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PLoS One. 2010; 5(5): e10676.
Published online 2010 May 21. doi:  10.1371/journal.pone.0010676
PMCID: PMC2873961

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

Brian Gratwicke, Editor

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.

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

We occupied fish stations at 25 locations during a rapid assessment (RAP) survey, 2–16 January 2006; including: 20 ichthyocide stations, 12 roving visual surveys, two hook & line stations, and five by-catch stations from lobster traps; two fish-ichthyocide stations were occupied at two additional locations on 20 June 2007 (Figure 1). In 2007, Toller assessed the benthic communities and fish assemblages based in part on 40 visual surveys of the fish fauna at an area on the eastern side of Saba Bank.

Figure 1
Bathymetric map of Saba Bank with fish stations marked.

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.

A non-metric multidimensional-scaling (MDS) ordination of the 12 rotenone and roving stations based on a Bray-Curtis similarity matrix of incidence data was used to illustrate similarities and differences of fish assemblages found on the four atoll habitat types: fore reef, patch reef, lagoonal, Small Bank (Figure 2). There were significant differences among habitat types (ANOSIM Global R = 0.96, P = 0.001). Differences between fore-reef stations and patch-reef stations were most pronounced (ANOSIM, Pairwise R = 0.92, P = 0.002). Fore-reef assemblages ranged from 39 to 60 species per station while the patch-reef stations had 26 and 32 species. Fore-reef stations were 50% similar to each other, and patch-reef stations were up to 60% similar. Fore-reef assemblages were not significantly different than Small Bank, or the lagoonal habitat (ANOSIM, Pairwise R = 0.973, P = 0.11). Low sample size (one station with 39 species) in the lagoonal habitat limits this comparison.

Figure 2
MDS ordination of fish-survey stations illustrates high similarity of assemblages within fore-reef sites and significant differences (ANOSIM, P<0.05) among habitats.

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

Figure 3
Ginglymostoma cirratum, underwater photo by Juan Sanchez.

Squalidae—dogfish sharks

Squalus cubensis Howell Rivero, 1936Cuban dogfish; F; Figure 4

Figure 4
Squalus cubensis, 475 mm TL, photo by W Toller.

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, 1957lined lantern shark; USNM, T

Dasyatidae—whiptail stingrays

Dasyatis americana Hildebrand & Schroeder, 1928southern stingray; V

Muraenidae—morays

Anarchias similis (Lea, 1913)pygmy moray; USNM, I

Enchelycore carychroa (Böhlke & Böhlke, 1976)chestnut moray; USNM, I, F; Figure 5

Figure 5
Enchelychore carychroa, 175 mm TL, photo by JT Williams.

Enchelycore nigricans (Bonnaterre, 1788)viper moray; USNM, I

Gymnothorax conspersus Poey, 1867saddled moray; USNM, F; Figure 6

Figure 6
Gymnothorax conspersus, 730 mm TL, photo by W Toller.

Gymnothorax maderensis(Johnson, 1862)sharktooth moray; USNM, F; Figure 7

Figure 7
Gymnothorax maderensis, 300 mm TL, photo by W Toller.

Gymnothorax miliaris (Kaup, 1856)goldentail moray; USNM, I, O; Figure 8

Figure 8
Gymnothorax miliaris, 60.3 mm TL, photo by JT Williams.

Gymnothorax moringa (Cuvier, 1829)spotted moray; USNM, I, O, V; Figure 9

Figure 9
Gymnothorax moringa, 189.8 mm TL, photo by JT Williams.

Gymnothorax polygonius Poey, 1876polygon moray; USNM, F; Figure 10

Figure 10
Gymnothorax polygonius, 810 mm TL, photo by W Toller.

Gymnothorax vicinus (Castelnau, 1855)purplemouth moray; USNM, I, O; Figure 11

Figure 11
Gymnothorax vicinus, 48.2 mm TL, photo by JT Williams.

Monopenchelys acuta (Parr, 1930)redface moray; USNM, I; Figure 12

Figure 12
Monopenchelys acuta, 132.4 mm TL, photo by JT Williams.

Uropterygius macularius (Lesueur, 1825)marbled moray; USNM, I

Ophichthidae—snake eels

Ahlia egmontis (Jordan, 1884)key worm eel; USNM, I; Figures 13, ,1414

Figure 13
Ahlia egmontis, 183.0 mm TL, photo by JT Williams.
Figure 14
Ahlia egmontis, 183.0 mm TL, close-up of head, photo by JT Williams.

Aprognathodon platyventris Böhlke, 1967stripe eel; USNM, I; Figures 15, ,1616

Figure 15
Aprognathodon platyventris, 149.2 mm TL, photo by JT Williams.
Figure 16
Aprognathodon platyventris, 149.2 mm TL, close-up of head, photo by JT Williams.

Myrichthys breviceps (Richardson, 1848) sharptail eel; O

Myrichthys ocellatus (Lesueur, 1825)goldspotted eel; USNM, I; Figures 17, ,1818

Figure 17
Myrichthys ocellatus, 383 mm TL, photo by JT Williams.
Figure 18
Myrichthys ocellatus, 383 mm TL, close-up of head, photo by JT Williams.

Congridae—conger eels

Bathycongrus thysanochilus (Reid, 1934)conger eel; USNM, T

Conger esculentus Poey, 1866grey conger; USNM, F

Chlopsidae—false morays

Kaupichthys hyoproroides (Strömman, 1896)false moray; USNM, I; Figure 19

Figure 19
Kaupichthys hyoproroides, 79.4 mm TL, photo by JT Williams.

