Reef-associated Bony Fishes of the Greater Caribbean: a Checklist; Version 4

      Reef-associated Bony Fishes of the Greater Caribbean: A Checklist (VERSION 4)

 

 Date: October 22, 2021.                                 DOI 10.5281/zenodo.5592149

 

D Ross Robertson     

Smithsonian Tropical Research Institute, Balboa, Republic of Panama. Robertsondr@si.edu

Luke Tornabene   

School of Aquatic and Fishery Sciences, and the Burke Museum of Natural History and Culture, University of Washington, Seattle, WA 98105, U.S.A.    Luke.Tornabene@gmail.com

 

This fourth edition of the checklist includes bony fishes belonging to families present in the Greater Caribbean biogeographic region that have species known to associate with shallow (0 – 40 m depth) coral and rocky reefs. That region extends from Cape Hatteras in the southeastern USA, through the Gulf of Mexico and the Caribbean Sea to somewhere around Guyana and includes the West Indies and Bermuda.

The classification of families and spelling of scientific names generally follows Eschmeyer’s Catalog of Fishes. (https://www.calacademy.org/scientists/projects/eschmeyers-catalog-of-fishes

The objective of this checklist is to provide a database of the entire regional fauna of reef-associated bony fishes that includes information on its taxonomic composition, and the conservation (IUCN Red List) status and various ecological characteristics of listed species. This is intended to facilitate comparisons of the structure of equivalent regional faunas in different parts of the globe, and for examining local variation in faunal structure (and faunal inventories) within the Greater Caribbean. Over the past decade the IUCN Red List (https://www.iucnredlist.org/) has produced a comprehensive set of assessments of Greater Caribbean fishes. Those include assessments for >95% of the species in the present checklist. Those assessments include much information on habitat usage by those fishes that was relevant to the construction of the present checklist. That information was supplemented with information from our own research and other published sources. Many of the reef-associated fishes known from the Greater Caribbean also occur in Brazil. Information used here was extracted from 185 publications about Greater Caribbean reef fishes and 152 publications on Brazilian reef fishes.

Greater Caribbean reefs down to depths of ~250-300 m have reef-fish faunas dominated by members of typical shallow-reef families of bony fishes (Baldwin et al. 2018). Here we focus exclusively on species belonging to those families, because those are what are traditionally considered to be reef fishes. In the Greater Caribbean this reef-fish fauna includes 992 species of native fishes belonging to 342 genera and 83 families, 92% of which are demersal or benthic forms, and 8% pelagic species.

Members of a few families of bony fishes that are found only on deep-water reefs (i.e. typically below the 150 m lower limit of the mesophotic zone) and have no representatives that live on shallow reefs, are not included here: Acropomatidae, Anomalopidae, Aulopidae, Berycidae, Caproidae, Gempylidae, Grammicolepidae, Polymixiidae, Polyprionidae, Scombropidae, Sebastidae, Symphysanodontidae, Trachichthyidae, and Triacanthodidae. Between them those 14 families have a total of ~41 species in the Greater Caribbean, only some of which are known to associate with very deep reefs.

The list is restricted to native species and excludes non-native species with resident populations in the Greater Caribbean, notably Pterois volitans and Neopomacentrus cyanomos.

Reef-associated fishes include only demersal and benthic species that use consolidated hard substrata (coral- and rocky reefs) but also demersal and benthic species that use 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, whereas Demersal species use and rely on both the bottom and the near-bottom water column. These two categories both relate to the behavior of fishes during the day, which, in some cases, changes at night. For example, Apogonids and Holocentrids are benthic during the day, but demersal when they become active at night, and day-active demersal species often hide within reef substrata at night (e.g. some Thalassoma bifasciatum rest inside tubular sponges, while Halichoeres bivittatus buries in sand). Reef-associated fishes also include Pelagic species that live in the water column, facultatively associate with reefs, are regularly seen over and immediately adjacent to them, and have trophic interactions with organisms on reefs.

Oyster reefs are estuarine and have an associated estuarine bony-fish fauna (e. g. see Harding & Mann 2001; La Peyre et al. 2019) that differs greatly in taxonomic composition from the faunas of marine reefs. Hence, we have not included species that associate with oyster reefs unless they also associate with rocky- or coral reefs. Species found almost exclusively in fresh- to brackish water also are excluded entirely from the list. Finally, we also exclude a few taxa of oceanic shallow-water fishes that may be seen in the general vicinity reefs: Exocoetidae, Xiphiidae, Istiophoridae and large (Thunnus) tunas. 

