, F; Figure 187
Antigonia combatia Berry & Rathjen 1959—shortspine boarfish; UF, T
The shortspine boarfish record from Saba Bank is based on a trawl collection with vouchered museum specimens at UF.
Bothidae—lefteye flounders
Bothus lunatus (Linnaeus, 1758)—peacock flounder; O
Bothus ocellatus (Agassiz, 1831)—eyed flounder; UF, USNM, I; Figure 188
Trichopsetta ventralis (Goode & Bean, 1885)—sash flounder; UF, T
Paralichthyidae—sand flounders
Citharicthys dinoceros Goode & Bean, 1886—spined whiff; USNM, T
Cynoglossidae—tonguefishes
Symphurus arawak Robins & Randall, 1965—Caribbean tonguefish; USNM, I; Figure 189
Symphurus ommaspilus Böhlke, 1961—ocellated tonguefish; USNM, I; Figure 190
Balistidae—triggerfishes
Balistes vetula Linnaeus, 1758—queen triggerfish; USNM, O, V
Canthidermis sufflamen (Mitchill, 1815)—ocean triggerfish; O, V
Melichthys niger (Bloch, 1786)—black durgon; USNM, O, V
Xanthichthys ringens (Linnaeus, 1758)—sargassum triggerfish; USNM, I; Figure 191
Monacanthidae—filefishes
Aluterus scriptus (Osbeck, 1765)—scrawled filefish; USNM, F, V
Cantherhines macrocerus (Hollard, 1853)—whitespotted filefish; USNM, F; Figure 192
Cantherhines pullus (Ranzani, 1842)—orangespotted filefish; O, V
Monacanthus ciliatus (Mitchill, 1818)—fringed filefish; USNM, I, V; Figure 193
Monacanthus tuckeri Bean, 1906—slender filefish; USNM, I; Figure 194
Ostraciidae—boxfishes
Acanthostracion polygonia Poey, 1876—honeycomb cowfish; USNM, F, O, V
Acanthostracion quadricornis (Linnaeus, 1758)—scrawled cowfish; USNM, F, O
Lactophrys bicaudalis (Linnaeus, 1758)—spotted trunkfish; F
Lactophrys trigonus (Linnaeus, 1758)—trunkfish; USNM, I, V; Figure 195
Lactophrys triqueter (Linnaeus, 1758)—smooth trunkfish; USNM, F, OBS, VIS; Figure 196
Tetraodontidae—puffers
Canthigaster rostrata (Bloch, 1786)—sharpnose puffer; USNM, I, O, V; Figure 197
Sphoeroides spengleri (Bloch, 1785)—bandtail puffer; USNM, I, O; Figure 198
Diodontidae—porcupinefishes
Diodon hystrix Linnaeus, 1758—porcupinefish; O
Discussion
We document the occurrence of 270 species of fishes at Saba Bank. The diversity of fishes at Saba Bank is comparable (Table 1) to that of the oceanic atolls of Colombia (273 species), the islands in the Mona Passage of Puerto Rico (261 species) and Buck Island Reef National Monument (BIRNM; 262 species). The relatively high diversity of fishes at Saba Bank exists despite the lack of emergent land at the bank. There is no shallow-water shore-fish fauna represented on the bank due to the absence of a high-energy shoreline. These habitats typically add significantly to the fish diversity of Caribbean habitats. For example, the tube blennies (family Chaenopsidae) are a group of shorefishes typically found in fairly shallow coastal waters. According to Williams$^4$, there are approximately 22 recognized species of tube blennies known to ​occur in the central Caribbean, but only three of these species were found on Saba Bank. In addition, the absence of mangrove vegetation and apparent lack of sea-grass beds also limits the fish fauna of Saba Bank. In their quantitative study of a number of Saba Bank habitats, Toller et al. $^5$ attribute the apparent lack or rarity of a number of fish species found on Saba Bank to the absence of those habitats as nurseries for the juvenile stages of these fish species. Nevertheless, diverse habitat types exist on Saba Bank ranging from coral reefs and algal flats to soft-bottom lagoon areas and scoured hard, flat, pavement-like zones. These diverse habitats support a highly diverse, fish fauna and include a number of undescribed, new species along with species rarely encountered elsewhere in the Caribbean.
