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Research Article
The marine fishes of St Eustatius Island, northeastern Caribbean: an annotated, photographic catalog
expand article infoDavid Ross Robertson, Carlos J. Estapé§, Allison M. Estapé§, Ernesto Peña, Luke Tornabene|, Carole C. Baldwin
‡ Smithsonian Tropical Research Institute, Balboa, Panama
§ Unaffiliated, Islamorada, United States of America
| University of Washington, Seattle, United States of America
¶ National Museum of Natural History, Smithsonian Institution, Washington, United States of America
Open Access

Abstract

Sint Eustatius (Statia) is a 21 km2 island situated in the northeastern Caribbean Sea. The most recent published sources of information on that island’s marine fish fauna is in two non-governmental organization reports from 2015–17 related to the formation of a marine reserve. The species-list in the 2017 report was based on field research in 2013–15 using SCUBA diving surveys, shallow “baited underwater video surveys” (BRUVs), and data from fishery surveys and scientific collections over the preceding century. That checklist comprised 304 species of shallow (mostly) and deep-water fishes. In 2017 the Smithsonian Deep Reef Observation Project surveyed deep-reef fishes at Statia using the crewed submersible Curasub. That effort recorded 120 species, including 59 new occurrences records. In March-May 2020, two experienced citizen scientists completed 62 SCUBA dives there and recorded 244 shallow species, 40 of them new records for Statia. The 2017–2020 research effort increased the number of species known from the island by 33.6% to 406. Here we present an updated catalog of that marine fish fauna, including voucher photographs of 280 species recorded there in 2017 and 2020. The Statia reef-fish fauna likely is incompletely documented as it has few small, shallow, cryptobenthic species, which are a major component of the regional fauna. A lack of targeted sampling is probably the major factor explaining that deficit, although a limited range of benthic marine habitats may also be contributing.

Keywords

biodiversity, checklist, faunal completeness, faunal structure, reef-associated bony fishes, SCUBA surveys, submersible surveys

Introduction

Sint Eustatius island, known locally as Statia, is a 21 km2 island in the northeastern Caribbean, and is one of the Leeward Islands in the Lesser Antilles. Until recently there were very few published accounts relating to the marine-fish fauna of Statia. The most comprehensive are represented by two non-governmental organization (NGO) environmental reports to the Statia government by van Kuijk et al. (2015) and Davies and Piontek (2016, 2017). Those two reports referred to only one older scientific publication, by Metzelaar (1919), relating to the fish fauna of that island, among other islands of the Dutch Caribbean. Davies and Piontek (2017) combined their own results from visual surveys with information from BRUV (Baited Remote Underwater Video) surveys by van Kuijk et al. (2015), and a variety of historical scientific collections and fisheries surveys to produce a general list of 307 species (modified to 304, see below), which included both deep- and shallow-water species. In this paper we use the results of deep-reef research using a crewed submersible in 2017 and shallow SCUBA surveys in 2020 to add to the checklist of the island’s marine fish fauna. We also include voucher photographs of most of the species observed and collected during those two surveys. In addition to representing vouchers for the species records, the photographs are intended for use by managers, citizen scientists, recreational divers and fishers who want to identify fishes they see and catch at Statia. Hopefully they will also stimulate future documentation of previously unreported species there. Finally, we compare aspects of the ecological structure of the Statia fauna to that of the regional, Greater Caribbean fauna to assess how complete the faunal inventory is for Statia.

Materials and methods

Study area

As one of the Dutch Caribbean islands, Statia sits among Saba, Sint Marten and St Kitts and Nevis (Figure 1) and shares a 200-m insular shelf with the last two islands (Suppl. material 1: Figure S1). Statia is surrounded by a narrow 200-m shelf, which is most extensive on the leeward, western side (Figure 2). The island has a limited diversity of marine habitats. It lacks large, deep embayments, particularly on the western side, that would otherwise provide sheltered locations for development of fringing and back-reef areas. Statia has little well-developed coral reef and most reef areas are of relatively low relief. Due to the general degree of exposure of the entire island to ocean swells it lacks any mangroves and has little in the way of seagrass beds, which are now dominated by a non-native species of Halodule (van Kuijk et al. 2015; Hoeksema 2016).

Figure 1. 

Location of Sint Eustatius. The Caribbean Sea, with the location of Sint Eustatius island indicated in the inset. Source: Hoetjes and Carpenter (2010: fig. 1).

The Caribbean Sea, with the location of Sint Eustatius island indicated in the inset. Source: Hoetjes and Carpenter (2010: fig. 1).

Data sources

Published species lists

A comprehensive set of species records came from two NGO studies, which were included in a report by Hoeksema (2016). van Kuijk et al. (2015) recorded 107 species during “baited underwater video surveys” (BRUVs) at 104 sites in shallow water (<30 m deep) scattered around all sides of the island in 2013. Davies and Piontek (2016, 2017) recorded 206 species during 38 of their own shallow, roving-diver surveys in 2015, and augmented that list with a list of species they extracted from historical literature, museum records (from major online aggregators, see below), photographs of fishes caught at the island that they obtained from various sources, and fisheries surveys. They added the species recorded by van Kuijk et al. (2015) to those they had seen and extracted from other sources to produce a combined list of 307 species.

Research in 2017 and 2020

In 2017 the Smithsonian Institution’s Deep Reef Observation Project (DROP) worked with the crewed submersible Curasub to make collections and observations on deep-reef fishes at Statia, to complement similar prior work at the Antillean islands of Dominica and Curaçao (e.g., Baldwin et al. 2018). The submersible was launched close to shore from the tender vessel R/V Chapman and towed by a surface boat to locations along the outer reef slope off the southwest coast where the shallow reef flat transitioned to the slope (~ 40–50 m). Eleven submersible dives were made off the southwestern edge of the island’s 200 m platform (see Figure 2, and Suppl. material 2: Table S1). Each dive lasted approximately five hours and reached a maximum depth of 143–305 m, depending on the habitat at that particular site. Submersible surveys follow the methods used by Baldwin et al. (2018). Dives were roving surveys with the submersible facing the reef and moving laterally while slowly descending the slope. Periodically, stops were made to collect specimens using an anesthetic (quinaldine in ethanol) ejection system attached to the sub’s manipulator arms, coupled with a suction pump attached to one arm that emptied into a holding chamber. On five of the eleven dives visual records of fishes were obtained by CB and LT, who were seated in the front of the submersible and linked their sightings of identifiable fishes to depth measurements recorded from a digital depth gauge inside the submersible. High-definition video was also recorded on five dives from a camera mounted on the front of the sub. Five scuba-based collection dives to a maximum depth of 20 m were also made by LT and CB, who were targeting sponge-associated gobies. A total of 210 specimens was collected, and 6475 individuals were recorded from visual observations during the SCUBA and submersible dives by DROP. Some of those specimens represent undescribed species or belong to groups with uncertain taxonomy.

Figure 2. 

Study sites at Sint Eustatius Island. Location of dive sites during 2017 and 2020: Black stars indicate submersible dives, blue stars 2017 SCUBA dives, red stars 2020 SCUBA dives (some individual stars indicate multiple dives in very close proximity), purple star an intertidal snorkeling site, and the red outline shows limits of the shore-diving area in 2020. See Suppl. material 2: Table S1 for georeferenced date on dive sites. Generalized 20 m, 30 m, 200 m and 500 m isobaths in blue; other lines indicate marine and terrestrial reserve areas. (Base map from Statiaparks, openstreemap.org, CC-BY-SA 2.0 with bathymetry data corrected from CARMABI/WWF/E.Imms (https://www.dcbd.nl/document/bathymetry-map-seas-surrounding-st-eustatius-saba-and-st-maarten, accessed 10 July 2020)

Two of the authors, CJE and AME, are citizen scientists with extensive experience photographing reef fishes at various sites in the Greater Caribbean. In 2020 they spent two months (mid-March to mid-May) living at Statia and SCUBA diving daily to obtain photographic vouchers of the fishes they observed. They made 62 dives, each of approximately one-hour duration, at depths between 1–30 m on both hard-reef, sand, rubble and seagrass habitats, as well as on sunken wrecked ships. Half of those dives were nearshore in a restricted area, as, during the second half of their stay at the island, they lacked dive-boat support and were able to dive only from the shoreline (see Figure 2, and Suppl. material 2: Table S1). During those dives CJE and AME accumulated photographs of the great majority of fish species they saw. They also obtained recent photographs of a few species taken by local divers and fishers at Statia that they did not see or photograph themselves.

Online aggregators

In addition, we also assessed information provided by three major aggregators of online georeferenced location data on marine fishes (GBIF https://www.gbif.org/, OBIS https://obis.org/, and FishNet2 http://www.fishnet2.net/search.aspx, all accessed on 7 May 2020), searching for records in ~ 120-km2 quadrat based on Admiralty Chart 487G that encompassed Statia and the surrounding shelf area: the area bounded by 17.433°N to 17.533°N and – 62.933°W to – 63.033°W. That quadrat contained almost 100 km2 of marine habitat. That area is a little larger than and centered on the area shown in Figure 2. Those sites regularly update the information they contain and might have had additional records to those found by Davies and Piontek (2017).

Location of dive sites during 2017 and 2020: Black stars indicate submersible dives, blue stars 2017 SCUBA dives, red stars 2020 SCUBA dives (some individual stars indicate multiple dives in very close proximity), purple star an intertidal snorkeling site, and the red outline shows limits of the shore-diving area in 2020. See Suppl. material 2: Table S1 for georeferenced date on dive sites. Generalized 20 m, 30 m, 200 m and 500 m isobaths in blue; other lines indicate marine and terrestrial reserve areas. (Base map from Statiaparks, openstreemap.org, CC-BY-SA 2.0 with bathymetry data corrected from CARMABI/WWF/E.Imms (https://www.dcbd.nl/document/bathymetry-map-seas-surrounding-st-eustatius-saba-and-st-maarten, accessed 10 July 2020)

The structure of the Statia reef-fish fauna

Zoogeography

Members of the entire Statia fauna as currently known (Table 1; hereafter Statia20) were assessed in terms of their global and local geographical ranges, as follows: (a) Endemism – we noted whether each is a Greater Caribbean endemic, or is distributed more widely in the tropical western Atlantic (i.e., to the north and south of the Greater Caribbean, or on both sides of the Atlantic, or in the Indo-Pacific as well as the Atlantic). (b) Geographical range size – we noted which species have small geographical ranges within the Greater Caribbean, which we defined as ranges that span no more than one third of the area of that region (based on maps of their ranges in Robertson and Van Tassell 2019).

