Coralliozetus cardonae: mode D-XVIII,11-12 sA-19-20 P-12-13 (D-XVII-IXX,10-13 sA-18-24) pop counts vary
Emblemariopsis carib: mode D-XX,11 sA-20 P-13 (D-IXX-XXI,10-12 sA-19-21) new sp 2010, Haiti to SE Caribbean
Emblemariopsis ruetzleri: mode D-XX,11-12 sA-20 P-14 (D-XX-XXI,10-13 sA-19-21) MAB, Panama, USVI
Emblemariopsis leptocirris: mode D-XX,11-13 sA-20-22 P-13 (D-IXX-XXI,10-13 sA-20-22) widespread
Emblemariopsis dianae: mode D-XX,11-12 sA-20 P-13 (D-XVIII-XX,10-12 sA-19-21) Belize mid-shelf, Honduras
Emblemariopsis occidentalis: mode D-XX,11-12 sA-20-21 P-13 (D-IXX-XXI,11-13 sA-20-21) Bahamas (species?)
Emblemariopsis arawak: mode D-XX,12-13 sA-21 P-13 (D-IXX-XXI,12-13 sA-20-22) new sp 2010, Antilles plus
Emblemariopsis tayrona: mode D-XX-XXI,11-12 sA-20-21 P-13 (D-IXX-XXI,11-13 sA-18-22) Colombia, eastward
Emblemariopsis ramirezi: mode D-XX-XXI,11-12 sA-20-21 P-13; Venezuela and S. Antilles only
Emblemariopsis bahamensis: mode D-XX-XXI,12-13 sA-21-22 P-13 (D-XX-XXI,12-13 sA-21-23) Bahamas, Antilles
Emblemariopsis bottomei: mode D-XX-XXI,12-13 sA-21 P-13 (D-XX-XXI,12-13 sA-21) SE Caribbean
Emblemariopsis pricei: mode D-XXI,13 sA-22-23 P-14 (D-XX-XXII,12-14 sA-21-24 P-13-15) Belize & Honduras
Emblemariopsis randalli: mode D-XXI,12-13 sA-21 P-14 (D-XX-XXI,10-13 sA-20-22) Venezuela coastal
Emblemariopsis diaphana: mode D-XX-XXI,13 sA-21-22 P-13 (D-XX-XXI,12-14 sA-21-23) Florida Keys, GOM
Emblemaria vitta: mode D-IXX,13 sA-19 P-13; Navassa (also Bahamas, Antilles, BZ/Hond); pop counts vary
Emblemaria piratula: mode D-IXX,14-15 sA-21 P-13 (D-XVII-XX,13-16 sA-20-21) deep, Gulf of Mexico
Emblemaria pandionis: mode D-XX-XXI,14-15 sA-21-23 P-13 (D-IXX-XXII,13-17 sA-20-23) population counts vary
Emblemaria caycedoi: mode D-XXI,14-15 sA-22 P-13 (D-IXX-XXI,14-15 sA-22-23) SW Caribbean
Emblemaria diphyodontis: mode D-XXI-XXII,14-15 sA-23 P-13 (D-XX-XXII,13-15 sA-22-24) Colombia, VZ, SE Car
Emblemaria biocellata: mode D-XXII,14-15 sA-22-23 P-13; deep-water, Colombia to Suriname
Emblemaria culmenis: mode D-XXII,15 sA-24 P-13; deep-water, Venezuela (culmenis not "E. culmensis")
Emblemaria caldwelli: mode D-XXII,14 sA-22 P-14 (D-XXI-XXIII,13-15 sA-21-23) reef walls, Bahamas, Belize
Emblemaria hyltoni: mode D-XXI-II,14-15 sA-23 P-14 (D-XXI-XXIII,14-16 sA-22-23) Bay Islands of Honduras
Protemblemaria punctata: mode D-XX,15-16 sA-23 P-14 (D-IXX-XXI,13-17 sA-22-23) NE Venezuela
Ekemblemaria nigra: mode D-XXI,17 sA-24 P-14 (D-XX-XXII,15-18 sA-23-25) Panama & Colombia
Hemiemblemaria simulus: mode D-XXII,16-17 sA-22-23 P-14 (D-XX-XXIII,16-17 sA-22-23) wide, not E or S Car
Lucayablennius zingaro: mode D-IXX,20 sA-23 P-13 (D-XVIII-XX,19-21 sA-22-23) widespread, not FL