Kaupichthys nuchalis Böhlke, 1967collared eel; USNM, I; Figure 20

Figure 20
Kaupichthys nuchalisi, 61.6 mm TL, photo by JT Williams.

Moringuidae—spaghetti eels

Moringua edwardsi (Jordan & Bollman, 1889)spaghetti eel; USNM, I; Figure 21

Figure 21
Moringua edwardsi, 258 mm TL, photo by JT Williams.

Synodontidae—lizardfishes

Saurida brasiliensis Norman 1935largescale lizardfish; UF, T

Saurida normani Longley 1935shortjaw lizardfish; UF, T

Synodus intermedius (Spix & Agassiz, 1829)sand diver; USNM, UF, F, O; Figure 22

Figure 22
Synodus intermedius, 316 mm SL, photo by W Toller.

Synodus poeyi Jordan, 1887offshore lizardfish; UF, T

Synodus saurus (Linnaeus, 1758)Atlantic lizardfish; UF, I, V

Synodus synodus (Linnaeus, 1758)red lizardfish; UF, USNM, I; Figure 23

Figure 23
Synodus synodus, 42.5 mm SL, photo by JT Williams.

Trachinocephalus myops (Forster, 1801)snakefish; V

Ophidiidae—cusk-eels

Brotula barbata (Bloch & Schneider, 1801)bearded brotula; USNM, O; Figure 24

Figure 24
Brotula barbata, 628 mm TL, photo by W Toller.

Neobythites ocellatus Günther 1887ocellate cusk-eel; USNM, T

Neobythites unicolor Nielsen & Retzer 1994unicolor cusk-eel; USNM, T

Ophidion antipholus Lea & Robins, 2003longnose 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

Figure 25
Parophidion schmidti, 69.3 mm TL, photo by JT Williams.

Petrotyx sanguineus (Meek & Hildebrand, 1928)redfin brotula; USNM, I; Figure 26

Figure 26
Petrotyx sanguineus, 62.2 mm TL, photo by JT Williams.

Bythitidae—viviparous brotulas

Ogilbia sabaji Moller, Schwarzhans & Nielsen, 2005Sabaj coralbrotula; USNM, I; Figure 27

Figure 27
Ogilbia sabaji, 27.2 mm SL, photo by JT Williams.

Antennariidae—frogfishes

Antennarius pauciradiatus Schultz, 1957 dwarf frogfish; USNM, I; Figures 28, ,2929

Figure 28
Antennarius pauciradiatus, 24.9 mm SL, photo by JT Williams.
Figure 29
Antennarius pauciradiatus, 24.9 mm SL, close-up of head, photo by JT Williams.

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, 1989Suttkus 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, 1876Atlantic batfish; UF, T

Halieutichthys aculeatus (Mitchill, 1818)pancake batfish; UF, T

Ogcocephalus pumilus Bradbury, 1980dwarf 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

Figure 30
Bryx randalli, 24.3 mm SL, photo by JT Williams.

Micrognathus crinitus (Jenyns, 1842) banded pipefish; USNM, I; Figures 31, ,3232

Figure 31
Micrognathus crinitus, 102.8 mm SL, photo by JT Williams.
Figure 32
Micrognathus crinitus, 102.8 mm SL, close-up of head, photo by JT Williams.

Aulostomidae—trumpetfishes

Aulostomus maculatus Valenciennes, 1837trumpetfish; USNM, I, O, V

Holocentridae—squirrelfishes

Holocentrus adscensionis (Osbeck, 1765)squirrelfish; USNM, I, O, V; Figure 33

Figure 33
Holocentrus ascensionus, 195.0 mm SL, photo by JT Williams.

Holocentrus rufus (Walbaum, 1792)longspine squirrelfish; USNM, I, O, V; Figure 34

Figure 34
Holocentrus rufus, 153.2 mm SL, photo by JT Williams.

Myripristis jacobus Cuvier, 1829blackbar soldierfish; USNM, I, O; Figure 35

Figure 35
Myripristis jacobus, 114.7 mm SL, photo by JT Williams.

Neoniphon marianus (Cuvier, 1829)longjaw squirrelfish; USNM, I; Figure 36

Figure 36
Neoniphon marianus, 98.2 mm SL, photo by JT Williams.

Plectrypops retrospinis (Guichenot, 1853)cardinal soldierfish; USNM, I; Figure 37

Figure 37
Plectrypops retrospinis, 71.5 mm SL, photo by JT Williams.

Sargocentron coruscum (Poey, 1860)reef squirrelfish; USNM, I, O; Figures 38, ,3939

Figure 38
Sargocentron coruscum, juvenile, 31.1 mm SL, photo by JT Williams.
Figure 39
Sargocentron coruscum, 81.7 mm SL, photo by JT Williams.

Scorpaenidae—scorpionfishes

Scorpaena albifimbria Evermann & Marsh, 1900coral scorpionfish; USNM, I; Figure 40

Figure 40
Scorpaena albifimbria, 37.4 mm SL, photo by JT Williams.

Scorpaena bergii Evermann & Marsh, 1900goosehead scorpionfish; USNM, I

Scorpaena grandicornis Cuvier, 1829plumed scorpionfish; USNM, I; Figure 41

Figure 41
Scorpaena grandicornis, 65.1 mm SL, photo by JT Williams.

Scorpaena inermis Cuvier, 1829 mushroom scorpionfish; USNM, I, T; Figures 42, ,4343

Figure 42
Scorpaena inermis, 40.1 mm SL, red morph, photo by JT Williams.
Figure 43
Scorpaena inermis, 65.9 mm SL, yellow morph, photo by JT Williams.

