scopula virgulata , the streaked wave , is a moth of the family geometridae .

it is found from most of europe to central asia and northern mongolia .

the wingspan is 20 – 22 millimetres ( 0.79 – 0.87 in ) .

adults are on wing from late july to august in one generation per year .

the larvae feed on carex and inula species .

larvae can be found from august to june .

it overwinters in the larval stage .

scopula virgulata

scopula ( scopula ) virgulata ( denis & schiffermuller , 1775 ) = geometra virgulata denis & schiffermüller , 1775 = geometra strigaria hübner , [ 1799 ] .
recently recorded for southern spain : j . gastón , f . morente & v . redondo : scopula donovani ( distant , 1892 ) , un nuevo geométrido para europa continental , descubierto en andalucía ( espana ) ( lepidoptera : geometridae : sterrhinae , scopulini ) . boletín de la sociedad entomológica aragonesa ( s . e . a . ) , nº 53 ( 31 / 12 / 2013 ) : 289–291 .
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there is some doubt as to the authenticity of this species as a british moth . there is a record of two individuals from kent in the late 19th century that some believe to be erroneous .
the species is distributed from france into eastern europe and beyond , where it occurs sparingly in open habitats .
there is a single generation , from june to august , and the larvae feed on a range of grasses and low plants .
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2011 - 01 - 26 by & van nieukerken , dr erik j . karsholt , dr ole & by
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austria , belgium , france , germany , spain , italy , latvia , lithuania , poland , portugal , romania , slovakia and the soviet union - the european part , finland , france , czech republic , switzerland , sweden , estonia , yugoslavia .
regions of the russian federation : the volga - don , east caucasus , gorno - altaisk , the european north - west , the european central black earth , the european central european south taiga , transbaikalia , western caucasus , kaliningrad , karelia , krasnoyarsk , nizhny - amur , prealtay , of baikal , pribaikalskiy , seaside , sakhalin , amur medium , medium - volzhsky , mid - ural , sredneobskaya , tuva , south west siberian , south ural , south yakutia .
austria , belarus , belgium , france , germany , spain ( mainland ) , italy ( mainland ) , latvia , lithuania , poland , portugal ( mainland ) , russia , romania , slovakia , slovenia , ukraine , finland , france ( mainland ) , croatia , czech republic , switzerland , sweden , estonia .
[ 10 ] de jong , y . s . d . m . ( ed . ) ( 2011 ) fauna europaea version 2 . 4 ( faunaeur . org )
[ 28 ] moths and butterflies of europe and north africa ( leps . it ) , 2012
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curators : konstantin efetov , vasiliy feoktistov , svyatoslav knyazev , evgeny komarov , stan korb , alexander zhakov .
species catalog enables to sort by characteristics such as expansion , flight time , etc . .
a taxon identifier is composed of name , author , year and attribute , all separated by a blank . these are all extracted from the original publication .
the name is reproduced exactly as proposed in the original publication . the name of a genus is made up of one word and species made up of two words ( genus and species ) separated by a blank .
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created by dicky sick ki yu 1997 - 2012 please send me information about errors and omissions ( contact information ) with supporting references , possibly with pdf or hard copy .
- azores , balearic is . , british i . , bulgaria , canary is , channel is . , corsica , crete , croatia , cyprus ,
the final instar larva is light brown with wide darker dorsal line and thinner subdorsal lines . spriracles show up as small black dots .
this moth is an appendix a moth and as such is an adventive species added to british list based on the record of two alleged individuals caught in gravesend kent back in 1870 . as such occurrence in the british isles is unconfirmed .
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- austria - near vienna - s . v . rossica djakonov , 1926 - population of st petersburg area - greyer , with quite strong contrast s . v . virgata
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joint project of the zoologische staatssammlung münchen ( dr . axel hausmann ) and apollo books , stenstrup ( peder skou )
this website aims to present an actual update to the book series
the geometrid moths of europe
( a hausmann ed . ) . so far four volumes have been published , two are in preparation :
gme5 ( ennominae i ) , skou & sihvonen ( in prep . 2014 ) : 144 species
gme6 ( ennominae ii ) , müller & erlacher ( in prep . 2015 ) : 172 species
( highres photo , labels : amur ; photo : i . kostjuk , university kiev )
recorded in the polar urals ( # pers . comm . ) , new for the fauna of europe . locus typicus in eastern siberia .
recorded in central spain ( exposito 2006 ) , new for the fauna of europe . locus typicus in eastern algeria .
was described as separate species from central spain ( exposito 2006 ) , quoting slight differences in male genitalia ( saccus ) . molecular data ( coi barcoding ) show these minor differences to fall under the variation of
recorded in astrachan desert , southern european russia ( anikin pers . comm . ) , new for the fauna of europe . locus typicus in southern caucasus .
recorded on samos island ( fritsch & skou pers . comm . ) , new for the fauna of europe . locus typicus in turkey .
( 2013 ; alexanor , 25 ( 6 ) , 2012 ( 2013 ) : 361 - 384 ) . recorded in northern portugal ( locus typicus ) and possibly in adjacent regions of spain .
from southern spain ( gaston & redondo 2005 ) . the justification of the species right for these sierra nevada populations merits further attention and analysis .
recorded in southern spain ( müller 2010 : shilap 38 ( 150 ) : 159 - 163 ) , new for the fauna of europe . locus typicus in morocco .
recorded on lampedusa island ( fiumi & guidi 2011 : quad . stud . not . stor . nat . romagna 32 : 199 - 205 ) , new for the fauna of europe . locus typicus in north africa .
upgraded from synonymy to species rank by can ( 2009 : zootaxa 2314 ) , based on correlated differences in male genitalia and dna barcodes . distribution : bulgaria to turkey . locus typicus in bulgaria .
perizoma barrassoi , holotype , italy , majella , leg . n . zahm )
newly described from central italy , majella mountains . male genitalia see zahm , cieslak & hausmann ( 2006 : mitt . münchn . ent . ges . 95 : 31 - 35 ) . so far , only the male holotype is known . the female paratype , mentioned in the original description , is misidentified and is an unusually small specimen of colostygia aquaeata .
one specimen collected in bashkiria ( southern urals ) : mironov , v . g . , trofimova , t . a . 2007 . new species of eupithecia curtis , 1825 ( lepidoptera : geometridae ) for the fauna of southern urals . - eversmannia , 10 : 39 - 41 . urltoken
upgraded from synonymy of e . absinthiata to species rank in : hausmann a , haszprunar g , hebert pdn ( 2011 ) : dna barcoding the geometrid fauna of bavaria ( lepidoptera ) : successes , surprises , and questions . plos one 6 ( 2 ) : e17134 . doi : 10 . 1371 / journal . pone . 0017134 .
described from greece ( pindos ) , sister species of e . conterminata : w eidlich , m . ( 2008 ) : beitrag zur lepidopteren - fauna des notia pindos ( tringia - massiv , lakmos - gebirge und athamano - gebirge ) in griechenland mit beschreibung von zwei neuen arten sowie angaben zur kocherfliegen - ( trichoptera ) und schnakenfauna ( diptera : tipulidae ) — entomofauna , zeitschrift für entomologie , bd . 29 , heft 27 , 469 - 504 .
described from southern spain , to be synonymized with e . dodoneata ( v . mironov ) : weisert , f . 2005 . eine neue art fur europa : eupithecia andrea sp . n . ( lepidoptera : geometridae : larentiinae ) . - z . arb . gem . ost . ent . 57 : 47 - 49 .
patrice leraut ( 2009 ) : moths of europe , vol . 2 : geometrid moths . - n . a . p . editions , 808 pp .
cf comprehensive book reviews in nota lepid . 32 ( 2 ) : 158 - 162 and r . a . r . e . xix ( 1 ) : 34 - 39
( leraut 2008 ) surprised the scientific community by the interesting discovery that this genus splits into several species , such as by the fact that the study was not based on type studies , neither on sufficient material ( strongly biased towards france ) and furthermore draws wrong nomenclatorial conclusions . as a consequence the paper caused additional work to rectify the taxonomy of the group (
) . such kind of scientific replication / rectification is not the way one likes to do , and it could have been avoided through contact and correspondence . the new book on european geometrids ( leraut 2009 ) , extends the same methodology to the whole family geometridae at a european scale . the following explanations may be helpful as immediate information to the whole community and will soon be published in a journal in order to recieve validity through a nomenclatural act .
the book has its strengths in its conception as richly illustrated identification guide in small pocket format , but nevertheless covering most european taxa of geometrids . preparing such a book costs a lot of time , which has to be acknowledged ,
see preface for some scientific shortcomings , e . g . insufficiently studied type specimens and collections ( just basing on material from mnhn / paris and a few french collections ) such as poor contact to the international scientific community of geometridologists , as e . g . organized in the forum herbulot and its periodical meetings ,
