hypoplectrus nigricans , the black hamlet , is a hamlet in the family serranidae .

it is native to shallow parts of the central western atlantic ocean and caribbean sea .

it grows to about 15 cm ( 6 in ) in total length .

it is a simultaneous hermaphrodite , with a breeding strategy known as egg trading .

one fish acts as a female and lays a batch of eggs which the other fertilises .

the following night , the roles are reversed .

hypoplectrus nigricans

f st values for pair - wise comparisons of hypoplectrus nigricans allopatric populations , based on analysis of aflp data .
froese , rainer and pauly , daniel , eds . ( 2008 ) .
hypoplectrus nigricans
in fishbase . december 2008 version .
results of the f st outlier analyses between belize , honduras , and panama in hypoplectrus puella , h . nigricans , and h . unicolor
hypoplectrus nigricans is a hamlet from the western atlantic . it occasionally makes its way into the aquarium trade . it grows to a size of 15 cm in length .
hypoplectrus floridae n . sp . and hypoplectrus ecosur n . sp . , two new barred hamlets from the gulf of mexico ( pisces : serranidae ) : more than 3 % different in coi mtdna sequence from the caribbean hypoplectrus species flock
hypoplectrus floridae n . sp . and hypoplectrus ecosur n . sp . , two new barred hamlets from the gulf of mexico ( pisces : serranidae ) : more than 3 % different in coi mtdna sequence from the caribbean hypoplectrus species flock
aguilar - perera , a . 2004 . variations in morphology and coloration in the black hamlet , hypoplectrus nigricans ( teleostei : serranidae ) . caribbean journal of science , 40 : 150 - 154 .
numbers and types of polymorphic aflp loci resulting from pair - wise comparisons of hypoplectrus populations .
a review of the caribbean hamlets ( serranidae , hypoplectrus ) with description of two new species .
hypoplectrus nigricans isolate m63hn . b nadh dehydrogenase subunit 5 ( nd5 ) gene , partial cds ; nadh dehydrogenase subunit 6 ( nd6 ) gene , complete cds ; and trna - glu gene , partial sequence ; mitochondrial genes for mitochondrial products
hypoplectrus nigricans isolate m62hn . b nadh dehydrogenase subunit 5 ( nd5 ) gene , partial cds ; nadh dehydrogenase subunit 6 ( nd6 ) gene , complete cds ; and trna - glu gene , partial sequence ; mitochondrial genes for mitochondrial products
f st values for pair - wise comparisons of hypoplectrus chlorurus allopatric populations , based on analysis of aflp data .
f st values for pair - wise comparisons of hypoplectrus puella allopatric populations , based on analysis of aflp data .
f st values for pair - wise comparisons of hypoplectrus unicolor allopatric populations , based on analysis of aflp data .
victor , b . c . 2012 . hypoplectrus floridae n . sp . and hypoplectrus ecosur n . sp . , two new barred hamlets from the gulf of mexico ( pisces : serranidae ) : more than 3 % different in coi mtdna sequence from the caribbean hypoplectrus species flock . journal of the ocean science foundation , 5 : 1 - 19 .
citation :
black hamlets , hypoplectrus nigricans ~ marinebio . org .
marinebio conservation society . web . accessed monday , july 9 , 2018 . < urltoken > . last update : 1 / 14 / 2013 2 : 22 : 00 pm ~ contributor ( s ) : marinebio
the blue hamlet hypoplectrus gemma and its supposed model , the blue chromis chromis cyanea . photos : a & b - dr robertson .
this study is based on nine samples including three sympatric species ( the barred hamlet hypoplectrus puella , the black hamlet hypoplectrus nigricans , and the butter hamlet hypoplectrus unicolor ) from three locations ( belize , honduras , and panama ) , with 14 individuals per sample ( total 126 individuals ) . this sampling design provides the opportunity to explore the population genomic patterns of local adaptation ( between allopatric populations within species ) and speciation ( between sympatric species ) within a single system , and to repeat comparisons both taxonomically ( in three species for local adaptation ) and geographically ( in three populations for speciation ) .
domeier , m . l . 1994 . speciation in the serranid fish hypoplectrus . bulletin of marine science , 54 : 103 - 141 .
f st estimates among belize , honduras , and panama in hypoplectrus puella , h . nigricans , and h . unicolor at 10 microsatellite loci , 97 , 962 snps , and at the three repeated outliers identified in this study . n sample size , n / a data not available , – coverage below filtering criteria for these snps in these populations
fischer , e . 1980 . speciation in the hamlets ( hypoplectrus : serranidae ) : a continuing enigma . copeia , 1980 : 649 - 659 .
the yellowbelly hamlet hypoplectrus aberrans and its supposed model , the cocoa damselfish stegastes variabilis . photos : a - f charpin ; b - dr robertson .
the yellowtail hamlet hypoplectrus chlorurus and its supposed model , the yellowtail damselfish microspathodon chrysurus . photos : a - c shipley ; b - dr robertson .
the tan hamlet hypoplectrus randallorum and its supposed model , the threespot damselfish stegastes planifrons . photos : a - p lobel ; b - dr robertson .
