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The marine fishes of St Eustatius Island, northeastern

Caribbean: an annotated, photographic catalog

David Ross Robertson1, Carlos J. Estapé2, Allison M. Estapé2,

Ernesto Peña1, Luke Tornabene3, Carole C. Baldwin4

1  Smithsonian Tropical Research Institute, Balboa, Panama 2  150 Nautilus Drive, Islamorada, Florida,

33036, USA 3 School of Aquatic and Fishery Sciences and the Burke Museum of Natural History and Culture, University of Washington, Seattle, WA 98107, USA 4 Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA

Corresponding author: David Ross Robertson (robertsondr@si.edu)

Academic editor: K. Piller  |  Received 9 September 2020  |  Accepted 23 November 2020  |  Published 30 December 2020

http://zoobank.org/3EDC5293-BB45-4741-A2AD-8AC36BA69CC3

Citation: Robertson DR, Estapé CJ, Estapé AM, Peña E, Tornabene L, Baldwin CC (2020) The marine fishes of St Eustatius Island, northeastern Caribbean: an annotated, photographic catalog. ZooKeys 1007: 145–180. https://doi. org/10.3897/zookeys.1007.58515

Abstract

Sint Eustatius (Statia) is a 21 km2 island situated in the northeastern Caribbean Sea. The most recent

pub-lished 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 sur-veys” (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 pho-tographs 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.

https://zookeys.pensoft.net

Copyright David Ross Robertson et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Launched to accelerate biodiversity research

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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 ma-rine 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

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Figure 1. Location of Sint Eustatius. The Caribbean Sea, with the location of Sint Eustatius island

indi-cated in the inset. Source: Hoetjes and Carpenter (2010: fig. 1).

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 bathym-etry 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)

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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).

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 in-cluded 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 lit-erature, museum records (from major online aggregators, see below), photographs of fishes caught at the island that they obtained from various sources, and fisheries sur-veys. 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, de-pending 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

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individu-als 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.

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 accu-mulated 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

on-line 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

al-most 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 in-dicate 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

cor-rected 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

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Tab

le

1.

U

pdated checklist of marine fishes fr

om S

int E

ustatius I

sland, 2020. K

ey to column headings and entries:

DR

OP –

CP = collected and photographed;

C collected only; V = visual obser vation only; Estapé – P = photographed by

CJE and AME; (P)

photographed by 3 rd par ties; V = visual obser vation only by CJE and AME. N ew – species is a ne w r ecor d r esulting fr om 2017–20 r esear

ch, and its sour

ce. O ther sour ces of species r ecor ds ar e v an K uijk et al. 2015 (vK15) , D avies and P iontek 2017 (DP17), GBIF , and OBIS . DR OP in GBIF indicates r ecor d in GBIF is deriv ed fr om 2017 DR

OP collection specimens deposited in the fish

collection of the US N ational M useum of N atural H istor y. F ishN et 2 r ecor ds ar

e not indicated separately because all such r

ecor ds ar e included b y GBIF . not applicable to non-nativ e Pter ois v olitans . P late –

number indicates supplemental plate containing the v

oucher photograph of that species.

Z

oogeography (Z

oo)-G

lobal geographic range of species; GC = G

reater Caribbean endemic; NW

A = GC plus temperate eastern USA;

W

A = GC plus B

razil;

TA =

W

A plus central or East

Atlantic; P AC = P acific; EP = East P acific; IWP = I ndo-w est P acific; P AN = P antr opical or Cir cumglobal. R ange –

extent of geographic range – L = range limited,

not mor

e than one thir

d of the G

reater Caribbean; r

emainder ar

e mor

e widely distributed in that r

egion.

D

eep –

species entir

ely or largely r

estricted to depths belo

w

40 m.

Ye

s indicates a species conforms to the heading of the column;

? indicates insufficient data.

Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep ACANTHURIDAE Acanthur us chir urgus (B loch, 1787) D octorfish V P Ye s Ye s Ye s Ye s 1 GC Acanthur us coer uleus B

loch & Schneider

, 1801 Blue T ang V P Ye s Ye s Ye s Ye s 1 GC Acanthur us tr actus P oey , 1860 N or thern O cean S urgeonfish V P Ye s Ye s Ye s Ye s 1 GC ACHIRIDAE Gymnachir us nudus Kaup , 1858 Flabb y S ole Estapé P 1 GC ACR OPOMA TIDAE Synagr ops bellus (G oode & B ean, 1896) Blackmouth B ass Ye s WA Ye s AET OBA TIDAE Aetobatus narinari (E uphrasen, 1790)

Spotted Eagle Ray

(P) Ye s Ye s 1 WA

ANTENNARIIDAE Antennarius multiocellatus

(V alenciennes, 1837) Longlur e F rogfish P Ye s 1 WA H istrio histrio (Linnaeus, 1758) Sargassumfish (P) Ye s 1 PA N

APOGONIDAE Apogon aur

olineatus (M owbray , 1927) Bridle Car dinalfish Ye s GC Apogon maculatus (P oey , 1860) Flamefish P Ye s 1 GC Apogon pillionatus

Bohlke & Randall, 1968

Br oadsaddle Car dinalfish DR OP V GC Apogon planifr ons Longley & H ildebrand, 1940 Pale Car dinalfish Estapé P 1 WA Apogon pseudomaculatus Longley , 1932 Twospot Car dinalfish DR OP C DR OP WA Apogon quadrisquamatus Longley , 1934 Saw cheek Car dinalfish P Ye s 1 WA Apogon to wnsendi (B reder , 1927) Belted Car dinalfish P Ye s 1 WA Astr apogon puncticulatus (P oey , 1867) Blackfin Car dinalfish Estapé V WA Astr apogon stellatus (Cope, 1867) Conchfish Ye s WA Par oncheilus affinis (P oey , 1875) Bigtooth Car dinalfish V P Ye s 1 TA

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep Phaeoptyx conklini (S ilv ester , 1915) Fr eckled Car dinalfish Estapé P 1 GC Phaeoptyx pigmentaria (P oey , 1860) D usky Car dinalfish Ye s 1 TA AR GENTINIDAE Argentina stewar ti Cohen & A tsaides, 1969 Ye s GC Ye s G lossanodon p ygmaeus Cohen, 1958 Pygmy Argentine DR OP CP 1 WA Ye s

ATHERINIDAE Atherina harringtonensis

G oode, 1877 Reef S ilv erside Ye s GC Atherinomor us stipes (M üller & Tr oschel, 1848) H ar dhead S ilv erside Estapé P 1 WA AUL OST OMIDAE Aulostomus maculatus V alenciennes, 1841 Atlantic T rumpetfish P Ye s Ye s Ye s 1 GC