the description of infrasubspecific forms is not conform with the code ( iczn ) of zoological nomenclature . all the presented ' names ' are unavailable and need to be ignored in nomenclature ,
many species are stated as being ' closely related ' but without close relationship in the phylogenetic sense . a few examples , standing for many more :
many of the proposed taxonomical and nomenclatural changes are proposed without justification or explanation , basing on personal opinion ( ' is in my view a separate species ' , ' does not appear to me an authentic species ' ) and without presenting a clear statement of differential or common traits . in some cases just the formal change is done without any comment at all , e . g .
( lucas , 1937 ) , bona sp . , stat . rev . ( p . 776 ) . hopefully all this lacking information will be published elsewhere in the near future . the book does not have a synopsis with all taxonomical changes , therefore all these changes are addressed and briefly discussed here on this website .
furthermore there are many apparent taxonomical and methodological errors , partly corrected here , partly requiring further study .
( p . 34 , pl 1 fig 13 and index ) : incorrect subsequent spelling . correct :
. genus can be accepted , cf . revision of leraut ( 2002 ) .
not justified with facts , subjectively presented (
it is clear
) without indication of characters or synapomorphies . in contrast to leraut ' s opinion , the shape of uncus and the much shorter terminal process of valva distinguish
species , thus the arrow on fig . 21b does not indicate a differential feature . study of the fine revision in inoue ( 1943 ) would have given a correct view on the things and on
rungs , 1950
which is not in conformity with the code ( iczn ) concerning authorship . no justification , diagnosis or geographical background given , thus not acceptable .
appear weak and require confirmation by morphology of other material and dna barcoding . preliminarily to be accepted .
without examining material ( not figured ) and ignoring differential anylsis in gme1 . to be maintained at species rank .
without justifying or presenting arguments for this view . to be maintained at species rank .
: downgraded to synonymy , justified only by the statement that such habitus forms
are found elsewhere as isolated individuals
. though this is no reason to draw into question a subspecific taxon , that change may be preliminarily accepted until detailed examination about gene - flow is available .
: downgraded to synonymy , justified only by the statement that such habitus forms are found also in morocco . reputely ,
is founded only on characters of wing pattern , but hausmann ( 2004 ) presents also differences in male pectination and female genitalia , ignored by leraut ( 2009 ) . to be raised again to subspecies rank ( stat . n . ) .
despite citation of differential characters but without bringing arguments for synonymy (
to me does not appear an authentic species
) . to be raised to species rank again ( stat . n . ) .
: raised from synonymy to subspecies rank , vaguely stating some differences in male genitalia , but without describing these differences and without examining the possibility of clinous distribution of characters . still awaiting a thorough analysis and well - done differential analysis , downgraded again to synonymy .
: fig . 32 : the differential feature of valva costa is almost inexistent on the left valva , but dramatically overdrawn on the right side . the indicated structure on the aedeagus of
, though mentioning differential features in habitus and genitalia . to be raised again to species rank ( stat . n . ) .
as separate taxon 1 . 5 % genetical distance ( coi , pairwise distance , k2 ) from populations of nominate subspecies in italy . differences in habitus ( near to constant ) slight differences and male and female genitalia are outlined in hausmann ( 2004 ) . therefore better to be raised again to subspecies rank ( stat . n . ) .
: new evidence ( ounap et al . 2008 ) from a well done revision revealing this genus to be a sterrhine genus , not a larentiine .
: distribution in france ( correctly ) drawn into doubt in the text , but occurrence in south - eastern france marked on the map .
to species rank vaguely stating differences in male genitalia , but without describing these differences and without examining the possibility of clinous distribution of characters . (
genitalia are different
) . still awaiting a thorough analysis and well - done differential analysis , downgraded again to synonymy . furthermore on the map the distribution of
to species rank vaguely stating differences in genitalia , but without describing these differences (
genitalia are different
) . downgraded again to the above mentioned synonymy basing on the analysis presented in hausmann ( 2004 ) . furthermore , inconsistently , on the map
on p . 741 is erroneous , because basing on the misidentified specimen on pl . 134 fig 7 .
could be a separate species
. own research on such individual forms with forewing tip pointed and postmedial line clearly visible not revealing any constant difference in genitalia and neither genetically different ( coi barcode fragment , n = 25 ) from the other forms . therefore to be downgraded to synonymy .
* the ' vernacular ' name
libyan wave
( pl . 135 fig . 18 ) for
( guenée , 1858 ) is misleading , because that species is definitely not distributed in libya , but in the lebanon and nearby countries .
without presenting facts ( e . g . figures of the examined material ) , basing on the erroneous opinion to have examined
the type
and completely ignoring the recently published comprehensive revision of the species group ( hausmann 2005 ) with lectotype designation . taxonomic change not acceptable .
to synonymy . to be maintained at subspecies rank ( cf hausmann 2004 ) .
with statement to have
examined the type
, referring , however , to a paralectotype of mnhn , paris . examination ( brushing of male abdomen and control of the particular shape of valva costa ) of the lectotype in coll herbulot / zsm confirming the proposed synonymy .
: pl . 141 fig 14 : very untypical , without dissection misidentification not excluded .
: pl . 142 , fig . 18 : misidentification not excluded in specimens from southern france without dissection .
. the type has been
examined
but it remains unclear whether it was dissected . the figure of the
type
( which kind of type ? ) shows a specimen which is more reminiscent of
the taxon remains unclear until the genitalia of the name bearing type specimen will be controlled . until then it should be kept in synonymy of
to species rank without any comment . in fact this is a possible view , considering the different length of cerata , but the situation is not that easy when considering also the populations of
the length of cerata underlies a complicated pattern of variation and polymorphisms . requiring a detailed integrated revision including molecular and morphological data .
glossotrophia confinaria , g . alba , g . mentzeri , g . rufomixtaria , g . sacraria , g . asellaria
. such taxonomic position confirmed by own molecular research ( hausmann unpublished data ; hausmann & hebert 2009 ) .
: ( p . 786 and pl . 153 fig 15 ) : possibly misidentified . the species - group around
needs thorough analysis and revision , which is in progress ( r . trusch , karlsruhe ) .
- lythria : positioned between ennominae and larentiinae , though belonging to sterrhinae ( cf ounap et al . 2008 ) .
- lythria plumularia : reported from pyrenees but these old data are doubtful . the species is not mentioned by redondo et al . ( 2009 ) . in collections some very old specimens from
hisp . m .
[ southern spain ] may be mislabelled , perhaps .
- lythria sanguinaria numantaria herrich schäffer , 1852 raised to subspecies rank by leraut ( 2009 ) , reputedly due to ‘differences in genitalia’ herewith downgraded to synonymy ( stat . n . ) , because of the lack of such constant differential features
- cataclysme riguata : wrong map : no occurrence in scandinavia and england ( cf text ) ! but present on sicily .
- cataclysme festivata “from central asia” raised to species rank by leraut . cidaria riguata var . festivata , however , was described basing on syntypes from central asia across altai , saisan to amur ! since leraut did not check the type material , the drawn conclusions are obsolete .
- pl . 77 , fig . 15 cataclysme festivata from “alexander gebirge” ( not “gebiets” ) probably referring to c . shirniensis ebert , 1965 or c . plurilinearia leech , 1897 .
- cataclysme uniformata : raised to species rank earlier by viidalepp ( 7 march 2009 ; cf . mazel 2010 ) , accepted .
- scotopteryx subvicinaria : reputedly recorded in europe , but basing on erroneous data , this is not a european element . the presented ‘differential feature ( postmedial dent ) ’ is not valid for discrimination , neither significant nor constant .
- pl . 78 , fig . 18 ( macedonia ) : misidentified : s . vicinaria rather than s . subvicinaria .
- scotopteryx bipunctaria : records for morocco ( rungs 1981 ) erroneous according to leraut 2009 and attributed to s octodurensis . evidence not presented .
- scotopteryx aelptes : reputedly recorded in south - east europe together with s . olympia , but european ‘records’ of scotopteryx aelptes referring to s . olympia .