( of plectropoma nigricans poey , 1852 ) froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
the shy hamlet hypoplectrus guttavarius and its supposed model , the rock beauty angelfish holacanthus tricolor . photos : a - f charpin ; b – j lyle .
f st values for pair - wise comparisons of hypoplectrus sympatric morphotype populations and for pair - wise same morphotype comparisons allopatric populations , based on analysis of aflp data .
lobel , p . s . 2011 . a review of the caribbean hamlets ( serranidae , hypoplectrus ) with description of two new species . zootaxa , 3096 : 1 - 17 .
distinct geographic variation in the coloration is present in h . unicolor , h . nigricans , and h . indigo , but not in the “model” of each . h . puella and h . nigricans display geographic variation in coloration that suggests that , if their coloration is cryptic , they are more cryptic at some locations than others . thus the limited information on geographic variation in hamlet coloration indicates it occurs in both “mimic” and non - mimic hamlets and is not related to mimicry , nor , perhaps , to crypsis .
domeier , m . l . , 1994 . speciation in the serranid fish hypoplectrus . bull . mar . sci . 54 ( 1 ) : 103 - 141 . ( ref . 26407 )
graves j . e . & rosenblatt r . h . 1980 . genetic relationships of the color morphs of the serranid fish hypoplectrus unicolor . evolution , 34 ( 2 ) , 240 - 245 .
fischer [ 45 ] made intensive observations on h . nigricans and noted no behavioral interactions between it and the damselfishes or other species indicative of mimicry . he suggested that the coloration resemblance between h . nigricans and the two stegastes species is coincidental , and the result of independent selection for background - matching crypsis in each taxon . aguilar - perera [ 48 ] described geographic variation in color , shape and size of h . nigricans , with fish in the northwest caribbean being uniformly black , with short , blunt fins , while those in puerto rico are grey with yellow eyes , and have longer , more pointed fins ( see also [ 33 ] , [ 40 ] ) . if the black pattern is cryptic [ 45 ] , then the geographic variation described by aguilar - perera [ 48 ] indicates that the species may be less cryptic at some locations than others .
research hypoplectrus nigricans » barcode of life ~ bioone ~ biodiversity heritage library ~ cites ~ cornell macaulay library [ audio / video ] ~ encyclopedia of life ( eol ) ~ esa online journals ~ fishbase ~ florida museum of natural history ichthyology department ~ gbif ~ google scholar ~ itis ~ iucn redlist ( threatened status ) ~ marine species identification portal ~ ncbi ( pubmed , genbank , etc . ) ~ ocean biogeographic information system ~ plos ~ siris ~ tree of life web project ~ unep - wcmc species database ~ worms
puebla o , bermingham e , guichard f ( 2008 ) population genetic analyses of hypoplectrus coral reef fishes provide evidence that local processes are operating during the early stages of marine adaptive radiations . molecular ecology 17 : 1405–1415 .
garcía - machado e . , chevalier - monteagudo , p . p . & solignac , m . 2004 . lack of mtdna differentiation among hamlets ( hypoplectrus , serranidae ) . marine biology , 144 : 147 - 152 .
only comparisons with suitable sample sizes are shown . see table s7 for loci and outlier numbers for all comparisons . abbreviations represent the morphotypes included in each comparison : c = h . chlorurus , n = h . nigricans , p = h . puella , u = h . unicolor , v = veracruz white .
observations of behavioral interactions relating to mimicry have been made on only two of the seven hamlets originally described as mimics by randall and randall [ 3 ] and thresher [ 42 ] , h . nigricans and h . unicolor . in a detailed study of the behavioral ecology of h . nigricans , fischer [ 45 ] found no support for mimicry by this species , and suggested there was a coincidental similarity related to background matching coloration . h . unicolor and h . indigo are the only hamlets actually known to behave in a manner consistent with a mimetic relationship . how do information on hamlets in general relate to the nine predictions of the mimicry hypothesis ?
citation : holt bg , côté im , emerson bc ( 2011 ) searching for speciation genes : molecular evidence for selection associated with colour morphotypes in the caribbean reef fish genus hypoplectrus . plos one 6 ( 6 ) : e20394 . urltoken
abstract nine of the sixteen species of the western atlantic genus hypoplectrus ( serranidae ) are currently recognized to be distributed in the gulf of mexico . hypoplectrus atlahua n . sp . is only known from tuxpan banks and isla lobos , veracruz . it differs from the only other similarly colored species , h . nigricans ( i . e black hamlet ) in the number of gill rakers on the first arch , snout length , upper jaw length , pectoral and pelvic fins lengths , and coloration . it is a dark brown species , lacking nose spots , mask , caudal fin dark spots , peduncle saddle , and pectoral pigmentation , but has iridescent violet speckles and lines on cheeks and chest , and also a well - defined but rather small violet dot over the dorsal flat spine on the rear edge of the opercle .
representative images of the hypoplectrus colour morphotypes at different locations included in aflp outlier detection analysis . n . b . images for samples obtained in u . s . virgin islands not available as all individuals were released immediately after capture within this location .
aguilar - perera , a . & gonzález - salas , c . 2010 . distribution of the genus hypoplectrus v ( teleostei : serranidae ) in the greater caribbean region : support for a color - based speciation . marine ecology , 31 : 375 - 387 .