BALISTIDAE Balistes capriscus

Gmelin, 1789 G ray T riggerfish P Ye s 1 TA Balistes v etula Linnaeus, 1758 Q ueen T riggerfish P Ye s Ye s Ye s 1 TA C anthider mis sufflamen (M itchill, 1815) O cean T riggerfish V Ye s WA M elichthys niger (B loch, 1786) Black D urgon P Ye s Ye s Ye s Ye s 1 PA N Xanthichthys ringens (Linnaeus, 1758) Sargassum T riggerfish DR OP V WA BEL ONIDAE

Platybelone argalus argalus

(Lesueur , 1821) K eeltail N eedlefish Ye s WA Tylosur us cr ocodilus (P ér on & Lesueur , 1821) H oundfish P Ye s 1 PA N BLENNIIDAE Entomacr odus nigricans G ill, 1859 Pearl B lenny P Ye s 1 GC H ypleur ochilus pseudoaequipinnis B ath, 1994 Oyster B lenny Estapé P 1 WA H ypleur ochilus springeri Randall, 1966 O rangespotted B lenny Estapé P 1 GC H ypsoblennius exstochilus Bohlke, 1959 Longhorn B lenny (P) Ye s 2 GC O phioblennius macclur ei (S ilv ester , 1915) Redlip B lenny P Ye s Ye s 2 GC Par ablennius mar mor eus (P oey , 1876) Seaw eed B lenny P Ye s 2 WA BO THIDAE Bothus lunatus (Linnaeus, 1758) Peacock F lounder P Ye s Ye s 2 TA Bothus ocellatus (Agassiz, 1831) Ey ed F lounder P Ye s 2 WA Chascanopsetta lugubris Alcock, 1894 Pelican F lounder Ye s TA,IWP Ye s

CALLIONYMIDAE Callionymus bair

di (J or dan, 1888) Lancer D ragonet P Ye s 2 WA Foetor epus species DR OP CP 13 WA ? ? Ye s CAPR OIDAE Antigonia capr os Lo w e, 1843 D eepbody Boarfish DR OP V TA,IWP Ye s

CARANGIDAE Alectis ciliaris

(B loch, 1787) African P ompano Ye s PA N

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep C ar anx bar tholomaei (C uvier , 1833) Yello w J ack P Ye s 2 TA C ar anx cr ysos (M itchill, 1815) Blue R unner P Ye s 2 TA C ar anx hippos (Linnaeus, 1766) C rev alle J ack Ye s WA C ar anx latus Agassiz, 1831 H orse-ey e J ack P Ye s Ye s 2 TA C ar anx lugubris P oey , 1860 Black J ack V Ye s Ye s PA N C ar anx r uber (B loch, 1793) Bar J ack V P Ye s Ye s Ye s Ye s 2 WA D ecapter us macar ellus (C uvier , 1833) M acker el Scad P Ye s 2 PA N D ecapter us punctatus (C uvier , 1829) Round Scad P Ye s 2 TA Elagatis bipinnulata (Q uo y & G aimar d, 1825) Rainbo w R unner P Ye s 2 PA N Selar cr umenophthalmus (B loch, 1793) Bigey e Scad P Ye s 2 PA N Seriola riv oliana V alenciennes, 1833 Almaco J ack P Ye s Ye s 2 PA N Tr achinotus falcatus (Linnaeus, 1758) Permit P Ye s 2 WA Tr achinotus goodei Jor dan & E vermann, 1896 Palometa P Ye s 2 WA CAR CHARHINIDAE C ar char hinus leucas (M üller & H enle, 1839) Bull S har k Ye s PA N C ar char hinus limbatus (M üller & H enle, 1839) Blacktip S har k Ye s Ye s PA N C ar char hinus per ezii (P oey , 1876) Reef S har k V Ye s Ye s WA G aleocer do cuvier (P er on & Lesueur , 1822) Tiger S har k Ye s PA N N egaprion br evir ostris (P oey , 1868) Lemon S har k P Ye s 2 TA,EP CENTR OP HORIDAE C entr ophor us gr anulosus (B

loch & Schneider

, 1801) Large G ulper S har k Ye s TA,IWP Ye s

CHAENOPSIDAE Acanthemblemaria asper

a (Longley , 1927) Roughhead B lenny P Ye s 2 GC Acanthemblemaria maria Bohlke, 1961 Secr etar y B lenny P Ye s Ye s 2 GC Acanthemblemaria spinosa M etz elaar , 1919 Spinyhead B lenny P Ye s Ye s 2 GC Chaenopsis limbaughi R

obins & Randall, 1965

Yello wface P ikeblenny P Ye s 2 GC Emblemaria pandionis E vermann & M arsh, 1900 Sailfin B lenny P Ye s Ye s 2 GC Emblemaria vitta W illiams, 2002 Ribbon B lenny Estapé (P) 2 GC Emblemariopsis bahamensis S tephens, 1961 Blackhead B lenny Estapé P 3 GC L Emblemariopsis carib V ictor , 2010 Carib B lenny Estapé P 3 GC L CHAET ODONTIDAE Chaetodon capistr atus Linnaeus, 1758 Four ey e B utterflyfish V P Ye s Ye s Ye s Ye s 3 GC Chaetodon ocellatus B loch, 1787 Spotfin B utterflyfish P Ye s Ye s Ye s 3 WA Chaetodon sedentarius P oey , 1860 Reef B utterflyfish V Ye s Ye s Ye s WA Chaetodon striatus Linnaeus, 1758 Banded B utterflyfish V P Ye s Ye s Ye s Ye s 3 WA Pr ognathodes aculeatus (P oey , 1860) Longsnout B utterflyfish C P Ye s Ye s Ye s Ye s 3 WA

