Reef-associated Bony Fishes of the Greater Caribbean: a Checklist; Version 3

      Reef-associated Bony Fishes of the Greater Caribbean: A Checklist (VERSION 3)

 

                                                      Date: November 18, 2020

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

https://zenodo.org/record/4279301#.X7UULWhKg2w

DOI: 10.5281/zenodo.4279301

This third edition of the checklist includes bony fishes belonging to families present in the Greater Caribbean biogeographic region that have members known to associate with shallow (0 – 40 m depth) reefs. That region extends from southeastern North America, through the Gulf of Mexico and the Caribbean Sea to somewhere around 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 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 96.7% 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.

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 those 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 reef-associated fishes typically found only on deep-water reefs (i.e. below the 150 m lower limit of the mesophotic zone), are not included on this list, among them the Acropomatidae, Aulopidae, Berycidae, Caproidae, Gempylidae, Grammicolepidae, Polymixiidae, Polyprionidae, Scombropidae, Sebastidae, Symphysanodontidae, Trachichthyidae, and Tricanthodidae.

Reef-associated fishes include demersal and benthic species that use consolidated hard substrata (coral- and rocky 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, whereas Demersal species use and rely on both the bottom and the near-bottom water column. These two categories relate to the behavior of fishes during the day, which, in some cases, changes at night. For example,  Apogonids, Holocentrids and Anomalopids 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.

Simple systems of habitat characterization like that 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). 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, with clearcut descriptions of actual behavior. Otherwise virtually everything could end up classed as reef associated.

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 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 (Ackerman & 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. While individuals of some cryptobenthic species, such as moray eels and cardinal-fishes 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. 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). In this edition we have added one family to the Core CRF list (see below).

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.

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).

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   

 

Numbers of species of Reef-associated Bony Fishes in different categories in the Table

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Number of Reef-associated Bony Fish taxa:    83 Families, 338 Genera, 981 Species

Of those:

Number of pelagic species: 78

                       Red List:  0 ND; 4 DD, 72 LC, 0 NT, 2 VU, 0 EN, 0 CR

                       Number of endemics:10

                       Number of small-range species: 13

                       Number of very-small range species: 1

Number of non-pelagic species: 903

Of the non-pelagic Reef-associated species:

                       Red List:  32 ND; 91 DD, 725 LC, 12 NT, 28 VU, 14 EN, 1CR

                       Number of endemics: 497

                       Number of small-range species: 291

                       Number of very-small range species: 59    

             Demersal species, number of: 316

                       Number of demersal species using hard bottoms: 260 total, 119 hard bottoms only

                       Number of demersal species using soft bottoms: 197 total, 56 soft bottoms only

                       Number of demersal species using both hard & soft bottoms: 141     

            Benthic species, number of: 587

                       Number of benthic species using hard bottoms: 433 total, 278 hard bottoms only

                       Number of benthic species using soft bottoms: 309 total, 154 soft bottoms only

                       Number of benthic species using both hard & soft bottoms: 155   

          Cryptobenthic species, number of: 535 total, 227 ≤ 5cm TL, 376 ≤ 10cm TL.

                       Red List:  20 ND; 69 DD, 422 LC, 2 NT, 15 VU, 7 EN, 0 CR

                       Number of endemics:365

                       Number of small-range species: 216

                       Number of very-small range species: 47

          Core CRF species, number of: 414 total, 216 ≤ 5cm TL, 362 ≤ 10cm TL

                       Red List:  19 ND; 67 DD, 305 LC, 2 NT, 14 VU, 7 EN, 0 CR

                       Number of endemics:331

                       Number of small-range species: 192

                       Number of very-small range species: 43                                                  

        Shallow species, number of: 769

                       Number of Shallow Demersal species: 272 total, 9 ≤ 5cm TL, 21 ≤ 10cm TL

                       Number of Shallow Benthic species: 497 total, 189 ≤ 5cm TL, 328 ≤ 10cm TL

                       Number of Shallow Cryptobenthic species: 455 total, 189 ≤ 5cm TL, 318 ≤ 10cm TL

                       Number of Shallow Core CRF species: 355 total, 195 ≤ 5cm TL, 308 ≤ 10cm TL

      Deep species, number of: 134

                       Number of Deep Demersal species: 44 total, 3 ≤ 5cm TL, 7 ≤ 10cm TL

                       Number of Deep Benthic species: 90 total, 38 ≤ 5cm TL, 58 ≤ 10cm TL

                       Number of Deep Cryptobenthic species: 80 total, 38 ≤ 5cm TL, 58 ≤ 10cm TL

                       Number of Deep Core CRF species: 59 total, 41 ≤ 5cm TL, 54 ≤ 10cm TL 

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Acknowledgements:

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.

 

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