aguilar - perera , a . & tuz - sulub , a . n . 2010 . hypoplectrus gemma ( teleostei : serranidae ) is not endemic to southern florida waters . pan - american journal of aquatic sciences , 5 ( 1 ) : 143 - 146 .
holt , b . g . , côté , i . m . & emerson , b . c . 2010 . signatures of speciation ? distribution and diversity of hypoplectrus ( teleostei : serranidae ) colour morphotypes . global ecology and biogeography , 19 : 432 - 441 .
puebla , o . , bermingham , e . , guichard , f . & whiteman , e . 2007 . colour pattern as a single trait driving speciation in hypoplectrus coral reef fishes ? proceedings . biological sciences / the royal society , 274 : 1265 - 1271 .
ramon , m . l . , lobel , p . s . & sorenson , m . d . 2003 . lack of mitochondrial genetic structure in hamlets ( hypoplectrus spp . ) : recent speciation or ongoing hybridization ? . molecular ecology 12 : 2975 - 2980 .
the significant repeated outliers were detected above consistently low , but significant , genetic divergence between pairs of morphotypes ( table 2 ) . our f st values , derived from the most comprehensive taxonomic and geographic sampling to date , are in agreement with previous analyses using aflps [ 15 ] and microsatellites [ 11 ] , [ 12 ] . these results suggest that hypoplectrus morphotypes do represent more than simple colour variants of a single species and that selection acting on a limited number of loci is indirectly resulting in low level significant isolation in other parts of the hypoplectrus genome .
holt , b . g . , côté , i . m . & emerson , b . c . 2011 . searching for speciation genes : molecular evidence for selection associated with colour morphotypes in the caribbean reef fish genus hypoplectrus . plos one , 6 ( 6 ) : e20394 .
del moral flores , l . , tello - musi , j . & martíinezpérez , j . 2011 . descripción de una nueva especie del género hypoplectrus ( actinopterigi : serranidae ) del sistema arrecifal veracruzano , suroeste del golfo de méxico . revista de zoología , 22 : 1 - 10 .
our results show evidence for consistent selection between at least two hypoplectrus morphotypes ( h . chlorurus and h . puella ) and highlight three aflp loci that show good evidence for being directly affected . selection between colour forms appears to play a role in maintaining this complex of morphotypes ; however , this has not resulted in consistent divergence for the vast majority of the loci studied . the low but significant genetic differentiation between morphotypes suggests that hypoplectrus colour forms do represent more than just colour variants of a single species . however , incipient speciation cannot be assumed and these results are consistent with the possibility of an evolutionarily stable colour polymorphism .
the repeated outlier loci identified are likely to be linked , either physically or otherwise , to genes coding for hypoplectrus colour pattern . an aflp outlier analysis , similar to ours , on intertidal snails [ 25 ] was followed up by sequencing of both outlier and non - outlier loci using bacterial artificial chromosome libraries [ 26 ] , allowing speculation regarding the phenotypic effects of outlier loci . applying these techniques to the repeated outliers uncovered in this study could result in a fundamental step forward in understanding exactly how hypoplectrus morphotypes differ from each other , potentially identifying regions of the genome under selection . comparative analysis of divergent and non - divergent loci may consequently prove to be informative regarding the history of polymorphism in this genus .
hypoplectrus chlorurus , h . nigricans , h . puella and h . unicolor each provided suitable sample sizes for repeated outlier analysis . a total of 10 loci were identified using the dfdist method as being potentially under selection between morphotypes at either the 99 % or the 95 % level ( fig . 3 , table 4 ) . the frequency of repeated outliers showed the opposite pattern to that shown by general outlier frequencies , with sympatric morphotype population comparisons producing more repeated outlier loci than allopatric same morphotype comparisons . the numbers of 95sel and 99sel loci were significantly higher than expected under neutral conditions ( both p < 0 . 001 ) . the numbers of 95geo and 99geo loci did not significantly differ from null expectations , however , the relatively small sample sizes for h . puella in these analyses is likely to have reduced the probability of detecting loci in disequilibrium in these comparisons involving this species .
in this study we use established outlier detection methods to search for molecular signatures of selection on specific loci between interbreeding populations . through analysis of amplified fragment length polymorphisms ( aflps ) we specifically considered whether individual loci are consistently associated with individual morphotypes across different locations . the aflp technique has a specific advantage over mtdna and microsatellite markers that have been so far employed for population genetic analysis of hypoplectrus [ 11 ] , [ 12 ] ; by efficiently producing hundreds of markers , it has higher potential to detect polymorphism across the genome . our study took advantage of this feature to search for direct molecular evidence of selection between hypoplectrus morphotypes across different locations . this approach has an advantage over methods such as bulk segregate analysis or qtl mapping in that it can be applied to samples from natural populations and does not require laboratory breeding , which has so far proven to be unfeasible for these fish [ 7 ] .
using a genome - wide scanning approach , we found evidence for consistent selection occurring between pairs of h . chlorurus and h . puella , and between h . puella and h . unicolor pairs , despite observing low to moderate overall divergence among colour forms . three aflp loci were identified as being under selection between morphotypes in pair - wise comparisons of sympatric morphotypes at two different locations , using both the dfdist and bayescan methods . these loci represent the first evidence for selection between morphotypes influencing the hypoplectrus genome across sampling locations .