- plates 78 - 81 : several specimens of scotopteryx sister species complexes may merit dissection to verify their species identity .
- three species , xanthorhoe uralensis choi , 2003 , x . pseudannotinata vasilenko , 2007 and x . pseudomajorata vasilenko , 2002 not mentioned at all , but x . derzhavini and x . majorata erroneously listed under x . spadicearia , though all five being closely related to the incursata species - group and partly synonym to each other .
- xanthorhoe decoloraria : map with distribution all over germany wrong , the species is restricted here to the southernmost parts in the alps . similarly the distribution in south - eastern europe is drawn far too wide .
- xanthorhoe quadrifasiata : map with distribution all over spain and italy wrong , the species is restricted here to the pyrenees resp . the alps . occurrence in sicily erroneous .
- ‘genus’ odonthorhoe ( with mentioned species alexandraria , tauaria , fidonaria , tianschanica ) validated at genus rank without analysis , just stating it to be ‘distinct’ , though retained as synonym in scoble ( 1999 ) and other recent revisions . herewith again downgraded to synonym of xanthorhoe .
- pl . 85 , fig . 9 : “turkish raker” ( o . alexandraria ) is a misleading name for a species definitely not distributed in turkey , but in turkestan .
- catarhoe basochesiata : existence of a second , allopatric vicariant species , c . hortulanaria in the eastern mediterranean ignored here ( cf . hausmann 1995 ; 1997 ) , greece records on the map and in the text have to be assigned to c . hortulanaria . the species is later ( under c . permixtaria ) mentioned as unknown to the author .
- catarhoe mazeli viidalepp , 2008 regarded as synonym of c . putridaria ( light coloured form ) without basing on a morphological analysis . herewith raised again to species rank , quoting differences in genitalia of both sexes ( see original description ) ( cf mazel 2010 ) .
- pl . 85 , fig . 5 misidentified , referring to c . mazeli rather than to c . putridaria
- epirrhoe pupillata : on the map the distributions throughout scandinavia and germany are wrong , in both regions it is restricted to the southernmost parts .
- epirrhoe latevittata : mentioned , under e . timozzaria , as unknown to the author . better to be compared with e . alternata where is should be subordinated as subspecies or synonym ( cf . gme3 ; foeu database ) .
- camptogramma bilineata bistrigata : downgraded from species rank to subspecies of c . bilineata . acceptable .
- entephria byssata : validated at species rank by leraut , without mentioning the senior synonym and valid name entephria punctipes . herewith downgraded again to synonymy of entephria punctipes ( synonymy previously established in hausmann et al . 2004 fauna of europe ) .
- entephria cyanata : map misleading , occurrence in germany and poland restricted to southernmost mountains ( alps , tatra ) , not distributed in the netherlands .
- entephria nobiliaria vs . flavata pl . 87 figs 16 - 18 : the figured moths would merit control of identification ( e . g . by dna ) , because presenting untypical differences in habitus .
- coenotephria salicata vs . “salicata ablutaria” pl . 90 figs 12 - 13 : the figured moths would merit control of identification ( e . g . by dna ) , because presenting untypical habitus features .
- coenotephria salicata ablutaria : downgraded from species rank to subspecies of salicata by leraut , ignoring extensive studies with rearings , larval morphology , long series of dissections ( e . g . published by rezbanyai - reser in several articles , e . g . 2008 ) . large genetical distance ( e . g . salicata from slowenja highland versus ablutaria from slowenja lowland 5 . 8 % ; salicata from switzerland highland versus ablutaria from north italian lowland 5 . 4 % ) supporting and confirming separate species status , therefore ( again ) raised to species rank herewith ( stat . n . ) .
- two different ( female ) genitalia figs 157b and 158c both reputedly showing “ coenotephria salicata ” .
- nebula nebulata : map misleading , occurrence in germany and poland restricted to southernmost mountains ( alps , tatra ) .
- nebula achromaria : map misleading , occurrence in germany and poland restricted to southernmost mountains ( alps , tatra ) .
- nebula ibericata : the species is common in early spring and in autumn , not in july . distribution ( text and map ) in sicily erroneous , in morocco replaced by nebula numidiata ( mentioned in leraut under n . nebulata ) .
- eulithis pyropata : map misleading , occurrence in germany and poland restricted to north - easternmost mountains of both countries , in sweden only in the south - easternmost part .
- chloroclysta miata : flight period is ususally from september to may , not from “april to october” .
- dysstroma latefasciata : flight period later , from mid - july to september rather than “june - july” .
- cidaria distinctata staudinger , 1892 : no parentheses required around author and year .
- cidaria ochracearia leech , 1897 : no parentheses required around author and year .
- plemyria rubiginata : bivoltinism suggested ( “may , july - august” ) , but the species is univoltine , usually from early june to early august .
- thera firmata : distribution on corsica ( subsp . tyrrhenica tautel & billi , 2009 ) not mentioned in text and on map .
- thera firmata tavoilloti : reputedly only
weakly differentiated
, suggesting it to be an altitudinal form . raised again to subspecies rank by mazel 2010 .
- thera ulicata : reputedly occurring on corsica ( text ) , but not figured on map . probably referring to subsp . tyrrhenica tautel & billi , 2009 ( see under t . firmata ) . map indicating occurrence all over italy and on balkan peninsula , referring to t . firmata , too .
- thera obeliscata : reputedly occurring in north africa ( map ) , data probably erroneous .
- “ thera variata cembrae” : cembrae downgraded from species rank to subspecies of variata without presenting any reason or analysis for that nor quoting several recent analyses in literature . herewith raised again to species rank , quoting striking genetical distance ( detailed analysis in gme3 ) to t . variata which is about 6fold the distance between variata and britannica .
- fig . 165 / 166 : the illustration of genitalia of a “ thera sp . nov . ( in press ) ” without any corresponding reference in the text and without any geographical indication is not very helpful . given the variability of thera genitalia , the indicated “differences” do not represent valuable differential features .
- thera callidaria ( p . 629 ) and pl . 95 , fig . 18 ( type ) : the taxon callidaria was so far retained to be valid at species rank in the genus antilurga ( scoble 1999 ) . leraut transfers it to thera without giving any reason or analysis . the photograph of the type , grace to its publication by leraut , however , clearly allows to put it in ( senior ! ) synonymy with mattia adlata . therefore , this species should be cited as “ mattia callidaria ( joannis , 1891 ) ( syn . mattia adlata syn . n . ) .
- pl . 96 fig . 4 , “singular spanish carpet” ( polythrena coloraria ) is a misleading english vernacular name for a species definitely not distributed in spain , but in north - eastern russia .
- pl . 96 fig . 9 , possibly misidentified and referring to lampropteryx suffumata rather than to electrophaes corylata . awaiting dissection or dna barcoding .
- eustroma reticulata : map misleading , occurrence in spain and italy restricted to northernmost mountains ( alps , pyrenees ) .
- electrophaes corylata : map misleading , occurrence in spain and italy restricted to northernmost mountains ( alps , pyrenees ) .
- colostygia aptata : ' occurrence ' on sicily ( however erroneous ) mentioned in the text but lacking on map .
- colostygia austriacaria : ' occurrence ' in bulgaria ( however erroneous ) mentioned in the text but lacking on map .
- colostygia sericeata : mentioned under c . tempestaria though ( very ) closely related to c . multistrigata .
- colostygia wolfschlaegerae and c . fitzi : mentioned under c . tempestaria though ( very ) closely related to c . olivata .
- colostygia corydalaria : mentioned as bona species on p . 631 under c . tempestaria ignoring its identity as pseudobaptria bogumilaria ( rebel 1904 ) ( cf mironov 2003 ; hausmann et al in fauna of europe 2004 ) . the same taxon is again mentioned on p . 655 as pseudobaptria corydalaria separately from p . bogumilaria .
- colostygia ‘larentiaria’ ( laetaria ) : the name larentiaria is introduced by leraut as ( reputed ) senior synonym for the name colostygia laetaria which was in continuous use for more than 100 years . apparently the type specimen has not been examined . without such an accurate and documented examination of the type specimen the taxonomical change cannot be accepted and the taxon larentiaria must be treated as junior synonym to c . kollariaria ( cf . analysis in gme3 ) despite its erroneous subordination under laetaria in scoble ( 1999 ) .
- colostygia laetaria ( ‘larentiaria’ ) : occurrence in carpathians ( mentioned in text but lacking on map ) erroneous .
- coenocalpe millierata erroneously subordinated as subspecies under c . lapidata , stating their genitalia as being identical with those of c . lapidata , ignoring several constant differential features , e . g . basal lobe of sacculus larger than in c . lapidata , sacculus projection stronger curved , costal projection shorter ( detailed analysis in gme3 ) . herewith raised again to species rank ( stat . n . ) .