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep Pr ognathodes guyanensis (D urand, 1960) G uyana B utterflyfish DR OP V GC Ye s CHA UNA CIDAE Chaunax suttkusi Car uso, 1989 Pale-cavity G aper Ye s TA Ye s CHIMAERIDAE Chimaer a cubana H ow ell Riv er o, 1936 C uban Chimaera Ye s GC Ye s H ydr olagus alber ti B igelo w & Schr oeder , 1951 G ulf Chimaera Ye s GC Ye s CHL OPSIDAE Chilor hinus suensonii L utken, 1852 Seagrass E el Ye s WA CHL OR OP HTHALMIDAE Chlor ophthalmus agassizi Bonapar te, 1840 Shor tnose G reeney e Ye s TA Ye s Par asudis tr uculenta (G oode & B ean, 1895) Longnose G reeney e Ye s WA Ye s CIRRHITIDAE Ambly cirr hitus pinos (M owbray , 1927) Redspotted H awkfish P Ye s Ye s 3 WA CL UP EIDAE H ar engula clupeola (C uvier , 1829) False P ilchar d Ye s WA H ar engula humer alis (C uvier , 1829) Redear S ar dine Ye s GC Jenkinsia lampr otaenia (G osse, 1851) Dwar f H erring Ye s GC O pisthonema oglinum (Lesueur , 1818) Atlantic Thr ead H erring Ye s WA Sar dinella aurita V alenciennes, 1847 Spanish S ar dine Ye s TA

CONGRIDAE Ariosoma balearicum

(D elar oche, 1809) Bandtooth Conger Estapé (P) 3 TA H eter oconger longissimus G unther , 1870 Br own G ar den E el P Ye s Ye s Ye s 3 WA Xenomystax bidentatus (R eid, 1940) Twopatched-teeth Conger Ye s TA Ye s COR YP HAENIDAE Cor yphaena hippur us Linnaeus, 1758 D olphinfish Ye s PA N CR URIRAJIDAE C rurir aja r ugosa B igelo w & Schr oeder , 1958 Rough Leg S kate Ye s GC Ye s CYNOGL OSSIDAE Symphur us marginatus (G oode & B ean, 1886) M argined T onguefish Ye s WA Ye s DA CT YL OPTERIDAE D actylopter us v olitans (Linnaeus, 1758) Flying G urnar d P Ye s Ye s Ye s 3 TA DASYA TIDAE H ypanus americanus H ildebrand & Schr oeder , 1928 Southern S tingray P Ye s Ye s Ye s 3 WA DIODONTIDAE Chilomy cter us antillar um Jor dan & R utter , 1897 W eb B urrfish P Ye s Ye s 3 WA Chilomy cter us schoepfii (W albaum, 1792) Striped B urrfish Ye s N WA

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep Diodon holocanthus Linnaeus, 1758 Balloonfish P Ye s 3 PA N Diodon hystrix Linnaeus, 1758 Por cupinefish P Ye s Ye s Ye s 3 PA N DIRETMIDAE Dir etmus argenteus Johnson, 1864 Silv er S pinyfish Ye s PA N Ye s

ECHENEIDAE Echeneis naucr

ates Linnaeus, 1758 Shar ksucker P Ye s Ye s Ye s 3 PA N Echeneis neucr atoides Z uie w, 1786 Whitefin S har ksucker Estapé P 3 N WA Remor a r emor a (Linnaeus, 1758) Remora Ye s PA N EP HIPP IDAE Chaetodipter us faber (B roussonet, 1782) Atlantic S padefish Ye s WA ETMOPTERIDAE Etmopter us hillianus (P oey , 1861) Caribbean Lantern S har k Ye s N WA Ye s Etmopter us r obinsi Schofield & B urgess, 1997 W est I ndian Lantern S har k Ye s GC Ye s FISTUL ARIIDAE Fistularia tabacaria Linnaeus, 1758 Bluespotted Cornetfish P Ye s Ye s 3 TA

GERREIDAE Eucinostomus jonesii

(G unther , 1879) Slender M ojarra Ye s WA Eucinostomus lefr oyi (G oode, 1874) M ottled M ojarra P Ye s 3 WA G err es ciner eus (W albaum, 1792) Yello wfin M ojarra Ye s WA GINGL YMOST OMA TIDAE G inglymostoma cirr atum (Bonnaterr e, 1788) N urse S har k (P) Ye s Ye s Ye s 3 TA

GOBIESOCIDAE Derilissus lombar

dii S par ks & G ruber , 2012 Tailspot Clingfish DR OP CP 3 GC Ye s

GOBIIDAE Antilligobius nikkiae

V

an

Tassell & Colin, 2012

Sabr e G ob y DR OP CP DR OP 3 GC Ye s Bathygobius antilliensis T ornabene, B aldwin & P ez old, 2010 Antilles F rillfin Estapé P 3 GC Cor yphopter us dicr us Bohlke & R obins, 1960 Colon G ob y P Ye s 3 WA Cor yphopter us eidolon Bohlke & R obins, 1960 Pallid G ob y P Ye s 3 GC Cor yphopter us glaucofr aenum G ill, 1863 Bridled G ob y Ye s WA Cor yphopter us hyalinus Bohlke & R obins, 1962 G lass G ob y P Ye s 4 GC Cor yphopter us kuna V ictor , 2007 K una G ob y Estapé P 4 GC Cor yphopter us liper nes Bohlke & R obins, 1962 Peppermint G ob y P Ye s 4 GC Cor yphopter us personatus (J or

dan & Thompson, 1905)

M asked G ob y V P Ye s 4 GC Cor yphopter us thrix Bohlke & R obins, 1960 Bar tail G ob y P Ye s 4 WA Cor yphopter us tor tugae (J or dan, 1904) Sand G ob y P Ye s 4 GC Cor yphopter us v enezuelae Cer vigon, 1966 Sand-Cany on G ob y Estapé P 4 GC

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep Ctenogobius saepepallens (G ilber t & Randall, 1968) D ash G ob y Estapé P 4 GC Elacatinus chancei (B eebe & H ollister , 1933) Shor tstripe G ob y C P Ye s Ye s 4 GC L Elacatinus ev elynae (Bohlke & R obins, 1968) Shar knose G ob y P Ye s Ye s 4 GC G enus 1 species 5 DR OP CP 13 GC? ? Ye s G enus 1 species 6 DR OP CP 13 GC? ? Ye s G enus 2 species 1 DR OP CP 13 GC? ? Ye s G insburgellus no vemlineatus (F owler , 1950) N inelined G ob y Estapé P 4 GC G natholepis thompsoni Jor dan, 1904 G oldspot G ob y V P Ye s Ye s 4 TA Lythr ypnus elasson Bohlke & R obins, 1960 Dwar f G ob y DR OP/ Estapé C P 4 GC M icr ogobius carri F owler , 1945 Seminole G ob y Estapé P 4 WA N es longus (N ichols, 1914) O rangespotted G ob y P Ye s 4 GC Palatogobius gr andoculus G reenfield, 2002 Bigey e G ob y DR OP CP DR OP 4 GC Ye s Palatogobius incendius T ornabene, R ober tson & B aldwin, 2017 Ember G ob y DR OP C DR OP GC Ye s Pinnichthys aimoriensis V an T assell & T ornabene, 2016 Thiony ’s G ob y DR OP CP 4 GC Ye s Priolepis hipoliti (M etz elaar , 1922) Rusty G ob y P Ye s 4 WA Pter eleotris helenae (Randall, 1968) H ov ering D ar tfish V P Ye s 4 GC Risor r uber (R osen, 1911) Tusked G ob y C P Ye s Ye s 4 WA Tigrigobius dilepis (R

obins & Bohlke, 1964)