Cryptobenthic fishes are species that are “visually and/or behaviorally cryptic” due to their form and coloration, and to their maintaining “a close association with the benthos” (Depczynski & Bellwood, 2003), by living directly on or within it. While cryptobenthic species are a major component of the diversity of reef fishes, they are strongly under-represented in visual surveys of reef-fish assemblages made by divers. The diversity and the numerical abundance of cryptobenthic fishes is revealed only through the use of chemical piscicides (Ackerman & Bellwood 2000; Willis 2001; Smith-Vaniz et al. 2006; Robertson & Smith-Vaniz 2008; Alzate et al. 2014,) or anaesthetics (Kovacic et al. 2012; Robertson & Smith-Vaniz 2010) that flush such fishes out of the substratum for collection and identification. The list indicates which species we have classed as cryptobenthic. While individuals of some cryptobenthic species, such as Muraenids and Apogonids may regularly be seen during the day on reefs while hiding in the substratum, we do not know what fraction of their populations those visible individuals represent (Willis 2001; Alzate et al 2014). On the other hand, species such as the tiny Glass- and Masked Gobies (Coryphopterus hyalinus and C. personatus) that school in the water just above the substratum, and the Garden Eels (Heteroconger spp.) that extend their long, slender bodies to feed in the water column above their burrows, we do not class as cryptobenthic because they can be censused visually.

Studies of cryptobenthic reef-fishes often emphasize that many such species derive their crypticity in part from being very small (Miller 1979; Depczynski and Bellwood 2003; Beldade et al 2006; Kovacic et al 2012; Brandl et al 2018). The list indicates which cryptobenthic reef-fishes are small, with a maximum Total Length (TL) </= 5 cm (Depczynski & Bellwood, 2003) and </= 10 cm (Miller 1979; Beldade et al 2006; Kovacic et al 2012). Brandl et al. (2018, 2019) classed the members of 17 families (only 13 of which are in the Greater Caribbean) that have relatively large numbers of small, cryptobenthic species, share many life-history characteristics and are important for energy flow in reef ecosystems as Core families of Cryptobenthic Reef-fishes (Core CRFs). Members of those families (Apogonidae, Blenniidae, Bythitidae, Callionymidae, Chaenopsidae, Dactyloscopidae, Gobiidae, Gobiesocidae, Grammatidae, Labrisomidae, Opistognathidae, Syngnathidae, Tripterygiidae), which Brandl et al. (2018, 2019) also referred to as microbenthic reef fishes, are identified in the list. Shortly before Brandl et al (2018) defined the group of Core CRFs, Moller et al (2016) split the Bythitidae into two families, Bythitidae and Dinematichthyidae. We have included both those families as Core CRFs as both have small, cryptobenthic species found on shallow reefs. While many cryptobenthic species in other families also are small, Brandl et al’s. (2018) definition of core CRFs was aimed at the family level, thus excluding some small cryptic species in speciose families that have many large, mobile members (e. g. the Labridae and Serranidae).

Shallow- and deep-reef fishes: Shallow-water species we consider to be those with populations restricted to or concentrated at depths above 40m. Deep-reef species are members of the same shallow-reef families that have populations concentrated in or restricted to depths greater than 40m. This depth breakpoint corresponds to the approximate breakpoint between shallow and deeper (mesophotic and altiphotic) faunal depth-zones on reefs, and the approximate depth breakpoint between research on shallow-water reef fishes based on open-circuit SCUBA, and research at greater depths that rely on mixed gas and Closed-Circuit Rebreather (CCR) technical diving (e.g. Pinheiro et al 2016; Baldwin et al 2018; García-Hernández et al 2018; Stefanoudis et al 2019).  We classified as shallow some species that have shallow populations at some sites in the Greater Caribbean and deep populations in others (e. g. Gramma melacara), and others in which juveniles are common in shallow water and adults are restricted to deep water (e. g. Lutjanus buccanella).

Simple systems of habitat characterization like those used here cannot capture all the nuances of how different fishes use a variety of habitats and microhabitats. Inevitably there are borderline cases of species that could be classified in either of two alternative habitat categories (e. g. benthic or demersal, deep or shallow). There also are borderline cases relating to whether a species should be regarded as reef-associated or not. A single observation of a species passing nearby during a dive on a reef does not make that a reef-associated species. In addition, geographic variation in whether some species on this list do or do not associate with reefs at any location complicates the situation. We have tried to be conservative in classing species as reef-associated, by relying as much as possible on multiple primary records of such behavior by those species and clearcut descriptions of actual behavior. Otherwise virtually everything could end up classed as reef associated.