Acanthemblemaria rivasi: mode D-XXII,12 sA-22 P-13 (D-XXI-XXII,11-13 sA-22-23) CR, Panama, SAm mainl
Acanthemblemaria maria: mode D-XXII-XXIII,13-14 sA-23-25 P-13 (D-XXI-XXIII,12-14 sA-22-25) not FL or SAm
Acanthemblemaria spinosa: mode D-XXI,14-15 sA-23-24 P-13 (D-XX-XXII,13-16 sA-22-25) not FL, Pan, SAm mainl
Acanthemblemaria harpeza: mode D-XXI,14-15 sA-24 P-13 (D-XXI,14-15 sA-23-24) Navassa
Acanthemblemaria sp.: mode D-XX-XXI,15 sA-23 P-13 (D-XX-XXI,15 sA-23) Panama
Acanthemblemaria paula: mode D-IXX,16-17 sA-23-24 P-13 (D-XVIII-XXI,15-19 sA-22-25) Belize
Acanthemblemaria aspera: mode D-XX-XXI,15-16 sA-23-24 P-13 (D-IXX-XXII,14-17 sA-21-25) wide, not SE Car
Acanthemblemaria johnsoni: mode D-XXIV,12-13 sA- P-13; NE Venezuela to Tobago
Acanthemblemaria medusa: mode D-XXI-XXII,16 sA-26 P-13 (D-XXI-XXIII,15-17 sA-25-27) SE Caribbean
Acanthemblemaria betinensis: mode D-XXIII,15 sA-24 P-13 (D-XXII-XXV,14-16 sA-23-25) C. Rica, Pan, Colombia
Acanthemblemaria greenfieldi: mode D-XXII-XXIII,17 sA-27-28 P-13 (D-XXI-XXIV,16-19 sA-26-29) Belize W Car isl
Acanthemblemaria chaplini: mode D-XXII-XXIII,17-18 sA-27-28 P-13 (D-XXI-XXIII,17-21 sA-26-29) Bahamas FL Cuba
Acanthemblemaria cubana: mode D-XXI-XXII,19-20 sA-26-28 P-13; Cuba (22,20,28) and Panama (21,19,26)
Stathmonotus tekla: mode D-XLI-XLII sA-23-24 P-8-9 segcaud 11 (D-XXXIX-XLIV sA-22-25) Bah, N & W Caribbean
Stathmonotus stahli: mode D-XLII-XLIII sA-24 P-8-9 segcaud 12 (D-XLI-XLV sA-23-25) Lesser Antilles, SE Carib
Stathmonotus gymnodermis: mode D-XLII-XLIV sA-23-24 P-8-9 sc 11 (D-XLI-XLVI sA-21-26) counts vary, low North
Stathmonotus hemphilli: mode D-L sA-27 P-4-5 (D-XLV-LIII sA-23-29) counts vary with population
Chaenopsis roseola: mode D-XVII-XVIII,26-28 sA-29-30 P-13; E. Gulf of Mexico, deep
Chaenopsis megalops: mode D-XVII-XVIII,35-36 sA-36 P-13; Colombia, deep
Chaenopsis resh: mode D-XVIII-IXX,35-36 sA-35-36 P-13 (D-XVII-XX,34-36 sA-35-37); NE Venezuela
Chaenopsis limbaughi: mode D-XVIII-XXI,31-36 sA-34-37 P-12-13; widespread
Chaenopsis ocellata: mode D-XVIII-XX,33-36 sA-34-37 P-12-13; widespread
Chaenopsis stephensi: mode D-XVII,28-30 sA-30-31 P-13; Venezuela & 275 m at Arrowsmith Bank off Yucatan
Most chaenopsid larvae are unremarkable small blennioid larvae with pointed snouts, although Chaenopsis larvae are longer and pike-like. Almost all chaenopsid larvae identified to date have the same limited pattern of melanophores: a ventral midline series including one or two around the isthmus and a long anal row (sometimes extending onto the ventral caudal peduncle), along with the internal retroperitoneal (Stathmonotus hemphilli has additional internal melanophores).