- horisme radicaria ; sister species of h . tersata , position of horisme scorteata rather aside h . predotai than between both sister species tersata and radicaria . fig 168 : size of corpus bursae is not a valuable differential feature . figs 169a and 169b : shown differential features invalid , the identification of the illustrated genitalia seems doubtful , possibly they are even inversed .
- horisme vitalbata staudingeri : locus typicus in amur region , thus easternmost asia , not central asia .
- melanthia procellata : map and text misleading , occurrence in spain and italy restricted to northernmost mountains ( alps , northern apennines , pyrenees , asturia ) .
- pareulype “lasithiotica nevadensis” : combined with p . lasithiotica ( nominate subspecies unknown to leraut , or both ) ignoring müller ( 1996 ) , hausmann & aistleitner ( 1998 ) , and redondo ( 2009 ) where treated as subspecies of berberata quoting absence of constant differences in genitalia and genetical identity , detailed analysis in gme3 . to be re - combined with p . berberata .
- spargania luctuata : map and text misleading , no occurrence in spain and italy south of alps and pyrenees .
- rheumaptera hastata : map and text misleading , no occurrence in spain and italy south of alps and pyrenees .
- synonymy of genera hydria and rheumaptera stated without any analysis of characters ; there are also arguments for separating both lineages , e . g . coremata , shape of juxta , sternum a8 a . s . o . .
- rheumaptera cervinalis : flight period indicated as “april - june ( after wintering ) , july - september” . this species , however , overwinters as pupa and a summer generation occurs very rarely .
- rheumaptera montivagata incertata ( mentioned both under r . cervinalis and r . montivagata ) : downgraded to subspecies of r . montivagata , ignoring constant structural differences in genitalia and male hindleg ( cf gme3 ) . to be raised again to species rank ( as e . g . in scoble 1999 ; hausmann et al . 2004 ) .
- rheumaptera gudarica : so far known only from eastern spain , teruel . occurrence in morocco ( leraut 2009 ; figured under pl . 104 , fig 8 from ifrane ) requiring confirmation , exact data should be published , e . g . whether the figured genitalia on fig . 173b are from that specimen from morocco .
- triphosa tauteli : the presented differential features in male genitalia underly artificial variation due to pressure on the genitalia when being embedded . according to redondo ( 2009 ) and hausmann et al . ( 2004 ) , t . dyriata is distributed also in spain . spanish populations should be analyzed in detail ( integrative taxonomic analysis ) to check if there are clinous character transitions between t . dyriata and t . tauteli . the taxon “ herzeti ” ( “ lot” and “jura” according to leraut 2009 ) must sink to synonymy , or the description of a large number of “subspecies” would be necessary from other regions when applying the same criteria .
- triphosa “sabaudiata agnata” : downgraded from species rank ( scoble 1999 ) to subspecies of t . sabaudiata without presenting reasons , analyses or statistical details . to be maintained at species rank until detailed analysis is performed .
- philereme transversata : flight period indicated as “may - july” . this species , however , usually flies from mid - june to mid - august , sometimes until early september .
- euphyia frustata : the taxa griseoviridis ( corsica ) and fulvocinctata ( spain ) are said to be “definitely not a separate subspecies” . however , because of the existence of several differential features in male and female genitalia ( shape of uncus , valva and antrum ) , both taxa should be maintained at subspecies rank .
- euphyia maximiliana : erroneously suggested to be a synonym of e . frustata .
- epirrita christyi : author and year stated as “prout , 1899” though “allen , 1906” is generally accepted as valid reference . text states absence from corsica , map erroneously shows occurrence on corsica .
- solitanea mariae : flight period stated as “july - august” . in italy , however , the species is bivoltine : early may to mid - june ; late july to mid - september , in the lowlands until late october . distribution in the text stated as “corsica : calabria” , despite wide distribution on apennine peninsula .
- chesias capriata : formally treated at species rank , following hausmann et al . ( 2004 ) , but expressing certains doubts about the rank . morphology , habitus and genetics confirming species rank .
- chesias plumbeata : raise from subspecies of ch . rufata to species rank without analysis and without mentioning any differential feature . to be maintained at subspecies rank of ch . rufata until not submitted to a sound morphological or integrative analysis .
- chesias linogrisearia : rank is confusingly presented as “form or species” , possibly a senior synonym of ch . rufata cinereata . species rank earlier postulated by scoble ( 1999 ) , now confirmed by a large genetical distance from ch . rufata by > 5 % .
- the taxa pinkeri and zuellichi are suggested to be synonyms of ch . angeri without giving any detail or argument . to be maintained in the systematic position as presented by hausmann et al . ( 2004 ) : ch . rufata pinkeri with zuellichi as synonym .
- carsia sororiata : map misleading , occurrence in germany and poland restricted to southernmost mountains .
- aplocera fraudulentata : downgraded from species rank to subspecies of a . plagiata without analysis and without discussing the existing differential features . examined specimens , however revealing strongly different genitalia and strongly differing dna barcodes ( coll . gelbrecht , coll . herbulot ; gelbrecht pers . comm . , own data ) . to be maintained at species rank .
- aplocera cretica ( “wingspan 27 - 28 mm” ) is said to distinctly smaller than a . plagiata but , largest a . cretica of first generation reach 38 mm wingspan . for flight time only october is mentioned , but the species is bivoltine , april to october .
- ‘ aplocera ’ fulgurata ( india ) : transferred from genus docirava to aplocera without morphological analysis and without discussing any differential feature . in gme3 a differential analysis between docirava and aplocera will be given . to be maintained in genus docirava .
- aplocera columbata : map misleading , occurrence in greece restricted to north - easternmost part , in ukraine restricted to crimea ( outside the map ) .
- docirava dervenaria and d . mundulata : generic systematics questioned under the species aplocera columbata without morphological analysis and without discussing any differential feature . in gme3 a differential analysis between docirava and aplocera will be given . both species to be maintained in genus docirava .
- docirava erubescens ( staudinger , 1892 ) ( turkey , armenia ) : reputedly with palumbata mentzer , 1981 as a new synonym . the former described from northern turkey ( amasia ) and figured in leraut from ‘ middle east’ ( ? ) . the latter described from northern iran ( elburs ) and so far retained as falling under d . mundulata . awaiting verification , examination of types still pending .
- schistostege nubilaria : map misleading , occurrence in poland ( if any ) restricted to south - easternmost part .
- lithostege duponcheli : on map ( not in text ) erroneously reported from central pyrenees . on peninsular italy restricted to the north - westernmost parts , and a very few localities in tuscany and southern italy .
- lithostege cinerata : no occurrence in europe , data referring to l . clarae : presented by hausmann et al . 2004 ( faeu database ) under the name ‘ cinerata ’ before the description of clarae .
- pl . 126 , figs 13 - 15 ( lithostege cinerata ) : 13 - 14 probably not conspecific with 15 , awaiting verification by dissection and / or dna barcoding .
- lithostege odessaria : distribution stated as “ ukraine , middle east” , the former probably wrong ( type locality odessa probably referring to an old name of a caucasian village ) , the latter misleading , as the species may be restricted to caucasus and transcaucasus .
- lithostege fissurata ( mentioned under l . odessaria ) : occurrence in europe ( malta : hausmann & seguna 2005 ) neglected .
- venusia cambrica : map and text misleading , no occurrence in spain and italy south of alps and pyrenees .
- hydrelia flammeolaria : map erroneously suggests occurrence on ireland , and iberian peninsula south of pyrenees .
- pterapherapteryx sexalata : map and text misleading , no occurrence in spain and italy south of alps and pyrenees .
- venusia cambrica : map and text misleading , no occurrence in italy north of central apennines .
- trichopteryx polycommata : map and text misleading , no occurrence in spain and italy south of alps and pyrenees .
- acasis viretata : map and text misleading , absent from southern half of iberian peninsula .
acknowledgements : we are deeply grateful to paul hebert and his competent team for offering us the possibility to integrate molecular data ( coi barcodes ) into our morphological analyses . these analyses are performed in the framework of the international ibol program ( campaigns geometridae , lepidoptera global and fauna of europe ) .
and allies : cf remarks to the presented , confusing data in mazel 2010 .
encyclo . co . uk , online since 2007 , is a search engine for english meanings and definitions . the website aims to publish all wordlists , big and small , on the internet , making it much easier to find the word you need . follow us on facebook