Two color morphs were collected at Saba Bank, a red morph (Figure 42) and a yellow morph (Figure 43).

Scorpaena plumieri Bloch, 1789spotted scorpionfish; USNM, I

Scorpaenodes caribbaeus Meek & Hildebrand, 1928reef scorpionfish; USNM, I; Figure 44

Figure 44
Scorpaenodes caribbaeus, 30.1 mm SL, photo by JT Williams.

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, 1885bandtail searobin; UF, T

Symphysanodontidae—slopefishes

Symphysanodon berryi Anderson, 1970slope bass; USNM, F; Figure 45

Figure 45
Symphysanodon berryi, top 75 mm SL, lower 63 mm SL, photo by W Toller.

Serranidae—sea basses

Alphestes afer (Bloch, 1793)mutton hamlet; F, O; Figure 46

Figure 46
Alphestes afer, underwater photo by W Toller.

Cephalopholis cruentata (Lacepède, 1802)graysby; USNM, I, F, O, V; Figure 47

Figure 47
Cephalopholis cruentata, 101.1 mm SL, photo by JT Williams.

Cephalopholis fulva (Linnaeus, 1758)coney; UF, USNM, I, F, O, V; Figure 48

Figure 48
Cephalopholis fulva, 121.1 mm SL, photo by JT Williams.

Diplectrum bivittatum (Valenciennes, 1828) dwarf sand perch; UF, T

Epinephelus flavolimbatus Poey, 1865yellowedge grouper; F; Figure 49

Figure 49
Epinephelus flavolimbatus, 652 mm SL, photo by W Toller.

Epinephelus guttatus (Linnaeus, 1758)red hind; UF, USNM, F, I, O, T, V; Figure 50

Figure 50
Epinephelus guttatus, 283.2 mm SL, photo by JT Williams.

Epinephelus morio (Valenciennes, 1828)red grouper; F

Epinephelus niveatus Valenciennes, 1828snowy grouper; USNM, F; Figure 51

Figure 51
Epinephelus niveatus, 526 mm SL, photo by W Toller.

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

Figure 52
Hypoplectrus nigricans, 67.2 mm SL, photo by JT Williams.

Hypoplectrus puella (Cuvier, 1828)barred hamlet; O, V

Liopropoma rubre Poey, 1861peppermint basslet; USNM, I; Figure 53

Figure 53
Liopropoma rubre, 30.0 mm SL, photo by JT Williams.

Mycteroperca interstitialis (Poey, 1860)yellowmouth grouper; F; Figure 54

Figure 54
Mycteroperca interstitialis, 328 mm SL, photo by W Toller.

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

Figure 55
Paranthias furcifer, 148.4 mm SL, photo by JT Williams.

Pseudogramma gregoryi (Breder, 1927)reef bass; USNM, I; Figure 56

Figure 56
Pseudogramma gregoryi, 43.5 mm SL, photo by JT Williams.

Rypticus bistripinus (Mitchill, 1818) freckled soapfish; USNM, I; Figure 57

Figure 57
Rypticus bistripinus, 46.2 mm SL, photo by JT Williams.

Rypticus saponaceus (Bloch & Schneider, 1801)greater soapfish; USNM, I; Figure 58

Figure 58
Rypticus saponaceus, 44.1 mm SL, photo by JT Williams.

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, 1861spotted soapfish; USNM, I; Figure 59

Figure 59
Rypticus subbifrenatus, 54.0 mm SL, photo by JT Williams.

Schultzea beta (Hildebrand, 1940)school bass; UF, USNM, I

Serranus baldwini (Evermann & Marsh, 1899)lantern bass; UF, USNM, I, O, V; Figure 60

Figure 60
Serranus baldwini, 41.6 mm SL, photo by JT Williams.

Serranus maytagi Robins & Starck, 1961 maytag bass; UF

Serranus notospilus Longley, 1935saddle bass; USNM, F; Figure 61

Figure 61
Serranus notospilus, 150 mm SL, photo by W Toller.

Serranus tabacarius (Cuvier, 1829)tobaccofish; UF, O, V

Serranus tigrinus (Bloch, 1790)harlequin bass; USNM, I, O, V; Figure 62

Figure 62
Serranus tigrinus, 71.1 mm SL, photo by JT Williams.

Serranus tortugarum Longley, 1935chalk bass; UF, USNM, I, O; Figure 63

Figure 63
Serranus tortugarum, 51.8 mm SL, photo by JT Williams.

Grammatidae—basslets

Gramma loreto Poey, 1868fairy basslet; USNM, I, O, V; Figure 64

Figure 64
Gramma loreto, 38.4 mm SL, photo by JT Williams.

Opistognathidae—jawfishes

Opistognathus aurifrons (Jordan & Thompson, 1905)yellowhead jawfish; USNM, I, O, V; Figure 65

Figure 65
Opistognathus aurifrons, 44.1 mm SL, photo by JT Williams.

Opistognathus whitehursti (Longley, 1927)dusky jawfish; USNM, I; Figures 66, ,6767

Figure 66
Opistognathus whitehursti, juvenile with large black spot in dorsal fin, 22.1 mm SL, photo by JT Williams.
Figure 67
Opistognathus whitehursti, adult, 44.0 mm SL, photo by JT Williams.