 

 

CHANGES IN VERSION 4 FROM VERSION 3:

1. Based on additional information about which species associate with reefs, as well as taxonomic rearrangements, the numbers of taxa on Version 4 of the list changed to 83 Families, 342 Genera and 992 Species, from 83 Families, 338 Genera and 981 Species in Version 3.
2. All species names now have the associated description authority.
3. Names of taxa added and of species that have been renamed or re-classified as reef associated in this version are indicated in BOLD RED.
4. The family Malacanthidae traditionally has comprised two subfamilies, the Latilinae and Malacanthinae, with the former including only deep living species in the Greater Caribbean. Those two were recently raised to families. Because of the sister relationship between those to taxa and their previous status we continue to include both among the GC reef-associated fauna.
5. The family Anomalopidae was removed from the list because its single Greater Caribbean member is cryptobenthic in deep water during the day, and daytime activity and depth range is the focus of this list (see above). Unlike the situation with the Malacanthids the near relatives of the Anomalopids are all deep-water taxa.

 

 

CHANGES IN VERSION 3 FROM VERSION 2

1. This edition incorporates data from additional published research on fishes in the Greater Caribbean, plus information on species that occur in Brazil as well as the Greater Caribbean that we reviewed in ~70 publications that deal with reef-associated fishes in Brazil. That additional information has increased the number of reef-associated bony fishes: to 83 families, 338 genera and 981 species.
2. Shortly before Brandl et al (2018) defined the group of Core CRFs, Moller et al (2016) split the Bythitidae into two families, Bythitidae and Dinematichthyidae. In this edition we have included both those families as Core CRFs as the dinematichthyids include many small, cryptobenthic species found on shallow reefs.
3. Three additional types of data are given for each species on the list. These include (1) whether or not a species is endemic to the Greater Caribbean (i.e has 90% or more of its range within that area, the northern and southern limits of which have yet to be precisely defined), and (2) whether or not it has a small or very small geographic range within that region. Small ranges are those no larger than 1/3 of the Greater Caribbean. Very small ranges are those no larger than the area occupied by the Mesoamerican reef system, which extends from northern Honduras and along the Caribbean coast of Mexico and represents ~5% of the area of the Greater Caribbean. Species for which insufficient information is available on range size, but which seem likely to have small ranges, are indicated with a query. Note that non-endemics can have small Greater Caribbean ranges that represent part of larger ranges that extend well outside that region, e.g. to Brazil. (3) The IUCN Red List extinction-risk category is also given for each species that has been assessed by that process. Red List categories include DD (Data Deficient); LC (Least Concern); NT (Near Threatened); VU (Vulnerable); E (Endangered); CR (Critically Endangered). Species not yet assessed by the Red List are indicated by ND (no data).

 

CHANGES IN VERSION 2 FROM VERSION 1:

1. In Version 1 all non-pelagic species were referred to as demersal. In Version 2 non-pelagic species are coded as either Benthic or Demersal (see above for definitions).
2. Species have been divided into two depth classes, based on their depth ranges: Shallow species are those commonly found above 40m depth and Deep Species are those entirely or largely restricted to depths below 40m. The choice of this 40 m separation depth was based on it  being the approximate lower limit of SCUBA-based research on shallow-water reef fishes, and also being the approximate breakpoint between shallow and deeper (mesophotic and rariphotic) faunal depth-zones on reefs, with shallow- and deep-reef fish assemblages being dominated by different members of the same set of families indicated in the table presented here (e.g. Pinheiro et al 2016; Baldwin et al 2018; García-Hernández et al 2018; Stefanoudis et al 2019).