substantial inter - morphotype gene flow may at first seem to contradict the results of extensive spawning observations , in which only ∼1 % of matings occurred between different morphotypes [ 7 ] , [ 13 ] . however , even this level of interbreeding could be consistent with our observed f st values given the likely size of the majority of hypoplectrus morphotype global populations . the possibility of infrequent hybridization preventing speciation has been suggested for other systems [ 28 ] , [ 29 ] , [ 30 ] , whereby equilibrium between convergence and divergence is maintained by occasional gene flow .
hypoplectrus is a genus of small , predatory groupers endemic to the tropical northwest atlantic . it has 16 named ‘species’ [ 3 ] , [ 39 ] - [ 42 ] , eight of which have been proposed as aggressive mimics of different reef fishes [ 3 ] , [ 42 ] , [ 43 ] . below i present information on the only four species for which there are behavioral observations relating to the mimicry hypothesis . relevant information on the coloration and behavior of five other species of “mimic” hamlets and their “models” is summarized in appendix s1 , together with general information on the coloration of hamlets .
two “mimetic” hamlets , h . nigricans and h . indigo , feed heavily on fish : ∼30–40 % and 90 % of their identifiable stomach contents respectively [ 37 ] , [ 43 ] , [ 45 ] . in addition , fishes also make up about 10–25 % of the identifiable stomach contents of h . chlorurus , h . puella and h . unicolor [ 37 ] , [ 43 ] . the resemblance of the “mimetic” hamlet that specializes in preying on fish , h indigo , to its potential model , the fish it eats , is sufficiently vague that it was not among the original group of hamlets labeled as mimics due to their similarity to other fishes . caribbean reef fishes , including potential targets of some “mimic” hamlets , can readily distinguish those hamlets from their models [ 42 ] , and recognize the different ecological and threat status of each .
the relatively high level of genetic isolation demonstrated for the two sympatric morphotypes sampled in mexico is surprising , as there was no a priori reason to suspect this comparison to differ from the others . this comparison produced an f st of 0 . 212 , far larger than the level of isolation measured in all of other pair - wise comparisons ( all other f st < 0 . 09 ) . other unusual features of this region include the fact that just two morphotypes are abundant ( bgh , personal observations ) and one is an endemic morphotype , the veracruz white . the only other study to consider hamlets from the gulf of mexico [ 10 ] found that veracruz whites tend to feed at a higher trophic level than the sympatric h . nigricans population . this species pair may be an interesting focus for future research to determine what has driven the divergence between the two morphotypes in veracruz .
the overall mean f st values among morphotypes across the region and among sampling locations were similar : 0 . 051 and 0 . 052 , respectively ( for pair - wise comparisons , see table 2 ) . these values will be influenced by regional variation in the numbers of morphotypes sampled . the mean f st between sympatric morphotypes was 0 . 058 , with all pair - wise comparisons being significantly greater than zero , with the exception of the comparison of two morphotypes from honduras . within morphotypes , geographical population structure had a mean f st value of 0 . 060 ; however , there was large variation between morphotypes , ranging from 0 . 020 among h . chlorurus populations to 0 . 114 among h . nigricans populations . f st values for all sympatric inter - morphtype comparisons and all allopatric intra - morphotype comparisons are presented in tables s2 , s3 , s4 , s5 and s6 .
according to thresher [ 42 ] h . nigricans and s . adustus show parallel geographic variation in coloration , with the jamaican population of both having yellow bellies , and pelvic , anal and tail fins . however , at montego bay , jamaica , ∼50 km from thresher’s study site , s . adustus have the same uniform grey - brown color they have elsewhere in the caribbean area ( drr pers obs , at montego bay , florida , bermuda , the bahamas , panama , curacao , venezuela , barbados , and puerto rico ) . further , the jamaican coloration described for the hamlet and damselfish by thresher [ 42 ] fits other hamlets ( h . aberrans or h . chlorurus ) and stegastes variabilis . when color differences are the defining characteristic of most hamlet “species” ( see appendix s1 ) , whether to call hamlets with different color patterns intraspecific geographic variants rather than different species becomes a semantic issue .
samples were collected using scuba , from coral reef sites at eight sampling locations ( i . e . countries ) distributed across the caribbean basin ( fig . 1 ) . we refer to all individuals of a particular colour form at a particular location as a ‘morphotype population’ and all individuals of a particular colour form from throughout the hypoplectrus distribution as a ‘global morphotype population’ . fish were sampled by using micro - spears or “hook and line” whereby baited , barbless hooks were offered to fish . hooked fish were released after fin clipping . fin clips were removed from the dorsal and anal fins and placed in ethanol and stored at 4°c . the morphotypes sampled and number of individuals collected varied among locations , depending on local abundance ( table 1 ) .
the webpage text is licensed under a creative commons attribution - noncommercial 4 . 0 license
froese , r . & d . pauly ( editors ) . ( 2018 ) . fishbase . world wide web electronic publication . , available online at urltoken [ details ]
mceachran , j . d . ( 2009 ) . fishes ( vertebrata : pisces ) of the gulf of mexico , pp . 1223–1316 in : felder , d . l . and d . k . camp ( eds . ) , gulf of mexico–origins , waters , and biota . biodiversity . texas a & m ; press , college station , texas . [ details ]
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there is an ongoing discussion on the validity of the hamlets as biological species ( domeier 1994 ) . hamlet
species
are defined primarily on differences in color patterns . they are known to interbreed and hybridize freely with very little consistent genetic differentiation amongst them .
