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Corresponding author: D Ross Robertson ( drr@stri.org ) Academic editor: Kyle Piller
© 2016 D Ross Robertson, Horacio Perez-España, Enrique Nuñez Lara, Francisco Puc Itza, Nuno Simões.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Robertson DR, Perez-España H, Lara EN, Itza FP, Simoes N (2016) The fishes of Cayo Arcas (Campeche Bank, Gulf of Mexico): an updated checklist. ZooKeys 640: 139-155. https://doi.org/10.3897/zookeys.640.10862
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Cayo Arcas is a small, offshore reef complex on the southwest corner of Campeche Bank, Gulf of Mexico. The only published information (from 2000) on the fishes of that reef refers to 37 species. Here additional information is added, some from unpublished observations during the 1980s, as well as author observations made during 2013 and 2016. These bring the checklist of that reef’s fishes up to 162 species. The possible effects of the limited number of fish habitats available at Cayo Arcas on the composition of its fish fauna are discussed. The Indo-Pacific damselfish Neopomacentrus cyanomos (Bleeker, 1856) was first recorded in the Atlantic in mid-2013, on shoreline reefs in the southwest corner of the Gulf of Mexico. Recently reviewed underwater photographs show that N. cyanomos also was present at Cayo Arcas in mid-2013, 350 km from the first-record site. Hence it evidently had a substantial population in the southwest Gulf of Mexico in 2013, and must have arrived in there long before that year.
Endemic species, invasive damselfish species, reef-fishes, reef-habitat, southwest Gulf of Mexico
The southwest Gulf of Mexico has relatively few coral reefs. Only a small number of these are offshore reefs on the broad, shallow Campeche Bank that extends 200+ km north from the Yucatan Peninsula. These include Alacran Reef, and a set of about 10 small submerged banks and emergent reefs scattered along the western edge of that bank (http://www.gulfbase.org/reef/). The reef-fish faunas of most of those offshore reefs are not well documented. Only two of them have substantial published checklists available: Alacran Reef, a large emergent reef in the center of the bank and the largest reef in the region (see
Here an updated checklist for the fishes of the Cayo Arcas reef complex is presented that draws on both old and new information. The object of publishing this checklist is to improve understanding of the distributions of reef fishes in the southwest Gulf of Mexico, which hosts a number of endemic reef fishes, has a different marine environment to the rest of that gulf (
This updated list is not complete, because the dives on which it is based did not provide comprehensive coverage of fishes across the full range of habitats available. Furthermore, because the list is based on diver observations, cryptic fishes that live within the reef matrix or within fringing soft sediments undoubtedly are under-represented. The only really effective way to comprehensively sample such fishes, which represent as much as half of any reef-associated fauna in the neotropics, is with the use of small ichthyoside stations (