 

This division into shallow- and deep species was made because levels of research effort on deep-reef members of the families on the list vary widely between regions and between locations within a region. There has been much more research and collecting of deep-reef fishes, particularly of small, cryptobenthic forms, using crewed submersibles specially equipped for such collecting (Gilmore 2016) in the Greater Caribbean than many other areas. That includes work at widely scattered sites in the region between the late 1970s and 2010 by the Johnson Sea Link I and II submarines, on the Atlantic and Gulf of Mexico coasts of Florida, the Bahamas, Caribbean Mexico, Cuba, Jamaica, Puerto Rico, the US Virgin Islands, Bonaire and Curacao. In addition, this activity includes observations by the Nekton submersibles at Jamaica and Belize, and the Alvin at the Bahamas (again during the 1970s). During the 2010s, this effort was renewed by observations and collecting by the Smithsonian Institution’s DROP program using the submersibles Curasub at Curacao, Bonaire, Dominica and St Eustatius, and the Idabel at Roatan.No similarly extensive set of submersible-based studies and collections of deep-reef fishes exists for any other equivalent biogeographic region in the tropics. Hence, the focus of quantitative comparisons with reef-associated fish faunas of most other regions (or locations within the Greater Caribbean that lack deep-reef research) is likely to be on shallow-water reefs.

 

1. A few, currently unnamed species of reef-associated Gobiids (6 species in two, currently unnamed, genera), Grammatids (2 species of Lipogramma), Labrids (2 species of Decodon) and Serranids (1 species of Baldwinella) for which sufficient ecological information is available have been added to the list.

 

1. Several deep-living reef-associated species listed in Version 1 as members of the Scorpaenidae have been removed from the list, because systematic rearrangements have placed them in other families (Setarchidae and Sebastidae) that, in the study area, lack shallow members.

 

1. The family Ophidiidae has about 78 species in 28 genera and three subfamilies in the study are. We include on the list only 41 members of 7 genera known to have shallow-water, reef-associated representatives or representatives that live within the depth range (0-~250m) of reefs dominated by shallow taxa. The remainder are species that live in very deep or oceanic waters and are irrelevant to the reef-associated fish fauna considered here.

 

1. Habitat classifications of a few species of carangids and lutjanids as pelagic or demersal in version 1 have been changed, based on additional information.

 

Hosting of this database by Zenodo will allow it to be updated through the production of new versions as new information becomes available.

New information, comments or queries relating to the classification or species list should be directed to robertsondr@si.edu.

A compilation of further information about the biological characteristics of fishes on this list can be found in the website: Robertson DR, Van Tassell J (2019) Shorefishes of the Greater Caribbean: online information system. Version 2.0. https://biogeodb.stri.si.edu/caribbean/en/pages

 

References.

Ackerman JL, Bellwood DR (2000) Reef fish assemblages: a re-evaluation using enclosed rotenone stations Marine Ecology Progress Series 206:227-237.  doi:10.3354/meps206227

Alzate, Zapata FA, Giraldo A (2014) A comparison of visual and collection-based methods for assessing community structure of coral reef fishes in the Tropical Eastern Pacific. Revista Biologica Tropical 62 (Supplement 1): 359–371.

Baldwin CC, Tornabene L, Robertson DR (2018) Below the mesophotic. Scientific Reports 8:4920   DOI:10.1038/s41598-018-23067-1

Beldade R, Erzini K, Goncalves EJ (2006) Composition and temporal dynamics of a temperate rocky
cryptobenthic fish assemblage. Journal of the Marine Biological Association of the U.K. 86: 1221-1228.

Brandl SJ, Goatley CHR, Bellwood DR, Tornabene L (2018) The hidden half: ecology and evolution of cryptobenthic fishes on coral reefs. Biological reviews of the Cambridge Philosophical Society 93: 1846 -1873.  https://doi.org/10.1111/brv.12423

Brandl SJ, Tornabene L, Goatley CHR,  Casey JM,  Morais RA,  Côté IM,  Baldwin CC, Parravicini V,  Schiettekatte MD, Bellwood  DR (2019) Demographic dynamics of the smallest marine vertebrates fuel coral reef ecosystem functioning. Science 364: 1189-1192  DOI: 10.1126/science.aav3384

Depczynski M, Bellwood DR (2003) The role of cryptobenthic reef fishes in coral reef trophodynamics. Marine Ecology Progress Series 256: 183–191

García-Hernández JE, Sanchez PJ, Hammerman NM and Schizas NV (2018) Fish, coral, and sponge assemblages associated with altiphotic and mesophotic reefs along the Guánica Biosphere Reserve continental shelf edge, southwest Puerto Rico. Frontiers in Marine Science 5:303. doi:10.3389/fmars.2018.00303

Gilmore, RG Jr (2016) You can’t catch a fish with a robot. Gulf and Caribbean Research 27: ii-xiv DOI: 10.18785/gcr.2701.11

Harding JM, Mann R (2001) Oyster reefs as fish habitat: opportunistic use of restored reefs by transient fishes. Journal of Shellfish Research 20: 951-959.