anderson , w . , carpenter , k . e . , gilmore , g . , milagrosa bustamante , g . & robertson , r .
this widely distributed species is common and abundant where it occurs over shallow reefs .
there are no known major threats . therefore , it is listed as least concern .
is distributed in the western atlantic from southeast florida , the bahamas , in the gulf of mexico from the florida keys and from tuxpan , mexico along the northern yucatan to northwestern cuba , in the caribbean throughout the antilles to tobago , and along central and south america from mexico to santa marta , colombia and in the offshore islands off venezuela ( r . robertson pers . comm . 2014 ) . the depth range is three to 67 m ( lieske and myers 1994 ) , but is most abundant in less than 10 m depth .
anguilla ; antigua and barbuda ; aruba ; bahamas ; barbados ; belize ; bonaire , sint eustatius and saba ; cayman islands ; colombia ; costa rica ; cuba ; curaçao ; dominica ; dominican republic ; grenada ; guadeloupe ; guatemala ; haiti ; honduras ; jamaica ; martinique ; mexico ; montserrat ; nicaragua ; panama ; puerto rico ; saint barthélemy ; saint kitts and nevis ; saint lucia ; saint martin ( french part ) ; saint vincent and the grenadines ; sint maarten ( dutch part ) ; trinidad and tobago ; turks and caicos islands ; united states ; venezuela , bolivarian republic of ; virgin islands , british ; virgin islands , u . s .
this species is common throughout its range . in a study conducted off san blas , panama , it was the most abundant hamlet on shallow reef flats ( fischer 1980 ) .
this species inhabits shallow reefs over hard or soft corals ( lieske and myers 1994 , smith 1997 ) . members of this genus are known to be synchronous hermaphrodites with individuals alternating male and female
roles
during spawning ( fischer 1981 ) . its maximum length 15 . 2 cm tl ( randall 1996 ) .
this species rarely occurs in the aquarium trade and is not preferred ( r . robertson pers . comm . 2012 ) .
there are no known major threats . it is potentially a prey item of the invasive lionfish , however , only juveniles are consumed and it is not likely that this will drive significant population declines on a global level ( l . rocha pers . comm . 2014 ) .
anderson , w . , carpenter , k . e . , gilmore , g . , milagrosa bustamante , g . & robertson , r . 2015 .
to make use of this information , please check the < terms of use > .
greek , hypo = under + greek , plektron = sting , spur ( ref . 45335 )
marine ; reef - associated ; depth range 3 - 13 m ( ref . 9710 ) . tropical
western atlantic : belize to panama ; throughout the rest of the caribbean . absent off the coast of venezuela . antilles ( ref . 26938 ) .
maturity : l m ? range ? - ? cm max length : 15 . 2 cm tl male / unsexed ; ( ref . 13442 )
body blackish ( the intensity of the black can vary from bluish to brownish ) . all fins pigmented , including the pectorals .
a solitary species ( ref . 26340 ) inhabiting shallow reefs ( ref . 9710 ) . feeds on fishes and crustaceans ( ref . 26180 ) . often found near the bottom around soft and hard corals . occasionally been seen spawning with the h . chlorurus ( ref . 26938 )
) : 27 . 9 - 28 . 5 , mean 28 ( based on 130 cells ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5000 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 01778 ( 0 . 00692 - 0 . 04567 ) , b = 3 . 03 ( 2 . 81 - 3 . 25 ) , in cm total length , based on lwr estimates for this ( sub ) family - body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 4 . 0 ±0 . 66 se ; based on food items .
resilience ( ref . 69278 ) : high , minimum population doubling time less than 15 months ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : low vulnerability ( 22 of 100 ) .
confused by a class within a class or an order within an order ? please see our brief essay .
to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
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claro , rodolfo , and lynne r . parenti / claro , rodolfo , kenyon c . lindeman , and l . r . parenti , eds .
nelson , joseph s . , edwin j . crossman , h . espinosa - pérez , l . t . findley , c . r . gilbert , et al . , eds .
full author list : nelson , joseph s . , edwin j . crossman , héctor espinosa - pérez , lloyd t . findley , carter r . gilbert , robert n . lea , and james d . williams
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
this is a clip of a video that i took following black hamlets around during the time of the day that they mate .
a solitary species ( ref . 26340 ) inhabiting shallow reefs ( ref . 9710 ) . feeds on fishes and crustaceans ( ref . 26180 ) . often found near the bottom around soft and hard corals . occasionally been seen spawning with the h . chlorurus ( ref . 26938 )
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occurrence describes how often the species is found on surveys within its distribution . it is calculated as the % of reef sites surveyed by rls divers across all the ecoregions in which the species has been observed
abundance is calculated as the average number of individuals recorded per rls transect , where present .
