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Published July 10, 2020 | Version 1.0
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Reef-associated Bony Fishes of the Greater Caribbean: a Checklist

  • 1. Smithsonian Tropical Research Institute, Panama
  • 2. University of Washington, Seattle, WA USA

Description

REEF-ASSOCIATED BONY FISHES OF THE GREATER CARIBBEAN: A CHECKLIST

DOI  10.5281/zenodo.3939070

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 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) found only on deep-water reefs (i.e. below the mesophotic zone), are not included in the list.

Reef-associated fishes include demersal species found on or in hard substrata (coral-, rock- and oyster reefs), and demersal species found on or in soft bottoms (sand, gravel, mud, seagrass and macroalgal beds growing on sediment, estuaries and mangroves) immediately adjacent to or within the matrices of reefs. They also include some pelagic species that, although they do not use reefs for shelter, facultatively associate with reefs, and are seen over and immediately adjacent to them, and derive food from them. A habitat-use category (hard bottom, soft bottom, pelagic) is assigned to each reef-associated species on the list, with demersal species that use both reef- and soft-bottom habitats being assigned both.

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 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.

Species in each family that are classed as reef-associated have their names highlighted in yellow in the database.

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).

               

            Numbers of species in different categories in the Checklist

____________________________________________________________________________________

Number of Reef-associated Bony Fish taxa:    76 Families, 303 Genera, 887 Species


Of the Reef-associated species:

      Number of demersal species: 843

      Number of demersal species using hard bottoms: 678 total, 398 hard bottoms only

      Number of demersal species using soft bottoms: 445 total, 165 soft bottoms only

      Number of demersal species using both hard & soft bottoms: 280

      Number of demersal species using a single bottom type: 563

      Number of pelagic species: 44

      Number of Cryptobenthic species: 536 total, 226 </= 5cm TL, 374 </= 10cm TL

      Number of Core CRF species: 378 total, 222 </= 5cm TL, 328 </= 10cm TL  

____________________________________________________________________________________

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/page

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.

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.  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.

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.

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.

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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