anarta myrtilli , the beautiful yellow underwing , is a moth in the noctuidae family .

it is found in most of europe including scandinavia , britain , france , germany , switzerland , spain , portugal , italy , and russia .

the wingspan is 20 – 22 mm .

forewing dull dark fuscous purple , with the lines slightly paler ; the stigmata obscure ; a subtriangular whitish blotch on base of vein 2 ; hindwing orange with broad black terminal border ; the costa and inner margin narrowly black and the base of wing often smoky blackish ; this , the type form , occurring in sweden , the north of england and scotland , and other northern localities is very different from the usual bright red form , which is the ab . rufescens tutt .

in this the forewing is a mixture of bright red , and olive brown or olive yellow ; the transverse lines being more or less strongly whitish , the stigmata red brown with pale rings , and the white spot on vein 2 distinct ; ab . peralbata ab .

nov. [ warren ] is an extreme form of this , in which the white lines are strongly developed , : and the central area is milk white from costa to inner margin , including the white blotch on vein 2 ; in the hindwing the yellow is ampler , the black of the costal , and inner margins and the basal suffusion being reduced ; - ab.albivena haw. , described from east anglian specimens , has the forewing suffused with olive brown , wing remaining normal ; while in alpina ractzer not only is the forewing olive brown , but the hindwing shows only a dull yellowish median band crossed by black veins ; - and again in olivacea fuchs the yellow of the hindwing is suffused with olive brown , while the coloration of the forewing remains of the normal bright red ; in ab . nigrescens ab .

nov. [ warren ] occurring at hyeres , s.france , the usual red fusion is almost entirely replaced by black ; lastly , in subsp. citrina subsp. nov. [ warren ] ] from cintra , portugal , the whole forewing is suffused with blackish , leaving only the white blotch on vein 2 conspicuous , and the orange of the hindwing , both above and below , is pale lemon yellow ; as the insect is decidedly larger than average typical myrtilli , it may prove a distinct species ; at present i have seen only one - taken in the spring of 1909 by mr - n. c. rothschild , and now in the tring museum .

adults are on wing between may to august depending on the location .

there are probably two generations per year .

the larvae feed on calluna vulgaris and erica tetralix .