rhynchactis is a genus of deep-sea anglerfish in the family gigantactinidae , containing three species found worldwide at depths greater than 400 m ( 1,300 ft ) .

adult female rhynchactis reach a standard length ( sl ) of 11 – 13 cm ( 4.3 – 5.1 in ) and have a dark-colored , streamlined body and a relatively small head bearing a very long illicium ( the " fishing rod " formed by the first ray of the dorsal fin ) .

unlike almost all other deep-sea anglerfishes , the illicium bears no bioluminescent esca ( the " lure " ) at the tip .

the mouth is almost devoid of teeth , and the inside of both jaws are covered by numerous white glands that are unique to this genus .

the lack of an esca , greatly reduced dentition , and glands inside the mouth all point to rhynchactis having a highly specialized mode of feeding , the nature of which has yet to be deciphered .

as in other deep-sea anglerfishes , there is enormous sexual dimorphism with males being much smaller than females and lacking an illicium , though they do not appear to be parasitic as in some families .

reproduction is oviparous , with the larvae having a rounded shape and enlarged pectoral fins .

rhynchactis

there are no species - specific conservation efforts at this time for rhynchactis macrothix .
revision of the deepsea anglerfish genus rhynchactis regan ( lophiiformes : gigantactinidae ) , with descriptions of two new species .
revision of the deep - sea anglerfish genus rhynchactis regan ( lophiiformes : gigantactinidae ) , with descriptions of two new apecies .
species of the family gigantactinidae are meso - bathypelagic anglerfish ( pietsch in press ) . species in the genus rhynchactis have a wide distribution in all three major oceans of the world between approximately 32°n and 12°s ( bertelsen and pietsch . 1998 ) . rhynchactis macrothrix is known to inhabit waters as shallow as 300 m and as deep as 2 , 000 m ( pietsch 2009 ) .
bertelsen , e . , and t . w . pietsch . 1998 . revision of the deepsea anglerfish genus rhynchactis regan ( lophiiformes : gigantactinidae ) , with descriptions of two new species . copeia , 1998 ( 3 ) : 583590 .
bertelsen , e . & t . w . pietsch . 1998 . revision of the deepsea anglerfish genus rhynchactis regan ( lophiiformes : gigantactinidae ) , with descriptions of two new species . copeia 1998 ( 3 ) : 583 - 590 .
bertelsen , e . , and t . w . pietsch . 1998 . revision of the deepsea anglerfish genus rhynchactis regan ( lophiiformes : gigantactinidae ) , with descriptions of two new species . copeia , 1998 ( 3 ) : 583 - 590 .
bertelsen , e . and t . w . pietsch , 1998 . revision of the deepsea anglerfish genus rhynchactis regan ( lophiiformes : gigantactinidae ) , with descriptions of two new species . copeia 1998 ( 3 ) : 583 - 590 . ( ref . 28611 )
color of females in preservative uniformly dark brown to black over entire external surface of head , body , and fins ( except for unpigmented transparent parts of escal bulb , escal appendages , and stem of illicium ; for ontogenetic and specific variation , see species accounts below ) ; oral cavity more or less pigmented . skin of metamorphosed males weakly pigmented in rhynchactis , varying among species - groups of gigantactis from unpigmented to dark brown or black . subdermal pigment of larvae , males , and juvenile females varying between genera and among species groups of gigantactis from absent ( e . g . , gigantactis longicirra ) to dense , dorsal and peritoneal groups of melanophores ( e . g . , rhynchactis ) .
justification : rhynchactis macrothix is distributed in all three major oceans of the world between approximately 32°n and 12°s . it is a meso - bathypelagic species , residing between 300 m and 2 , 000 m . little is known about its population or ecology . there are no known major threats . based on the limited information , r . macrothix is assessed as data deficient .
the whipnose anglers are placed in 2 genera , gigantactis brauer , 1902 , with 18 species , and rhynchactis regan , 1925 , with 3 . the rare , primarily tropical rhynchactis is known just outside our area ( off madagascar ) from a single female of r . macrothrix bertelsen and pietsch , 1998 and is not included here . six species of gigantactis have been found in our area . females of gigantactis reach about 41 cm sl , and adults of both sexes are primarily bathypelagic . metamorphosed males are non - parasitic and reach 22 mm sl ; none are yet known from our area . parr ( 1927 ) described the first male of gigantactis but placed it in a new genus , laevoceratias , family aceratiidae . bertelsen ( 1951 ) tentatively placed it in the family diceratiidae . the gigantactinids were last reviewed by bertelsen et al . ( 1981 ) who examined all the then known material and recognized 17 species . kharin ( 1984 ) described g . balushkini from off japan .
the fishing
lure ,
or illicium , in ceratioids is tipped with a luminescent organ , the esca , except in the caulophrynidae , and in the gigantactinid genus rhynchactis . the illicium of rhynchactis macrothrix , a species that has been taken just outside our area , is tipped with a slight bulbous swelling , but this is not an esca ( bertelsen and pietsch , 1998 ) . often the esca is adorned with spinules , papillae and a variety of filaments and appendages . illicium length is measured from its base only to the distal end of the bulb of the esca . other light organs include the hyoid barbels of linophryne and the dorsal midbody caruncles in ceratiids . tooth counts are a summation of both sides of each jaw . free - living metamorphosed males have a unique toothy bone articulating with the anterior ends of both jaws called the upper and lower denticulars ; their denticles form into rows and are counted . scales are often modified into skin spinules ( = prickles or dermal spines elsewhere ) , or greatly enlarged bucklers ( himantolophidae ) .
both genera of the family are represented in all three major oceans of the world . females of gigantactis have been reported from as far north as southern greenland , with the southernmost record close to 50°s in the atlantic sector of the southern ocean . the known distribution of rhynchactis is restricted to tropical and subtropical regions between about 30°n and 10°s . the great majority of the recorded metamorphosed specimens of the family have been captured in nets fished in maximum depths exceeding 1000 m . gigantactinids thus appear to be among the deepest - living ceratioids , with greatest abundance between 1000 and at least 2500 m .
ceratioid anglerfishes differ further from their shallow - water relatives in having a bacterial light - organ that serves as bait to attract prey , a structure technically called the “esca”—exceptions among members of the suborder include the monotypic family neoceratiidae ( bertelsen , 1951 ) , the three species of the gigantactinid genus rhynchactis ( bertelsen et al . , 1981 ; bertelsen and pietsch , 1998 ) , and the five members of the family caulophrynidae ( pietsch , 1979 ) . parr ( 1927 ) was the first to recognize the diagnostic value of the external morphology of escae in ceratioids , pointing out the need for a closer examination of individual variation in the structure of this organ . since that time , differences in the number , shape , and size of escal appendages and filaments , as well as variation in external escal pigment patterns , have been , for the most part , the sole basis on which new species have been described ( e . g . , see pietsch , 1974 ; bertelsen et al . , 1981 ; bertelsen and krefft , 1988 ) .
bertelsen , e . & t . w . pietsch . 1977 . results of the research cruises of frv
walther herwig
to south america . xlvii . ceratioid anglerfishes of the family oneirodidae collected by the frv
walther herwig .
arch . fischwiss . , 27 ( 3 ) : 171 - 189 .  . 1983 . the ceratioid anglerfishes of australia . recs . australian mus . , 35 : 77 - 99 .  . 1984 . results of the research cruises of frv
walther herwig
to south america . lxiii . a resurrection of the ceratioid anglerfish ceratias tentaculatus ( norman , 1930 ) with notes on the occurrence of the species of ceratias in the atlantic ocean ( pisces : lophiiformes ) . arch . fischwiss . , 35 ( 1 / 2 ) : 43 - 51 .  . 1996 . revision of the ceratioid anglerfish genus lasiognathus ( lophiiformes : thaumatichthyidae ) . copeia , 1996 ( 2 ) : 401 - 409 .  . 1998 . revision of the deepsea anglerfish genus rhynchactis regan ( lophiiformes : gigantactinidae ) , with descriptions of two new species . copeia , 1998 ( 3 ) : 583 - 590 .