Ecological structure

The research during 2017–2020 was aimed at documenting the reef-associated bony fishes of Statia. For analyses of the structure of the Statia20 fauna we assigned those species to the following ecological groups (following Robertson and Tornabene 2020): Reef-associated fishes include demersal and benthic species that use hard substrata (coral- and rock reefs), and soft bottoms (sand, gravel, mud, seagrass and macroalgal beds growing on sediment, estuaries and mangroves) immediately adjacent to or within the matrices of reefs. Benthic species are restricted to living on and in the bottom, while demersal species use both the bottom and the near-bottom water column. Cryptobenthic fishes are visually and/or behaviorally cryptic due to their form and coloration, and to their maintaining a close association with the benthos, directly on or within it. Small size (here maximum total length (TL) ≤10 cm) also is thought to be important for crypsis among such species. Core families of cryptobenthic reef fishes (Core CRFs) (see Brandl et al. 2018, 2019) found in the western Atlantic include the Apogonidae, Blenniidae, Bythitidae, Callionymidae, Chaenopsidae, Dactyloscopidae, Gobiesocidae, Gobiidae, Grammatidae, Labrisomidae, Opistognathidae, Syngnathidae, Tripterygiidae. To these families we added the Dinematichthyidae, which was split from the Bythitidae by Møller et al. (2016) shortly before Brandl et al. (2018) assembled their list of Core CRF families, and contains many shallow, reef-associated species. Species in the list are divided into two depth classes, based on their depth ranges: shallow species are those commonly found above 40 m depth, and deep species are those entirely or largely restricted to depths below 40 m.

In the Greater Caribbean region reef-associated bony fishes comprise ~ 900 species from 304 genera in 76 families (Robertson and Tornabene 2020). Reef-fish faunas of deep reefs down to ~ 250 m are dominated by the same set of families that are common on shallow reefs (Baldwin et al. 2018). At the regional level ~ 95% of those reef-associated species are non-pelagic, demersal and benthic forms, which were the focus of the 2017–2020 research at Statia. The relative abundance of the different ecological groups in the Statia20 fauna was compared to: (a) that of the regional fauna to assess similarities and differences; (b) that of the Statia fauna of Davies and Piontek (2017) (hereafter Statia17) to assess any changes; and (c) that of the Saba Exclusive Economic Zone (EEZ) (which includes Statia) (hereafter Saba17) prior to the 2017–2020 research to assess the identity and ecotypes of species that, although they are not on the Statia20 list, do occur very near Statia. Finally, we compare the relative abundances of the different ecogroups in the Statia20 fauna to those at one of the best sampled reefs in the Greater Caribbean, which has the largest published fauna: Alligator Reef in the Florida Keys (see Williams et al. 2010). The Alligator reef faunal checklist was recently updated and expanded (Starck et al. 2017; Estapé et al. 2020; hereafter Alligator20), and, hence, should provide a useful comparison.

A list of reef-associated fishes known from Alligator Reef was extracted from the list in Starck et al. (2017), and Estapé et al (2020) by comparing it to the checklist of regional reef-associated fishes of Robertson and Tornabene (2020). A faunal list for the Saba EEZ (see Suppl. material 1: Figure S1) was obtained by using the “Species List Assembly” tool in Robertson and Van Tassell (2019) (https://biogeodb.stri.si.edu/caribbean/en/research/index/list), as follows: within the tool the following combination of factors was selected – all species/ political area/ Saba EEZ. The confirmed species on the list generated (those with actual records within that EEZ) were then used here. A few species represented solely by data from the 2017–20 research at Statia that were on the Saba EEZ list generated by that tool were excluded from that list for the present comparisons.

Results

Modifications to the list of Davies and Piontek (2017)

We reduced the number of species on the list of Davies and Piontek (2017) (which is unchanged from that of Davies and Piontek 2016) from 307 to 304 through three deletions. Those included Emblemariopsis occidentalis Stephens, 1970, Pterois miles (Bennett, 1828) and Enneanectes pectoralis (Evermann & Marsh, 1899). Those authors recorded E. occidentalis and provided a photograph (on p 75 of Davies and Piontek 2016) of the fish they gave this name. However, E. occidentalis is now known to be restricted to the Bahamas (B Victor pers. comm., 26 May 2020). Authors CJE and AME photographed two species of this genus at Statia, E. bahamensis and E. carib. While E. carib (and E. occidentalis) has a simple ocular cirrus, E. bahamensis lacks such a cirrus. As the fish in Davies and Piontek’s (2016) photograph clearly has an ocular cirrus it cannot be E. bahamensis. B Victor (pers. comm., 26 May 2020) examined that photograph and concluded it is of either E. carib or possibly E leptocirris Stephens 1970, which has an ocular cirrus and is known from the Puerto Rican plateau, 185 km from Statia. Hence, we deleted E. occidentalis from the list but did not include E. leptocirris due to the uncertain identification of that photograph. The Indo-west Pacific lionfish P. volitans apparently is a hybrid of two Indo-west Pacific species, and the West Atlantic population of this lionfish appears to be composed almost entirely of P. volitans (Wilcox et al. 2018). Hence, we excluded P. miles from the list as it is unlikely to be present at Statia and any such an occurrence has not been confirmed genetically. Davies and Piontek (2017) included both Enneanectes pectoralis and E. jordani on the list. However, we excluded E. pectoralis as it recently has been shown to be a synonym of E. jordani (see Victor 2017). In addition, we changed the names for two of Davies and Piontek’s (2017) species: Davies and Piontek (2017) recorded L. campechanus (Poey, 1860), which is now known to be restricted to the Gulf of Mexico and US area. The taxonomic separation of L. purpureus, which ranges from the Caribbean to Brazil, from L. campechanus was recently confirmed by da Silva et al. (2020). Davies and Piontek (2017) recorded S. mitsukurii Jordan & Snyder, 1903. However, the Greater Caribbean population was recently renamed S. clarkae (see Ehemann et al. 2019) and S. mitsukurii is now regarded as restricted to the Eastern Atlantic and Indo-west Pacific. Those changes reduced the Statia17 list from 307 to 304 species.

Additions from other sources

The Van Kuijk et al. (2015) list of 106 species contained one species (Chilomycterus schoepfi) not included by Davies and Piontek (2017) in their list. FishNet2 supplies data based on museum records to GBIF and all 34 species records from FishNet2 were also in the GBIF list and are not separately indicated in Table 1. The GBIF list included 103 species, and, after discounting the 27 DROP2017 collection records included therein, none of the 76 remaining species represented “new” records that are not on the Davies and Piontek (2017) list. OBIS, which also supplies data to GBIF, produced 37 records, 13 of which (all common, widely distributed species) were not in the GBIF list, but only one of which (Coryphaena hippurus) was not in any other database.

DROP recorded a total of 120 species, 59 of which were not in any other list, except for two new records it shared with the Estapé 2020 list. Eight of those 59 records are of species that have yet to be described and named. The Estapé 2020 list includes 244 records, 40 of them new, plus two other new additions they share with DROP. Summing the deletions and additions from various sources produced a total of 406 species for the Statia20 checklist (see Table 1).

Table 1.

Updated checklist of marine fishes from Sint Eustatius Island, 2020. Key to column headings and entries: DROP – CP = collected and photographed; C collected only; V = visual observation only; Estapé – P = photographed by CJE and AME; (P) photographed by 3rd parties; V = visual observation only by CJE and AME. New – species is a new record resulting from 2017–20 research, and its source. Other sources of species records are van Kuijk et al. 2015 (vK15), Davies and Piontek 2017 (DP17), GBIF, and OBIS. DROP in GBIF indicates record in GBIF is derived from 2017 DROP collection specimens deposited in the fish collection of the US National Museum of Natural History. FishNet 2 records are not indicated separately because all such records are included by GBIF. NA- not applicable to non-native Pterois volitans. Plate – number indicates supplemental plate containing the voucher photograph of that species. Zoogeography (Zoo)- Global geographic range of species; GC = Greater Caribbean endemic; NWA = GC plus temperate eastern USA; WA = GC plus Brazil; TA = WA plus central or East Atlantic; PAC = Pacific; EP = East Pacific; IWP = Indo-west Pacific; PAN = Pantropical or Circumglobal. Range – extent of geographic range – L = range limited, not more than one third of the Greater Caribbean; remainder are more widely distributed in that region. Deep – species entirely or largely restricted to depths below 40 m. Yes indicates a species conforms to the heading of the column; ? indicates insufficient data.