The absence of cranial, nuchal, cheek, otic, dorsal and caudal-fin melanophores rules out most other blennioid larvae. The exception is the labrisomid genus Starksia, some of whose larvae share the limited melanophore pattern, but they are easily distinguished by having many fewer dorsal-fin soft rays (at most 9) and fewer melanophores in the anal row (fewer than 19 total pvm). Chaenopsid larvae have 3 to 6 procurrent caudal-fin rays, fewer than the labrisomids Labrisomus and Malacoctenus and triplefins Enneanectes, but overlapping with the labrisomids Starksia and Paraclinus.
The chaenopsids have, with few exceptions, relatively uniform larvae and thus fin-ray counts are needed for most genus and species identifications by eye. Although adults can have distinctive morphologies, especially in the shape and ornamentation of the head and variously extended fin elements, larvae are adapted to a pelagic life and do not show any of these distinguishing features. They are all slender and have pointed snouts and virtually all share the basic limited melanophore pattern, a ventral midline series, until they begin to develop their species-specific markings during transition.
Although there are many chaenopsid genera and a proliferation of species, the family can be broken down into four basic groups. There are the tiny and mostly transparent glass blennies of the Emblemariopsis group (including Coralliozetus) that have very small-settling larvae and the lowest fin-ray counts in the family (E. bottomei, by Michael Brogan, at left). Their counts overlap somewhat with the highest-count labrisomids and the lowest of the Acanthemblemaria species. The emblemariopsids have only 3 or 4 procurrent caudal-fin rays (larvae can appear to have 2).
The next group, the sailfin-type blennies of Emblemaria, are generally larger and have higher fin-ray counts than the emblemariopsids. Most species of Emblemaria have greatly elongated dorsal fins in both males and females. The third group comprises Acanthemblemaria, typically spiny-headed blennies with blunt snouts and bushy cirri, which are also larger (although still tiny fishes) and have higher fin-ray counts, especially of anal-fin rays, and 4-6 procurrent caudal-fin rays (A. maria, secretary blenny, by Les Wilk, at right). The final group is made up of the remainder- an eclectic collection of several genera, mostly monotypic, with longer eel-like bodies and relatively high fin-ray counts.
Extensive DNA sequencing shows many widespread chaenopsid species to be made up of sets of closely-related species that can be hard to distinguish, even as adults (i.e. cryptic species). The chaenopsids, unlike most other reef fishes, have benthic eggs and short larval lives which promote reproductive isolation and genetic divergence within the region. As a result, there can be a proliferation of cryptic species and lineages and quite complex phylogeography. The larvae and juveniles of many cryptic species would be expected to be almost identical and are sometimes pooled into a type for the species complex in the descriptions below.