rhynchactis has undergone such a drastic reduction and loss of parts that clearly it is the more derived of the two gigantactinid genera . within the genus gigantactis , there are four morphological trends that seem to characterize anglerfish evolution in general ( pietsch , 1972 , 1974 : 87 , 88 ) : ( 1 ) an increase in the length of the illicium , ( 2 ) a decrease in the number of median - fin rays , ( 3 ) a loss of jaw teeth , and ( 4 ) an increase in morphological complexity of the luring apparatus , in this case , reflected in a general tendency to increase the number of distal filaments of the esca and filaments of the illicium . giagantactis longicirra appears to be the least derived member of the genus , having the shortest illicium , the greatest number of longitudinal series of dentary teeth , the highest dorsal - ray count , and the least number of distal escal filaments . members of the g . macronema group ( g . macronema , g . microdontis , and g . ios ) are the most derived , having the longest illicium , the fewest series and total number of dentary teeth , the lowest dorsal - ray counts , and numbers distal , escal filaments . the remaining species of the genus are more or less intermediate in specialization . members of the g . vanhoeffeni group , include g . vanhoeffeni , g . meadi , g . gibbsi , g . gracilicauda , and g . paxtoni , are united in sharing a relatively short illicium and a similar escal morphology . members of the g . gargantua group , including g . gargantua , g . watermani , and g . herwigi , likewise share a similar escal morphology but are also united on the basis of having a relatively long illicium and an elongation of the second and seventh caudal - fin rays .
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the gigantactinidae is one of the most well - defined and highly specialized families of deep - sea anglerfishes . the females are readily distinguished from those of the other ten families of the suborder by having an elongate streamlined shape , a relatively small head and slender caudal peduncle , five pectoral radials , and a greatly prolonged illicium that reaches a length between one and four times standard length . the family includes 22 species in two genera .
as is the case with nearly all ceratioid groups , little is known about the ecology of the gigantactinids . however , recent evidence suggests that this group lives a benthic lifestyle and may even swim upside down while foraging , drifting motionless as the esca lures prey off the bottom ( moore , 2002 ) . rov jason , of woods hole oceanographic insitutue , captured a whipnose anglerfish ( gigantactis sp . ) on video ( hawaii - 2 observatory video clips ) swimming ( see clip fish _ gig _ 1 _ 1 ) and apparently foraging upside down ( see clip fish _ gig _ 1 _ 3 ) .
metamorphosed females of the gigantactinidae differ from those of all other ceratioid families in lacking the vomer , mesopterygoid , and an ossified scapula ; in having the preopercle reduced to a short narrow strut of bone ; the interopercle reduced , without ligamentous connection to the angular ; the caudal fin emarginate ( except in the largest known females of gigantactis kreffti and g . macronema ) , with nine caudal - fin rays , the ventralmost ray reduced and embedded within skin surrounding the adjacent ray ; the pterygiophore of the illicium exceptionally large , the compressed posterior end butting up against the supraoccipital ; and five pectoral radials ( bertelsen et al . , 1981 ) .
free - living male of gigantactis male group i , 14 . 5mm , zmuc p921533 ( after bertelsen et al . , 1981 ) . © 1981 , e . bertelsen et al . , k . elsman
metamorphosed gigantactinid males differ from those of all other ceratioid families in having the following combination of character states : eyes minute , only 3 - 5 % sl in most specimens ; olfactory organs large , depth 8 - 10 % sl in most specimens ; anterior nostrils close together , directed anteriorly ; premaxillae degenerate , but maxillae well developed ; jaw teeth absent ; denticular teeth all or nearly all mutually free ; upper denticular teeth 3 - 6 ( rarely 2 ) , not connected to pterygiophore of illicium ; lower denticular teeth 4 - 7 ( rarely 3 ) ; hyomandibular with a single head ; branchiostegal rays 6 ( rarely 7 ) ; pectoral radials 5 ; fin - ray counts as given for metamorphosed females ; pelvic bones absent ; skin naked or densely covered with dermal spinules ; free - living , no evidence that males ever become parasitic , but temporarily attached males are unknown ( see pietsch , 2005 ) .
in addition to the sexual dimorphism common to all metamorphosed ceratioids , gigantactinid males differ further from females of the family in having a symphysial cartilage , a vomer , and a basibranchial ossification ; they differ also in having fully developed frontals , parietals , opercular bones , and ceratobranchials ( bertelsen et al . , 1981 , figs . 11d - f , 14 , 15 ) .
gigantactinid larvae differ from those of all other ceratioid families in having exceptionally large pectoral fins ( length 45 - 55 % sl ) , comparable only to those of the caulophrynidae ; they differ further in having the following combination of character states : short , nearly spherical ; skin highly inflated ; pelvic fins absent ; fin - ray counts as given for metamorphosed females ; sexual dimorphism evident , females with a small , club - shaped illicial rudiment protruding from head ; metamorphosis beginning at 8 - 10 mm sl , metamorphosal stages 9 - 20 mm sl ( bertelsen , 1981 , 62 , 1984 : 326 , fig . 168a - b ) .
gigantactinids are among the largest known ceratioids . while some of the 30 or so females of gigantactis greater than 200 mm have relatively large ovaries , none contain ripe or ripening eggs . eggs larger than about 0 . 5 mm in diameter have not been found . at the same time , none of the more than 175 metamorphosed females in collections around the world are parasitized by males , thus it is assumed that males remain free - living . the largest known male , a member of the gigantactis male group i measures 22 mm and has ripe testes .
bertelsen , e . 1951 . the ceratioid fishes . ontogeny , taxonomy , distribution and biology . dana rept . , 39 , 276 pp .
bertelsen , e . , t . w . pietsch , and r . j . lavenberg . 1981 . ceratioid anglerfishes of the family gigantactinidae : morphology , systematics , and distribution . nat . hist . mus . l . a . co . , contri . sci . , 332 , vi , 74 pp .
brauer , a . 1902 . diagnosen von neuen tiefseefischen , welche von der valdivia - expedition gesammelt sind . zool . anz . , 25 , 668 ( 4 ) : 277298 .
moore , j . a . 2002 . upside - down swimming behaviour in a whipnose anglerfish ( teleost : ceratioidei : gigantactinidae ) . copeia , 2002 ( 4 ) : 11441146 .
pietsch , t . w . 1972 . ergebnisse der forschungsreisen des ffs
walther herwig
nach südamerika . xix . systematics and distribution of ceratioid fishes of the genus dolopichthys ( family oneirodidae ) , with the description of a new species . arch . fischwiss . , 23 ( 1 ) : 128 .
pietsch , t . w . 1974 . osteology and relationships of ceratioid anglerfishes of the family oneirodidae , with a review of the genus oneirodes lütken . nat . hist . mus . l . a . co . , sci . bull . , 18 , 113 pp .
pietsch , t . w . 2005 . dimorphism , parasitism , and sex revisited : modes of reproduction among deep - sea ceratioid anglerfishes ( teleostei : lophiiformes ) . ichthyol . res . , 52 : 207236 .
regan , c . t . 1912 . the classification of the teleostean fishes of the order pediculati . ann . mag . nat . hist . , ser . 8 , 9 ( 28 ) : 277289 .
regan , c . t . 1925 . new ceratioid fishes from the n . atlantic , the caribbean sea , and the gulf of panama , collected by the
dana .
ann . mag . nat . hist . , ser . 8 , 8 ( 62 ) : 561567 .
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pietsch , theodore w . 2005 . gigantactinidae . whipnose seadevils . version 06 november 2005 ( under construction ) .
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greek , ' rhynch ' = beak or snout + greek , ' aktis ' = ray ( in reference to the illicium that arises near the tip of the snout ) ( ref . 86949 )
marine ; bathypelagic ; depth range 0 - 2250 m ( ref . 28611 ) . deep - water
maturity : l m ? range ? - ? cm max length : 11 . 3 cm sl ( female )
dorsal soft rays ( total ) : 4 ; anal soft rays : 4 . length of illicium 210 % sl , tip with 4 tiny filaments ; irregular series of 19 filaments on distal 14 % of length of illicium , five longest 2 - 4 % sl , all gradually tapering from base to tip ( ref . 86949 ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 6250 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 01000 ( 0 . 00244 - 0 . 04107 ) , b = 3 . 04 ( 2 . 81 - 3 . 27 ) , in cm total length , based on all lwr estimates for this body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 9 ±0 . 2 se ; based on size and trophs of closest relatives
vulnerability ( ref . 59153 ) : low vulnerability ( 10 of 100 ) .
marine ; bathypelagic ; depth range 300 - 2000 m ( ref . 86949 ) . deep - water