The Cayo Arcas reef complex is situated at 20.21°N, -91.98°W, 145 km from the mainland. This complex comprises reefs fringing a cluster of three sand cays that are spread over an area of ~4 km by ~2.6 km (Figure
Mangroves and beds of seagrasses (turtlegrass, Thalassia testudinum Koenig 1805, and manatee-grass, Syringodium filiforme Kutzing in Hohenacker, 1860, represent major ancillary habitats adjacent to many reefs in the tropical northwest Atlantic. Those habitats are used by many reef fishes, often as nursery habitats. While turtlegrass often forms large beds within which dense growths of flat, strip-like blades provide ample shelter for fish, manatee-grass has thin, thread-like blades and a sparser growth pattern that provide lower quality shelter for reef fishes. There are no mangroves at Cayo Arcas. The www.gulfbase.org marine species list for reefs in the Mexican section of the Gulf of Mexico indicates that turtlegrass does occur at Cayo Arcas (see p 27 of http://www.gulfbase.org/checklist/pdfs/marine-spp-list.pdf). That gulfbase list cites
The earliest information available on Cayo Arcas fishes comes from an unpublished Master’s thesis by
Table
Preliminary checklist of fishes observed at Cayo Arcas, Campeche Bank. Abundance and distribution in 2016: Rare (1 seen during expedition); Uncommon (< 5); Occasional (~ 20, at multiple dive sites); Common, widespread (scores, at most/all dive sites); Locally common (scores to hundreds at 1–2 dive sites); Abundant, widespread (hundreds to thousands at most/all dive sites); Platform (present on oil-loading platform 1.5 km from reef complex). Abundance 2013: Mean no. fish/transect by HP-E and ENL (see methods); Abundance 1984-86: G&C 2000 =
Taxon | Abundance and distribution 2016 | Abundance 2013 HP-E/ENL | Abundance 1984-86 G&C 2000/ G 1988 |
---|---|---|---|
UROTRYGONIDAE | |||
Urobatis jamaicensis | Rare | ||
DASYATIDAE | |||
Hypanus americana* | Uncommon | 0/0.1 | |
AETOBATIDAE* | |||
Aetobatus narinari | 0/0.1 | ||
MEGALOPIDAE | |||
Megalops atlanticus | Locally common (schools); platform | 0/0.2 | |
MURAENIDAE | |||
Gymnothorax funebris | Uncommon | ||
Gymnothorax moringa | Rare | 0/0.1 | |
ATHERINIDAE | |||
Atherina harringtonensis * | Locally common (large schools) | ||
Atherinomorus stipes | Locally common (large schools) | ||
HOLOCENTRIDAE | |||
Holocentrus adscensionis | Common, widespread | 0.097/0.250 | |
Holocentrus rufus | Occasional, widespread | 0/0.9 | |
Myripristis jacobus | Uncommon | ||
Neoniphon vexillarium | Uncommon | ||
AULOSTOMIDAE | |||
Aulostomus maculatus | Uncommon | 0.042/0.125 | 0/0.2 |
SCORPAENIDAE | |||
Pterois volitans | Occasional, widespread; platform (species note) | ||
SERRANIDAE | |||
Cephalopholis cruentata | Common, widespread; platform | 0.042/0.125 | 0/0.2 |
Cephalopholis fulva | 0.014/0.042 | 0/0.3 | |
Epinephelus adscensionis | Common, widespread | 0.055/0.083 | 0/1.1 |
Epinephelus guttatus | Common, widespread | 0.111/ 0.333 | 0.05/1.0 |
Hypoplectrus aberrans | Locally common | ||
Hypoplectrus ecosur | Locally common | ||
Hypoplectrus gemma | 0.014/0 | ||
Hypoplectrus indigo | Uncommon | ||
Hypoplectrus maculiferus | Rare | ||
Hypoplectrus nigricans | Occasional, widespread | 0.028/0.083 | 0/0.1 |
Hypoplectrus puella | Occasional, widespread | 0.097/0.250 | 0/0.4 |