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head and body deep , stro ngly compressed ; forehead straight ; snout relatively short ; top jaw protrusible ; rear of top jaw exposed when mouth closed , without accessory bone above it ; teeth fixed ; preoperculum angular , serrated , several small forward pointing spines on lower edge near the corner ; gill rakers 17 - 23 ; dorsal fin x , 14 - 17 , no notch after spines , membranes between spines not indented ; pelvics long , reach to or beyond anus ; tail fin slightly forked ; lateral line scales 48 - 53 ; soft dorsal and anal fins mostly scaleless .
head and body black , grey or dark brown ; all fins except the pectorals are colored as the body ; pectorals are black in black bodied fish , clear in those with grey bodies ; there may be a bluish cast on various parts of the body and fins .
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warning : the ncbi web site requires javascript to function . more . . .
format summary genbank genbank ( full ) fasta asn . 1 xml insdseq xml tinyseq xml feature table accession list gi list gff3
finds sub - sequence or patterns in the sequence and highlights the matching region . the tool works with standard single letter nucleotide or protein codes including ambiguities and can match prosite patterns in protein sequences . more . . .
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western atlantic : belize to panama ; throughout the rest of the caribbean . absent off the coast of venezuela . antilles ( ref . 26938 ) .
15 . 2 cm tl ( male / unsexed ; ( ref . 13442 ) )
depth range based on 4 specimens in 1 taxon . environmental ranges depth range ( m ) : 5 - 10 graphical representation depth range ( m ) : 5 - 10 note : this information has not been validated . check this * note * . your feedback is most welcome .
reef - associated ; marine ; depth range 3 - 13 m ( ref . 9710 )
depth : 3 - 13m . from 3 to 13 meters . habitat : reef - associated .
inhabits shallow reefs ( ref . 9710 ) . often found near the bottom around soft and hard corals . feeds on fishes and crustaceans . carnivore ( ref . 57616 ) .
the following is a representative barcode sequence , the centroid of all available sequences for this species . there are 4 barcode sequences available from bold and genbank .
below is a sequence of the barcode region cytochrome oxidase subunit 1 ( coi or cox1 ) from a member of the species .
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marine ; occupy small territories at the foot of , or near hard or soft coral reefs .
body and fins are a mat dark blue brown to black . pelvic fins are long . length 3 - 4 . 5in . , max . 6in .
claro , r . , 1994 características generales de la ictiofauna . p . 55 - 70 . in r . claro ( ed . ) ecología de los peces marinos de cuba . instituto de oceanología academia de ciencias de cuba and centro de investigaciones de quintana roo . humann , p . and deloach , n . ( 2002 ) reef fish identification . florida , caribbean , bahamas . new world publications , inc . jacksonville , florida , usa . lieske , e . and r . myers , 1994 collins pocket guide . coral reef fishes . indo - pacific & caribbean including the red sea . haper collins publishers , 400 p . ogden , j . c . , j . a . yntema , and i . clavijo , 1975 an annotated list of the fishes of st . croix , u . s . virgin islands . spec . publ . no . 3 . smith , c . l . , 1997 national audubon society field guide to tropical marine fishes of the caribbean , the gulf of mexico , florida , the bahamas , and bermuda . alfred a . knopf , inc . , new york . 720 p .
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copyright : © 2011 holt et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
funding : this work was funded by the national environment research council ( studentship s ner / sj2004 / 13064 and lsmsf application no . ek76 - 02 / 05 ) , with the additional funding by smgf and the sydney l . wright fellowship ( contribution number 1700 ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
whether speciation can occur in the absence of geographic barriers remains controversial [ 1 ] . in this context colour polymorphisms , which occur in a wide range of taxa , provide excellent opportunities for studies of intraspecific evolutionary divergence ( e . g . [ 2 ] ) . many different mechanisms have been implicated in the origin and maintenance of colour polymorphism , including sexual selection , mimicry , predation , crypsis and genetic drift [ 3 ] . as all these mechanisms may also contribute to reproductive isolation , colour polymorphisms are often considered as systems potentially undergoing speciation ( e . g . [ 4 ] ) .
1 = bermuda , 2 = curaçao , 3 = dominican republic , 4 = honduras , 5 = mexico , 6 = panama , 7 = puerto rico , 8 = u . s . virgin islands .
the aflp procedure was based on vos et al . , [ 16 ] . dna was isolated from ≈0 . 5 cm 2 sub - samples of fin clips , using a phenol - chloroform extraction method [ 17 ] . extractions were checked for concentration and purity to ensure isolated dna was suitable for the production of clear repeatable aflp profiles . for each sample , 100 ng of dna ( 10 µl solution ) was added to 1 . 4 µl restriction digestion mixture containing 1 u ecori and 1 u msei restriction enzymes , along with 1 . 1 µl 10×ta reaction buffer ( 100 mm tris - ac ph 7 . 9 , 100 mm mgac , 500 mm kac , 10 mm dithiothreitol ( dtt ) ) and 3 µg bsa . dna was digested for three hours at 37°c . digested dna was added to 5 . 5 µl of ligation mastermix containing 1 µl 5× t4 ligase buffer , 0 . 5 u t4 ligase , 25 pmol of ecori adaptor and 25 pmol of mse adaptor ( table s1 ) . this solution was incubated overnight at 16°c and gel electrophoresis used to ensure that the dna was fully digested . 2 µl of diluted digestion mixture was used for the pre - selective pcr , which was run in 10 µl total solution containing 5 pmol of primer eco p and 5 pmol of mse p ( table s1 ) , along with 1 µl 10× pcr buffer , 15 nmol mgcl 2 , 2 nmol dntp and 0 . 25 u taq dna polymerase . the pre - selective pcr amplification comprised an initial denaturation step of two minutes at 94°c , followed by 20 cycles of denaturation at 94°c for 20 seconds , annealing at 56°c for 30 seconds and final extension at 72°c for two minutes .