Kovacic M, Patzner RA, Schliewen U (2012) A first quantitative assessment of the ecology of cryptobenthic fishes in the Mediterranean Sea. Marine Biology 159: 2731–2742.

La Peyre MK, Aguilar Marshall D, Miller LS, Humphries AT (2019) Oyster Reefs in Northern Gulf of Mexico Estuaries Harbor Diverse Fish and Decapod Crustacean Assemblages: A Meta-Synthesis.
Frontiers in Marine Science. 6:666. doi: 10.3389/fmars.2019.0066

Miller PJ (1979) Adaptiveness and implications of small size in Teleosts. Symposia of the Zoological Society of London. 44: 263–306

Møller PR, Knudsen W, Schwarzhans W, Nielsen JG (2016) A new classification of viviparous brotulas (Bythitidae) – with family status for Dinematichthyidae – based on molecular, morphological and fossil data. Molecular Phylogenetics and Evolution 100: 391-408. https://doi.org/10.1016/j.ympev.2016.04.008

Pinheiro H, Goodbody-Gringley G, Jessup M, Shepherd B, Chequer A, Rocha L (2016) Upper and lower mesophotic coral reef fish communities evaluated by underwater visual censuses in two Caribbean locations. Coral Reefs 35: 139–151 DOI 10.1007/s00338-015-1381-0

Robertson DR, Smith-Vaniz WF (2008) Rotenone: An essential but demonized tool for assessing marine fish diversity. Bioscience 58:165-170.

Robertson DR, Smith-Vaniz WF (2010) Use of clove oil in collecting coral reef fishes for research. Marine Ecology Progress Series.401:295-302

Smith-Vaniz WF, Jelks HL, Rocha LA (2006) Relevance of cryptic fishes in biodiversity assessments: A case study at Buck Island National Monument, St. Croix.  Bulletin of Marine Science 79:17–48

Stefanoudis PV, Gress E, Pitt JM, Smith SR, Kincaid T, Rivers M, Andradi-Brown DA, Rowlands G, Woodall LC and Rogers AD (2019) Depth-Dependent Structuring of Reef Fish Assemblages From the Shallows to the Rariphotic Zone. Frontiers in Marine Science. 6:307 doi: 10.3389/fmars.2019.00307

Willis TJ (2001). Visual census methods underestimate density and diversity of cryptic reef fishes. Journal of Fish Biology 59: 1408-1411.

      Reef-associated Bony Fishes of the Greater Caribbean: A Checklist (VERSION 4)

 

 Date: October 22, 2021

 

D Ross Robertson     

Smithsonian Tropical Research Institute, Balboa, Republic of Panama. Robertsondr@si.edu

Luke Tornabene   

School of Aquatic and Fishery Sciences, and the Burke Museum of Natural History and Culture, University of Washington, Seattle, WA 98105, U.S.A.    Luke.Tornabene@gmail.com

 

This fourth edition of the checklist includes bony fishes belonging to families present in the Greater Caribbean biogeographic region that have species known to associate with shallow (0 – 40 m depth) coral and rocky reefs. That region extends from Cape Hatteras in the southeastern USA, through the Gulf of Mexico and the Caribbean Sea to somewhere around Guyana and includes the West Indies and Bermuda.

The classification of families and spelling of scientific names generally follows Eschmeyer’s Catalog of Fishes. (https://www.calacademy.org/scientists/projects/eschmeyers-catalog-of-fishes

The objective of this checklist is to provide a database of the entire regional fauna of reef-associated bony fishes that includes information on its taxonomic composition, and the conservation (IUCN Red List) status and various ecological characteristics of listed species. This is intended to facilitate comparisons of the structure of equivalent regional faunas in different parts of the globe, and for examining local variation in faunal structure (and faunal inventories) within the Greater Caribbean. Over the past decade the IUCN Red List (https://www.iucnredlist.org/) has produced a comprehensive set of assessments of Greater Caribbean fishes. Those include assessments for >95% of the species in the present checklist. Those assessments include much information on habitat usage by those fishes that was relevant to the construction of the present checklist. That information was supplemented with information from our own research and other published sources. Many of the reef-associated fishes known from the Greater Caribbean also occur in Brazil. Information used here was extracted from 185 publications about Greater Caribbean reef fishes and 152 publications on Brazilian reef fishes.