Chaenopsid blennies are tiny fishes and rarely noticed underwater, yet they are one of the most speciose families of marine fishes in the New World. Interestingly, they are also one of the few marine fish families to be limited to the Americas. They have a variety of fanciful common names including sailfin, flagfin, glass, banner, secretary, tube, pike, wrasse, and arrow blennies, among others- few of which are sufficiently specific. Almost all chaenopsid blennies are hole-dwelling, often with precise habitat requirements and sometimes narrow geographic ranges. They live in most habitats around Caribbean reefs, including coral structure, algae, sand, grassbeds, rocks, tidepools, and pilings. There are numerous genera and identifications can be difficult. A number of ill-defined species and species-complexes with many cryptic species and lineages are being discovered as DNA-sequencing of the group progresses. In general, chaenopsid larvae are not common and mostly undescribed. Larval chaenopsids can be recognized by their long thin body, large round eyes, a long and continuous dorsal fin with numerous slender spines and fewer soft rays (in all but 2 genera), a very short and narrow caudal peduncle, pelvic fins thoracic (in front of the pectoral fins) with only 2 or 3 long strand-like rays (not markedly curled-up over the body), no head spines, no silvery peritoneal lining, and light markings. The markings vary little and typically consist of a simple ventral-midline series: around the isthmus and a row along the base of the anal fin. These larval characters are generally shared with the larvae of the closely related scaled blennies of the family Labrisomidae, which well outnumber chaenopsids in most larval collections. Labrisomid larvae are very similar to larval chaenopsids and are distinguished mainly by having fewer dorsal-fin rays (although there is a small overlap): regional labrisomids have 12 or fewer dorsal-fin soft rays (with a rare 13) and fewer than 32 total dorsal-fin elements (with rare exceptions), while most chaenopsids have 13 or more dorsal-fin soft rays and more than 32 total dorsal-fin elements. In addition, labrisomid larvae (other than Starksia) are larger at the same stage of development, have additional melanophores, and more procurrent caudal-fin rays. The Caribbean chaenopsids with low fin-ray counts that can overlap with some labrisomids comprise the Emblemariopsis species, Coralliozetus cardonae, and rarely Emblemaria vitta; these larvae can be distinguished from labrisomids by size, markings, and lower procurrent caudal-fin ray counts. The basic taxonomic features separating the two families, i.e. scales on labrisomids and scales absent on chaenopsids (along with a set of osteological characters), are useless for larval stages. The distinctive genus Stathmonotus is still considered chaenopsid even though their dorsal fin is made up of all spines (and they can have scales). Some labrisomids of Paraclinus also have a dorsal fin made up of all spines; fortunately the larvae of the two genera are easily distinguished by their obvious morphological differences. Chaenopsid larvae broadly resemble those of other blennioid families of reef fishes. They can be distinguished easily from larvae of the true blennies (family Blenniidae), which have fewer dorsal-fin spines than soft rays, blunt snouts at all stages, and are heavily marked, while larval chaenopsid blennies have more dorsal-fin spines than soft rays (except in the snake-like Chaenopsis and Lucayablennius zingaro, which still have many more spines than true blennies), have pointed snouts (except Stathmonotus), and are very lightly marked. Larvae of the blennioid triplefins (family Tripterygiidae) have three separate dorsal fins and distinctive melanophores on the dorsal caudal peduncle. Stargazer larvae (family Dactyloscopidae) have relatively foreshortened anterior bodies, curled-up pelvic fins, and very few procurrent caudal-fin rays. Larval chaenopsids are superficially similar to the larvae of gobies and scarids, which can have a similar anal-fin row of melanophores and are the same size as chaenopsid larvae. However, those larvae notably have many fewer dorsal-fin spines, short and/or fused pelvic fins or just stubs, and narrowed or oddly-shaped eyes, while chaenopsids have long spinous dorsal fins, large round eyes, and, at least in later stages, long thread-like pelvic fins. Larval gerreids (mojarras) are also common and have an anal-fin row, however they have silvery abdominal linings and short dorsal and anal fins. Larval grunts (Haemulidae) often have an anal-fin row of melanophores, but they have spiny heads, short dorsal and anal fins, and characteristic tail spots.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:824
Specimens with Barcodes:821
Species With Barcodes:59
The blennioid family Chaenopsidae includes the pike-blennies, tube-blennies, and flagblennies, all perciform marine fish. The family is strictly tropical, ranging from North to South America. Members are also present in waters off Japan, Taiwan and Korea. Fourteen genera and 90 species are represented, the largest being the sarcastic fringehead, Neoclinus blanchardi, at 30 cm (12 in) in length; most are much smaller, and the group includes perhaps the smallest of all vertebrates, Acanthemblemaria paula, measuring just 1.3 cm (0.51 in) long as an adult.
With highly compressed bodies, some may be so elongated as to appear eel-like; chaenopsids are scaleless and lack lateral lines. Their heads are rough and may be armed with spines. There may be 17 to 28 spines in the dorsal fin, with two in the anal fin.
The habit of taking up home in abandoned worm tubes has earned some species in this family the name "tube-blenny". Many will also inhabit empty clam shells, which also serve as nesting sites; males are known to guard the brood. Some species have dorsal fins which are significantly higher towards the head, explaining the moniker "flagblenny". Crustaceans make up the bulk of the chaenopsid diet.
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