Species in families English common name New DROP Estapé vK15 DP17 GBIF OBIS Plate Zoo Range Deep
ACANTHURIDAE
Acanthurus chirurgus (Bloch, 1787) Doctorfish V P Yes Yes Yes Yes 1 GC
Acanthurus coeruleus Bloch & Schneider, 1801 Blue Tang V P Yes Yes Yes Yes 1 GC
Acanthurus tractus Poey, 1860 Northern Ocean Surgeonfish V P Yes Yes Yes Yes 1 GC
ACHIRIDAE
Gymnachirus nudus Kaup, 1858 Flabby Sole Estapé P 1 GC
ACROPOMATIDAE
Synagrops bellus (Goode & Bean, 1896) Blackmouth Bass Yes WA Yes
AETOBATIDAE
Aetobatus narinari (Euphrasen, 1790) Spotted Eagle Ray (P) Yes Yes 1 WA
ANTENNARIIDAE
Antennarius multiocellatus (Valenciennes, 1837) Longlure Frogfish P Yes 1 WA
Histrio histrio (Linnaeus, 1758) Sargassumfish (P) Yes 1 PAN
APOGONIDAE
Apogon aurolineatus (Mowbray, 1927) Bridle Cardinalfish Yes GC
Apogon maculatus (Poey, 1860) Flamefish P Yes 1 GC
Apogon pillionatus Bohlke & Randall, 1968 Broadsaddle Cardinalfish DROP V GC
Apogon planifrons Longley & Hildebrand, 1940 Pale Cardinalfish Estapé P 1 WA
Apogon pseudomaculatus Longley, 1932 Twospot Cardinalfish DROP C DROP WA
Apogon quadrisquamatus Longley, 1934 Sawcheek Cardinalfish P Yes 1 WA
Apogon townsendi (Breder, 1927) Belted Cardinalfish P Yes 1 WA
Astrapogon puncticulatus (Poey, 1867) Blackfin Cardinalfish Estapé V WA
Astrapogon stellatus (Cope, 1867) Conchfish Yes WA
Paroncheilus affinis (Poey, 1875) Bigtooth Cardinalfish V P Yes 1 TA
Phaeoptyx conklini (Silvester, 1915) Freckled Cardinalfish Estapé P 1 GC
Phaeoptyx pigmentaria (Poey, 1860) Dusky Cardinalfish Yes 1 TA
ARGENTINIDAE
Argentina stewarti Cohen & Atsaides, 1969 Yes GC Yes
Glossanodon pygmaeus Cohen, 1958 Pygmy Argentine DROP CP 1 WA Yes
ATHERINIDAE
Atherina harringtonensis Goode, 1877 Reef Silverside Yes GC
Atherinomorus stipes (Müller & Troschel, 1848) Hardhead Silverside Estapé P 1 WA
AULOSTOMIDAE
Aulostomus maculatus Valenciennes, 1841 Atlantic Trumpetfish P Yes Yes Yes 1 GC
BALISTIDAE
Balistes capriscus Gmelin, 1789 Gray Triggerfish P Yes 1 TA
Balistes vetula Linnaeus, 1758 Queen Triggerfish P Yes Yes Yes 1 TA
Canthidermis sufflamen (Mitchill, 1815) Ocean Triggerfish V Yes WA
Melichthys niger (Bloch, 1786) Black Durgon P Yes Yes Yes Yes 1 PAN
Xanthichthys ringens (Linnaeus, 1758) Sargassum Triggerfish DROP V WA
BELONIDAE
Platybelone argalus argalus (Lesueur, 1821) Keeltail Needlefish Yes WA
Tylosurus crocodilus (Péron & Lesueur, 1821) Houndfish P Yes 1 PAN
BLENNIIDAE
Entomacrodus nigricans Gill, 1859 Pearl Blenny P Yes 1 GC
Hypleurochilus pseudoaequipinnis Bath, 1994 Oyster Blenny Estapé P 1 WA
Hypleurochilus springeri Randall, 1966 Orangespotted Blenny Estapé P 1 GC
Hypsoblennius exstochilus Bohlke, 1959 Longhorn Blenny (P) Yes 2 GC
Ophioblennius macclurei (Silvester, 1915) Redlip Blenny P Yes Yes 2 GC
Parablennius marmoreus (Poey, 1876) Seaweed Blenny P Yes 2 WA
BOTHIDAE
Bothus lunatus (Linnaeus, 1758) Peacock Flounder P Yes Yes 2 TA
Bothus ocellatus (Agassiz, 1831) Eyed Flounder P Yes 2 WA
Chascanopsetta lugubris Alcock, 1894 Pelican Flounder Yes TA,IWP Yes
CALLIONYMIDAE
Callionymus bairdi (Jordan, 1888) Lancer Dragonet P Yes 2 WA
Foetorepus species DROP CP 13 WA? ? Yes
CAPROIDAE
Antigonia capros Lowe, 1843 Deepbody Boarfish DROP V TA,IWP Yes
CARANGIDAE
Alectis ciliaris (Bloch, 1787) African Pompano Yes PAN
Caranx bartholomaei (Cuvier, 1833) Yellow Jack P Yes 2 TA
Caranx crysos (Mitchill, 1815) Blue Runner P Yes 2 TA
Caranx hippos (Linnaeus, 1766) Crevalle Jack Yes WA
Caranx latus Agassiz, 1831 Horse-eye Jack P Yes Yes 2 TA
Caranx lugubris Poey, 1860 Black Jack V Yes Yes PAN
Caranx ruber (Bloch, 1793) Bar Jack V P Yes Yes Yes Yes 2 WA
Decapterus macarellus (Cuvier, 1833) Mackerel Scad P Yes 2 PAN
Decapterus punctatus (Cuvier, 1829) Round Scad P Yes 2 TA
Elagatis bipinnulata (Quoy & Gaimard, 1825) Rainbow Runner P Yes 2 PAN
Selar crumenophthalmus (Bloch, 1793) Bigeye Scad P Yes 2 PAN
Seriola rivoliana Valenciennes, 1833 Almaco Jack P Yes Yes 2 PAN
Trachinotus falcatus (Linnaeus, 1758) Permit P Yes 2 WA
Trachinotus goodei Jordan & Evermann, 1896 Palometa P Yes 2 WA
CARCHARHINIDAE
Carcharhinus leucas (Müller & Henle, 1839) Bull Shark Yes PAN
Carcharhinus limbatus (Müller & Henle, 1839) Blacktip Shark Yes Yes PAN
Carcharhinus perezii (Poey, 1876) Reef Shark V Yes Yes WA
Galeocerdo cuvier (Peron & Lesueur, 1822) Tiger Shark Yes PAN
Negaprion brevirostris (Poey, 1868) Lemon Shark P Yes 2 TA,EP
CENTROPHORIDAE
Centrophorus granulosus (Bloch & Schneider, 1801) Large Gulper Shark Yes TA,IWP Yes
CHAENOPSIDAE
Acanthemblemaria aspera (Longley, 1927) Roughhead Blenny P Yes 2 GC
Acanthemblemaria maria Bohlke, 1961 Secretary Blenny P Yes Yes 2 GC
Acanthemblemaria spinosa Metzelaar, 1919 Spinyhead Blenny P Yes Yes 2 GC
Chaenopsis limbaughi Robins & Randall, 1965 Yellowface Pikeblenny P Yes 2 GC
Emblemaria pandionis Evermann & Marsh, 1900 Sailfin Blenny P Yes Yes 2 GC
Emblemaria vitta Williams, 2002 Ribbon Blenny Estapé (P) 2 GC
Emblemariopsis bahamensis Stephens, 1961 Blackhead Blenny Estapé P 3 GC L
Emblemariopsis carib Victor, 2010 Carib Blenny Estapé P 3 GC L
CHAETODONTIDAE
Chaetodon capistratus Linnaeus, 1758 Foureye Butterflyfish V P Yes Yes Yes Yes 3 GC
Chaetodon ocellatus Bloch, 1787 Spotfin Butterflyfish P Yes Yes Yes 3 WA
Chaetodon sedentarius Poey, 1860 Reef Butterflyfish V Yes Yes Yes WA
Chaetodon striatus Linnaeus, 1758 Banded Butterflyfish V P Yes Yes Yes Yes 3 WA
Prognathodes aculeatus (Poey, 1860) Longsnout Butterflyfish C P Yes Yes Yes Yes 3 WA
Prognathodes guyanensis (Durand, 1960) Guyana Butterflyfish DROP V GC Yes
CHAUNACIDAE
Chaunax suttkusi Caruso, 1989 Pale-cavity Gaper Yes TA Yes
CHIMAERIDAE
Chimaera cubana Howell Rivero, 1936 Cuban Chimaera Yes GC Yes
Hydrolagus alberti Bigelow & Schroeder, 1951 Gulf Chimaera Yes GC Yes
CHLOPSIDAE
Chilorhinus suensonii Lutken, 1852 Seagrass Eel Yes WA
CHLOROPHTHALMIDAE
Chlorophthalmus agassizi Bonaparte, 1840 Shortnose Greeneye Yes TA Yes
Parasudis truculenta (Goode & Bean, 1895) Longnose Greeneye Yes WA Yes
CIRRHITIDAE
Amblycirrhitus pinos (Mowbray, 1927) Redspotted Hawkfish P Yes Yes 3 WA
CLUPEIDAE
Harengula clupeola (Cuvier, 1829) False Pilchard Yes WA
Harengula humeralis (Cuvier, 1829) Redear Sardine Yes GC
Jenkinsia lamprotaenia (Gosse, 1851) Dwarf Herring Yes GC
Opisthonema oglinum (Lesueur, 1818) Atlantic Thread Herring Yes WA
Sardinella aurita Valenciennes, 1847 Spanish Sardine Yes TA
CONGRIDAE
Ariosoma balearicum (Delaroche, 1809) Bandtooth Conger Estapé (P) 3 TA
Heteroconger longissimus Gunther, 1870 Brown Garden Eel P Yes Yes Yes 3 WA
Xenomystax bidentatus (Reid, 1940) Twopatched-teeth Conger Yes TA Yes
CORYPHAENIDAE
Coryphaena hippurus Linnaeus, 1758 Dolphinfish Yes PAN
CRURIRAJIDAE
Cruriraja rugosa Bigelow & Schroeder, 1958 Rough Leg Skate Yes GC Yes
CYNOGLOSSIDAE
Symphurus marginatus (Goode & Bean, 1886) Margined Tonguefish Yes WA Yes
DACTYLOPTERIDAE
Dactylopterus volitans (Linnaeus, 1758) Flying Gurnard P Yes Yes Yes 3 TA
DASYATIDAE
Hypanus americanus Hildebrand & Schroeder, 1928 Southern Stingray P Yes Yes Yes 3 WA
DIODONTIDAE
Chilomycterus antillarum Jordan & Rutter, 1897 Web Burrfish P Yes Yes 3 WA
Chilomycterus schoepfii (Walbaum, 1792) Striped Burrfish Yes NWA