Greater Caribbean reefs down to depths of ~250-300 m have reef-fish faunas dominated by members of typical shallow-reef families of bony fishes (Baldwin et al. 2018). Here we focus exclusively on species belonging to those families, because those are what are traditionally considered to be reef fishes. In the Greater Caribbean this reef-fish fauna includes 992 species of native fishes belonging to 342 genera and 83 families, 92% of which are demersal or benthic forms, and 8% pelagic species.

Members of a few families of bony fishes that are found only on deep-water reefs (i.e. typically below the 150 m lower limit of the mesophotic zone) and have no representatives that live on shallow reefs, are not included here: Acropomatidae, Anomalopidae, Aulopidae, Berycidae, Caproidae, Gempylidae, Grammicolepidae, Polymixiidae, Polyprionidae, Scombropidae, Sebastidae, Symphysanodontidae, Trachichthyidae, and Triacanthodidae. Between them those 14 families have a total of ~41 species in the Greater Caribbean, only some of which are known to associate with very deep reefs.

The list is restricted to native species and excludes non-native species with resident populations in the Greater Caribbean, notably Pterois volitans and Neopomacentrus cyanomos.

Reef-associated fishes include only demersal and benthic species that use consolidated hard substrata (coral- and rocky reefs) but also demersal and benthic species that use 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, whereas Demersal species use and rely on both the bottom and the near-bottom water column. These two categories both relate to the behavior of fishes during the day, which, in some cases, changes at night. For example, Apogonids and Holocentrids are benthic during the day, but demersal when they become active at night, and day-active demersal species often hide within reef substrata at night (e.g. some Thalassoma bifasciatum rest inside tubular sponges, while Halichoeres bivittatus buries in sand). Reef-associated fishes also include Pelagic species that live in the water column, facultatively associate with reefs, are regularly seen over and immediately adjacent to them, and have trophic interactions with organisms on reefs.

Oyster reefs are estuarine and have an associated estuarine bony-fish fauna (e. g. see Harding & Mann 2001; La Peyre et al. 2019) that differs greatly in taxonomic composition from the faunas of marine reefs. Hence, we have not included species that associate with oyster reefs unless they also associate with rocky- or coral reefs. Species found almost exclusively in fresh- to brackish water also are excluded entirely from the list. Finally, we also exclude a few taxa of oceanic shallow-water fishes that may be seen in the general vicinity reefs: Exocoetidae, Xiphiidae, Istiophoridae and large (Thunnus) tunas. 

Cryptobenthic fishes are species that are “visually and/or behaviorally cryptic” due to their form and coloration, and to their maintaining “a close association with the benthos” (Depczynski & Bellwood, 2003), by living directly on or within it. While cryptobenthic species are a major component of the diversity of reef fishes, they are strongly under-represented in visual surveys of reef-fish assemblages made by divers. The diversity and the numerical abundance of cryptobenthic fishes is revealed only through the use of chemical piscicides (Ackerman & Bellwood 2000; Willis 2001; Smith-Vaniz et al. 2006; Robertson & Smith-Vaniz 2008; Alzate et al. 2014,) or anaesthetics (Kovacic et al. 2012; Robertson & Smith-Vaniz 2010) that flush such fishes out of the substratum for collection and identification. The list indicates which species we have classed as cryptobenthic. While individuals of some cryptobenthic species, such as Muraenids and Apogonids may regularly be seen during the day on reefs while hiding in the substratum, we do not know what fraction of their populations those visible individuals represent (Willis 2001; Alzate et al 2014). On the other hand, species such as the tiny Glass- and Masked Gobies (Coryphopterus hyalinus and C. personatus) that school in the water just above the substratum, and the Garden Eels (Heteroconger spp.) that extend their long, slender bodies to feed in the water column above their burrows, we do not class as cryptobenthic because they can be censused visually.