Hypoplectrus randallorum | Rare | ||
Hypoplectrus unicolor | Rare | ||
Mycteroperca bonaci | Common, widespread; platform | 0/1.0 | |
Mycteroperca interstitialis | Locally common | ||
Mycteroperca microlepis | Uncommon | ||
Mycteroperca phenax | Uncommon | 0.014/0.042 | |
Mycteroperca tigris | Occasional, widespread | 0.014/0.042 | 0/0.4 |
Paranthias furcifer | Locally common (aggregations); platform | ||
Serranus baldwini | Rare | ||
Serranus tabacarius | Locally common | ||
Serranus tigrinus | Common, widespread | 0.083/0.167 | 0/0.7 |
Serranus tortugarum | Locally common (aggregations) | ||
GRAMMATIDAE | |||
Gramma loreto | Common, widespread | 0.139/ 0.208 | 0.10/0.7 |
MALACANTHIDAE | |||
Malacanthus plumieri | 0/0.1 | ||
OPISTOGNATHIDAE | |||
Opistognathus aurifrons | Locally common (aggregations) | 0.028/0 | |
RACHYCENTRIDAE | |||
Rachycentron canadum | Rare; platform only | ||
CARANGIDAE | |||
Caranx bartholomaei | Uncommon (small schools) | ||
Caranx crysos | Common, widespread (aggregations) | 0/0.3 | |
Caranx latus | Common, widespread (aggregations) | 0.028/0 | |
Caranx lugubris | Uncommon (aggregation); platform only | ||
Caranx ruber | Locally common (aggregations) | 0.305/0.208 | 0.05/0.7 |
Decapterus macarellus | Locally common (school) | ||
Trachinotus carolinus | Rare | ||
LUTJANIDAE | |||
Lutjanus analis | Locally occasional | 0/0.1 | |
Lutjanus apodus | (Species note) | 0.05/0.1 | |
Lutjanus buccanella | Occasional (juveniles) | ||
Lutjanus griseus | Common, widespread; platform | 0/1.2 | |
Lutjanus jocu | Common, widespread | 0.028/0.083 | |
Lutjanus mahogoni | Occasional, widespread | 0/0.1 | |
Lutjanus synagris | Occasional (juveniles) | ||
Ocyurus chrysurus | Abundant, widespread; platform | 2.619/7.333 | 0.05/1.1 |
GERREIDAE | |||
Gerres cinereus | Uncommon | 0.014/0.042 | 0/0.1 |
HAEMULIDAE | |||
Anisotremus virginicus | Locally common (school) | 0.05/0 | |
Emmelichthyops atlanticus | Locally common (schools) | ||
Haemulon aurolineatum | Common, widespread (aggregations) | 0.014/0 | 0.05/0 |
Haemulon carbonarium | Uncommon | 0.05/0 | |
Haemulon chrysargyreum | Locally common (schools) | 0.05/0.4 | |
Haemulon flavolineatum | Common, widespread | 1.067/1.125 | 0.73/2.4 |
Haemulon macrostomum | Uncommon | 0/0.7 | |
Haemulon melanurum | Uncommon | ||
Haemulon plumierii | Uncommon | 0.083/0 | 0.05/0 |
Haemulon sciurus | Uncommon | 0.05/0.4 | |
Haemulon striatum | Locally common (large schools) | ||
Haemulon vittatum * | Locally common (large schools) | 1.386/0 | 12.83/1.7 |
SPARIDAE | |||
Calamus calamus | Common, widespread | 0.055/0 | |
Calamus nodosus | Common, widespread | ||
SCIAENIDAE | |||
Equetus punctatus | Uncommon | ||
Pareques acuminatus | Uncommon | ||
MULLIDAE | |||
Mulloidichthys martinicus | Common, widespread (schools) | 0.014/0.083 | 0/2.4 |
Pseudupeneus maculatus | Locally common | 0.028/0 | |
PEMPHERIDAE | |||
Pempheris schomburgkii | Locally common (aggregations) | ||
KYPHOSIDAE* | 3.672/1.542 | ||
Kyphosus bigibbus | Occasional | ||
Kyphosus cinerascens | Common, widespread | ||
Kyphosus sectatrix | Common, widespread | 0/0.5 | |
Kyphosus vaigiensis | Common, widespread | ||
CHAETODONTIDAE | |||
Chaetodon aculeatus | 0/0.2 | ||