O rangesided G ob y P Ye s 4 GC Tigrigobius multifasciatus (S teindachner , 1876) G reenbanded G ob y Estapé P 4 GC L Varicus cephalocellatus G ilmor e, Van Tassell & B aldwin, 2016 O cellated S plit-F in G ob y DR OP CP DR OP 4 GC L Ye s Varicus v eliguttatus V an Tassell, B aldwin & G ilmor e, 2016 Spotted-S ail G ob y DR OP CP DR OP 4 GC Ye s GRAMMA TIDAE G ramma linki S tar ck & Colin, 1978 Yello w cheek B asslet DR OP CP DR OP 5 GC G ramma lor eto P oey , 1868 Fair y B asslet P Ye s Ye s 5 GC Lipogr amma evides R

obins & Colin, 1979

Banded B asslet DR OP CP DR OP 5 GC Ye s Lipogr amma klayi Randall, 1963 Bicolor B asslet DR OP CP 5 GC Ye s Lipogr amma levinsoni B aldwin, N onaka & R ober tson, 2016 H ourglass B asslet DR OP CP 5 GC Ye s Lipogr amma r egia R

obins & Colin, 1979

Ro yal B asslet DR OP CP DR OP 5 GC Ye s Lipogr amma trilineata Randall, 1963 Thr eeline B asslet DR OP CP DR OP 5 GC Ye s GRAMMICOLEP IDIDAE G rammicolepis br achiusculus P oey , 1873 Thorny T inselfish Ye s PA N Ye s HAEMULIDAE Anisotr emus surinamensis (B loch, 1791) Black M argate P Ye s Ye s 5 WA Br achygenys chr ysarg yr eum (G unther , 1859) Smallmouth G runt P Ye s Ye s Ye s 5 GC H aemulon album C uvier , 1830 M argate P Ye s Ye s 5 WA

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep H aemulon aur olineatum C uvier , 1830 Tomtate P Ye s Ye s Ye s 5 WA H aemulon carbonarium P oey , 1860 Caesar G runt P Ye s Ye s Ye s Ye s 5 GC H aemulon flav olineatum (D esmar est, 1823) Fr ench G runt P Ye s Ye s Ye s Ye s 5 GC H aemulon macr ostomum G unther , 1859 Spanish G runt Ye s GC H aemulon melanur um (Linnaeus, 1758) Cottonwick P Ye s 5 WA H aemulon parr a (D esmar est, 1823) Sailors Choice Ye s WA H aemulon plumierii (Lacepede, 1801) White G runt P Ye s 5 WA H aemulon sciur us (S haw , 1803) Bluestriped G runt (P) Ye s Ye s 5 GC H aemulon striatum (Linnaeus, 1758) Striped G runt V V Ye s WA H aemulon vittatum (P oey , 1860) Boga P Ye s 5 GC HAL OSA URIDAE H alosaur us o venii Johnson, 1864 Stripejaw H alosaur Ye s TA,IWP Ye s HEMIRAMP HIDAE H emir amphus br asiliensis (Linnaeus, 1758) Ballyhoo P Ye s 5 WA HOL OCENTRIDAE Cor niger spinosus Agassiz, 1831 Spiny cheek S oldierfish DR OP V TA Ye s H olocentr us adscensionis (O sbeck, 1765) Squirr elfish V P Ye s Ye s Ye s 5 TA H olocentr us r ufus (W albaum, 1792) Longspine S quirr elfish V P Ye s Ye s Ye s 5 GC M yripristis jacobus C uvier , 1829 Blackbar S oldierfish V P Ye s Ye s 5 TA N eoniphon cor uscum (P oey , 1860) Reef S quirr elfish P Ye s 5 GC N eoniphon marianus (C uvier , 1829) Longjaw S quirr elfish C P Ye s Ye s 5 GC N eoniphon v exillarium (P oey , 1860) D usky S quirr elfish P Ye s 5 GC O stichthys tr achypoma (G unther , 1859) Bigey e S oldierfish DR OP CP DR OP 6 WA Ye s Plectr ypops r etr ospinis (G uichenot, 1853) Car dinal S oldierfish Estapé P 6 WA ISTIOP HORIDAE Istiophor us platypter us (S haw , 1792) Sailfish Ye s TA M akair a nigricans Lacepede, 1802 Blue M arlin Ye s PA N KYP HOSIDAE Kyphosus bigibbus Lacepede, 1801 G ray S eachub Estapé P 6 TA/IWP Kyphosus ciner ascens (F orsskal, 1775) Topsail S eachub P Ye s 6 PA N Kyphosus sectatrix (Linnaeus, 1766) Bermuda Chub P Ye s 6 PA N Kyphosus v aigiensis (Q uo y & G aimar d, 1825) Yello w Chub V Ye s PA N

LABRIDAE Labrinae Bodianus r

ufus (Linnaeus, 1758) Spanish H ogfish V P Ye s Ye s Ye s 6 WA Clepticus parr ae (B