pre - selective amplifications were checked using gel electrophoresis and then diluted to 1∶20 concentration by the addition of sterile distilled water . 1 µl of pre - selective pcr amplification was used as the template for the selective pcr , which was added to 9 µl pcr mixture containing 5 pmol of each selective primer ( table s1 ) , 1 µl 10× pcr buffer , 20 nmol mgcl 2 , 2 nmol dntp and 0 . 25 u taq dna polymerase . eco p selective primers were labelled with either fam or ned fluorescent labels for genotyping . pcr products were diluted 10× with sterile distilled water , and 5 µl of diluted product of fam - labelled samples were mixed with an equal quantity of ned - labelled samples for multiplex genotyping . 0 . 5 µl of the combined diluted product was then added to 9 . 45 µl hidi formamide and 0 . 05 µl abi rox size standard . prior to genotyping all samples were denatured at 95°c for three minutes and then quenched on ice for a further three minutes . control samples were added to every 96 - well pcr reaction plate to ensure repeatability between runs .
samples were genotyped using an abi 3730 capillary dna analyser . the fluorescent spectra of ned and fam labels do not overlap ; however , the possibility that the two combined samples interfered with each other ' s aflp profiles was investigated by genotyping several individuals , both singly and also after combining with an alternatively labelled sample . combined dye samples proved to be equally as reliable as single dye samples . primer combinations were chosen after screening 32 different primer combinations for profile quality ( i . e . whether the presence or absence of individual peaks could be clearly determined ) and polymorphism ( i . e . there appeared to be high variation in peak presence / absence between individual profiles ) . primer pairs showing the highest levels of polymorphism were selected for further analysis .
the frequencies of 95 % and 99 % outliers detected using dfdist were checked for the assumptions of parametric analysis and then compared with relevant neutral expectations . neutral expectations were assumed to be 5 % and 1 % of the total number of polymorphic loci for each comparison , for the 95 % and 99 % levels respectively . the pair - wise comparisons included in this analysis were all nine sympatric morphotype comparisons ( excluding morphotype populations with sample sizes of six or fewer individuals ) as well as nine randomly selected allopatric same - morphotype comparisons and nine randomly selected allopatric different - morphotype comparisons . the mean % of outlier loci for each type of morphotype population comparison was compared against neutral expectations .
the second analytical method we applied to our data was a fully bayesian approach developed by foll & gaggiotti [ 23 ] , which uses the software bayescan ( available at urltoken ) . this method extends the approach developed by beaumont & balding [ 24 ] , in order to allow the use of dominant markers ( such as aflps ) and to rigorously estimate the probability that each specific locus is subject to selection , rather than solely returning the probability of a locus being selectively neutral . to provide an independent test of the repeated outliers detected using dfdist , the same pair - wise comparisons were analysed using bayescan and loci showing probability values higher than 0 . 7 for the likelihood of being under directional selection were noted ( as per [ 23 ] ) . repeated dfdist outliers that the bayescan did not find to be under selection in either comparison are considered to be false positives , those which are found to be under selection in only one comparison may be false positives or , alternatively , may represent local selection that is not consistent across morphotype populations . outliers that are shown in both pair - wise comparisons using both methods are likely to be under selection between morphotypes .
a total of 528 hamlets were sampled from eight different locations representing five different colour morphotypes , with considerable variation among the numbers of each morphotype found at each location ( table 1 ) . representative images of the morphotypes sampled at each location are presented in figure s1 . a total of 436 scorable aflp loci were produced with an overall repeatability rate of 97 . 1 % ( see table s1 for primer combinations ) . of these , 423 loci were polymorphic , i . e . , they were scored differently in at least one individual .
global morphotypes ( i . e . all individuals of a given morph collected across all locations ) .
morphotype populations ( i . e . individuals of a given morph at a given sampling location ) .
left hand plots represent pairwise comparisons for populations of h . chlorurus and h . puella in curacao and panama . right hand plots represent pairwise comparisons for populations of h . puella and h . unicolor in curacao and panama . “sym” refers to sympatric comparisons of the two morphotypes concerned ( followed by the location ) , “allo” refers to allopatric comparisons of the two locations concerned ( followed by the morphotype ) . letters refer to loci repeated across sympatric comparisons ( see table 4 for full loci names and further details ) . dashed lines represent the simulated 95 % f st level , dotted lines represent the simulated 99 % f st level . repeated outlier comparisons that did not return any outliers repeated across sympatric comparisons are not shown .
values that were significantly higher than expected in the absence of selection between colour morphotypes , in pair - wise comparisons at two separate locations , using both dfdist and bayescan outlier detection methods .