Diodon holocanthus Linnaeus, 1758 Balloonfish P Yes 3 PAN
Diodon hystrix Linnaeus, 1758 Porcupinefish P Yes Yes Yes 3 PAN
DIRETMIDAE
Diretmus argenteus Johnson, 1864 Silver Spinyfish Yes PAN Yes
ECHENEIDAE
Echeneis naucrates Linnaeus, 1758 Sharksucker P Yes Yes Yes 3 PAN
Echeneis neucratoides Zuiew, 1786 Whitefin Sharksucker Estapé P 3 NWA
Remora remora (Linnaeus, 1758) Remora Yes PAN
EPHIPPIDAE
Chaetodipterus faber (Broussonet, 1782) Atlantic Spadefish Yes WA
ETMOPTERIDAE
Etmopterus hillianus (Poey, 1861) Caribbean Lantern Shark Yes NWA Yes
Etmopterus robinsi Schofield & Burgess, 1997 West Indian Lantern Shark Yes GC Yes
FISTULARIIDAE
Fistularia tabacaria Linnaeus, 1758 Bluespotted Cornetfish P Yes Yes 3 TA
GERREIDAE
Eucinostomus jonesii (Gunther, 1879) Slender Mojarra Yes WA
Eucinostomus lefroyi (Goode, 1874) Mottled Mojarra P Yes 3 WA
Gerres cinereus (Walbaum, 1792) Yellowfin Mojarra Yes WA
GINGLYMOSTOMATIDAE
Ginglymostoma cirratum (Bonnaterre, 1788) Nurse Shark (P) Yes Yes Yes 3 TA
GOBIESOCIDAE
Derilissus lombardii Sparks & Gruber, 2012 Tailspot Clingfish DROP CP 3 GC Yes
GOBIIDAE
Antilligobius nikkiae Van Tassell & Colin, 2012 Sabre Goby DROP CP DROP 3 GC Yes
Bathygobius antilliensis Tornabene, Baldwin & Pezold, 2010 Antilles Frillfin Estapé P 3 GC
Coryphopterus dicrus Bohlke & Robins, 1960 Colon Goby P Yes 3 WA
Coryphopterus eidolon Bohlke & Robins, 1960 Pallid Goby P Yes 3 GC
Coryphopterus glaucofraenum Gill, 1863 Bridled Goby Yes WA
Coryphopterus hyalinus Bohlke & Robins, 1962 Glass Goby P Yes 4 GC
Coryphopterus kuna Victor, 2007 Kuna Goby Estapé P 4 GC
Coryphopterus lipernes Bohlke & Robins, 1962 Peppermint Goby P Yes 4 GC
Coryphopterus personatus (Jordan & Thompson, 1905) Masked Goby V P Yes 4 GC
Coryphopterus thrix Bohlke & Robins, 1960 Bartail Goby P Yes 4 WA
Coryphopterus tortugae (Jordan, 1904) Sand Goby P Yes 4 GC
Coryphopterus venezuelae Cervigon, 1966 Sand-Canyon Goby Estapé P 4 GC
Ctenogobius saepepallens (Gilbert & Randall, 1968) Dash Goby Estapé P 4 GC
Elacatinus chancei (Beebe & Hollister, 1933) Shortstripe Goby C P Yes Yes 4 GC L
Elacatinus evelynae (Bohlke & Robins, 1968) Sharknose Goby P Yes Yes 4 GC
Genus 1 species 5 DROP CP 13 GC? ? Yes
Genus 1 species 6 DROP CP 13 GC? ? Yes
Genus 2 species 1 DROP CP 13 GC? ? Yes
Ginsburgellus novemlineatus (Fowler, 1950) Ninelined Goby Estapé P 4 GC
Gnatholepis thompsoni Jordan, 1904 Goldspot Goby V P Yes Yes 4 TA
Lythrypnus elasson Bohlke & Robins, 1960 Dwarf Goby DROP/ Estapé C P 4 GC
Microgobius carri Fowler, 1945 Seminole Goby Estapé P 4 WA
Nes longus (Nichols, 1914) Orangespotted Goby P Yes 4 GC
Palatogobius grandoculus Greenfield, 2002 Bigeye Goby DROP CP DROP 4 GC Yes
Palatogobius incendius Tornabene, Robertson & Baldwin, 2017 Ember Goby DROP C DROP GC Yes
Pinnichthys aimoriensis Van Tassell & Tornabene, 2016 Thiony’s Goby DROP CP 4 GC Yes
Priolepis hipoliti (Metzelaar, 1922) Rusty Goby P Yes 4 WA
Ptereleotris helenae (Randall, 1968) Hovering Dartfish V P Yes 4 GC
Risor ruber (Rosen, 1911) Tusked Goby C P Yes Yes 4 WA
Tigrigobius dilepis (Robins & Bohlke, 1964) Orangesided Goby P Yes 4 GC
Tigrigobius multifasciatus (Steindachner, 1876) Greenbanded Goby Estapé P 4 GC L
Varicus cephalocellatus Gilmore, Van Tassell & Baldwin, 2016 Ocellated Split-Fin Goby DROP CP DROP 4 GC L Yes
Varicus veliguttatus Van Tassell, Baldwin & Gilmore, 2016 Spotted-Sail Goby DROP CP DROP 4 GC Yes
GRAMMATIDAE
Gramma linki Starck & Colin, 1978 Yellowcheek Basslet DROP CP DROP 5 GC
Gramma loreto Poey, 1868 Fairy Basslet P Yes Yes 5 GC
Lipogramma evides Robins & Colin, 1979 Banded Basslet DROP CP DROP 5 GC Yes
Lipogramma klayi Randall, 1963 Bicolor Basslet DROP CP 5 GC Yes
Lipogramma levinsoni Baldwin, Nonaka & Robertson, 2016 Hourglass Basslet DROP CP 5 GC Yes
Lipogramma regia Robins & Colin, 1979 Royal Basslet DROP CP DROP 5 GC Yes
Lipogramma trilineata Randall, 1963 Threeline Basslet DROP CP DROP 5 GC Yes
GRAMMICOLEPIDIDAE
Grammicolepis brachiusculus Poey, 1873 Thorny Tinselfish Yes PAN Yes
HAEMULIDAE
Anisotremus surinamensis (Bloch, 1791) Black Margate P Yes Yes 5 WA
Brachygenys chrysargyreum (Gunther, 1859) Smallmouth Grunt P Yes Yes Yes 5 GC
Haemulon album Cuvier, 1830 Margate P Yes Yes 5 WA
Haemulon aurolineatum Cuvier, 1830 Tomtate P Yes Yes Yes 5 WA
Haemulon carbonarium Poey, 1860 Caesar Grunt P Yes Yes Yes Yes 5 GC
Haemulon flavolineatum (Desmarest, 1823) French Grunt P Yes Yes Yes Yes 5 GC
Haemulon macrostomum Gunther, 1859 Spanish Grunt Yes GC
Haemulon melanurum (Linnaeus, 1758) Cottonwick P Yes 5 WA
Haemulon parra (Desmarest, 1823) Sailors Choice Yes WA
Haemulon plumierii (Lacepede, 1801) White Grunt P Yes 5 WA
Haemulon sciurus (Shaw, 1803) Bluestriped Grunt (P) Yes Yes 5 GC
Haemulon striatum (Linnaeus, 1758) Striped Grunt V V Yes WA
Haemulon vittatum (Poey, 1860) Boga P Yes 5 GC
HALOSAURIDAE
Halosaurus ovenii Johnson, 1864 Stripejaw Halosaur Yes TA,IWP Yes
HEMIRAMPHIDAE
Hemiramphus brasiliensis (Linnaeus, 1758) Ballyhoo P Yes 5 WA
HOLOCENTRIDAE
Corniger spinosus Agassiz, 1831 Spinycheek Soldierfish DROP V TA Yes
Holocentrus adscensionis (Osbeck, 1765) Squirrelfish V P Yes Yes Yes 5 TA
Holocentrus rufus (Walbaum, 1792) Longspine Squirrelfish V P Yes Yes Yes 5 GC
Myripristis jacobus Cuvier, 1829 Blackbar Soldierfish V P Yes Yes 5 TA
Neoniphon coruscum (Poey, 1860) Reef Squirrelfish P Yes 5 GC
Neoniphon marianus (Cuvier, 1829) Longjaw Squirrelfish C P Yes Yes 5 GC
Neoniphon vexillarium (Poey, 1860) Dusky Squirrelfish P Yes 5 GC
Ostichthys trachypoma (Gunther, 1859) Bigeye Soldierfish DROP CP DROP 6 WA Yes
Plectrypops retrospinis (Guichenot, 1853) Cardinal Soldierfish Estapé P 6 WA
ISTIOPHORIDAE
Istiophorus platypterus (Shaw, 1792) Sailfish Yes TA
Makaira nigricans Lacepede, 1802 Blue Marlin Yes PAN
KYPHOSIDAE
Kyphosus bigibbus Lacepede, 1801 Gray Seachub Estapé P 6 TA/IWP
Kyphosus cinerascens (Forsskal, 1775) Topsail Seachub P Yes 6 PAN
Kyphosus sectatrix (Linnaeus, 1766) Bermuda Chub P Yes 6 PAN
Kyphosus vaigiensis (Quoy & Gaimard, 1825) Yellow Chub V Yes PAN
LABRIDAE
Labrinae
Bodianus rufus (Linnaeus, 1758) Spanish Hogfish V P Yes Yes Yes 6 WA
Clepticus parrae (Bloch & Schneider, 1801) Creole Wrasse V P Yes Yes Yes 6 GC
Decodon puellaris (Poey, 1860) Red Hogfish DROP CP DROP 6 WA Yes
Decodon species 2 DROP CP 13 GC Yes
Halichoeres bathyphilus (Beebe & Tee-Van,1932) Greenband Wrasse DROP V GC Yes
Halichoeres bivittatus (Bloch, 1791) Slippery Dick P Yes Yes 6 WA
Halichoeres cyanocephalus (Bloch, 1791) Yellowcheek Wrasse P Yes Yes 6 GC
Halichoeres garnoti (Valenciennes, 1839) Yellowhead Wrasse V P Yes Yes Yes 6 GC
Halichoeres maculipinna (Müller & Troschel, 1848) Clown Wrasse P Yes Yes 6 GC
Halichoeres pictus (Poey, 1860) Rainbow Wrasse P Yes 6 GC
Halichoeres poeyi (Steindachner, 1867) Blackear Wrasse P Yes Yes 6 WA
Halichoeres radiatus (Linnaeus, 1758) Puddingwife P Yes Yes 6 WA
Thalassoma bifasciatum (Bloch, 1791) Bluehead V P Yes Yes Yes 6 GC
Xyrichtys martinicensis Valenciennes, 1840 Rosy Razorfish P Yes Yes 6 GC
Xyrichtys novacula (Linnaeus, 1758) Pearly Razorfish P Yes 6 WA
Xyrichtys splendens Castelnau, 1855 Green Razorfish P Yes Yes Yes 6 GC