loch & Schneider

, 1801) C reole W rasse V P Ye s Ye s Ye s 6 GC

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep D ecodon puellaris (P oey , 1860) Red H ogfish DR OP CP DR OP 6 WA Ye s D ecodon species 2 DR OP CP 13 GC Ye s H alichoer es bathyphilus (B eebe & Tee-V an,1932) G reenband W rasse DR OP V GC Ye s H alichoer es bivittatus (B loch, 1791) Slipper y D ick P Ye s Ye s 6 WA H alichoer es cyanocephalus (B loch, 1791) Yello w cheek W rasse P Ye s Ye s 6 GC H alichoer es gar noti (V alenciennes, 1839) Yello whead W rasse V P Ye s Ye s Ye s 6 GC H alichoer es maculipinna (M üller & Tr oschel, 1848) Clo wn W rasse P Ye s Ye s 6 GC H alichoer es pictus (P oey , 1860) Rainbo w W rasse P Ye s 6 GC H alichoer es poeyi (S teindachner , 1867) Blackear W rasse P Ye s Ye s 6 WA H alichoer es r adiatus (Linnaeus, 1758) Puddingwife P Ye s Ye s 6 WA Thalassoma bifasciatum (B loch, 1791) Bluehead V P Ye s Ye s Ye s 6 GC Xyrichtys mar tinicensis V alenciennes, 1840 Rosy Raz orfish P Ye s Ye s 6 GC Xyrichtys no vacula (Linnaeus, 1758) Pearly Raz orfish P Ye s 6 WA Xyrichtys splendens Castelnau, 1855 G reen Raz orfish P Ye s Ye s Ye s 6 GC Scarinae Cryptotomus r oseus Cope, 1871 Bluelip P arr otfish P Ye s 6 WA Scar us coer uleus (B loch, 1786) Blue P arr otfish Ye s Ye s GC Scar us guacamaia C uvier , 1829 Rainbo w P arr otfish Ye s GC Scar us iseri (B loch, 1789) Striped P arr otfish P Ye s Ye s Ye s 6 GC Scar us taeniopter us D esmar est, 1831 Princess P arr otfish V P Ye s Ye s Ye s Ye s 6 GC Scar us v etula B

loch & Schneider

, 1801 Q ueen P arr otfish P Ye s Ye s Ye s Ye s 6 GC Sparisoma atomarium (P oey , 1861) G reenblotch P arr otfish P Ye s 6 GC Sparisoma aur ofr enatum (V alenciennes, 1840) Redband P arr otfish V P Ye s Ye s Ye s Ye s 7 GC Sparisoma chr ysopter um (B

loch & Schneider

, 1801) Redtail P arr otfish P Ye s Ye s Ye s 7 GC Sparisoma r adians (V alenciennes, 1840) Bucktooth P arr otfish P Ye s 7 WA Sparisoma r ubripinne (V alenciennes, 1840) Yello wtail P arr otfish P Ye s Ye s 7 GC Sparisoma viride (Bonnaterr e, 1788) Stoplight P arr otfish V P Ye s Ye s Ye s 7 GC

LABRISOMIDAE Brockius nigricinctus

H ow ell Riv er o, 1936 Spotcheek B lenny Estapé P 7 GC G obioclinus buccifer us P oey , 1868 Puffcheek B lenny Estapé P 7 GC G obioclinus gobio (V alenciennes, 1836) Palehead B lenny Estapé P 7 WA G obioclinus gupp yi (N orman, 1922) M imic B lenny Estapé P 7 WA Labrisomus nuchipinnis (Q uo y & G aimar d, 1824) H air y B lenny P Ye s Ye s 7 TA M alacoctenus aur olineatus S mith, 1957 G oldline B lenny P Ye s 7 GC M alacoctenus boehlkei S pringer , 1959 D iamond B lenny Ye s GC M alacoctenus er dmani S mith, 1957 Imitator B lenny Estapé P 7 GC

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep M alacoctenus macr opus (P oey , 1868) Rosy B lenny Estapé P 7 GC M alacoctenus triangulatus S pringer , 1959 Saddled B lenny P Ye s 7 GC LOBO TIDAE Lobotes surinamensis (B loch, 1790) Atlantic T ripletail Ye s TA/IWP LOP HIIDAE Lophiodes monodi Le D anois, 1971 Club-bait G oosefish Ye s GC Ye s

LUTJANIDAE Apsilus dentatus

G uichenot, 1853 Black S napper Ye s GC Ye s Etelis oculatus (V alenciennes, 1828) Q ueen S napper Ye s WA Ye s Lutjanus analis (C uvier , 1828) M utton S napper P Ye s Ye s 7 WA Lutjanus apodus (W albaum, 1792) Schoolmaster V P Ye s Ye s Ye s Ye s 7 GC Lutjanus buccanella (C uvier , 1828) Blackfin S napper V P Ye s 7 WA Lutjanus cyanopter us (C uvier , 1828) C ubera S napper P Ye s 7 WA Lutjanus griseus (Linnaeus, 1758) G ray S napper (P) Ye s Ye s 7 TA Lutjanus jocu (B

loch & Schneider

, 1801) D og S napper P Ye s Ye s 7 TA Lutjanus mahogoni (C uvier , 1828) M ahogany S napper V P Ye s Ye s Ye s Ye s 7 GC Lutjanus purpur eus (P oey , 1866) Caribbean R ed S napper Ye s TA Lutjanus synagris (Linnaeus, 1758) Lane S napper P Ye s Ye s Ye s 7 TA Lutjanus viv anus (C uvier , 1828) Silk S napper Ye s TA Ye s O cyur us chr ysur us (B loch, 1791) Yello wtail S napper P Ye s Ye s Ye s 7 TA Pristipomoides sp. 1 V WA ? ? Ye s MA CR OURIDAE G adomus ar cuatus (G oode & B ean, 1886) D oublethr ead G renadier Ye s TA Ye s G adomus dispar (V aillant, 1888) O nelong G renadier Ye s TA Ye s H ymenocephalus aterrimus G ilber t, 1905 N obear d G renadier Ye s WA / PA C Ye s H ymenocephalus billsam M arshall & I wamoto, 1973 Bigey e G renadier Ye s WA Ye s M alacocephalus laevis (Lo w e, 1843) Velv et G renadier Ye s PA N Ye s N ezumia aequalis (G ünther , 1878) Atlantic B lacktip G renadier Ye s TA Ye s Ventrifossa macr opogon M arshall, 1973 Longbear d G renadier Ye s WA / W PA C Ye s MAL ACANTHIDAE M alacanthus plumieri (B loch, 1786) Sand T ilefish V P Ye s Ye s Ye s 7 WA MEGAL OP IDAE M egalops atlanticus V alenciennes, 1847 Tarpon P Ye s 8 TA MERL UCCIIDAE Steindachneria argentea G oode & B ean, 1896 Luminous H ake Ye s GC Ye s

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep MONA CANTHIDAE Aluter us scriptus (O sbeck, 1765) Scrawled F ilefish P Ye s Ye s 8 PA N C anther hines macr ocer us (H ollar d, 1853) Whitespotted F ilefish P Ye s Ye s Ye s Ye s 8 WA C anther hines pullus (Ranzani, 1842) O rangespotted F ilefish P Ye s Ye s Ye s 8 TA M onacanthus ciliatus (M itchill, 1818) Fringed F ilefish P Ye s Ye s 8 TA M onacanthus tuckeri B ean, 1906 Slender F ilefish P Ye s Ye s 8 GC Stephanolepis setifer (B ennett, 1831) Pygmy F ilefish P Ye s Ye s 8 WA