Chaetodon capistratus | Occasional | ||
Chaetodon ocellatus | Common, widespread | 0.194/0.458 | 0/1.5 |
Chaetodon sedentarius | Common, widespread | 0.152/0.250 | |
POMACANTHIDAE | |||
Holacanthus bermudensis | Uncommon | ||
Holacanthus ciliaris | Common, widespread; platform | 0/0.4 | |
Holacanthus tricolor | 0.05/0 | ||
Pomacanthus arcuatus | Locally common | 0.055/0 | 0/1.2 |
Pomacanthus paru | Locally common | 0.014/0.042 | 0.05/0.5 |
CIRRHITIDAE | |||
Amblycirrhitus pinos | Rare; platform only | ||
POMACENTRIDAE | |||
Abudefduf saxatilis | Locally abundant, widespread | 1.108/1.167 | 7.16/2.1 |
Chromis cyanea | Locally common | 0.430/0 | 0.05/0.2 |
Chromis insolata | Locally common | 0.028/0.083 | 0/0.2 |
Chromis multilineata | Abundant, widespread | 47.056/ 90.750 | 28.93/3.2 |
Chromis scotti | Common, widespread; platform | 0.194/0 | |
Microspathodon chrysurus | Common, widespread | 0.152/0.375 | 1.14/2.3 |
Neopomacentrus cyanomos | Abundant, widespread (aggregations); platform. (species note) | Present/0 | |
Stegastes adustus | Abundant, widespread (species note) | 0.443/0.167 | |
Stegastes diencaeus | 0.055/0.167 | 1.14/1.4 | |
Stegastes leucostictus | Locally common | 0.222/0.417 | 0/0.3 |
Stegastes partitus | Abundant, widespread; platform | 3.603/4.667 | 0.05/0.9 |
Stegastes planifrons | Abundant, widespread | 0.679/1.500 | 12.93/3.7 |
Stegastes xanthurus* | Abundant, widespread (species note) | 0.679/0.542 | 0.05/0 |
LABRIDAE | |||
Bodianus pulchellus | Locally common; platform only | ||
Bodianus rufus | Common, widespread | 0.651/1.208 | 0/1.1 |
Clepticus parrae | Locally common, schools | 1.261/3.792 | 9.73/0.4 |
Halichoeres bivittatus | Very common, widespread | 0.443/0.083 | 0/0.9 |
Halichoeres burekae | Abundant, widespread (species note) | 5.085/0.083 | |
Halichoeres garnoti | Common, widespread | 2.245/4.452 | 0.05/0.2 |
Halichoeres maculipinna | Common, widespread | 0.402/1.208 | 1.36/1.3 |
Halichoeres poeyi | 0.319/0 | ||
Halichoeres radiatus | Common, widespread | 0.222/0.083 | 0/1.0 |
Lachnolaimus maximus | Uncommon | ||
Thalassoma bifasciatum | Abundant, widespread; platform | 10.752/11.833 | 1.22/2.1 |
Xyrichtys splendens | Uncommon; local | ||
SCARIDAE | |||
Cryptotomus roseus | Occasional | ||
Scarus coelestinus | Occasional | 0/0.6 | |
Scarus coeruleus | Common, widespread | 0.152/0 | 0.05/1.6 |
Scarus guacamaia | Occasional, widespread | 0.042/0 | 0/0.3 |
Scarus iseri | Common, widespread | 2.480/3.042 | |
Scarus taeniopterus | Common, widespread | 0.111/0.333 | 5.40/1.3 |
Scarus vetula | Common, widespread | 1.178/2.875 | 5.23/3.4 |
Sparisoma atomarium | Locally common | 0.055/0.167 | |
Sparisoma aurofrenatum | Common, widespread | 1.857/2.833 | 0/0.4 |
Sparisoma chrysopterum | Occasional, widespread | 0.014/0.042 | 0.05/1.7 |
Sparisoma radians | Common, widespread | ||
Sparisoma rubripinne | Locally common | 0.291/0.500 | 0/1.5 |
Sparisoma viride | Very common, widespread | 1.136/1.833 | 3.63/3.2 |
TRIPTERYGIIDAE | |||
Enneanectes boehlkei | Present (species note) | ||
BLENNIIDAE | |||
Entomacrodus nigricans | Uncommon; platform only | ||
Hypsoblennius invemar | Locally common; platform only | ||
Ophioblennius macclurei * | Common, widespread | 0.028/0 | 0/01.2 |