maturity : l m ? range ? - ? cm max length : 13 . 0 cm sl ( female )
dorsal soft rays ( total ) : 4 ; anal soft rays : 3 . length of illicium 109 - 144 % sl , distal tip bearing 3 - 4 small unpigmented filaments , some with tiny distal swelling ; distal 28 - 57 % illicial length bearing series of 11 - 20 secondary filaments , five longest filaments measuring 6 - 15 % sl ; each secondary filament with 1 - 2 unpigmented terminal filaments , some with tiny distal swelling ; large specimens ( 110 - 113 mm ) with secondary filaments darkly pigmented to base of terminal filaments , divided on each side by a narrow , well - defined longitudinal transparent band ( ref . 86949 ) .
) : 2 - 9 , mean 6 . 1 ( based on 368 cells ) .
bayesian length - weight : a = 0 . 01995 ( 0 . 00906 - 0 . 04395 ) , b = 3 . 01 ( 2 . 83 - 3 . 19 ) , in cm total length , based on all lwr estimates for this body shape ( ref . 93245 ) .
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anguilla ; antigua and barbuda ; aruba ; australia ; bahamas ; bangladesh ; barbados ; belize ; benin ; bermuda ; bonaire , sint eustatius and saba ( saba , sint eustatius ) ; brazil ; british indian ocean territory ; cameroon ; cape verde ; cayman islands ; china ; christmas island ; cocos ( keeling ) islands ; colombia ; comoros ; congo ; cook islands ; costa rica ; côte d ' ivoire ; cuba ; curaçao ; disputed territory ( paracel is . , spratly is . ) ; djibouti ; dominica ; dominican republic ; ecuador ( ecuador ( mainland ) , galápagos ) ; el salvador ; equatorial guinea ; fiji ; france ( clipperton i . ) ; french guiana ; french polynesia ; french southern territories ( mozambique channel is . ) ; gabon ; gambia ; ghana ; guadeloupe ; guam ; guatemala ; guinea ; guinea - bissau ; guyana ; haiti ; honduras ; india ( andaman is . , nicobar is . ) ; indonesia ; iran , islamic republic of ; jamaica ; japan ; kenya ; kiribati ( kiribati line is . , phoenix is . ) ; liberia ; madagascar ; malaysia ; maldives ; marshall islands ; martinique ; mauritania ; mauritius ; mayotte ; mexico ; micronesia , federated states of ; montserrat ; morocco ; mozambique ; myanmar ; nauru ; nicaragua ; nigeria ; northern mariana islands ; oman ; pakistan ; palau ; panama ; papua new guinea ; philippines ; portugal ( azores , madeira ) ; puerto rico ; saint helena , ascension and tristan da cunha ( ascension ) ; saint kitts and nevis ; saint lucia ; saint martin ( french part ) ; saint vincent and the grenadines ; samoa ; sao tomé and principe ; senegal ; seychelles ; sierra leone ; solomon islands ; somalia ; spain ( canary is . ) ; sri lanka ; suriname ; taiwan , province of china ; tanzania , united republic of ; thailand ; togo ; tokelau ; trinidad and tobago ; turks and caicos islands ; tuvalu ; united states ( hawaiian is . ) ; united states minor outlying islands ( howland - baker is . , johnston i . , wake is . ) ; venezuela , bolivarian republic of ; viet nam ; virgin islands , british ; virgin islands , u . s . ; wallis and futuna ; western sahara ; yemen
the gigantactinidae are known for their extreme sexual dimorphism . males of the gigantactinidae family are characterized by their small size . they are considered a dwarf male family with a maximum size of 2 . 2 cm , while the females are significantly larger than the males with the ability to reach 43 . 5 cm in size ( pietsch in press ) . in the gigantactinidae family males are known to attach to the females by using pincer - like denticles . in this particular family males do not permanently attach and are only attached for a small period of time ( pietsch 2005 ) .
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atp synthase or complex v ) produces atp from adp in the presence of a proton gradient across the membrane which is generated by electron transport complexes of the respiratory chain . f - type atpases consist of two structural domains , f
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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 .
of the five major taxa of lophiiform fishes , the deep - sea ceratioidei is the most phylogenetically derived ( bertelsen , 1984 ; pietsch , 1984 ) . ceratioidei are distributed throughout the world’s oceans below a depth of 300 m . with 160 species , it constitutes by far the most species - rich vertebrate taxon within the bathypelagic zone and below—more than twice as many families and genera and more than three times the number of species as the cetomimoidei , the next most species - rich deep - sea vertebrate taxon ( see paxton , 1998 ; herring , 2002 ) . at the same time , new species are being added to the suborder at a steady if not increasing rate .
the internal structure of ceratioid escae is infinitely more complex , involving a confusing array of bacteria - filled vesicles , light - absorbing pigment layers , reflecting tissues , tubular light - guiding structures , nerves , blood vessels , and smooth muscle fibers ( munk and bertelsen , 1980 ; munk , 1988 , 1998 , 1999 ; herring and munk , 1994 ; munk and herring , 1996 ; munk et al . , 1998 ) . there is some evidence also that ceratioid escae contain pheromone - producing secretory glands that function to attract a conspecific male ( munk , 1992 ) , but the true nature and adaptive significance of these structures and most of the other internal parts of escae are unknown .
in addition to the esca , all 23 currently recognized species of the ceratioid genus linophryne ( family linophrynidae ) bear an elaborate bioluminescent hyoid barbel , the light of which does not originate from symbiotic luminescent bacteria but rather from a complex array of intrinsic , intracellular , paracrystalline photogenic granules ; the bacteria - filled esca is ectodermal in origin , whereas the barbel light organ appears to be derived from the mesoderm ( hansen and herring , 1977 ) . this remarkable dual system , involving two entirely separate mechanisms of light production , is unique among animals .
in summary , ceratioid anglerfishes are among the most intriguing of all animals , possessing a host of spectacular morphological , behavioral , and physiological innovations found nowhere else . the suborder is taxonomically diverse : with 160 currently recognized species ( and many more certain to be discovered in the future ) , it forms a major contribution to the biodiversity of the deep - sea . it is exceedingly widespread geographically , occurring in deep waters of all major oceans and seas of the world , from high arctic latitudes to the southern ocean ; while some species appear to be almost cosmopolitan in distribution , many others have surprisingly small , restricted , vertical and horizontal ranges . their relative abundance , high species diversity , and trophic position as the top primary carnivores in meso - and bathypelagic communities make them important ecologically . their unique mode of reproduction has significant biomedical implications to the fields of endocrinology and immunology . yet , despite these many aspects of biological interest and importance , as well as a large amount of revisionary work published in the 1970s and early 1980s , including repeated attempts to resolve phylogenetic relationships , ceratioid anglerfishes have remained poorly known .
a suborder of lophiiforms unique and derived in lacking pelvic fins ( except in larval and newly metamorphosed caulophrynidae ) , pelvic bones reduced or absent ; extremely sexually dimorphic , males dwarfed , a fraction of size of females . spinous dorsal - fin of two spines ( the illicium and second cephalic spine ) , both supported by a single pterygiophore ; anteriormost dorsal - fin spine modified to serve as a lure , emerging from dorsal surface of cranium ( somewhat behind cranium in bufoceratias , from roof of mouth in thaumatichthyidae , absent in neoceratiidae ) ; second dorsal - fin spine reduced to a tiny remnant ( but relatively well developed in adolescent diceratiidae and ceratiidae , absent in neoceratiidae and linophrynidae ) , embedded beneath skin of head and lying on , or fused to , dorsal surface of pterygiophore just behind base of illicial bone ; postcephalic , spinous dorsal - fin absent ; palatine teeth absent ; interhyal without medial , posterolaterally directed process ; ceratobranchial v toothless , reduced to a slender rod - shaped element in most families , represented by tiny remnants in the gigantactinidae ; epibranchial i simple , without ligamentous attachment to epibranchial ii ; pharyngobranchial iv absent ; cleithral spine absent ; interopercle reduced , elongate and narrow ; subopercle with ascending process absent or reduced to a small spine or projection detached from opercle ; gill filaments present as holobranchs on gill arches ii and iii , extending as hemibranchs onto proximal end of arch i in some families ( i . e . , caulophrynidae , himantolophidae , melanocetidae , diceratiidae , some oneirodidae , centrophrynidae , and neoceratiidae ) , and as hemibranchs on arch iv ; pseudobranch absent ( tiny remnants present in some gigantactis ) ; swimbladder absent .