MUGILIDAE Mugil cur

ema V alenciennes, 1836 White M ullet Ye s TA

MULLIDAE Mulloidichthys mar

tinicus (C uvier , 1829) Yello w G oatfish V P Ye s Ye s Ye s 8 TA Pseudupeneus maculatus (B loch, 1793) Spotted G oatfish V P Ye s Ye s Ye s 8 WA

MURAENIDAE Echidna catenata

(B loch, 1795) Chain M oray P Ye s 8 WA Enchely cor e car ychr oa

Bohlke & Bohlke, 1976

Chestnut M oray Estapé (P) 8 TA Enchely cor e nigricans (Bonnaterr e, 1788) V iper M oray Estapé (P) 8 TA G ymnothor ax funebris Ranzani, 1839 G reen M oray P Ye s Ye s 8 TA G ymnothor ax miliaris (Kaup , 1856) G oldentail M oray P Ye s Ye s 8 TA G ymnothor ax moringa (C uvier , 1829) Spotted M oray P Ye s Ye s Ye s 8 TA G ymnothor ax vicinus (Castelnau, 1855) Purplemouth M oray (P) Ye s 8 TA NAR CINIDAE N ar cine bancr oftii (G riffith & S mith, 1834) Lesser E lectric Ray Ye s GC OGCOCEP HALIDAE Dibr anchus atlanticus P eters, 1876 Atlantic B atfish Ye s TA Ye s O gcocephalus cor niger B radbur y, 1980 Longnose B atfish DR OP CP 8 GC Zalieutes mcgintyi (F owler , 1952) Tricorn B atfish DR OP CP 8 GC Ye s OP HICHTHIDAE M yrichthys br eviceps (Richar dson, 1848) Sharptail E el Ye s WA M yrichthys ocellatus (Lesueur , 1825) G oldspotted E el Estapé P 8 WA O phichthus ophis (Linnaeus, 1758) Spotted S nake E el Ye s WA OP HIDIIDAE Br otula barbata (B

loch & Schneider

, 1801) Atlantic B ear ded B rotula DR OP CP DR OP 8 TA N eob ythites elongatus N ielsen & R etzler , 1994 Elongate C usk-eel Ye s GC Ye s Par ophidion schmidti (W

oods & Kanazawa, 1951)

D usky C usk-eel Estapé P 8 GC OP IST OGNA THIDAE O pistognathus aurifr ons (J or

dan & Thompson, 1905)

Yello whead J awfish P Ye s Ye s Ye s 8 WA O pistognathus macr ognathus P oey , 1860 Banded J awfish Ye s GC

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep O pistognathus maxillosus P oey , 1860 M ottled J awfish Estapé P 8 GC OSTRA CIIDAE Acanthostr acion polygonius P oey , 1876 H oney comb Co wfish V P Ye s Ye s Ye s 8 WA Acanthostr acion quadricor nis (Linnaeus, 1758) Scrawled Co wfish V P Ye s 9 TA Lactophr ys bicaudalis (Linnaeus, 1758) Spotted T runkfish P Ye s Ye s 9 TA Lactophr ys trigonus (Linnaeus, 1758) Tr unkfish P Ye s Ye s 9 TA Lactophr ys triqueter (Linnaeus, 1758) Smooth T runkfish P Ye s Ye s Ye s 9 WA

PARALICHTHYIDAE Citharichthys cor

nutus (G unther , 1880) H orned Whiff Ye s WA Ye s G astr opsetta fr ontalis B ean, 1895 Shrimp F lounder DR OP CP DR OP 9 GC Syacium micr ur um Ranzani, 1842 Channel F lounder P Ye s 9 WA PARAZENIDAE Cyttopsis r osea (Lo w e, 1843) Red D or y Ye s TA/IWP Ye s PEMP HERIDAE Pempheris schomburgkii M üller & Tr oschel, 1848 G lassy S w eeper P Ye s 9 WA PENT ANCHIDAE Apristur us canutus S pringer & H eemstra, 1979 H oar y Cat S har k Ye s GC Ye s G aleus antillensis S pringer , 1979 Antilles S awtail Catshar k Ye s GC L Ye s PER COP HIDAE Bembr ops ocellatus Thompson & S uttkus, 1998 O cellate D uckbill Ye s GC Ye s Bembr ops quadrisella Thompson & S uttkus, 1998 Saddleback D uckbill Ye s GC Ye s Chrionema squamentum (G insburg, 1955) Scaly chin F lathead DR OP CP DR OP 9 GC Ye s PERISTEDIIDAE Peristedion tr uncatum (G unther , 1880) Black Armor ed S ear obin Ye s WA Ye s POL YMIXIIDAE Polymixia lo w ei G unther , 1859 Bear dfish Ye s WA Ye s POMA CANTHIDAE C entr op yge argi W

oods & Kanazawa, 1951

Cher ubfish V P Ye s 9 GC H olacanthus ciliaris (Linnaeus, 1758) Q ueen Angelfish V P Ye s Ye s Ye s 9 WA H olacanthus tricolor (B loch, 1795) Rock B eauty V P Ye s Ye s Ye s Ye s 9 WA Pomacanthus ar cuatus (Linnaeus, 1758) G ray Angelfish Ye s Ye s Ye s WA Pomacanthus par u (B loch, 1787) Fr ench Angelfish V P Ye s Ye s Ye s Ye s 9 WA POMA CENTRIDAE Abudefduf saxatilis (Linnaeus, 1758) Sergeant M ajor P Ye s Ye s 9 TA Abudefduf taur us (M üller & Tr oschel, 1848) N ight S ergeant P Ye s 9 TA Chr omis cf . enchr ysur a 2 DR OP CP DR OP 13 WA Ye s

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep Chr omis cyanea (P oey , 1860) Blue Chr omis V P Ye s Ye s Ye s 9 GC Chr omis insolata (C uvier , 1830) Sunshinefish V P Ye s 9 GC Chr omis multilineata (G uichenot, 1853) Br own Chr omis V P Ye s Ye s Ye s 9 TA Chr omis scotti E mer y, 1968 Purple R eeffish DR OP V WA M icr ospathodon chr ysur us (C uvier , 1830) Yello wtail D amselfish P Ye s Ye s Ye s Ye s 9 WA Stegastes adustus (T roschel, 1865) D usky D amselfish P Ye s 9 GC Stegastes diencaeus (J or dan & R utter , 1897) Longfin D amselfish P Ye s 9 GC Stegastes leucostictus (M üller & Tr oschel, 1848) Beaugr egor y Ye s Ye s GC Stegastes par titus (P oey , 1868) Bicolor D amselfish V P Ye s Ye s Ye s 9 GC Stegastes planifr ons (C uvier , 1830) Thr eespot D amselfish P Ye s 9 GC Stegastes xanthur us (P oey , 1860) Cocoa D amselfish P Ye s 9 GC PRIA CANTHIDAE H eter opriacanthus cr uentatus (Lacepède, 1801) G lassey e S napper P Ye s Ye s 9 TA Priacanthus ar enatus C uvier , 1829 Bigey e V Ye s TA Pristigenys alta (G ill, 1862) Shor t B igey e DR OP V WA