Parablennius marmoreus | Locally common; platform | ||
Scartella cristata | Locally common; platform only | ||
LABRISOMIDAE | |||
Malacoctenus aurolineatus | Locally common | ||
Malacoctenus macropus | Common, widespread | ||
Malacoctenus triangulatus | Common, widespread | 0.097/0.083 | |
Starksia ocellata | Present (species note) | ||
GOBIIDAE | |||
Coryphopterus dicrus | Abundant, widespread | ||
Coryphopterus glaucofraenum | Abundant, widespread | 0.319/0 | |
Coryphopterus hyalinus/personatus | Abundant, widespread (species note) | 4.432/5.625 | |
Elacatinus oceanops | Common, widespread | 0.222/0.292 | |
Elacatinus xanthiprora | Uncommon | ||
Gnatholepis thompsoni | Abundant, widespread | 0.249/0.042 | |
PTERELEOTRIDAE | |||
Ptereleotris calliura | Locally common (aggregations) | ||
ACANTHURIDAE | |||
Acanthurus chirurgus | Common, widespread | 0.097/0.167 | 0.05/0.8 |
Acanthurus coeruleus | Common, widespread | 0.139/0.250 | 0.05/1.6 |
Acanthurus tractus * | Common, widespread (species note) | 0.291/0.875 | 1.09/1.2 |
SPHYRAENIDAE | |||
Sphyraena barracuda | Occasional, widespread | 0.014/0.042 | 0/1.1 |
BALISTIDAE | |||
Balistes capriscus | Rare | ||
Balistes vetula | 0.042/0.125 | ||
Canthidermis sufflamen | Occasional, widespread | 0/0.1 | |
Melichthys niger | Locally common | 0/0.2 | |
Xanthichthys ringens | 0/0.2 | ||
MONACANTHIDAE | |||
Aluterus scriptus | Occasional, widespread | 0/0.3 | |
Cantherhines pullus | Rare | 0.028/0 | |
Monacanthus tuckeri | Locally common | 0.028/0 | |
OSTRACIIDAE | |||
Lactophrys bicaudalis | Rare | 0/0.1 | |
Lactophrys triqueter | Occasional, widespread | 0.014/0.042 | 0/0.8 |
TETRAODONTIDAE | |||
Canthigaster rostrata | Common, widespread | 0.152/0.250 | 0.05/0.8 |
Sphoeroides testudineus | Rare | ||
DIODONTIDAE | |||
Diodon hystrix | Uncommon | 0/0.4 |
Pterois volitans (Linnaeus, 1758). This invasive Indo-Pacific lionfish was first reported on the USGS invasive species website at an oil platform near Cayo Arcas in late 2012 (http://nas.er.usgs.gov/queries/SpecimenViewer.aspx?SpecimenID=292473. Fewer than a dozen individuals of this species were seen by the group of eight divers during the August 2016 expedition. Those occurred at depths of between 8-20 m on the reef, and at 30 m on the oil platform. Lionfish were first reported at Alacran reef, 350 km northeast of Cayo Arcas, in 2010, and on coastal reefs ~450 km west of Cayo Arcas in 2012 (http://nas.er.usgs.gov/queries/collectioninfo.aspx?SpeciesID=963). At Alacran Reef lionfish seem to be concentrated at mesophotic depths (
Hypoplectrus nigricans (Poey, 1852). This species is known to vary geographically in color and shape (
Lutjanus apodus (Walbaum, 1792).
Neopomacentrus cyanomos. During a visit to Cayo Arcas in April 2016 NS found it to occur both on the reefs and the adjacent oil-loading platform, where it was superabundant (
Neopomacentrus cyanomos at Cayo Arcas, July 2013 (Photo HP-E); 11 N. cyanomos are visible, and can be identified by the presence of a black blotch on the shoulder, and a large white blotch on the rear edge of the dorsal fin; a single Chromis multilineata (without the shoulder- and dorsal-fin blotches) is present in the foreground immediately above the brain coral.