Priacanthidae—bigeyes

Heteropriacanthus cruentatus (Lacepède, 1801)glasseye snapper; O

Priacanthus arenatus Cuvier, 1829bigeye; USNM, I; Figure 68

Figure 68
Priacanthus arenatus, 67.4 mm SL, photo by JT Williams.

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

Figure 69
Apogon maculatus, 21.1 mm SL, photo by JT Williams.

Apogon pillionatus Böhlke and Randall, 1968 broadsaddle cardinalfish; UF, USNM, I

Apogon cf quadrisquamatus Longley, 1934 sawcheek cardinalfish; USNM, I; Figure 70

Figure 70
Apogon cf quadrisquamatus, 21.1 mm SL, photo by JT Williams (this specimen represents a new undescribed species.

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

Figure 71
Apogon robinsi, 35.5 mm SL, photo by JT Williams.

Apogon townsendi (Breder, 1927)belted cardinalfish; USNM, I; Figure 72

Figure 72
Apogon townsendi, 39.1 mm SL, photo by JT Williams.

Astrapogon puncticulatus (Poey, 1867) blackfin cardinalfish; USNM, I; Figure 73

Figure 73
Astrapogon puncticulatus, 40.0 mm SL, photo by JT Williams.

Phaeoptyx conklini (Silvester, 1916)freckled cardinalfish; USNM, I; Figure 74

Figure 74
Phaeoptyx conklini, 40.0 mm SL, photo by JT Williams.

Phaeoptyx pigmentaria (Poey, 1860)dusky cardinalfish; USNM, I

Malacanthidae—tilefishes

Caulolatilus cyanops Poey, 1866blackline tilefish; USNM, F; Figure 75

Figure 75
Caulolatilus cyanops, 288 mm SL, photo by W Toller.

Malacanthus plumieri (Bloch, 1786)sand tilefish; UF, O, V

Coryphaenidae-dolphinfishes

Coryphaena hippurus Linnaeus, 1758common 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

Figure 76
Alectis ciliaris, underwater photo by W Toller.

Caranx bartholomaei Cuvier, 1833yellow jack; O, V

Caranx crysos (Mitchill, 1815) blue runner; O

Caranx latus Agassiz, 1831horse-eye jack; O, V

Caranx lugubris Poey, 1860black jack; O, V

Caranx ruber (Bloch, 1793)bar jack; UF, F, O, V; Figure 77

Figure 77
Caranx ruber, 260.8 mm SL, photo by JT Williams.

Decapterus macarellus (Cuvier, 1833)mackerel scad; O; Figure 78

Figure 78
Decapterus macarellus, underwater photo by W Toller.

Elagatis bipinnulata (Quoy & Gaimard, 1825)rainbow runner; F, OBS

Selar crumenophthalmus (Bloch, 1793)bigeye scad; F; Figure 79

Figure 79
Selar crumenophthalmus, 240 mm SL, photo by W Toller.

Seriola dumerili (Risso, 1810)greater amberjack; F; Figure 80

Figure 80
Seriola dumerili, large specimen at top; Seriola rivoliana, two smaller specimens below, photo by W Toller.

Seriola rivoliana Valenciennes, 1833almaco jack; F; Figure 80

Lutjanidae—snappers

Apsilus dentatus Guichenot, 1853black snapper; USNM, F; Figure 81

Figure 81
Apsilus dentatus, 234 mm SL, photo by W Toller.

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

Figure 82
Lutjanus buccanella, 249 mm SL, photo by W Toller.

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

Figure 83
Lutjanus purpureus, 207 mm SL, photo by W Toller.

Lutjanus synagris (Linnaeus, 1758)lane snapper; USNM, F; Figure 84

Figure 84
Lutjanus synagris, 208 mm SL, photo by W Toller.

Lutjanus vivanus (Cuvier, 1828)silk snapper; USNM, F; Figure 85

Figure 85
Lutjanus vivanus, 311 mm SL, photo by W Toller.

Ocyurus chrysurus (Bloch, 1791)yellowtail snapper; UF, O, V

Pristipomoides aquilonaris (Goode & Bean, 1896)wenchman; USNM, F; Figure 86

Figure 86
Pristipomoides aquilonaris, 263 mm SL, photo by W Toller.

Rhomboplites aurorubens (Cuvier, 1829) vermilion snapper; USNM, F; Figure 87

Figure 87
Rhomboplites aurorubens, 227.3 mm SL, photo by JT Williams.

Lobotidae—tripletails

Lobotes surinamensis (Bloch, 1790)Atlantic tripletail; F; Figure 88

Figure 88
Lobotes surinamensis, 300 mm SL, photo by W Toller.

Haemulidae—grunts

Haemulon album Cuvier, 1830margate; F; Figure 89

Figure 89
Haemulon album, 342 mm SL, photo by W Toller.

Haemulon aurolineatum Cuvier, 1830tomtate; F, O; Figure 90

Figure 90
Haemulon aurolineatum, 175 mm SL, photo by W Toller.

Haemulon carbonarium Poey, 1860caesar grunt; F; Figure 91

Figure 91
Haemulon carbonarium, approximately 200 mm SL, photo by W Toller.

Haemulon flavolineatum (Desmarest, 1823)French grunt; O, V

Haemulon melanurum (Linnaeus, 1758)cottonwick; UF, USNM, F, I, O, V; Figure 92

Figure 92
Haemulon melanurum, 213.8 mm SL, photo by JT Williams.

Haemulon plumierii (Lacepède, 1801)white grunt; USNM, F, I, O, V; Figure 93

Figure 93
Haemulon plumierii, 244.7 mm SL, photo by JT Williams.