RHINCODONTIDAE Rhincodon typus

S mith, 1828 Whale S har k Ye s PA N

SCIAENIDAE Equetus lanceolatus

(Linnaeus, 1758) Jackknife-fish V P Ye s Ye s 10 WA Equetus punctatus (B

loch & Schneider

, 1801) Spotted D rum P Ye s Ye s 10 WA Par eques acuminatus (B

loch & Schneider

, 1801) H igh-hat P Ye s 10 WA U mbrina cor oides C uvier , 1830 Sand D rum Ye s WA

SCOMBRIDAE Acanthocybium solandri

(C uvier , 1832) W ahoo Ye s PA N Euthynnus alletter atus (Rafinesque, 1810) Little T unny P Ye s 10 TA Katsuwonus pelamis (Linnaeus, 1758) Skipjack T una Ye s PA N Scomber omor us cav alla (C uvier , 1829) King M acker el P Ye s 10 WA Scomber omor us r egalis (B loch, 1793) Cer o V P Ye s Ye s 10 WA Thunnus atlanticus (Lesson, 1831) Blackfin T una Ye s WA SCORP AENIDAE Pontinus castor P oey , 1860 Longsnout Scorpionfish DR OP CP DR OP 10 GC Ye s Pontinus nematophthalmus (G unther , 1860) Spinythr oat Scorpionfish DR OP CP 10 WA Ye s Pter ois v olitans (Linnaeus, 1758) Red Lionfish V P Ye s Ye s 10 NA NA NA Scorpaena plumieri B loch, 1789 Spotted Scorpionfish P Ye s Ye s 10 TA Scorpaenodes caribbaeus M eek & H ildebrand, 1928 Reef Scorpionfish Estapé P 10 WA

SERRANIDAE Alphestes afer

(B loch, 1793) M utton H amlet (P) Ye s 10 TA

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep Baldwinella viv anus (J or dan & S wain, 1885) 3 Red B arbier DR OP V WA Ye s Bathyanthias species A DR OP CP 13 GC L Ye s Bullisichthys caribbaeus Riv as, 1971 Pugnose B ass DR OP CP DR OP 10 GC Ye s C ephalopholis cr uentata (Lacepede, 1802) G raysb y V P Ye s Ye s Ye s Ye s 10 GC C ephalopholis fulv a (Linnaeus, 1758) Coney V P Ye s Ye s Ye s Ye s 10 WA Diplectr um bivittatum (V alenciennes, 1828) Dwar f S and P er ch Estapé P 10 WA Epinephelus adscensionis (O sbeck, 1765) Rock H ind V (P) Ye s 10 TA Epinephelus guttatus (Linnaeus, 1758) Red H ind V P Ye s Ye s Ye s Ye s 10 WA Epinephelus striatus (B loch, 1792) N assau G rouper Ye s GC G onioplectr us hispanus (C uvier , 1828) Spanish F lag DR OP V WA Ye s H ypoplectr us chlor ur us (C uvier , 1828) Yello wtail H amlet P Ye s Ye s 10 GC H ypoplectr us guttav arius (P oey , 1852) Shy H amlet P Ye s 10 GC H ypoplectr us indigo (P oey , 1851) Indigo H amlet DR OP V GC H ypoplectr us nigricans (P oey , 1852) Black H amlet Ye s GC H ypoplectr us puella (C uvier , 1828) Barr ed H amlet V P Ye s Ye s Ye s 10 GC H ypoplectr us species 1 Bluelip H amlet Estapé P 13 GC H ypoplectr us unicolor (W albaum, 1792) Butter H amlet Ye s GC Liopr opoma car mabi (Randall, 1963) Candy B asslet DR OP CP DR OP 10 WA Liopr opoma mo wbr ayi W

oods & Kanazawa, 1951

Cav e B asslet DR OP CP DR OP 10 GC Liopr opoma olneyi B aldwin & J ohnson, 2014 Yello w-S potted B asslet DR OP CP DR OP 10 GC L Ye s Liopr opoma r ubr e P oey , 1861 Peppermint B asslet P Ye s 11 GC M ycter oper ca interstitialis (P oey , 1860) Yello wmouth G rouper Ye s WA M ycter oper ca tigris (V alenciennes, 1833) Tiger G rouper Ye s WA M ycter oper ca v enenosa (Linnaeus, 1758) Yello wfin G rouper P Ye s Ye s 11 WA Par anthias fur cifer (V alenciennes, 1828) Atlantic C reolefish V P Ye s Ye s 11 TA Plectr anthias species A DR OP CP 13 GC L Ye s Pr onotogr ammus mar tinicensis (G uichenot, 1868) Roughtongue B ass DR OP CP DR OP 11 WA Ye s Rypticus bistrispinus (M itchill, 1818) Fr eckled S oapfish Ye s WA Rypticus saponaceus (B

loch & Schneider

, 1801) G reater S oapfish V P Ye s Ye s Ye s 11 TA Serr anus annularis (G unther , 1880) O rangeback B ass DR OP V WA Serr anus baldwini (E vermann & M arsh, 1899) Lantern B ass P Ye s Ye s Ye s 11 WA Serr anus flaviv entris (C uvier , 1829) Twinspot B ass Ye s WA Serr anus fuscula (P oey , 1861) Twospot S ea B ass DR OP CP DR OP 11 WA Ye s Serr anus lucioper canus P oey , 1852 C rosshatch B ass DR OP V GC Ye s Serr anus notospilus Longley , 1935 Saddle B ass DR OP V GC Serr anus phoebe Poey , 1851 Tattler V Ye s WA Ye s