Studies of cryptobenthic reef-fishes often emphasize that many such species derive their crypticity in part from being very small (Miller 1979; Depczynski and Bellwood 2003; Beldade et al 2006; Kovacic et al 2012; Brandl et al 2018). The list indicates which cryptobenthic reef-fishes are small, with a maximum Total Length (TL) </= 5 cm (Depczynski & Bellwood, 2003) and </= 10 cm (Miller 1979; Beldade et al 2006; Kovacic et al 2012). Brandl et al. (2018, 2019) classed the members of 17 families (only 13 of which are in the Greater Caribbean) that have relatively large numbers of small, cryptobenthic species, share many life-history characteristics and are important for energy flow in reef ecosystems as Core families of Cryptobenthic Reef-fishes (Core CRFs). Members of those families (Apogonidae, Blenniidae, Bythitidae, Callionymidae, Chaenopsidae, Dactyloscopidae, Gobiidae, Gobiesocidae, Grammatidae, Labrisomidae, Opistognathidae, Syngnathidae, Tripterygiidae), which Brandl et al. (2018, 2019) also referred to as microbenthic reef fishes, are identified in the list. Shortly before Brandl et al (2018) defined the group of Core CRFs, Moller et al (2016) split the Bythitidae into two families, Bythitidae and Dinematichthyidae. We have included both those families as Core CRFs as both have small, cryptobenthic species found on shallow reefs. While many cryptobenthic species in other families also are small, Brandl et al’s. (2018) definition of core CRFs was aimed at the family level, thus excluding some small cryptic species in speciose families that have many large, mobile members (e. g. the Labridae and Serranidae).

Shallow- and deep-reef fishes: Shallow-water species we consider to be those with populations restricted to or concentrated at depths above 40m. Deep-reef species are members of the same shallow-reef families that have populations concentrated in or restricted to depths greater than 40m. This depth breakpoint corresponds to the approximate breakpoint between shallow and deeper (mesophotic and altiphotic) faunal depth-zones on reefs, and the approximate depth breakpoint between research on shallow-water reef fishes based on open-circuit SCUBA, and research at greater depths that rely on mixed gas and Closed-Circuit Rebreather (CCR) technical diving (e.g. Pinheiro et al 2016; Baldwin et al 2018; García-Hernández et al 2018; Stefanoudis et al 2019).  We classified as shallow some species that have shallow populations at some sites in the Greater Caribbean and deep populations in others (e. g. Gramma melacara), and others in which juveniles are common in shallow water and adults are restricted to deep water (e. g. Lutjanus buccanella).

Simple systems of habitat characterization like those used here cannot capture all the nuances of how different fishes use a variety of habitats and microhabitats. Inevitably there are borderline cases of species that could be classified in either of two alternative habitat categories (e. g. benthic or demersal, deep or shallow). There also are borderline cases relating to whether a species should be regarded as reef-associated or not. A single observation of a species passing nearby during a dive on a reef does not make that a reef-associated species. In addition, geographic variation in whether some species on this list do or do not associate with reefs at any location complicates the situation. We have tried to be conservative in classing species as reef-associated, by relying as much as possible on multiple primary records of such behavior by those species and clearcut descriptions of actual behavior. Otherwise virtually everything could end up classed as reef associated.

CHANGES IN VERSION 4 FROM VERSION 3:

1. Based on additional information about which species associate with reefs, as well as taxonomic rearrangements, the numbers of taxa on Version 4 of the list changed to 83 Families, 342 Genera and 992 Species, from 83 Families, 338 Genera and 981 Species in Version 3.
2. All species names now have the associated description authority.
3. Names of taxa added and of species that have been renamed or re-classified as reef associated in this version are indicated in BOLD RED.
4. The family Malacanthidae traditionally has comprised two subfamilies, the Latilinae and Malacanthinae, with the former including only deep living species in the Greater Caribbean. Those two were recently raised to families. Because of the sister relationship between those to taxa and their previous status we continue to include both among the GC reef-associated fauna.
5. The family Anomalopidae was removed from the list because its single Greater Caribbean member is cryptobenthic in deep water during the day, and daytime activity and depth range is the focus of this list (see above). Unlike the situation with the Malacanthids the near relatives of the Anomalopids are all deep-water taxa.