Stegastes adustus (Troschel in Muller, 1865) and S. diencaeus (Jordan & Rutter, 1897). In 2016 S. adustus, a benthic-feeding, omnivorous damselfish, was common on all hard reef substrata not covered by live corals, at depths of 0.5-7 m. HP-E recorded S. adustus but not S. diencaeus, while ENL recorded both species. In their list
Halichoeres burekae Weaver & Rocha, 2007. This recently described species is endemic to the southwest and northwest Gulf of Mexico. Its known range extends from Alacran Reef on Campeche Bank to the Flower Garden Banks at the edge of the continental shelf off Texas (see http://biogeodb.stri.si.edu/caribbean/en/thefishes/species/4707; and http://maps.iucnredlist.org/map.html?id=187608). At Alacran Reef, 350 km northeast of Cayo Arcas,
Scarus guacamaia Cuvier, 1829. At Cayo Arcas one or two adults of this species were seen on each of eight of the 14 dives on the reef complex in 2016. S. guacamaia also was recorded by
Enneanects boehlkei Rosenblatt, 1960 and Starksia ocellata (Steindachner, 1876). One individual of each of these two cryptic species was collected as bycatch while using clove oil to anaesthetize Neopomacentrus cyanomos for collection.
Coryphopterus hyalinus Bohlke & Robins, 1962, and C. personatus (Jordan & Thompson, 1905). This pair of sister species (see
The present checklist includes 162 species from 41 families. The great majority of those species are widespread in the Greater Caribbean, with only Halichoeres burekae and Hypoplectrus gemma Goode and Bean, 1882 representing species that are entirely (or almost so) restricted to the Gulf of Mexico. Most of the geographically widely distributed species that were common in the 1980s and 2013 surveys at Cayo Arcas also were common there in 2016. There are several noteworthy features of the suite of species found at Cayo Arcas: Chromis multilineata (Guichenot, 1853), which is a common species widely distributed on reefs throughout the Greater Caribbean, was notably abundant in all surveys made between the 1980s and 2016. H. burekae, a western Gulf of Mexico endemic, was (perhaps) the most common labrid at Cayo Arcas, and was more abundant there than has been recorded anywhere else previously. This species is listed as Endangered (i.e. at a high risk of extinction) by the IUCN Red List (http://www.iucnredlist.org/details/187608/0), due to the small size of its geographic range and the paucity of reef habitat in that range. The abundance of this species at Cayo Arcas has substantial conservation significance for this species, as it indicates that the set of small offshore reefs scattered along the western side of Campeche Bank may be essential for its continued existence. None of those reefs is a yet designated a marine protected area.
Neopomacentrus cyanomos was first recorded in the Atlantic by
Two species that are commonly found on reefs nearby and more widely within the greater Caribbean were conspicuously absent/rare at Cayo Arcas: Lutjanus apodus and Stegastes diencaeus. L. apodus is one of the species thought to be strongly dependent on mangroves as nursery habitat in the Caribbean area (
Future, comprehensive faunal surveys should be made of the reef fish faunas not only of Cayo Arcas but also other, more poorly known emergent reefs and submerged banks near the outer edges of Campeche Bank to assess their reef fish faunas, to gain a better understanding of the biogeography of their fishes, and their importance for conservation, as well as assessing them as natural biogeographical experiments. Those surveys should also involve genetic analyses to determine the extent to which the regional fauna and faunas of individual reefs include cryptic endemic species.
Thanks to Quetzalli Hernandez, whose leadership of the Cayo Arcas expedition of August 2016 greatly facilitated work there, and to essential support by the captain (Héctor Agustin Torres Murillo) and crew of the Buque de Investigación Oceanográfico Arm “Hondo” (BI-06), Secretaria de Marina, Estados Unidos Mexicanos. The 2016 expedition was financed by grants to NS from the Harte Institute (Biodiversity of the southern Gulf of Mexico) and CONABIO (NE018; Actualización del conocimiento de la diversidad de especies de invertebrados marinos bentónicos de aguas someras (< 50m) del Sur del Golfo de México. Fieldwork in 2013 was funded by SEP-CONACyt grants 69747 to HP-E and 108083 to ENL.