anarta myrtilli

kari pihlaviita added the finnish common name
kanervakangasyökkönen
to
anarta myrtilli linnaeus 1761
.
beautiful yellow underwing ( anarta myrtilli ) - norfolk moths - the macro and micro moths of norfolk .
hans - martin braun added the german common name
heidekrauteule
to
anarta myrtilli linnaeus 1761
.
hans - martin braun added the german common name
heidekrauteulchen
to
anarta myrtilli linnaeus 1761
.
hans - martin braun added the german common name
heidekraut - bunteule
to
anarta myrtilli linnaeus 1761
.
hans - martin braun added the german common name
heidekraut - bunteulchen
to
anarta myrtilli linnaeus 1761
.
habitat : anarta myrtilli inhabits sunny calluna locations as heath bogs , silicate calluna heaths in the mountains or sandy heaths in the plane .
endangerment factors : in the lowlands , for example in the upper rhine valley , anarta myrtilli has already lost almost all locations as a result of the intensification of agriculture and the increasingly rampant overbuilding by industry , transportation routes and settlements . in the mountains , anarta myrtilli is currently threatened even less , though here also constantly habitat area is lost by reforestation , ski runs and other tourist facilities . in the northern black forest anarta myrtilli could benefit from the storm areas in recent decades , but that is currently canceled again with their gradual overgrowth .
remarks : anarta myrtilli is quite widespread in europe as a whole , but lacks many landscapes completely ( limestone areas , etc . ) . in the south , it is still more local and is missing e . g . in southern italy or greece .
id : forewing ground colour reddish - brown or brownish - grey , marbled greyish - white ; white blotch at dorsal margin of oval ; hindwing yellow with a broad black termen and narrow black costal and dorsal margins . anarta cordigera ( small dark yellow underwing ) has a dark grey forewing ground colour and a conspicuous white - edged kidney mark .
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gaden s . robinson , phillip r . ackery , ian j . kitching , george w . beccaloni and luis m . hernández
records of caterpillar hostplants are scattered through published and manuscript sources worldwide and are difficult to retrieve . many rearing records are never published and so are not accessible to other entomologists . but collected hostplant records form a valuable scientific resource that can be used eventually to answer broader biological questions about how lepidoptera and plants interact ( eg , letourneau , hagen & robinson , 2001 ) . it provides information of immediate relevance to agriculture , ecology , forestry , conservation and taxonomy .
hosts brings together an enormous body of information on what the world ' s butterfly and moth ( lepidoptera ) caterpillars eat . the web - based version presented here offers a synoptic data set drawn from about 180 , 000 records comprising taxonomically ' cleaned ' hostplant data for about 22 , 000 lepidoptera species drawn from about 1600 published and manuscript sources . it is not ( and cannot be ) exhaustive , but it is probably the best and most comprehensive compilation of hostplant data available .
we hope that it will be useful to a wide range of biologists and that it will act as a spur to further recording and analysis of caterpillar - plant interactions .
hosts can be searched in two ways , using text search and drill - down search modes .
in text search mode , use either lepidoptera or hostplant criteria or a combination of the two . hosts operates only using scientific names . it is not possible to search for the hostplants of the red admiral butterfly but a search for hostplants using its scientific name , vanessa atalanta , will be successful . enter the generic name ( vanessa ) in the - lepidoptera criteria - genus box ; enter atalanta in the species : box ; click search .
hint : leave the ' starts with ' command as the default and in the species entry box omit the last few letters of the species - name ( eg , atalant ) . this will get around the problem of variable gender - endings . hosts uses original orthography of species - group names as far as possible , but some checklists follow the convention of altering the species - group name to accord with the presumed gender of the generic name ( eg flava , flavus ) .
restrict or refine searches using additional criteria ; choosing ' usa ' from the drop - down location box and entering [ starts with ] ' urt ' in the hostplant family box will return hostplant records of vanessa atalanta from urticaceae in the usa .
hint : restricting location may deliver a very incomplete subset of records : in the previous example all records specified as from the nearctic region ( usa + canada ) would be missed .
drill - down search mode allows the user to home in from the starting - point of either the lepidoptera or the plant family . choose a family group from the drop - down box and allow time for all genera of that family in the database to load to the genus drop - down box ; choose a genus and wait for the species to load . the search button can be pressed at any time , but the record delivery limit may be exceeded at higher taxonomic levels . drill - down search allows the user to see by scrolling all the taxa that are represented in the database . following the previous example , choose nymphalidae , then choose vanessa from the genus drop - down box and then atalanta from the ten possible species of vanessa included in hosts .
the preliminary search results pages give family , genus and species of the caterpillar and its hostplant . note that many hostplants are recorded as a plant genus only . clicking the caterpillar name will give the full record . the full record listing gives , additionally , the author of the lepidoptera species , subspecific information on the insect and the plant , if available , together with details of larval damage . laboratory rearings where the food utilised may not be the natural hostplant are indicated . the status of the record ( whether considered true , erroneous or suspect ) is not currently implemented in this version of hosts .
leguminosae ( c ) - caesalpinioideae . leguminosae ( m ) - mimosoideae . leguminosae ( p ) - papilionoideae .
all lepidoptera groups are covered and subspecific taxa are differentiated . the original source of the record , original form of the names used ( prior to taxonomic ' cleaning ' and standardisation of nomenclature ) and validation and verification fields are not included . this information is , however , retained in the databases used to generate this site . while we have included species that do not feed on green plants , the known food substrates of these are not listed exhaustively . such species comprise detritophages and predators and include , for example , most tineidae and the stored - products pests . the published compilations from hosts ( see more detail - publications from hosts ) include record status and the sources of all records .
