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Reef-associated Bony Fishes of the Greater Caribbean: a Checklist; Version 2

Robertson, D Ross; Tornabene, Luke

      Reef-associated Bony Fishes of the Greater Caribbean: A Checklist (VERSION 2)

                                                      Date: August 6, 2020

D Ross Robertson

Smithsonian Tropical Research Institute, Balboa, Republic of Panama.

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.

DOI 10.5281/zenodo.3974538

     This checklist includes all species of bony fishes belonging to families that have at least one member known to associate with shallow reefs in the Greater Caribbean biogeographic region. That region extends from southeastern North America, through the Gulf of Mexico and the Caribbean Sea to Guyana, and includes the West Indies and Bermuda. 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 ecological characteristics of its species as described above. 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. Greater Caribbean reefs down to depths of ~250 m have reef-fish faunas dominated by species belonging to typical shallow-reef families (Baldwin et al. 2018). Some of those families have both shallow and deep-living members. All members of shallow-water families that have at least one reef-associated species are included on the list, regardless of their individual depth ranges. Members of a few families of demersal fishes (e.g. Acropomatidae, Symphysanodontidae, Percophidae, Trachichthyidae) found only on deep-water reefs (i.e. below the mesophotic zone), are not included in the list.

     Reef-associated fishes include demersal and benthic species that use hard substrata (coral-, rock- and oyster reefs), and 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, while Demersal species use and rely on both the bottom and the near-bottom water column. 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. Simple, tripartite systems like this cannot capture 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 categories.

     The name of each reef-associated species in the list is highlighted in yellow.

     Over the past decade the IUCN Red List ( has produced a comprehensive set of assessments of Greater Caribbean fishes. Those include assessments for almost all the named 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.

     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 and Bellwood, 2003), by living directly on or within it. Such crypsis relates to the daytime status of those species, as members of some nocturnally active families (e.g. Apogonids) that are cryptic during the day leave their daytime refuges to forage in the open at night, and members of day-active taxa often disappear into the substratum at night. While cryptobenthic species are a major component of the diversity of reef fishes, they typically 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 (Akermann & Bellwood 2000; Willis 2001; Smith-Vaniz et al 2006; Robertson & Smith-Vaniz 2008;  Alzate et al 2014,) or anaesthetics  (Kovacic et al 2012) that flush such fishes out of the substratum for collection and identification. The list indicates which species we have classed as cryptobenthic. Those are species that, due to cryptic characteristics, visual censuses do not necessarily determine their presence or abundance. For example, while some individuals of cryptic species, such as eels, may be noticed on reefs, we do not know what fraction of an eel’s population those visible individuals might 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, are not classed 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 200; 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 and Bellwood, 2003) and </= 10 cm (Miller 979; 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. 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).



  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). This classification relates to the behavior of fishes during the day, as some species that are benthic while they shelter in the substrate during the day become demersal when they move about actively in the water column at night (e.g. apogonids and some holocentrids).
  2. Species were 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 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 II submarine (the Atlantic and Gulf of Mexico coasts of Florida, the Bahamas, Cuba, Jamaica, Puerto Rico, Bonaire and Curacao), observations by the Nekton submersibles at Jamaica and Belize, and the Alvin at the Bahamas (both in 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.
  3. All species on the list, reef-associated and not, are coded as being benthic, demersal or pelagic; as using hard and soft bottoms (for benthic and demersal species); and as being deep or shallow.
  4. 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.
  5. Several deep-living reef-associated species listed in Version 1 as members of the Scorpaenidae have been removed from the list, due to systematic rearrangements having placed them in other families (Setarchidae and Sebastidae) that, in the study area, lack shallow members.
  6. 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.
  7. Habitat classifications of a few species of carangids and lutjanids as pelagic or not in version 1 have been changed, based on additional information.
  8. With the changes in inclusion of species the number of families is unchanged but the number of genera has increased from 303 in Version 1 to 306, and the number of species from 887 (including 44 pelagics) to 904 (including 49 pelagics).


     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

     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.               


Numbers of species of Reef-associated bony fishes in different categories in the table


Number of Reef-associated Bony Fish taxa:    76 Families, 306 Genera, 904 Species

Of the Reef-associated species:     

              Number of pelagic species: 49     

              Number of non-pelagic species: 855

Of the non-pelagic Reef-associated species:     

      Demersal species, number of: 301

              Number of demersal species using hard bottoms: 261 total, 123 hard bottoms only

              Number of demersal species using soft bottoms: 178 total, 40 soft bottoms only

              Number of demersal species using both hard & soft bottoms: 138           

       Benthic species, number of: 554

             Number of benthic species using hard bottoms: 434 total, 284 hard bottoms only

             Number of benthic species using soft bottoms: 270 total, 120 soft bottoms only

             Number of benthic species using both hard & soft bottoms: 150         

     Cryptobenthic species, number of: 533 total, 227 5cm TL, 374 10cm TL 

     Core CRF species, number of: 388 total, 231 5cm TL, 338 10cm TL                                                 

     Shallow species, number of: 724

             Number of Shallow Demersal species: 259 total, 9 ≤ 5cm TL, 23 10cm TL

             Number of Shallow Benthic species: 465 total, 189 5cm TL, 320 10cm TL

             Number of Shallow Cryptobenthic species: 454 total, 189 5cm TL, 316 10cm TL

             Number of Shallow Core CRF species: 330 total, 190 5cm TL, 284 10cm TL

     Deep species, number of: 131

             Number of Deep Demersal species: 42 total, 3 ≤ 5cm TL, 7 ≤ 10cm TL

             Number of Deep Benthic species: 89 total, 38 5cm TL, 58 10cm TL

             Number of Deep Cryptobenthic species: 79 total, 38 ≤ 5cm TL, 58 ≤ 10cm TL

             Number of Deep Core CRF species: 58 total, 41≤ 5cm TL, 54 10cm TL 



     Thanks to Simon Brandl for discussion about terminology and classification of species on the list, and to Christi Linardich for providing a database of IUCN Red List assessments.

Literature Cited:

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

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.

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

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.

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

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

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.


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