Scarinae
Cryptotomus roseus Cope, 1871 Bluelip Parrotfish P Yes 6 WA
Scarus coeruleus (Bloch, 1786) Blue Parrotfish Yes Yes GC
Scarus guacamaia Cuvier, 1829 Rainbow Parrotfish Yes GC
Scarus iseri (Bloch, 1789) Striped Parrotfish P Yes Yes Yes 6 GC
Scarus taeniopterus Desmarest, 1831 Princess Parrotfish V P Yes Yes Yes Yes 6 GC
Scarus vetula Bloch & Schneider, 1801 Queen Parrotfish P Yes Yes Yes Yes 6 GC
Sparisoma atomarium (Poey, 1861) Greenblotch Parrotfish P Yes 6 GC
Sparisoma aurofrenatum (Valenciennes, 1840) Redband Parrotfish V P Yes Yes Yes Yes 7 GC
Sparisoma chrysopterum (Bloch & Schneider, 1801) Redtail Parrotfish P Yes Yes Yes 7 GC
Sparisoma radians (Valenciennes, 1840) Bucktooth Parrotfish P Yes 7 WA
Sparisoma rubripinne (Valenciennes, 1840) Yellowtail Parrotfish P Yes Yes 7 GC
Sparisoma viride (Bonnaterre, 1788) Stoplight Parrotfish V P Yes Yes Yes 7 GC
LABRISOMIDAE
Brockius nigricinctus Howell Rivero, 1936 Spotcheek Blenny Estapé P 7 GC
Gobioclinus bucciferus Poey, 1868 Puffcheek Blenny Estapé P 7 GC
Gobioclinus gobio (Valenciennes, 1836) Palehead Blenny Estapé P 7 WA
Gobioclinus guppyi (Norman, 1922) Mimic Blenny Estapé P 7 WA
Labrisomus nuchipinnis (Quoy & Gaimard, 1824) Hairy Blenny P Yes Yes 7 TA
Malacoctenus aurolineatus Smith, 1957 Goldline Blenny P Yes 7 GC
Malacoctenus boehlkei Springer, 1959 Diamond Blenny Yes GC
Malacoctenus erdmani Smith, 1957 Imitator Blenny Estapé P 7 GC
Malacoctenus macropus (Poey, 1868) Rosy Blenny Estapé P 7 GC
Malacoctenus triangulatus Springer, 1959 Saddled Blenny P Yes 7 GC
LOBOTIDAE
Lobotes surinamensis (Bloch, 1790) Atlantic Tripletail Yes TA/IWP
LOPHIIDAE
Lophiodes monodi Le Danois, 1971 Club-bait Goosefish Yes GC Yes
LUTJANIDAE
Apsilus dentatus Guichenot, 1853 Black Snapper Yes GC Yes
Etelis oculatus (Valenciennes, 1828) Queen Snapper Yes WA Yes
Lutjanus analis (Cuvier, 1828) Mutton Snapper P Yes Yes 7 WA
Lutjanus apodus (Walbaum, 1792) Schoolmaster V P Yes Yes Yes Yes 7 GC
Lutjanus buccanella (Cuvier, 1828) Blackfin Snapper V P Yes 7 WA
Lutjanus cyanopterus (Cuvier, 1828) Cubera Snapper P Yes 7 WA
Lutjanus griseus (Linnaeus, 1758) Gray Snapper (P) Yes Yes 7 TA
Lutjanus jocu (Bloch & Schneider, 1801) Dog Snapper P Yes Yes 7 TA
Lutjanus mahogoni (Cuvier, 1828) Mahogany Snapper V P Yes Yes Yes Yes 7 GC
Lutjanus purpureus (Poey, 1866) Caribbean Red Snapper Yes TA
Lutjanus synagris (Linnaeus, 1758) Lane Snapper P Yes Yes Yes 7 TA
Lutjanus vivanus (Cuvier, 1828) Silk Snapper Yes TA Yes
Ocyurus chrysurus (Bloch, 1791) Yellowtail Snapper P Yes Yes Yes 7 TA
Pristipomoides sp.1 V WA? ? Yes
MACROURIDAE
Gadomus arcuatus (Goode & Bean, 1886) Doublethread Grenadier Yes TA Yes
Gadomus dispar (Vaillant, 1888) Onelong Grenadier Yes TA Yes
Hymenocephalus aterrimus Gilbert, 1905 Nobeard Grenadier Yes WA/PAC Yes
Hymenocephalus billsam Marshall & Iwamoto, 1973 Bigeye Grenadier Yes WA Yes
Malacocephalus laevis (Lowe, 1843) Velvet Grenadier Yes PAN Yes
Nezumia aequalis (Günther, 1878) Atlantic Blacktip Grenadier Yes TA Yes
Ventrifossa macropogon Marshall, 1973 Longbeard Grenadier Yes WA/WPAC Yes
MALACANTHIDAE
Malacanthus plumieri (Bloch, 1786) Sand Tilefish V P Yes Yes Yes 7 WA
MEGALOPIDAE
Megalops atlanticus Valenciennes, 1847 Tarpon P Yes 8 TA
MERLUCCIIDAE
Steindachneria argentea Goode & Bean, 1896 Luminous Hake Yes GC Yes
MONACANTHIDAE
Aluterus scriptus (Osbeck, 1765) Scrawled Filefish P Yes Yes 8 PAN
Cantherhines macrocerus (Hollard, 1853) Whitespotted Filefish P Yes Yes Yes Yes 8 WA
Cantherhines pullus (Ranzani, 1842) Orangespotted Filefish P Yes Yes Yes 8 TA
Monacanthus ciliatus (Mitchill, 1818) Fringed Filefish P Yes Yes 8 TA
Monacanthus tuckeri Bean, 1906 Slender Filefish P Yes Yes 8 GC
Stephanolepis setifer (Bennett, 1831) Pygmy Filefish P Yes Yes 8 WA
MUGILIDAE
Mugil curema Valenciennes, 1836 White Mullet Yes TA
MULLIDAE
Mulloidichthys martinicus (Cuvier, 1829) Yellow Goatfish V P Yes Yes Yes 8 TA
Pseudupeneus maculatus (Bloch, 1793) Spotted Goatfish V P Yes Yes Yes 8 WA
MURAENIDAE
Echidna catenata (Bloch, 1795) Chain Moray P Yes 8 WA
Enchelycore carychroa Bohlke & Bohlke, 1976 Chestnut Moray Estapé (P) 8 TA
Enchelycore nigricans (Bonnaterre, 1788) Viper Moray Estapé (P) 8 TA
Gymnothorax funebris Ranzani, 1839 Green Moray P Yes Yes 8 TA
Gymnothorax miliaris (Kaup, 1856) Goldentail Moray P Yes Yes 8 TA
Gymnothorax moringa (Cuvier, 1829) Spotted Moray P Yes Yes Yes 8 TA
Gymnothorax vicinus (Castelnau, 1855) Purplemouth Moray (P) Yes 8 TA
NARCINIDAE
Narcine bancroftii (Griffith & Smith, 1834) Lesser Electric Ray Yes GC
OGCOCEPHALIDAE
Dibranchus atlanticus Peters, 1876 Atlantic Batfish Yes TA Yes
Ogcocephalus corniger Bradbury, 1980 Longnose Batfish DROP CP 8 GC
Zalieutes mcgintyi (Fowler, 1952) Tricorn Batfish DROP CP 8 GC Yes
OPHICHTHIDAE
Myrichthys breviceps (Richardson, 1848) Sharptail Eel Yes WA
Myrichthys ocellatus (Lesueur, 1825) Goldspotted Eel Estapé P 8 WA
Ophichthus ophis (Linnaeus, 1758) Spotted Snake Eel Yes WA
OPHIDIIDAE
Brotula barbata (Bloch & Schneider, 1801) Atlantic Bearded Brotula DROP CP DROP 8 TA
Neobythites elongatus Nielsen & Retzler, 1994 Elongate Cusk-eel Yes GC Yes
Parophidion schmidti (Woods & Kanazawa, 1951) Dusky Cusk-eel Estapé P 8 GC
OPISTOGNATHIDAE
Opistognathus aurifrons (Jordan & Thompson, 1905) Yellowhead Jawfish P Yes Yes Yes 8 WA
Opistognathus macrognathus Poey, 1860 Banded Jawfish Yes GC
Opistognathus maxillosus Poey, 1860 Mottled Jawfish Estapé P 8 GC
OSTRACIIDAE
Acanthostracion polygonius Poey, 1876 Honeycomb Cowfish V P Yes Yes Yes 8 WA
Acanthostracion quadricornis (Linnaeus, 1758) Scrawled Cowfish V P Yes 9 TA
Lactophrys bicaudalis (Linnaeus, 1758) Spotted Trunkfish P Yes Yes 9 TA
Lactophrys trigonus (Linnaeus, 1758) Trunkfish P Yes Yes 9 TA
Lactophrys triqueter (Linnaeus, 1758) Smooth Trunkfish P Yes Yes Yes 9 WA
PARALICHTHYIDAE
Citharichthys cornutus (Gunther, 1880) Horned Whiff Yes WA Yes
Gastropsetta frontalis Bean, 1895 Shrimp Flounder DROP CP DROP 9 GC
Syacium micrurum Ranzani, 1842 Channel Flounder P Yes 9 WA
PARAZENIDAE
Cyttopsis rosea (Lowe, 1843) Red Dory Yes TA/IWP Yes
PEMPHERIDAE
Pempheris schomburgkii Müller & Troschel, 1848 Glassy Sweeper P Yes 9 WA
PENTANCHIDAE
Apristurus canutus Springer & Heemstra, 1979 Hoary Cat Shark Yes GC Yes
Galeus antillensis Springer, 1979 Antilles Sawtail Catshark Yes GC L Yes
PERCOPHIDAE
Bembrops ocellatus Thompson & Suttkus, 1998 Ocellate Duckbill Yes GC Yes
Bembrops quadrisella Thompson & Suttkus, 1998 Saddleback Duckbill Yes GC Yes
Chrionema squamentum (Ginsburg, 1955) Scalychin Flathead DROP CP DROP 9 GC Yes
PERISTEDIIDAE
Peristedion truncatum (Gunther, 1880) Black Armored Searobin Yes WA Yes
POLYMIXIIDAE
Polymixia lowei Gunther, 1859 Beardfish Yes WA Yes
POMACANTHIDAE
Centropyge argi Woods & Kanazawa, 1951 Cherubfish V P Yes 9 GC
Holacanthus ciliaris (Linnaeus, 1758) Queen Angelfish V P Yes Yes Yes 9 WA
Holacanthus tricolor (Bloch, 1795) Rock Beauty V P Yes Yes Yes Yes 9 WA
Pomacanthus arcuatus (Linnaeus, 1758) Gray Angelfish Yes Yes Yes WA
Pomacanthus paru (Bloch, 1787) French Angelfish V P Yes Yes Yes Yes 9 WA
POMACENTRIDAE
Abudefduf saxatilis (Linnaeus, 1758) Sergeant Major P Yes Yes 9 TA