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep Serr anus tabacarius (C uvier , 1829) Tobaccofish V P Ye s Ye s Ye s 11 WA Serr anus tigrinus (B loch, 1790) H arlequin B ass V P Ye s Ye s Ye s 11 GC Serr anus tor tugar um Longley , 1935 Chalk B ass V P Ye s Ye s 11 GC SET AR CHIDAE Setar ches guentheri Johnson, 1862 D eep water Scorpionfish Ye s TA/IWP Ye s SP ARIDAE C alamus bajonado (B

loch & Schneider

, 1801) Jolthead P orgy Ye s Ye s WA C alamus calamus (V alenciennes, 1830) Saucer ey e P orgy P Ye s Ye s 11 WA C alamus pennatula G uichenot, 1868 Pluma P orgy P Ye s 11 WA SP HYRAENIDAE Sphyr aena barr acuda (E dwar ds, 1771) G reat B arracuda V P Ye s Ye s Ye s Ye s 11 PA N Sphyr aena bor ealis D eKay , 1842 Sennet P Ye s 11 WA SP HYRNIDAE Sphyr na mokarr an (R üppell, 1837) G reat H ammerhead Ye s PA N SQ U ALIDAE Squalus clar kae Pfleger , G

rubbs, Cotton & D

aly-E ngel, 2018 G ulf D ogfish Ye s GC Ye s SYMP HYSANODONTIDAE Symphysanodon berr yi Anderson, 1970 Slope B ass DR OP CP DR OP 11 TA Ye s Symphysanodon octoactinus Anderson, 1970 Insular B unquelo vely DR OP CP DR OP 11 GC Ye s SYNGNA THIDAE Ampheliktur us dendriticus (B arbour , 1905) Seahorse P ipefish Estapé P 11 WA Br yx dunckeri (M etz elaar , 1919) Pugnose P ipefish Ye s WA Cosmocampus albir ostris (Kaup , 1856) Whitenose P ipefish Ye s WA H alicampus crinitus (J enyns, 1842) Banded P ipefish Estapé V WA H ippocampus er ectus Perr y, 1810 Lined S eahorse P Ye s 11 WA H ippocampus r eidi G insburg, 1933 Longsnout S eahorse P Ye s 11 GC

SYNODONTIDAE Synodus foetens

(Linnaeus, 1766) Inshor e Lizar dfish Ye s N WA Synodus inter medius (Agassiz, 1829) Sand D iv er P Ye s Ye s Ye s 11 WA Synodus synodus (Linnaeus, 1758) Red Lizar dfish P Ye s 11 TA Tr achinocephalus my ops (F orster , 1801) Snakefish P Ye s 11 TA TETRA ODONTIDAE C anthigaster jamestyleri M

oura & Castr

o, 2002 G oldface T ob y DR OP CP DR OP 11 GC C anthigaster r ostr ata (B loch, 1786) Sharpnose P uffer V P Ye s Ye s Ye s 11 GC Sphoer oides dorsalis Longley , 1934 M arbled P uffer DR OP/ Estapé CP P DR OP 12 GC Sphoer oides nephelus (G oode & B ean, 1882) Southern P uffer Ye s GC

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Species in families

English common name

Ne w DR OP Estapé vK15 DP17 GBIF OBIS Plate Zo o R ange D eep Sphoer oides spengleri (B loch, 1785) Bandtail P uffer P Ye s Ye s 12 WA TRA CHICHTHYIDAE H oplostethus occidentalis W oods, 1973 W estern R oughy Ye s WA Ye s TRIA CANTHODIDAE H ollar dia hollar di P oey , 1861 Reticulate S pikefish Ye s GC Ye s

TRIGLIDAE Bellator egr

etta (G oode & B ean, 1896) Str eamer S ear obin DR OP CP 12 GC Ye s TRIPTER YGIIDAE Enneanectes altiv elis R osenblatt, 1960 Lofty T riplefin Estapé P 12 GC Enneanectes boehlkei R osenblatt, 1960 Roughhead T riplefin Estapé P 12 GC Enneanectes jor dani (E vermann & M arsh, 1899) M imic T riplefin p Ye s 12 GC Enneanectes matador V ictor , 2013 M atador T riplefin Estapé p 12 GC N otes: 1. Pristipomoides. This is P. aquilonaris and/or P. macr ophthalmus . S

tatia is within the geographical range of both species.

2. Chr omis cf . enchr ysur a is an undescribed species r ecor ded as C. enchr ysur

a in the GBIF database, wher

e it is a DR OP entr y. 3. The Baldwinellaviv anus ” population fr

om the Caribbean likely is a separate species fr

om

B. viv

anus

, which was described fr

om specimens collected on the nor

th coast of C uba. Photograph cr edits: B B ro wn: A. nikkiae , B. barbata , C. jamestyleri , D. puellaris , D. lombar dii , F oetor epus sp, G. linkii , L. mo wbr ayi , L. klayi , L. r egia , P . gr andoculus , S. fuscula , V . cephalocellatus , Z. mcg yntii ; M and R B entley: A. narinari , E. car ychr oa , E. adscensionis , G. cirr atum , L. griseus , H. sciur us ; M H ar terink: A. balearicum , E. carib , E. vitta , E. nigricans , G. vicinus , H. exstochilus ; M P istor (STENAP A): A. afer , H. histrio

; all other photographs ar

e b

y the two sets of coauthors during their r

espectiv

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– 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) Geo-graphical range size – we noted which species have small geoGeo-graphical 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 macroal-gal 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 bot-tom, 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, Syngnathi-dae, Tripterygiidae. To these families we added the DinematichthyiSyngnathi-dae, 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 com-mon 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 re-search 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

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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 gener-ated 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 re-stricted 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 pho-tograph 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 (Wil-cox et al. 2018). Hence, we excluded P. miles from the list as it is unlikely to be pre-sent 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

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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.

mitsuku-rii 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

sch-oepfi) 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, ex-cept 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).

Photographic plates

The 13 photographic plates (Suppl. materials 4–16: 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

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inside the Atlantic represented only a quarter of the fauna (Table 1, Figure 3). The rela-tive 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.

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. Figure 3. Percentages of the Sint Eustatius marine fish fauna represented by groups of species with

dif-ferent 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.

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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.

Ecology – Reef-associated bony fishes

The Statia20 fauna of such species is 38.3% larger than the Statia17 fauna, with num-bers 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 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

Notes: see methods for classification of ecotypes. For lists of species in the Saba EEZ and Statia2020, and their ecotypic classifications see Suppl. material 2: Table S1. Pterois volitans and Pristipomoides spp are excluded from Suppl. material 2: Table S1 and the calculations in Table 2. The former is non-native and the specific identity of Pristipomoides at Statia is uncertain.

References

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