the bayescan analysis showed mixed support for the repeated outliers identified using dfdist . three of the outliers repeated across different comparisons of h . chlorurus and h . puella were also identified as being repeatedly under selection using bayescan . the remaining seven dfdist outliers were either only identified in one comparison ( four outliers ) or not identified in any comparison ( three outliers ) as being under selection between morphotypes by this analysis ( table 4 ) .
for the majority of loci , differentiation between sympatric morphotypes appears to be no more consistent than differentiation shown between allopatric populations of the same morphotype ( table 3 ) . overall , pair - wise comparisons of sympatric morphotype populations produced more outliers than expected under neutral conditions but this difference was not significant . by comparison , analysis of allopatric populations composed of the same two morphotypes did reveal a significantly higher number of 99 % outliers than expected under neutral conditions , although the number of 95 % outliers did not significantly exceed null expectations .
the hamlet system has attracted considerable attention as a potential case study for speciation in the marine environment . however , the identification of loci specifically associated with certain morphotypes does not necessarily imply that divergence between morphotypes is currently ongoing . an alternative explanation for these results is that assortative mating , linked to the simultaneous hermaphroditism in these fish [ 8 ] , is maintaining divergence between morphotypes but inter - morphotype reproductive isolation is not fully complete and some successful interbreeding between morphotypes is limiting further divergence . gene flow between morphotypes would also explain the lack of association between mitochondrial sequence haplotypes and morphotypes shown in other studies [ 11 ] , [ 14 ] , [ 27 ] .
we would like to thank terry burke , andy krupa , raj whitlock and all at the sheffield molecular genetics facility ( smgf ) , who provided first rate advice and facilities for a large part of the aflp work . we would also like to thank oscar puebla and dr e bermingham for their help in the field and for agreeing to review an early draft of this manuscript . we thank dr e whiteman and the staff at magueyes laboratories ( puerto rico ) , carmabi ( curaçao ) , the national aquarium ( dominican republic ) , centre of ecology & fisheries ( mexico ) and b . b . s . r . ( bermuda ) for their support in the field . special thanks to yolanda leon , thad murdoch , horacio perez , joanna pitt , and monica vega for their invaluable assistance .
conceived and designed the experiments : bgh imc bce . performed the experiments : bgh . analyzed the data : bgh bce . wrote the paper : bgh imc bce .
alexander hj , breden f ( 2004 ) sexual isolation and extreme morphological divergence in the cumaná guppy : a possible case of incipient speciation . journal of evolutionary biology 17 : 1238–1254 .
gray sm , mckinnon js ( 2007 ) linking color polymorphism maintenance and speciation . trends in ecology & evolution 22 : 71–79 .
pierotti mer , seehausen o ( 2007 ) male mating preferences pre - date the origin of a female trait polymorphism in an incipient species complex of lake victoria cichlids . journal of evolutionary biology 20 : 240–248 .
bierne n , bonhomme f , david p ( 2003 ) habitat preference and the marine - speciation paradox . proceedings of the royal society b - biological sciences 270 : 1399–1406 .
palumbi sr ( 1992 ) marine speciation on a small planet . trends in ecology & evolution 7 : 114–118 .
( teleostei : serranidae ) colour morphotypes . global ecology and biogeography 19 : 432–441 .
species complex reveals no evidence for dietary niche divergence . marine ecology - progress series 357 : 283–289 .
mccartney ma , acevedo j , heredia c , rico c , quenoville b , et al . ( 2003 ) genetic mosaic in a marine species flock . molecular ecology 12 : 2963–2973 .
coral reef fishes ? proceedings of the royal society b - biological sciences 274 : 1265–1271 .
barreto fs , mccartney ma ( 2008 ) extraordinary aflp fingerprint similarity despite strong assortative mating between reef fish color morphospecies . evolution 62 : 226–233 .
vos p , hogers r , bleeker m , reijans m , vandelee t , et al . ( 1995 ) aflp - a new technique for dna fingerprinting . nucleic acids research 23 : 4407–4414 .
sambrook j , fritsch ef , maniatis t ( 1989 ) molecular cloning : a laboratory manual . new york : cold spring harbor press .
whitlock r , hipperson h , mannarelli m , butlin rk , burke t ( 2008 ) an objective , rapid and reproducible method for scoring aflp peak - height data that minimizes genotyping error . molecular ecology resources 8 : 725–735 .
vekemans x ( 2002 ) aflp - surv version 1 . 0 . distributed by the author . laboratoire de génétique et ecologie , végétale , université libre de bruxelles , belgium .
beaumont ma , nichols ra ( 1996 ) evaluating loci for use in the genetic analysis of population structure . proceedings of the royal society of london series b : biological sciences 263 : 1619–1626 .
zhivotovsky la ( 1999 ) estimating population structure in diploids with multilocus dominant dna markers . molecular ecology 8 : 907–913 .
nosil p , egan sp , funk dj ( 2008 ) heterogeneous genomic differentiation between walking - stick ecotypes : “isolation by adaptation” and multiple roles for divergent selection . evolution 62 : 316–336 .
foll m , gaggiotti o ( 2008 ) a genome - scan method to identify selected loci appropriate for both dominant and codominant markers : a bayesian perspective . genetics 180 : 977–993 .