Abudefduf taurus (Müller & Troschel, 1848) Night Sergeant P Yes 9 TA
Chromis cf. enchrysura2 DROP CP DROP 13 WA Yes
Chromis cyanea (Poey, 1860) Blue Chromis V P Yes Yes Yes 9 GC
Chromis insolata (Cuvier, 1830) Sunshinefish V P Yes 9 GC
Chromis multilineata (Guichenot, 1853) Brown Chromis V P Yes Yes Yes 9 TA
Chromis scotti Emery, 1968 Purple Reeffish DROP V WA
Microspathodon chrysurus (Cuvier, 1830) Yellowtail Damselfish P Yes Yes Yes Yes 9 WA
Stegastes adustus (Troschel, 1865) Dusky Damselfish P Yes 9 GC
Stegastes diencaeus (Jordan & Rutter, 1897) Longfin Damselfish P Yes 9 GC
Stegastes leucostictus (Müller & Troschel, 1848) Beaugregory Yes Yes GC
Stegastes partitus (Poey, 1868) Bicolor Damselfish V P Yes Yes Yes 9 GC
Stegastes planifrons (Cuvier, 1830) Threespot Damselfish P Yes 9 GC
Stegastes xanthurus (Poey, 1860) Cocoa Damselfish P Yes 9 GC
PRIACANTHIDAE
Heteropriacanthus cruentatus (Lacepède, 1801) Glasseye Snapper P Yes Yes 9 TA
Priacanthus arenatus Cuvier, 1829 Bigeye V Yes TA
Pristigenys alta (Gill, 1862) Short Bigeye DROP V WA
RHINCODONTIDAE
Rhincodon typus Smith, 1828 Whale Shark Yes PAN
SCIAENIDAE
Equetus lanceolatus (Linnaeus, 1758) Jackknife-fish V P Yes Yes 10 WA
Equetus punctatus (Bloch & Schneider, 1801) Spotted Drum P Yes Yes 10 WA
Pareques acuminatus (Bloch & Schneider, 1801) High-hat P Yes 10 WA
Umbrina coroides Cuvier, 1830 Sand Drum Yes WA
SCOMBRIDAE
Acanthocybium solandri (Cuvier, 1832) Wahoo Yes PAN
Euthynnus alletteratus (Rafinesque, 1810) Little Tunny P Yes 10 TA
Katsuwonus pelamis (Linnaeus, 1758) Skipjack Tuna Yes PAN
Scomberomorus cavalla (Cuvier, 1829) King Mackerel P Yes 10 WA
Scomberomorus regalis (Bloch, 1793) Cero V P Yes Yes 10 WA
Thunnus atlanticus (Lesson, 1831) Blackfin Tuna Yes WA
SCORPAENIDAE
Pontinus castor Poey, 1860 Longsnout Scorpionfish DROP CP DROP 10 GC Yes
Pontinus nematophthalmus (Gunther, 1860) Spinythroat Scorpionfish DROP CP 10 WA Yes
Pterois volitans (Linnaeus, 1758) Red Lionfish V P Yes Yes 10 NA NA NA
Scorpaena plumieri Bloch, 1789 Spotted Scorpionfish P Yes Yes 10 TA
Scorpaenodes caribbaeus Meek & Hildebrand, 1928 Reef Scorpionfish Estapé P 10 WA
SERRANIDAE
Alphestes afer (Bloch, 1793) Mutton Hamlet (P) Yes 10 TA
Baldwinella vivanus (Jordan & Swain, 1885)3 Red Barbier DROP V WA Yes
Bathyanthias species A DROP CP 13 GC L Yes
Bullisichthys caribbaeus Rivas, 1971 Pugnose Bass DROP CP DROP 10 GC Yes
Cephalopholis cruentata (Lacepede, 1802) Graysby V P Yes Yes Yes Yes 10 GC
Cephalopholis fulva (Linnaeus, 1758) Coney V P Yes Yes Yes Yes 10 WA
Diplectrum bivittatum (Valenciennes, 1828) Dwarf Sand Perch Estapé P 10 WA
Epinephelus adscensionis (Osbeck, 1765) Rock Hind V (P) Yes 10 TA
Epinephelus guttatus (Linnaeus, 1758) Red Hind V P Yes Yes Yes Yes 10 WA
Epinephelus striatus (Bloch, 1792) Nassau Grouper Yes GC
Gonioplectrus hispanus (Cuvier, 1828) Spanish Flag DROP V WA Yes
Hypoplectrus chlorurus (Cuvier, 1828) Yellowtail Hamlet P Yes Yes 10 GC
Hypoplectrus guttavarius (Poey, 1852) Shy Hamlet P Yes 10 GC
Hypoplectrus indigo (Poey, 1851) Indigo Hamlet DROP V GC
Hypoplectrus nigricans (Poey, 1852) Black Hamlet Yes GC
Hypoplectrus puella (Cuvier, 1828) Barred Hamlet V P Yes Yes Yes 10 GC
Hypoplectrus species 1 Bluelip Hamlet Estapé P 13 GC
Hypoplectrus unicolor (Walbaum, 1792) Butter Hamlet Yes GC
Liopropoma carmabi (Randall, 1963) Candy Basslet DROP CP DROP 10 WA
Liopropoma mowbrayi Woods & Kanazawa, 1951 Cave Basslet DROP CP DROP 10 GC
Liopropoma olneyi Baldwin & Johnson, 2014 Yellow-Spotted Basslet DROP CP DROP 10 GC L Yes
Liopropoma rubre Poey, 1861 Peppermint Basslet P Yes 11 GC
Mycteroperca interstitialis (Poey, 1860) Yellowmouth Grouper Yes WA
Mycteroperca tigris (Valenciennes, 1833) Tiger Grouper Yes WA
Mycteroperca venenosa (Linnaeus, 1758) Yellowfin Grouper P Yes Yes 11 WA
Paranthias furcifer (Valenciennes, 1828) Atlantic Creolefish V P Yes Yes 11 TA
Plectranthias species A DROP CP 13 GC L Yes
Pronotogrammus martinicensis (Guichenot, 1868) Roughtongue Bass DROP CP DROP 11 WA Yes
Rypticus bistrispinus (Mitchill, 1818) Freckled Soapfish Yes WA
Rypticus saponaceus (Bloch & Schneider, 1801) Greater Soapfish V P Yes Yes Yes 11 TA
Serranus annularis (Gunther, 1880) Orangeback Bass DROP V WA
Serranus baldwini (Evermann & Marsh, 1899) Lantern Bass P Yes Yes Yes 11 WA
Serranus flaviventris (Cuvier, 1829) Twinspot Bass Yes WA
Serranus fuscula (Poey, 1861) Twospot Sea Bass DROP CP DROP 11 WA Yes
Serranus luciopercanus Poey, 1852 Crosshatch Bass DROP V GC Yes
Serranus notospilus Longley, 1935 Saddle Bass DROP V GC
Serranus phoebe Poey, 1851 Tattler V Yes WA Yes
Serranus tabacarius (Cuvier, 1829) Tobaccofish V P Yes Yes Yes 11 WA
Serranus tigrinus (Bloch, 1790) Harlequin Bass V P Yes Yes Yes 11 GC
Serranus tortugarum Longley, 1935 Chalk Bass V P Yes Yes 11 GC
SETARCHIDAE
Setarches guentheri Johnson, 1862 Deepwater Scorpionfish Yes TA/IWP Yes
SPARIDAE
Calamus bajonado (Bloch & Schneider, 1801) Jolthead Porgy Yes Yes WA
Calamus calamus (Valenciennes, 1830) Saucereye Porgy P Yes Yes 11 WA
Calamus pennatula Guichenot, 1868 Pluma Porgy P Yes 11 WA
SPHYRAENIDAE
Sphyraena barracuda (Edwards, 1771) Great Barracuda V P Yes Yes Yes Yes 11 PAN
Sphyraena borealis DeKay, 1842 Sennet P Yes 11 WA
SPHYRNIDAE
Sphyrna mokarran (Rüppell, 1837) Great Hammerhead Yes PAN
SQUALIDAE
Squalus clarkae Pfleger, Grubbs, Cotton & Daly-Engel, 2018 Gulf Dogfish Yes GC Yes
SYMPHYSANODONTIDAE
Symphysanodon berryi Anderson, 1970 Slope Bass DROP CP DROP 11 TA Yes
Symphysanodon octoactinus Anderson, 1970 Insular Bunquelovely DROP CP DROP 11 GC Yes
SYNGNATHIDAE
Amphelikturus dendriticus (Barbour, 1905) Seahorse Pipefish Estapé P 11 WA
Bryx dunckeri (Metzelaar, 1919) Pugnose Pipefish Yes WA
Cosmocampus albirostris (Kaup, 1856) Whitenose Pipefish Yes WA
Halicampus crinitus (Jenyns, 1842) Banded Pipefish Estapé V WA
Hippocampus erectus Perry, 1810 Lined Seahorse P Yes 11 WA
Hippocampus reidi Ginsburg, 1933 Longsnout Seahorse P Yes 11 GC
SYNODONTIDAE
Synodus foetens (Linnaeus, 1766) Inshore Lizardfish Yes NWA
Synodus intermedius (Agassiz, 1829) Sand Diver P Yes Yes Yes 11 WA
Synodus synodus (Linnaeus, 1758) Red Lizardfish P Yes 11 TA
Trachinocephalus myops (Forster, 1801) Snakefish P Yes 11 TA
TETRAODONTIDAE
Canthigaster jamestyleri Moura & Castro, 2002 Goldface Toby DROP CP DROP 11 GC
Canthigaster rostrata (Bloch, 1786) Sharpnose Puffer V P Yes Yes Yes 11 GC
Sphoeroides dorsalis Longley, 1934 Marbled Puffer DROP/ Estapé CP P DROP 12 GC
Sphoeroides nephelus (Goode & Bean, 1882) Southern Puffer Yes GC
Sphoeroides spengleri (Bloch, 1785) Bandtail Puffer P Yes Yes 12 WA
TRACHICHTHYIDAE
Hoplostethus occidentalis Woods, 1973 Western Roughy Yes WA Yes
TRIACANTHODIDAE
Hollardia hollardi Poey, 1861 Reticulate Spikefish Yes GC Yes
TRIGLIDAE
Bellator egretta (Goode & Bean, 1896) Streamer Searobin DROP CP 12 GC Yes
TRIPTERYGIIDAE
Enneanectes altivelis Rosenblatt, 1960 Lofty Triplefin Estapé P 12 GC
Enneanectes boehlkei Rosenblatt, 1960 Roughhead Triplefin Estapé P 12 GC
Enneanectes jordani (Evermann & Marsh, 1899) Mimic Triplefin p Yes 12 GC
Enneanectes matador Victor, 2013 Matador Triplefin Estapé p 12 GC