The highly diverse, fish fauna so far reported appears to substantially under represent the species richness of fishes for Saba Bank. The actual species-accumulation curve of rotenone collections and visual surveys combined do not reach an asymptote (Figure 201). The expected species-accumulation curves (Chao2 and Jack1 estimators in EstimateS) predict total fish-species richness somewhere between 320 and 411 species.
Of the 270 species we report (Table 1) from Saba Bank, 132 (49%) were observed during visual surveys. This result is comparable to the findings for Navassa Island where 41% of the fish species were detected by visual surveys $^6$, for the Mona Passage islands with 43% of the fish species detected visually $^7$, and for BIRNM with 44% of the fish species detected visually $^8$. A higher percentage of fish species was detected visually at Saba Bank than at Navassa, Mona, or BIRNM. This could possibly be a result of the higher number of visual surveys carried out at Saba Bank than at Navassa or Mona. Although the BIRNM results are based on a higher number of visual surveys (70), a comparable percentage of the fauna was detected visually at BIRNM. BIRNM ichthyocide surveys (58) appear to demonstrate the ability of ichthyocide collections to more thoroughly sample the fauna (see below). Visual censuses have numerous biases as discussed in the methods for the Pelican Cays study $^9$. Despite the slight differences in methods and sampling designs between the published fish species-richness studies in different parts of the Caribbean, the methods utilized produce consistent and comparable results.
Ichthyocide collections at Saba Bank yielded specimens representing 155 fish species (57% of the total fish fauna). At Navassa Island where there were fewer visual surveys than at Saba Bank, over 70% of the fish species were collected with ichthyocide $^6$, at the Mona Passage islands, 61% of the fish species were collected with ichthyocide $^7$, and at BIRNM, 87% of the fish species were collected with ichthyocide despite their being more visual surveys in the BIRNM study $^8$. By occupying 58 (27 at Saba Bank) ichthyocide stations covering a broader diversity of ecological habitats, the BIRNM study appears to have obtained a more thorough sampling of the resident species. The lower number of species taken at Saba Bank with ichthyocide is at least partially due to the absence of shallow habitat. Shallow habitat is required by many of the small cryptic fish species that are normally detected only with the use of ichthyocide and these species have not been found at Saba Bank. The prevalence of strong currents across Saba Bank further limits the effectiveness of ichthyocide, which is only effective when relatively high concentrations of ichthyocide remain in one place (preferably in a confined area) for more than 15 to 20 minutes. The ichthyocide-collecting methods employed at Saba Bank were the same as those used at Navassa Island and Belize. The BIRNM ichthyocide collecting methods differed from ours only in that the BIRNM study used a block net in addition to collecting outside the net. Nevertheless, the BIRNM methods were comparable to ours because the BIRNM study included those species collected outside the block nets in their results. There are inherent biases in ichthyocide collecting due to unpredictability of environmental parameters, such as currents, temperature, degree of confinement of the sampling area, ability of larger species to swim away, and variable assays of active rotenone in the powdered Derris root (batches often vary from 5% to over 11% active rotenone due to natural variation in rotenone concentration among the roots of different plants). Despite these biases, ichthyocide collections yield comparable results as described above.
Neither visual surveying nor ichthyocide collecting alone are capable of providing a comprehensive survey of coastal (or submerged atoll in the case of Saba Bank) fish species. A ​combination of visual surveys, ichthyocide sampling using SCUBA, and various fishing techniques must be employed to effectively assess fish-species richness in marine habitats shallow enough to be accessible to SCUBA divers.
10.1371/journal.pone.0010676.t001
The number of fish species living on Saba Bank is undoubtedly higher than 270 as indicated by Chao2 and Jack1 estimators. As most parts of Saba Bank are deeper than 25 m, sampling with ichthyocides using SCUBA is limited by the reduced bottom time at these depths. As a result we focused primarily on the rim (shallowest parts) of the submerged atoll to maximize bottom time for collecting. Future sampling with ichthyocides applied by divers utilizing rebreathers, supplemented with trawl and dredge sampling would allow collecting from the outer slopes and would certainly yield additional new and interesting species of fishes from this submerged atoll.