CHANGES IN VERSION 3 FROM VERSION 2

1. This edition incorporates data from additional published research on fishes in the Greater Caribbean, plus information on species that occur in Brazil as well as the Greater Caribbean that we reviewed in ~70 publications that deal with reef-associated fishes in Brazil. That additional information has increased the number of reef-associated bony fishes: to 83 families, 338 genera and 981 species.
2. Shortly before Brandl et al (2018) defined the group of Core CRFs, Moller et al (2016) split the Bythitidae into two families, Bythitidae and Dinematichthyidae. In this edition we have included both those families as Core CRFs as the dinematichthyids include many small, cryptobenthic species found on shallow reefs.
3. Three additional types of data are given for each species on the list. These include (1) whether or not a species is endemic to the Greater Caribbean (i.e has 90% or more of its range within that area, the northern and southern limits of which have yet to be precisely defined), and (2) whether or not it has a small or very small geographic range within that region. Small ranges are those no larger than 1/3 of the Greater Caribbean. Very small ranges are those no larger than the area occupied by the Mesoamerican reef system, which extends from northern Honduras and along the Caribbean coast of Mexico and represents ~5% of the area of the Greater Caribbean. Species for which insufficient information is available on range size, but which seem likely to have small ranges, are indicated with a query. Note that non-endemics can have small Greater Caribbean ranges that represent part of larger ranges that extend well outside that region, e.g. to Brazil. (3) The IUCN Red List extinction-risk category is also given for each species that has been assessed by that process. Red List categories include DD (Data Deficient); LC (Least Concern); NT (Near Threatened); VU (Vulnerable); E (Endangered); CR (Critically Endangered). Species not yet assessed by the Red List are indicated by ND (no data).

CHANGES IN VERSION 2 FROM VERSION 1:

1. In Version 1 all non-pelagic species were referred to as demersal. In Version 2 non-pelagic species are coded as either Benthic or Demersal (see above for definitions).
2. Species have been divided into two depth classes, based on their depth ranges: Shallow species are those commonly found above 40m depth and Deep Species are those entirely or largely restricted to depths below 40m. The choice of this 40 m separation depth was based on it  being the approximate lower limit of SCUBA-based research on shallow-water reef fishes, and also being the approximate breakpoint between shallow and deeper (mesophotic and rariphotic) faunal depth-zones on reefs, with shallow- and deep-reef fish assemblages being dominated by different members of the same set of families indicated in the table presented here (e.g. Pinheiro et al 2016; Baldwin et al 2018; García-Hernández et al 2018; Stefanoudis et al 2019).

 

This division into shallow- and deep species was made because levels of research effort on deep-reef members of the families on the list vary widely between regions and between locations within a region. There has been much more research and collecting of deep-reef fishes, particularly of small, cryptobenthic forms, using crewed submersibles specially equipped for such collecting (Gilmore 2016) in the Greater Caribbean than many other areas. That includes work at widely scattered sites in the region between the late 1970s and 2010 by the Johnson Sea Link I and II submarines, on the Atlantic and Gulf of Mexico coasts of Florida, the Bahamas, Caribbean Mexico, Cuba, Jamaica, Puerto Rico, the US Virgin Islands, Bonaire and Curacao. In addition, this activity includes observations by the Nekton submersibles at Jamaica and Belize, and the Alvin at the Bahamas (again during the 1970s). During the 2010s, this effort was renewed by observations and collecting by the Smithsonian Institution’s DROP program using the submersibles Curasub at Curacao, Bonaire, Dominica and St Eustatius, and the Idabel at Roatan.No similarly extensive set of submersible-based studies and collections of deep-reef fishes exists for any other equivalent biogeographic region in the tropics. Hence, the focus of quantitative comparisons with reef-associated fish faunas of most other regions (or locations within the Greater Caribbean that lack deep-reef research) is likely to be on shallow-water reefs.

1. A few, currently unnamed species of reef-associated Gobiids (6 species in two, currently unnamed, genera), Grammatids (2 species of Lipogramma), Labrids (2 species of Decodon) and Serranids (1 species of Baldwinella) for which sufficient ecological information is available have been added to the list.
2. Several deep-living reef-associated species listed in Version 1 as members of the Scorpaenidae have been removed from the list, because systematic rearrangements have placed them in other families (Setarchidae and Sebastidae) that, in the study area, lack shallow members.
3. The family Ophidiidae has about 78 species in 28 genera and three subfamilies in the study are. We include on the list only 41 members of 7 genera known to have shallow-water, reef-associated representatives or representatives that live within the depth range (0-~250m) of reefs dominated by shallow taxa. The remainder are species that live in very deep or oceanic waters and are irrelevant to the reef-associated fish fauna considered here.
4. Habitat classifications of a few species of carangids and lutjanids as pelagic or demersal in version 1 have been changed, based on additional information.

 

Hosting of this database by Zenodo will allow it to be updated through the production of new versions as new information becomes available.

New information, comments or queries relating to the classification or species list should be directed to robertsondr@si.edu.

A compilation of further information about the biological characteristics of fishes on this list can be found in the website: Robertson DR, Van Tassell J (2019) Shorefishes of the Greater Caribbean: online information system. Version 2.0. https://biogeodb.stri.si.edu/caribbean/en/pages

 

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