Photographic plates

The 13 photographic plates (Suppl. materials 416: Plates S1–S13) include images of 280 species, 69% of those on the Statia20 list. In addition, Davies and Piontek (2017) provided images of Chimaera cubana, which are not included in the supplemental plates. Of the plate images, 40 species come from DROP collections, 226 were taken by CJE and AME and 14 were provided to them by local divers and fishers at Statia (Table 1). Images are available from other sources for all remaining species listed in Table 1 (except the seven species of macrourids), on their individual species pages at https://biogeodb.stri.si.edu/caribbean/en/pages.

Structure of the Statia20 reef-associated bony fish fauna

Global geographical ranges

Greater Caribbean endemics represent the largest group of species in the Statia fauna, and, together with more widely ranging western Atlantic endemics, constitute almost three quarters of the species. Trans-Atlantic species and species found outside as well as inside the Atlantic represented only a quarter of the fauna (Table 1, Figure 3). The relative abundances of species with different types of large-scale geographic ranges are very similar to those of species in the well documented fauna of nearby St. Croix (Smith-Vaniz and Jelks 2014). Species found in Brazil constituted one third of the Statia fauna, while those extending northwards from the Greater Caribbean represented only 1%, a reflection of the greater effects of temperature limitation on northward extension of ranges as compared to effects of the Amazon-Orinoco outflow on limitation of range extension much further south of the Greater Caribbean.

Figure 3. 

Percentages of the Sint Eustatius marine fish fauna represented by groups of species with different global geographical ranges. GC = Greater Caribbean endemics; NWA = GC plus temperate eastern USA; WA = GC plus Brazil; TA = WA plus central or East Atlantic; and A&P = species found in both the Atlantic and various parts of the Indo-Pacific.

Extent of geographical ranges within the Greater Caribbean

The vast majority of species are widely distributed within the Greater Caribbean, with only nine (2.25%) of them having ranges limited to a restricted part of the Caribbean. Among those nine, five are deep-living species, and five belong to Core CRF families (Table 1). The four shallow species with restricted ranges are all Core CRFs. None of the species were micro-endemics, restricted to Statia or that island plus immediately surrounding islands, and no micro-endemics are known to exist in that general area.

Ecology – Depth

The number of deep species increased from 44 on the Statia17 list to 86 in the Statia20 fauna (Table 1), representing an increase from 14.5% in the former to 21.2% in the latter. Among the reef-associated bony fishes (Table 2) the number of deep species increased from 6 (2.7%) to 39 (11.7%) in those two lists.

Table 2.

Characteristics of assemblages of reef fishes at different locations in the Greater Caribbean region. Percentages of ecotypes in the entire regional fauna, the entire faunas from each local area, and within each of two depth subgroups refer to number of species as a % of the entire fauna and of each depth subgroup. Assemblages include those at Statia in 2017 and 2020 (Statia17 and Statia20), in the Saba EEZ in 2017 (Saba17), of species in the Saba17 fauna that are not currently known to occur at Statia (Saba17-Statia20), of the Saba EEZ in 2020 (Statia20 + Saba17), and of Alligator reef in 2020 (Alligator20). Small species are those with ≤ 10 cm maximum total length. Percentage values for individual sites that are greater than the regional value are shown in red, those below the regional value are in blue.

Region Statia20 Statia17 Saba17 Saba20 Alligator20 Saba17 – Statia20
ALL SPECIES (n) 903 306 220 377 427 427 121
Demersal species% 35.0 55.1 66.8 47.5 46.1 49.4 19.0
Benthic species% 65.0 43.1 33.2 52.5 53.9 50.6 81.0
Cryptobenthic species% 59.2 40.8 30.9 49.1 50.1 46.4 73.6
Small cryptobenthic species% 41.6 24.8 15.5 30.2 31.9 24.8 49.6
Core CRF species% 45.8 28.8 20.5 33.4 35.1 27.6 48.8
SHALLOW SPECIES% 85.1 87.3 97.3 93.4 88.0 95.3 90.1
Non-cryptic species% 40.8 59.6 68.1 50.3 47.6 58.6 23.9
Cryptobenthic species% 59.2 40.4 31.3 49.7 52.4 41.1 76.1
Small cryptobenthic species% 41.3 23.2 15.4 30.7 31.9 25.8 53.2
Core CRF species% 46.2 28.5 21.0 34.1 35.4 29.0 52.3
DEEP SPECIES% 14.9 12.7 2.7 6.6 12.0 4.7 9.9
Non-cryptic species% 40.3 56.5 83.3 60.0 54.9 75 50.0
Cryptobenthic species% 59.7 43.5 16.7 40.0 45.1 25 50.0
Small cryptobenthic species% 43.3 35.6 16.7 24.0 31.4 5 16.7
Core CRF species% 44.0 38.5 0 24.0 33.3 0 16.7

Ecology – Reef-associated bony fishes

The Statia20 fauna of such species is 38.3% larger than the Statia17 fauna, with numbers of shallow species increasing by 24.8% (from 214 to 267) and of deep species increasing 6.2-fold (from 6 to 39). This led to substantial increases in the relative abundance of deep-reef species, and of benthic, cryptobenthic, small cryptobenthic and core CRFs on both shallow and deep reefs. The Saba17 fauna was 71% larger than that of Statia17, with greater percentages of deep-reef, benthic, cryptobenthic, small cryptobenthic and Core CRFs. The Saba17 fauna was 23% larger than the Statia20 fauna and had a greater proportion of shallow species and fewer deep species, and higher proportions of shallow members of cryptobenthic, small cryptobenthic and Core CRF groups. Thirty-two percent of the Saba17 species were not in the Statia20 fauna. Those 121 species comprised mainly shallow cryptobenthic types, including small-cryptobenthic and Core-CRF species. When those are combined with the Statia20 fauna the resultant Saba20 fauna has substantial increases in the proportions of shallow cryptobenthic, small cryptobenthic and core CRF species compared to the Statia20 fauna. Relative to the regional fauna, however, the faunas of Statia17, Statia20, Saba17, and Saba20 all had deficits of deep species of all types and of shallow cryptobenthic species, including small- and Core-CRF species. The Alligator20 fauna of reef-associated species is the same size as the Saba20 fauna. It has the same characteristics as the Statia17 and Saba17 faunas: a large deficit of deep-reef fishes and deficits of shallow cryptobenthic species, including small- and Core-CRF species. Although there has been some collecting at Alligator reef of shallow cryptobenthic species there has been no submersible-based collecting there.