Haemulon striatum (Linnaeus, 1758)striped grunt; F; Figure 94

Figure 94
Haemulon striatum, 154 mm SL, photo by W Toller.

Inermiidae—bonnetmouths

Inermia vittata Poey, 1860boga; 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

Figure 95
Equetus punctatus, 146.8 mm SL, photo by JT Williams.

Pareques acuminatus (Bloch & Schneider, 1801)high-hat; USNM, I; Figure 96

Figure 96
Pareques acuminatus, 150.0 mm SL, photo by JT Williams.

Mullidae—goatfishes

Mulloidichthys martinicus (Cuvier, 1829)yellow goatfish; USNM, F, I, O; Figure 97

Figure 97
Mulloidichthys martinicus, 105 mm SL, photo by JT Williams.

Pseudupeneus maculatus (Bloch, 1793)spotted goatfish; USNM, I, O, V; Figure 98

Figure 98
Pseudupeneus maculatus, 218.8 mm SL, photo by JT Williams.

Chaetodontidae—butterflyfishes

Chaetodon capistratus Linnaeus, 1758foureye butterflyfish; USNM, I, O, V; Figure 99

Figure 99
Chaetodon capistratus, 99.4 mm SL; when younger, the specimen was apparently injured in the region of the fourth dorsal-fin spine (missing) and the area healed leaving a gap in the fin; photo by JT Williams.

Chaetodon ocellatus Bloch, 1787spotfin butterflyfish; USNM, F, O, V

Chaetodon sedentarius Poey, 1860reef butterflyfish; UF, O, V

Chaetodon striatus Linnaeus, 1758banded butterflyfish; USNM, I, O, V; Figure 100

Figure 100
Chaetodon striatus, 111.3 mm SL, photo by JT Williams.

Prognathodes aculeatus (Poey, 1860)longsnout butterflyfish; USNM, I, O, V; Figure 101

Figure 101
Prognathodes aculeatus, 62.9 mm SL, photo by JT Williams.

Pomacanthidae—angelfishes

Centropyge argi Woods & Kanazawa, 1951cherubfish; USNM, I, O, V; Figure 102

Figure 102
Centropyge argi, 33.8 mm SL, photo by JT Williams.

Holacanthus ciliaris (Linnaeus, 1758)queen angelfish; USNM, F, I, O, V

Holacanthus tricolor (Bloch, 1795)rock beauty; USNM, I, O, V; Figures 103, 104

Figure 103
Holacanthus tricolor, 19.3 mm SL, juvenile color pattern, photo by JT Williams.
Figure 104
Holacanthus tricolor, 40.1 mm SL, intermediate color pattern, photo by JT Williams.

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

Figure 105
Amblycirrhitus pinos, 22.6 mm SL, photo by JT Williams.

Pomacentridae-damselfishes

Chromis cyanea (Poey, 1860)blue chromis; USNM, I, O, V; Figure 106

Figure 106
Chromis cyanea, 67.5 mm SL, iridescent blue colors on body faded immediately after death, photo by JT Williams.

Chromis multilineata (Guichenot, 1853)brown chromis; USNM, I, O, V; Figure 107

Figure 107
Chromis multilineata, 56.3 mm SL, photo by JT Williams.

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

Figure 108
Stegastes partitus, 58.6 mm SL, black-tailed color morph, photo by JT Williams.
Figure 109
Stegastes partitus, 58.6 mm SL, yellow-tailed color morph, photo by JT Williams.

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

Figure 110
Stegastes planifrons, 82.7 mm SL, photo by JT Williams.

Labridae—wrasses

Bodianus rufus (Linnaeus, 1758)Spanish hogfish; O, V

Clepticus parrae (Bloch & Schneider, 1801)creole wrasse; USNM, I, O, V; Figure 111

Figure 111
Clepticus parrae, 59.8 mm SL, photo by JT Williams.

Doratonotus megalepis Günther, 1862dwarf wrasse; USNM, I, V; Figure 112

Figure 112
Doratonotus megalepis, 44.1 mm SL, photo by JT Williams.

Halichoeres bivittatus (Bloch, 1791)slippery dick; USNM, I, O, V; Figures 113, 114

Figure 113
Halichoeres bivittatus, 37.9 mm SL, juvenile/initial color phase, showing the black spot in the dorsal fin and a somewhat unusual orangish anal fin, photo by JT Williams.
Figure 114
Halichoeres bivittatus, 103.0 mm SL, terminal male color phase, with a distinctive red blotch on the side of the body above the pectoral fin, photo by JT Williams.

Halichoeres cyanocephalus (Bloch, 1791)yellowcheek wrasse; O, V

Halichoeres garnoti (Valenciennes, 1839)yellowhead wrasse; UF, USNM, I, O, V; Figures 115, 116, 117

Figure 115
Halichoeres garnoti, 46.4 mm SL, juvenile color phase, showing the characteristic blue stripe on a yellow body, photo by JT Williams.
Figure 116
Halichoeres garnoti, 118.1 mm SL, initial/terminal color phase, this specimen has almost completed the transition from the female initial phase into a terminal male, photo by JT Williams.
Figure 117
Halichoeres garnoti, 125.3 mm SL, terminal male color phase, this specimen has completed the transition from the female initial phase into a terminal male and shows the characteristic black bar and dark area over the caudal peduncle, photo by JT Williams. ...

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

Figure 118
Halichoeres pictus, 70.7 mm SL, juvenile color phase, photo by JT Williams.
Figure 119
Halichoeres pictus, 85.0 mm SL, terminal male color phase, photo by JT Williams.