Methods
The NMNH/SI Animal Care and Use Committee approved the methods and procedures utilized during the course of this biodiversity assessment project. All Saba Bank projects had collecting permits through the Convention on Trade in Endangered Species (CITES, where necessary) and the Saba Conservation Foundation (where CITES was not required).
Roving surveys were completed using SCUBA and lasted 60 minutes, bottom time permitting. All species encountered were listed on a slate while swimming in a haphazard pattern covering all bottom depths possible down to a maximum of 38 m. Other visual surveys are described in Toller et al $^5$. Collecting methods follow Collette et al $^6$. Species were photographed in aquaria after the fins were pinned out and brushed with formaldehyde solution. Tissue samples were taken from fresh specimens and the voucher specimens were preserved in a formaldehyde solution diluted with water to 3.75% formaldehyde. Large specimens were also injected with 37.5% formaldehyde before being soaked in the 3.75% formaldehyde solution.
After arrival at the Museum Support Center (MSC), National Museum of Natural History, specimens were transferred sequentially through water-diluted solutions of 25% ethanol, 50% ethanol, and finally into 75% ethanol for permanent archival storage. Specimens were then processed and cataloged into the USNM at the MSC in Suitland, MD.
To generate species-accumulation curves, a data matrix of presence/absence was constructed from 38 combined roving surveys, rotenone collections, and fish-habitat transects using species as variables and dives as observations (Table 2 provides details for each survey and collecting station). The matrix was employed for actual and expected species-accumulation curves (Mao Tau) in EstimateS v.8.0 software $^{10}$. Total expected richness was estimated using Chao2 and Jack1 estimators. ​Table 2. Collecting stations occupied on Saba Bank, 2006-2007.
10.1371/journal.pone.0010676.t002
A Bray-Curtis resemblance matrix was generated from a matrix of presence/absence data from 12 combined roving surveys and rotenone collections in order to produce a non-metric multidimensional scaling (MDS) ordination and group averaged hierarchical clustering dendrogram in PRIMER v. 6.1 software $^{11}$. The ANOSIM statistic was employed to test for a priori differences between habitat types and depth classes. A dendrogram based on group averaged hierarchical cluster techniques was used to illustrate the differences among depth classes (Figure 202). Color codes are derived from the similarity profile (SIMPROF) statistic in Primer 6.1. SIMPROF is less powerful than ANOSIM, intended for a posteriori tests of structure in the data. The test was employed here for representational purposes. A second data matrix of species presence and absence was introduced for an a posteriori test of genuine data structure among Caribbean localities, using the Bray-Curtis resemblance measure to determine the level ​of similarity among localities. Principal Components Analysis (PCA) was also conducted in PRIMER 6.1 in order to determine the fish species primarily responsible for differences among habitats.
Acknowledgments
This program was a joint initiative of the Department of Environment and Nature of the Netherlands Antilles and Conservation International. We are especially grateful to the many people who made the biodiversity survey possible, including Andy Caballero of Nature Foundation St. Maarten; David Kooistra, Stanley Peterson, and Jan Den Dulk of Saba Conservation Foundation; Shelley Lundvall, Saba Bank project manager; fishermen Leroy Peterson, Nicky Johnson and Armand Simmons. We thank Jerome Finan, Kris Murphy, and David G. Smith, National Museum of Natural History, Smithsonian Institution, for their help with this project. We thank Menno Van der Velde and Wim Schutten for collecting fishes at several ichthyocide stations. Jill Leger, Yap Films Inc, provided a video clip of fishes filmed at the shipwreck on Saba Bank. Emma L. Hickerson, Flower Garden Banks National Marine Sanctuary (FGBNMS), NOAA, kindly provided information on the number of fish species known to occur at the FGBNMS. Dr. JA Sanchez, University de los Andes, allowed us to use his photograph for Figure 3.
Author Contributions
Conceived and designed the experiments: MS. Performed the experiments: JTW KEC JLVT PCH WWT PJE MS. Analyzed the data: JTW KEC JLVT PCH PJE. Wrote the paper: JTW. Wrote portions of the paper: KEC PCH PJE. Helped prepare tables and check species identifications: JLVT. Prepared Fig. 1: PCH.
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