males with elements of cephalic dorsal - fin spines greatly reduced , remnants embedded beneath skin of head ; eyes and olfactory organs greatly enlarged in most taxa ; tip of snout and chin bearing hooked denticular teeth , used for attachment to females ; in some families and genera attachment becomes parasitic through fusion of male and female tissue .
females with body short and deep , almost spherical in most families , but elongate and somewhat laterally compressed in thaumatichthyidae , centrophrynidae , ceratiidae , gigantactinidae , neoceratiidae , and some oneirodidae ; head length usually greater than 40 % sl , but only about 25 % in elongate species ; mouth moderate to extremely large ( compared to other lophiiforms ) , length of premaxilla as little as 10 % sl ( in some gigantactinidae ) to as great as 40 % sl ( in some linophrynidae ) ; cleft of mouth nearly horizontal ( e . g . , thaumatichthyidae , centrophrynidae , and gigantactinidae ) or oblique to almost vertical ( e . g . , melanocetidae and ceratiidae ) .
sexual dimorphism extreme : males dwarfed , those of most genera reaching a maximum standard length of only 6 - 8 % of that of females , and less than 25 % of that of the largest females of genera for which mature females are known ; body elongate , head less than 40 % sl ; mouth small , length of premaxilla usually less than 15 % sl ; cleft of mouth horizontal . metamorphosed males of all families with hooked denticular teeth apparently adapted especially for gripping the skin of females ; becoming permanently attached and parasitic in caulophrynidae , ceratiidae , neoceratiidae , linophrynidae , and the oneirodid genera bertella and leptacanthichthys .
fin - ray counts the same for males and females , highly variable : dorsal rays 3 - 8 in most families , 11 - 13 in neoceratiidae , and 12 - 22 in caulophrynidae and melanocetidae ; anal rays 3 - 7 in most families , 5 - 19 in caulophrynidae , and 10 - 13 in neoceratiidae ; caudal rays nearly always 9 , but 8 in the ceratiid genus cryptopsaras and 10 in some specimens of neoceratiidae ; pectoral rays 12 - 30 . pelvic fins absent ( except in larval and early metamorphosis stages of caulophrynidae ) . rays of unpaired fins sometimes greatly elongate ( e . g . , in females of caulophrynidae and gigantactinidae ) and often not interconnected , or interconnected with extremely thin , transparent , membrane - like tissue . caudal fin rounded in nearly all families , but emarginate in some gigantactinidae ; caudal rays of some large individuals of ceratiidae terminating in spherical skin - covered ossifications .
coloration of males and females usually uniform dark brown to black over entire head , body , and fins ; dermis everywhere unpigmented and translucent in the linophrynid genus haplophryne .
in some ceratioid taxa , the male ' s attachment to the female is followed by fusion of epidermal and dermal tissues and , eventually , by a connection of the circulatory systems so that the male becomes permanently dependent on the female for blood - transported nutrients , while the host female becomes a kind of self - fertilizing hermaphrodite ( regan , 1925a , b , 1926 ; parr , 1930 ; regan and trewavas , 1932 ; bertelsen , 1951 ; pietsch , 1975 , 1976 ; munk and bertelsen , 1983 ; munk , 2000 ) . permanent attachment is usually accomplished by means of separate outgrowths from the snout and tip of the lower jaw of the male , both of which eventually fuse with the skin of the female . in some species a papilla of female tissue protrudes into the mouth of the male , sometimes appearing to completely occlude the pharynx . the heads of some males become broadly fused to the skin of the female , extending from the tip of the lower jaw to the rear of the skull , appearing as if embedded or absorbed by their mate , while in others , the male is carried at the tip of an elongate , cylindrical stalk of female tissue .
increasing considerably in size once fused , their volume becoming much greater than free - living males of the same species , and being otherwise completely unable to acquire nutrients on their own , the males are considered to be parasites . they apparently remain alive and reproductively functional as long as the female lives , participating in repeated spawning events . a single male per female appears to be the rule in some taxa , but in others multiple attachments are relatively common , with as many as eight coupled to a single host ( saruwatari et al . , 2001 ) .
since its discovery some 80 years ago ( saemundsson , 1922 ; regan , 1925a , b ) , the story of sexual parasitism in ceratioid anglerfishes has become a part of common scientific knowledge . however , the known facts concerning this remarkable reproductive mode have never been thoroughly and satisfactorily analyzed , despite the work of bertelsen ( 1951 ) and more recently of munk and bertelsen ( 1983 ) , munk ( 2000 ) , and pietsch ( 2005 ) . the physiological mechanisms ( endocrinological and immunological ) that allow for sexual parasitism , which could be of significant biomedical importance , have never been explored .
males of the caulophrynid genus robia , the single known female of which has dorsal - fin rays 6 and anal - fin rays 5 , are unknown .
males of the thaumatichthyid genus lasiognathus , the females of which have naked skin , are unknown .
males of eight of the 16 recognized oneirodid genera are unknown , including spiniphryne , the females of which have spinulose skin .
bertelsen , e . 1984 . ceratioidei : development and relationships . pp . 325334 , in : moser , h . g . , w . j . richards , d . m . cohen , m . p . fahay , a . w . kendall , jr . , and s . l . richardson ( editors ) , ontogeny and systematics of fishes , spec . publ . no . 1 , amer . soc . ichthy . herpet . , ix , 760 pp .
bertelsen , e . , and g . krefft . 1965 . on a rare ceratioid fish , linophryne lucifer collett , 1886 . vidensk . medd . fra dansk naturh . foren . , 128 : 293301 .
bertelsen , e . , and g . krefft . 1988 . the ceratioid family himantolophidae ( pisces , lophiiformes ) . steenstrupia , 14 ( 2 ) : 989 .
bertelsen , e . , and t . w . pietsch . 1975 . results of the research cruises of frv “walther herwig” to south america . xxxviii . osteology and relationships of the ceratioid anglerfish genus spiniphryne ( family oneirodidae ) . arch . fischwiss . , 26 ( 1 ) : 111 .
bertelsen , e . , and t . w . pietsch . 1977 . results of the research cruises of the frv “walther herwig” to south america . xlvii . ceratioid anglerfishes of the family oneirodidae collected by the frv “walther herwig . ” arch . fischwiss . , 27 ( 3 ) : 171189 .
bertelsen , e . , and t . w . pietsch . 1996 . revision of the ceratioid anglerfish genus lasiognathus ( lophiiformes : thaumatichthyidae ) , with the description of a new species . copeia , 1996 ( 2 ) : 401409 .
hansen , k . , and p . j . herring . 1977 . dual bioluminescent systems in the anglerfish genus linophryne ( pisces : ceratioidea ) . j . zool . , london , 182 : 103124 .
herring , p . j . 2000 . species abundance , sexual encounter and bioluminescent signaling in the deep sea . phil . trans . roy . soc . london , b , 355 : 12731276 .
herring p . j . 2002 . biology of the deep sea . oxford univ . press . 314 pp .
herring , p . j . , and o . munk . 1994 . the escal light gland of the deep - sea anglerfish haplophryne mollis ( pisces : ceratioidei ) , with observations on luminescence control . j . mar . biol . assn . u . k . , 74 : 747763 .
herring , p . j . , e . a . widder , and o . munk . 1994 . flashing anglerfish ; an unexpected signalling system . abstracts of the eighth deep sea biology symposium , monterey , california , 1997 : 52 .
hickling , c . f . 1925 . a new type of luminescence in fishes . j . mar . biol . assoc . u . k . , n . s . , 13 : 914937 .
kottelat , m . , and c . vidthayanon . 1993 . boraras micros , a new genus and species of minute freshwater fish from thailand ( teleostei : cyprinidae ) . ichthy . explor . freshwaters , 4 ( 2 ) : 161176 .
marshall , n . b . 1967a . the olfactory organs of bathypelagic fishes . symp . zool . soc . london , 19 : 5770 .
marshall , n . b . 1967b . the organization of deep - sea fishes . pp . 473479 , in : f . m . bayer , c . p . idyll , j . i . jones , f . f . koczy , a . a . myrberg , c . r . robins , f . g . w . smith , g . l . vos , e . j . f . wood , and a . c . jensen ( editors ) , proceedings of the international conference on tropical oceanography , 1724 november 1965 , miami beach , florida , studies in tropical oceanography , miami , vol . 5 .
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