Halichoeres poeyi (Steindachner, 1867)blackear wrasse; UF, USNM, I, O, V; Figure 120

Figure 120
Halichoeres poeyi, 87.9 mm SL, terminal male color phase, photo by JT Williams.

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

Figure 121
Thalassoma bifasciatum, 39.3 mm SL, juvenile/initial color phase, photo by JT Williams.
Figure 122
Thalassoma bifasciatum, 63.5 mm SL, terminal male color phase, photo by JT Williams.

Xyrichtys splendens Castelnau, 1855green razorfish; USNM, I, O; Figures 123, 124, 125, 126

Figure 123
Xyrichtys splendens, 15.1 mm SL, young juvenile color phase, photo by JT Williams.
Figure 124
Xyrichtys splendens, 22.1 mm SL, juvenile color phase, photo by JT Williams.
Figure 125
Xyrichtys splendens, 52.4 mm SL, initial color phase of female, photo by JT Williams.
Figure 126
Xyrichtys splendens, 72.5 mm SL, terminal male color phase, photo by JT Williams.

Scaridae—parrotfishes

Cryptotomus roseus Cope, 1871bluelip parrotfish; UF, USNM, I, O; Figure 127

Figure 127
Cryptotomus roseus, 61.5 mm SL, terminal male color phase, photo by JT Williams.

Scarus coelestinus Valenciennes, 1840midnight parrotfish; V

Scarus guacamaia Cuvier, 1829rainbow parrotfish; F; Figure 128

Figure 128
Scarus guacamaia, 647 mm SL, terminal male color phase, photo by W Toller.

Scarus iseri (Bloch, 1789)striped parrotfish; USNM, O, V

Scarus taeniopterus Desmarest, 1831princess parrotfish; USNM, I, O, V; Figure 129, 130

Figure 129
Scarus taeniopterus, 174.9 mm SL, initial female color phase, photo by JT Williams.
Figure 130
Scarus taeniopterus, 231.7 mm SL, terminal male color phase, photo by JT Williams.

Scarus vetula Bloch & Schneider, 1801queen parrotfish; O

Sparisoma atomarium (Poey, 1861)greenblotch parrotfish; USNM, I, O; Figure 131

Figure 131
Sparisoma atomarium, 13.7 mm SL, juvenile color phase, photo by JT Williams.

Sparisoma aurofrenatum (Valenciennes, 1840)redband parrotfish; USNM, I, O, V; Figures 132, 133, 134

Figure 132
Sparisoma aurofrenatum, 53.8 mm SL, juvenile color phase, photo by JT Williams.
Figure 133
Sparisoma aurofrenatum, 164.0 mm SL, initial female color phase, photo by JT Williams.
Figure 134
Sparisoma aurofrenatum, 164 mm SL, terminal male color phase, photo by JT Williams.

Sparisoma chrysopterum (Bloch & Schneider, 1801)redtail parrotfish; USNM, I, O; Figure 135

Figure 135
Sparisoma chrysopterum, 274 mm SL, terminal male color phase, photo by JT Williams.

Sparisoma radians (Valenciennes, 1840)bucktooth parrotfish; USNM, I, O, V; Figure 136

Figure 136
Sparisoma radians, 51.6 mm SL, juvenile color phase, photo by JT Williams.

Sparisoma viride (Bonnaterre, 1788)stoplight parrotfish; USNM, I, O, V; Figure 137

Figure 137
Sparisoma viride, 222.9 mm SL, initial female color phase, photo by JT Williams.

Tripterygiidae—triplefins

Enneanectes altivelis Rosenblatt, 1960 lofty triplefin; USNM, I; Figure 138

Figure 138
Enneanectes altivelis, 18.3 mm SL, photo by JT Williams.

Enneanectes atrorus Rosenblatt, 1960redeye triplefin; USNM, I

Enneanectes jordani (Evermann and Marsh, 1899) mimic triplefin; USNM, I

Dactyloscopidae-sand stargazers

Dactyloscopus tridigitatus Gill, 1859sand stargazer; USNM, I; Figures 139, 140

Figure 139
Dactyloscopus tridigitatus, 50.8 mm SL, photo by JT Williams.
Figure 140
Dactyloscopus tridigitatus, 50.8 mm SL, dorsal view, photo by JT Williams.

Gillellus uranidea Böhlke, 1968warteye stargazer; USNM, I; Figure 141

Figure 141
Gillellus uranidea, 27.7 mm SL, photo by JT Williams.

Platygillellus rubrocinctus (Longley, 1934)saddle stargazer; USNM, I; Figures 142, 143, 144

Figure 142
Platygillellus rubrocinctus, 24.1 mm SL, adult, photo by JT Williams.
Figure 143
Platygillellus rubrocinctus, 12.1 mm SL, lateral view of juvenile, photo by JT Williams.
Figure 144
Platygillellus rubrocinctus, 12.1 mm SL, dorsal view of juvenile, photo by JT Williams.

Blenniidae—combtooth blennies

Parablennius marmoreus (Poey, 1876)seaweed blenny; USNM, I

Labrisomidae—scaly blennies

Labrisomus gobio (Valenciennes, 1836)palehead blenny; USNM, I; Figure 145

Figure 145
Labrisomus gobio, 36.2 mm SL, photo by JT Williams.

Labrisomus haitiensis Beebe &Tee-Van, 1918longfin blenny; USNM, I; Figure 146

Figure 146
Labrisomus haitiensis, 18.8 mm SL, photo by JT Williams.

