
Reef-associated Bony Fishes of the Greater Caribbean: A Checklist (VERSION 5) Date: November 30, 2023 DOI 10.5281/zenodo.10225031 https://zenodo.org/records/10225031 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 Introduction to the Fifth Edition Changes to this edition include: 1. Rearrangements of species in different families following splitting up of some families (e.g. Cyclopsettidae split off from Paralichthyidae; Spratelloididae split off from Clupeidae) and reassignment of some genera from one family to another (e.g. Peristedion has been reassigned to Triglidae) following changes defined in Eschmeyer's Catalog of Fishes. 2. Some previously undescribed species in version four have been described and named (e.g. Polylepion species 1 is now Polylepion gilmorei). 3. Some species have been reclassified as reef associated based on additional information on habitat usage, photographs of fishes in natural habitat, such as those on iNaturalist (https://inaturalist.org ) and information on collection habitat (e.g. Gillellus inescatus). 4. Some species have been reclassified as non-endemic to the región, based on new information about occurrence in Brazil (e.g. Cephalopholis cruentatus), while others have been reclassified as endemics to the región based on taxonomic splits (e.g. Pareques acuminatus, Sphoeroides spengleri). 5. A few newly described, resurrected or otherwise recognized species (e.g. Rhonciscus paucus, Pareques lineatus, Sphoeroides camila) have been added to the list. Note: All changes and new data in the list are indicated in red and all reef-associated species have their names highlighted for ease of recognition. Also note the explanatory text added to the introduction to the Fourth Edition, below. Introduction to the Fourth Edition 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. Those non-endemics are considered to be residents in the Greater Caribbean if there are multiple records of them in that region, particularly records in different years and at different locations. However, species that associate with reefs in Brazil do not necessarily do so in the Greater Caribbean; Mola mola is an example. Information used here was extracted from 185 publications about Greater Caribbean reef-associated fishes and 152 publications on Brazilian reef-associated 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 84 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, Setarchidae, Symphysanodontidae, Trachichthyidae, and Triacanthodidae. Between them those 14 families have a total of ~43 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 introduced non-native species with resident populations in the Greater Caribbean, notably Pterois volitans and Neopomacentrus cyanomos. Reef-associated fishes include not 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, either contributing food to reefs (e. g. parasites removed from pelagic fishes such as jacks by reef fishes) or extracting food from them (e. g. by pelagics such as mackerels (Scomberomorus) preying on spawning reef fishes). 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)
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