Discussion

The efforts of van Kuijk et al. (2015) and Davies and Piontek (2017) substantially increased our knowledge of the known ichthyofauna of Statia, from 215 to 304 species. The information added through the research in 2017 and 2020 has produced a further significant increase, by 33.6%, to 406 species. While the size of the Statia17 fauna was similar to that known for other islands in the Caribbean (Williams et al. 2010; Davies and Piontek 2017) the Statia20 fauna is distinctly larger. That can be attributed to the combination of research on deep-reef fishes by DROP in 2017 and on shallow species by CJE and AME in 2020. Williams et al. (2010) compared the size of the Saba Bank fauna to the faunas of various Caribbean sites and two in the Florida Keys. The size of the large known fauna at one of those Florida sites, Alligator Reef, has increased by ~ 20% since the Williams et al. (2010) study (see Starck et al. 2017; Estapé et al. 2020). However, the current state of knowledge for the other Caribbean sites referred to by Davies and Piontek (2017) and Williams et al. (2010) is unclear.

Zoogeographically the two largest groups of species in the Statia20 fauna are Greater Caribbean endemics and western Atlantic endemics, and the smallest group is of species found in the Indo-Pacific as well as the Atlantic. This mixture is fairly representative of the Greater Caribbean fish fauna as a whole (Robertson and Cramer 2014), and similar to that of nearby St. Croix (Smith-Vaniz and Jelks 2014). The vast majority of the species in the Statia20 fauna are widely distributed in the Greater Caribbean. Among the very few (2.25%) with restricted ranges most information on range-size is available for the shallow species, which belong to two of the most speciose Core CRF families in the Greater Caribbean, the Gobiidae and Chaenopsidae. High levels of local endemism is a feature of some CRF taxa (Brandl et al. 2018) and regionally those two families have substantial proportions of species with restricted ranges, as defined here: 78.7% of 47 chaenopsids and 42.4% of 139 reef-associated gobies (see species maps in Robertson and Van Tassell 2019).

Most species recorded in the Statia17 fauna are readily visible reef fishes, demersal and non-cryptic benthic species commonly found on wider Caribbean reefs, and the proportions of cryptobenthic (particularly small ones) and deep-reef species were relatively low. Davies and Piontek (2017) recognized that both those groups were probably underrepresented in their checklist due to inadequate sampling. Aspects of data collection that affect the adequacy of sampling at a location include its spatial distribution, techniques used, and the depth of sampled habitats. Of all research efforts to date at Statia only the shallow BRUV sampling by van Kuijk et al. (2015) can be regarded as spatially representative, as it was well dispersed around both exposed and sheltered sides of the island. SCUBA-based sampling by Davies and Piontek (2017) and both DROP and the Estapés was largely limited to the more sheltered platform on the western side of the island, and the submersible sampling by DROP was restricted to one small area at the southwest corner of the island shelf. Hence, there are large areas of habitat on the seaward platform and on deep reefs around three quarters of the island that remain unsampled. Furthermore, roving SCUBA surveys are largely limited to providing information on larger, more readily visible demersal and pelagic species (Ackerman and Bellwood 2000; Smith-Vaniz et al. 2006; Alzate et al. 2014). BRUVs are similarly limited: only 10.3% of the 106 species recorded by van Kuijk et al. (2015) are cryptobenthic forms, and only 2.8% are small cryptobenthic species (see Suppl. materials 2, 3: Tables S1, S2).

Rotenone is an ichthyocide commonly used in small quantities by researchers to extract cryptobenthic fishes hiding within reef structures or buried in soft bottoms, and is an important tool for elucidating the contribution of such species to reef-fish faunal assessments (Ackerman and Bellwood 2000; Smith-Vaniz et al. 2006; Robertson and Smith-Vaniz 2008). Davies and Piontek (2017) indicated that sampling using ichthyocides to extract cryptobenthic species hiding within the matrix of the reef at Statia likely would increase the size of the fauna. Rotenone sampling has been employed on shallow reefs of Saba Bank by Williams et al. (2010), and can account for the large numbers of small cryptobenthic species encountered there that are not on the Statia20 checklist: 60% of the 142 species collected by Williams et al. (2010) at Saba bank using that ichthyocide are cryptobenthic forms. Given that that bank is very close to Statia (the two shallow platforms are < 20 km apart) and, since it lacks mangroves, seagrasses and intertidal habitats, the bank may have even lower habitat diversity than Statia. Hence, it seems quite likely that many of the cryptobenthic species, particularly the small ones, found on that bank will be encountered at Statia when appropriate sampling has been done. However, the increase in numbers of shallow cryptobenthic species at Statia from 2017 to 2020 does show that organized searching by skilled citizen scientists can contribute substantially to knowledge of cryptobenthic species. The activities of CJE and AME added 33 shallow cryptobenthic species to the checklist, 31% of the total and 85% of the new records for that ecogroup in the 2020 fauna, and equivalent to 49% of the number present in the Statia17 fauna (Tables 1, 2).

The DROP submersible-based sampling is the only organized research on deep-reef fishes conducted to date at Statia or in the Saba EEZ. It produced more than half the new records in the Statia 2020 fauna, including records of eight recently discovered species that currently lack scientific names. It dramatically increased the numerical and proportional abundance of deep-reef species in the general fauna and in the reef-associated component. A lack of such research at Saba bank and Alligator Reef accounts for the very low abundance of deep-reef fishes at those sites.

The proportional abundances of shallow cryptobenthic species, including small species and core CRFs, are also distinctly lower in the Statia20 fauna than the regional fauna. Even if all 121 reef-associated species in the Saba EEZ that are not known from Statia are assumed to be at Statia those proportions still remain below the regional levels. Some of that difference is probably due to sampling artifacts. However, the proportional abundances of those ecotypes in a local fauna like that of Statia, or Alligator Reef, may always be lower than the regional level. In the Greater Caribbean small cryptobenthic species, particularly Core-CRF species such as blennioids and gobies, often have small geographical ranges (see above), which are scattered in different parts of the region (see Robertson and Van Tassell 2019). While the regional level of the proportional abundance of such taxa is based on an aggregate of many such species from a large area, only a subset of species in those taxa will be found at any single site and their proportional contribution to local faunal richness most likely will be lower than the regional level. The Statia20 fauna includes 33.9% of the Greater Caribbean’s reef-associated fish fauna. That percentage rises to 47.3% in Saba20. Whether a tiny island with a small area of a limited range of habitats is likely to support many more species, and whether pelagic recruitment of reef fishes from nearby islands found around three sides of Statia helps sustain the Statia fauna are both debatable issues that bear on the size of its marine fish fauna.

Conclusions

The research reported in the present study substantially increased our knowledge of the size of the marine fish fauna of Statia and resulted in the discovery of a significant number of undescribed deep-reef species. Although that island fauna is now one of the best documented in the Greater Caribbean there is still much to do to provide a thorough assessment of its diversity. Collecting with ichthyocide (or anesthetics) is essential for effective sampling of the fauna of small, shallow cryptobenthic reef fishes present there, and sampling of both deep and shallow reef fishes needs to be more effectively distributed across the range of habitats present at the island. No single site in the Caribbean Sea has been subject to sufficiently thorough sampling to provide a clear understanding of the size of its entire marine fish fauna, the size of its reef-associated fish fauna, or even the size of its shallow, reef-associated fauna, let alone its deep-reef fish fauna.

Permits

Collecting by DROP was performed under Saba/Statia BES Permit No. 120317 to the Foundation Curacao Deep Reef Research Centre.

Animal-Care Permission

DROP collecting was approved by a Smithsonian Institution Animal Care and Use Committee, approval No. 2014-13 to CCB.

Acknowledgements

CJE and AME: We thank Mike Harterink, Marieke van de Wetering, Menno and Ingrid Walther, and the crew of the Scubaqua Dive Center; St. Eustatius National Parks Foundation (STENAPA); Sybolt and Marlise ten Hoopen, The Old Gin House Hotel; and Robert and Marilyn Bentley, Mike Harterink, and Marit Pistor (STENAPA) for photographs they provided of various species of fishes (marit.pistor@statiapark.org; mike@scubaqua.com; bentley.robertn@gmail.com).

LT and CCB: We thank Thomas Devine for organizing fieldwork and assisting with collections. Jenna Barrett helped transcribe data from submersible videos and generate maps of Statia, and Jordan Casey assisted with collections in the field. Barry B. Brown photographed many of the specimens collected by DROP that were brought to the surface alive.The DROP expedition was made possible through the help of Adriaan ‘Dutch’ Schrier, Bruce Brandt, Barbara van Bebber, and the rest of the staff of Substation Curacao and the crew of the R/V Chapman. Ocean Heritage Foundation/Curacao Sea Aquarium/Substation Curacao contribution number OHF/CSA/SC#48

References

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Supplementary materials

Supplementary material 1 

Figure S1

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Map of EEZ

Explanation note: Map of Saba EEZ.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (50.64 kb)
Supplementary material 2 

Table S1

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Dive site list

Explanation note: List of dive sites with dates and georeferenced coordinates.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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Supplementary material 3 

Table S2

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Occurences

Explanation note: Fish species occurrences at Saba and Statia.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (34.90 kb)
Supplementary material 4 

Plate S1

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (2.74 MB)
Supplementary material 5 

Plate S2

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (2.82 MB)
Supplementary material 6 

Plate S3

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (2.80 MB)
Supplementary material 7 

Plate S4

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (2.64 MB)
Supplementary material 8 

Plate S5

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (2.30 MB)
Supplementary material 9 

Plate S6

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (2.39 MB)
Supplementary material 10 

Plate S7

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (2.87 MB)
Supplementary material 11 

Plate S8

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (2.87 MB)
Supplementary material 12 

Plate S9

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (3.10 MB)
Supplementary material 13 

Plate S10

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (3.11 MB)
Supplementary material 14 

Plate S11

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (3.29 MB)
Supplementary material 15 

Plate S12

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (912.94 kb)
Supplementary material 16 

Plate S13

David Ross Robertson, Carlos J. Estapé, Allison M. Estapé, Ernesto Peña, Luke Tornabene, Carole C. Baldwin

Data type: Photographs

Explanation note: Voucher photographs of fishes.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
Download file (669.61 kb)