Malacoctenus boehlkei Springer, 1959diamond blenny; USNM, I; Figure 147

Figure 147
Malacoctenus boehlkei, 36.3 mm SL, photo by JT Williams.

Paraclinus grandicomis (Rosen, 1911)horned blenny; UF, USNM, I; Figure 148

Figure 148
Paraclinus grandicomis, 27.1 mm SL, photo by JT Williams.

Starksia atlantica Longley, 1934smootheye blenny; USNM, I; Figures 149, 150

Figure 149
Starksia atlantica, 9.5 mm SL, juvenile, photo by JT Williams.
Figure 150
Starksia atlantica, 15.1 mm SL, adult male, photo by JT Williams.

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, 1961blackcheek blenny; USNM, I; Figures 151, 152

Figure 151
Starksia lepicoelia, 20.2 mm SL, adult female, photo by JT Williams.
Figure 152
Starksia lepicoelia, 19.8 mm SL, adult male, photo by JT Williams.

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, 2003black spot blenny; USNM, I

Starksia nanodes Böhlke & Springer, 1961dwarf blenny; USNM, I; Figure 153

Figure 153
Starksia nanodes, 10.6 mm SL, adult male, photo by JT Williams.

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

Figure 154
Acanthemblemaria aspera, 14.0 mm SL, juvenile/female, photo by JT Williams.
Figure 155
Acanthemblemaria aspera, 19.7 mm SL, adult male, photo by JT Williams.
Figure 156
Acanthemblemaria aspera, 20.4 mm SL, yellow color morph (female?), photo by JT Williams.

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, 1900sailfin blenny; USNM, I; Figure 157

Figure 157
Emblemaria pandionis, 37.0 mm SL, adult female, photo by JT Williams.

Emblemariopsis cf signifer (Ginsburg, 1942)flagfin blenny; USNM, I; Figure 158

Figure 158
Emblemariopsis cf signifer, 16.2 mm SL, adult male, this specimen represents an undescribed species in the signifer species complex, photo by JT Williams.

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, 1955papillate clingfish; USNM, I; Figures 159, 160, 161

Figure 159
Acyrtus artius, 20.4 mm SL, dorsal view, photo by JT Williams.
Figure 160
Acyrtus artius, 20.4 mm SL, lateral view, photo by JT Williams.
Figure 161
Acyrtus artius, 20.4 mm SL, ventral view showing pelvic disk, photo by JT Williams.

Callionymidae—dragonets

Paradiplogrammus bairdi Jordan, 1888lancer dragonet; USNM, I, O; Figures 162, 163

Figure 162
Paradiplogrammus bairdi, 39.8 mm SL, lateral view of an adult male, photo by JT Williams.
Figure 163
Paradiplogrammus bairdi, 29.8 mm SL, dorsal view of a smaller adult male, photo by JT Williams.

Gobiidae-gobies

Coryphopterus dicrus Böhlke & Robins, 1960colon goby; USNM, I; Figure 164

Figure 164
Coryphopterus dicrus, 31.8 mm SL, photo by JT Williams.

Coryphopterus eidolon Böhlke & Robins, 1960pallid goby; USNM, I; Figure 165

Figure 165
Coryphopterus eidolon, 18.3 mm SL, photo by JT Williams.

Coryphopterus glaucofraenum Gill, 1863bridled goby; USNM, I, V; Figure 166

Figure 166
Coryphopterus glaucofraenum, 33.7 mm SL, photo by JT Williams.

Coryphopterus personatus (Jordan & Thompson, 1905)masked goby; USNM, I; Figure 167

Figure 167
Coryphopterus personatus, 19.0 mm SL, photo by JT Williams.

Coryphopterus thrix Böhlke & Robins, 1960bartail goby; USNM, I; Figure 168

Figure 168
Coryphopterus thrix, 26.8 mm SL, photo by JT Williams.

Elacatinus chancei (Beebe & Hollister 1933)shortstripe goby; USNM, I; Figure 169

Figure 169
Elacatinus chancei, 28.1 mm SL, photo by JT Williams.

Elacatinus evelynae (Böhlke & Robins, 1968)sharknose goby; USNM, I; Figure 170

Figure 170
Elacatinus evelynae, 27.7 mm SL, photo by JT Williams.

Elacatinus genie (Böhlke & Robins, 1968)cleaner goby; USNM, I

Evermannichthys metzelaari Hubbs, 1923 roughtail goby; USNM, I; Figure 171

Figure 171
Evermannichthys metzelaari, 21.0 mm SL, photo by JT Williams.

Gnatholepis thompsoni Jordan, 1904goldspot goby; USNM, I, V; Figure 172

Figure 172
Gnatholepis thompsoni, 31.5 mm SL, photo by JT Williams.

Lythrypnus elasson Böhlke & Robins, 1960dwarf goby; USNM, I; Figure 173

Figure 173
Lythrypnus elasson, 9.2 mm SL, adult male, photo by JT Williams.

Lythrypnus minimus Garzón & Acero P., 1988 pygmy goby; USNM, I; Figure 174

Figure 174
Lythrypnus minimus, 10.6 mm SL, adult male, photo by JT Williams.

Lythrypnus nesiotes Böhlke & Robins, 1960island goby; USNM, I; Figure 175

Figure 175
Lythrypnus nesiotes, 11.9 mm SL, adult male, photo by JT Williams.

Priolepis hipoliti (Metzelaar, 1922)rusty goby; USNM, I; Figures 176, 177

Figure 176
Priolepis hipoliti, 12.2 mm SL, female, photo by JT Williams.
Figure 177
Priolepis hipoliti, 18.7 mm SL, adult male, photo by JT Williams.

Psilotris batrachodes Böhlke, 1963toadfish goby; USNM, I; Figure 178

Figure 178
Psilotris batrachodes, 13.3 mm SL, adult, photo by JT Williams.

Psilotris boehlkei Greenfield, 1993yellowspot goby; I; Figure 179

Figure 179
Psilotris boehlkei, 26.5 mm SL, adult, photo by JT Williams.

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

Figure 180
Pycnomma roosevelti, 14.6 mm SL, adult, photo by JT Williams.

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

Figure 181
Risor ruber, 18.7 mm SL, adult, photo by JT Williams.

Acanthuridae—surgeonfishes

Acanthurus bahianus Castelnau, 1855ocean surgeon; USNM, I, O, V; Figure 182

Figure 182
Acanthurus bahianus, 128.2 mm SL, photo by JT Williams.

Acanthurus chirurgus (Bloch, 1787)doctorfish; USNM, I, O, V; Figure 183

Figure 183
Acanthurus chirurgus, 168.0 mm SL, photo by JT Williams.

Acanthurus coeruleus Bloch & Schneider, 1801blue tang; USNM, I, O, V; Figure 184

Figure 184
Acanthurus coeruleus, 146.2 mm SL, photo by JT Williams.

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

Figure 185
Euthynnus alletteratus, 355.1 mm FL, photo by JT Williams.

Scomberomorus regalis (Bloch, 1793)cero; USNM

Thunnus atlanticus (Lesson 1831)blackfin tuna; F; Figure 186

Figure 186
Thunnus atlanticus, 516 mm FL, photo by W Toller.

Caproidae—boarfishes

Antigonia capros Lowe, 1843deepbody boarfish; USNM, F; Figure 187

Figure 187
Antigonia capros, 37 mm SL, close-up of head, photo by W Toller.

Antigonia combatia Berry & Rathjen 1959shortspine 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

Figure 188
Bothus ocellatus, 95.8 mm SL, photo by JT Williams.

Trichopsetta ventralis (Goode & Bean, 1885)sash flounder; UF, T

Paralichthyidae—sand flounders

Citharicthys dinoceros Goode & Bean, 1886spined whiff; USNM, T

Cynoglossidae—tonguefishes

Symphurus arawak Robins & Randall, 1965Caribbean tonguefish; USNM, I; Figure 189

Figure 189
Symphurus arawak, 30.7 mm SL, photo by JT Williams.

Symphurus ommaspilus Böhlke, 1961ocellated tonguefish; USNM, I; Figure 190

Figure 190
Symphurus ommaspilus, 42.9 mm SL, photo by JT Williams.

Balistidae—triggerfishes

Balistes vetula Linnaeus, 1758queen 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

Figure 191
Xanthichthys ringens, 87.3 mm SL, photo by JT Williams.

Monacanthidae—filefishes

Aluterus scriptus (Osbeck, 1765)scrawled filefish; USNM, F, V

Cantherhines macrocerus (Hollard, 1853)whitespotted filefish; USNM, F; Figure 192

Figure 192
Cantherhines macrocerus, 273 mm SL, image flipped horizontally – right side is shown, photo by W Toller.

Cantherhines pullus (Ranzani, 1842)orangespotted filefish; O, V

Monacanthus ciliatus (Mitchill, 1818)fringed filefish; USNM, I, V; Figure 193

Figure 193
Monacanthus ciliatus, 33.4 mm SL, photo by JT Williams.

Monacanthus tuckeri Bean, 1906slender filefish; USNM, I; Figure 194

Figure 194
Monacanthus tuckeri, 32.3 mm SL, photo by JT Williams.

Ostraciidae—boxfishes

Acanthostracion polygonia Poey, 1876honeycomb 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

Figure 195
Lactophrys trigonus, 293.1 mm SL, photo by JT Williams.

Lactophrys triqueter (Linnaeus, 1758)smooth trunkfish; USNM, F, OBS, VIS; Figure 196

Figure 196
Lactophrys triqueter, 111.7 mm SL, photo by JT Williams.

Tetraodontidae—puffers

Canthigaster rostrata (Bloch, 1786)sharpnose puffer; USNM, I, O, V; Figure 197

Figure 197
Canthigaster rostrata, 50.9 mm SL, photo by JT Williams.

Sphoeroides spengleri (Bloch, 1785)bandtail puffer; USNM, I, O; Figure 198

Figure 198
Sphoeroides spengleri, 73.1 mm SL, photo by JT Williams.

Diodontidae—porcupinefishes

Chilomycterus antillarum Jordan & Rutter, 1897web burrfish; USNM, F; Figure 199

Figure 199
Chilomycterus antillarum, 180.0 mm SL, photo by JT Williams.

Diodon holocanthus Linnaeus, 1758balloonfish; USNM, I, V; Figure 200

Figure 200
Diodon holocanthus, 104.9 mm SL, photo by JT Williams.

Diodon hystrix Linnaeus, 1758porcupinefish; 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.

Table 1
Number of fishes recorded at well sampled sites in the Greater 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.

Figure 201
Actual species-accrual curve (black dots) for 38 dives collecting and identifying fish species on Saba Bank.

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.

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.

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.

Figure 202
Hierarchical clustering dendrogram of stations showing significant differences (black bars) among fish assemblages at deep stations versus middle-depth and shallow stations (SIMPROF, P<0.05).

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.

Footnotes

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.

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.

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