detailed published compilations from hosts are available in press . these books give greater detail than the website , together with comprehensive cross - indexes , record status and full bibliographies . they are indispensable tools for naturalists and professional entomologists .
robinson , g . s . , ackery , p . r . , kitching , i . j . , beccaloni , g . w . & hernández , l . m . 2001 . hostplants of the moth and butterfly caterpillars of the oriental region . 744 pp .
robinson , g . s . , ackery , p . r . , kitching , i . j . , beccaloni , g . w . & hernández , l . m . 2002 . hostplants of the moth and butterfly caterpillars of america north of mexico . 824 pp . [ memoirs of the american entomological institute , volume 69 . ]
beccaloni , g . w . , viloria , a . l . , hall , s . k . & robinson , g . s . 2008 . catalogue of the hostplants of the neotropical butterflies / catálogo de las plantas huésped de las mariposas neotropicales . m3m - monografías tercer milenio , volume 8 . zaragoza , spain : sociedad entomológica aragonesa ( sea ) / red iberoamericana de biogeografía y entomología sistemática ( ribes ) / ciencia y tecnología para el desarrollo ( cyted ) / natural history museum , london , u . k . ( nhm ) / instituto venezolano de investigaciones científicas , venezuela ( ivic ) . 1 - 536 pp . , 1 fig , 3 tabs .
gaden robinson was responsible for the overall project design and management of the hosts database , and for records of lepidoptera exclusive of butterflies and bombycoid moths . phillip ackery and george beccaloni were responsible for butterfly data , including data drawn from card catalogues developed by ackery , whilst ian kitching was responsible for hostplant data of bombycoid moths . luis m . hernández was responsible for abstracting in the latter two years of the project and for development of the bibliography for the hardcopy versions of the data .
we are extremely grateful to the many people who contributed their own rearing records of lepidoptera or personal accumulations of data for inclusion in the hosts database , particularly mike bigger ( uk ) , john w . brown ( usa ) , chris conlan ( usa ) , rob ferber ( usa ) , konrad fiedler ( germany ) , jeremy holloway ( uk ) , frank hsu ( usa ) , jurie intachat ( malaysia ) , alec mcclay ( canada ) , bill palmer ( australia ) , pierre plauzoles ( usa ) and the generous individuals who contributed rearing records through the worldwideweb and who are known to us only as an email address .
we are particularly grateful to julian donahue and the los angeles county museum of natural history for allowing us to include data on microlepidoptera from the card catalogue prepared by the late j . a . comstock and c . henne , and for access to manuscript records by noel mcfarland .
marian fricano ( santa clara university ) and aileen giovanello ( clark university , international internship 1996 ) made substantial contributions of abstracted data ; fran love ( north carolina ) painstakingly checked scanned texts and reformatted them for import to paradox .
much of the immense task of data abstracting and entry from printed and manuscript sources as well as preliminary editing and name - checking was carried out by volunteers . many of these were school students on work - experience placements during 1993 - 2000 from , initially , the coopers ' company and coborn school , upminster , and later from other schools in greater london : christopher andrewes , simon bennett ( 1994 nhm vacation studentship ) , steven bond , michael brownlow , emma causer , laurence cooper , ailsa cranfield ( 1998 nuffield studentship ) , emily dwiar , andrew enever , jane feehan , madeleine ferry , max friedman , edward gold , jennifer hodgkinson , christopher joint , fateha khatun ( 1996 nuffield studentship ) , james lowe , louisa marchant , gemma millward , christopher milne , carolyn oughton , william perkins , rebecca reith , eleanor resheph , clare sambidge , neil shaftain , stephen sloan , helen stevens , samuel tarry , david taylor and thomas yeatman .
we are indebted to all our helpers for their diligence , accuracy and patience , and for their unswerving faith that this daunting project would reach a conclusion .
our colleagues here and overseas provided substantial help , advice and assistance with the checking of names of lepidoptera and with many other aspects of this project : kim and david goodger and jeremy holloway ( macrolepidoptera families ) , martin honey ( noctuoidea ) , brian pitkin ( computing ) , malcolm scoble and linda pitkin ( geometroidea ) , klaus sattler ( gelechioidea ) , michael shaffer ( pyraloidea , thyridoidea , pterophoroidea ) , alma solis ( usda , washington - pyraloidea ) , fernley symons ( oxford university - technical support ) and kevin tuck ( tortricoidea ) . julie harvey and the staff of the bmnh entomology and general libraries provided sterling support in locating obscure source material and intuitively correcting bowdlerized references .
we thank the trustees of the loke wan tho memorial foundation for their generous support for this project .
robinson , g . s . , p . r . ackery , i . j . kitching , g . w . beccaloni & l . m . hernández , 2010 . hosts - a database of the world ' s lepidopteran hostplants . natural history museum , london . urltoken . ( accessed : 18 aug . 2010 ) .
brummitt , r . k . 1992 . vascular plant families and genera . [ vi ] + 804 pp . , royal botanic gardens , kew .
kartesz , j . t . 1994 . a synonymized checklist of the vascular flora of the unites states , canada and greenland . timber press , portland . 1 , checklist , lxi + 622 pp ; 2 , thesaurus , vii + 816 pp .
kitching , i . j . & cadiou , j . - m . 2000 . hawkmoths of the world : an annotated and illustrated revsionary checklist . xx + 500 pp . , cornell university press , ithaca .
kitching , i . j . & rawlins , j . e . 1999 . the noctuoidea . pp . 355 - 401 . in : kristensen , n . p . ( ed . ) lepidoptera , moths and butterflies . 1 . evolution , systematics and biogeography . handbook of zoology , 4 ( 35 ) . lepidoptera . x + 491 pp . de gruyter , berlin .
letourneau , d . k . , hagen , j . a . & robinson , g . s . 2001 . bt crops : evaluating benefits under cultivation and risks from escaped transgenes in the wild . pp . 33 - 98 . in : letourneau , d . k . & burrows , b . e . ( eds ) , genetically engineered organisms : assessing environmental and human health effects . [ viii ] + 438 pp . , crc press , boca raton .
mabberley , d . j . 1987 . the plant book . a portable dictionary of the higher plants . xii + 707 pp . , cambridge university press . [ the 1993 reprint edition is currently used in editing . ]
nielsen , e . s . , edwards , e . d . & rangsi , t . v . ( eds ) 1996 . checklist of the lepidoptera of australia . monographs on australian lepidoptera . 4 . xiv + 529 pp . , csiro , melbourne .
nye , i . w . b . ( ed . ) 1975 - 91 . the generic names of moths of the world . 1 : 568 pp . ( noctuoidea ( part ) - nye , i . w . b . , 1975 ) ; 2 : xiv + 228 pp . ( noctuoidea ( part ) - watson , a . , fletcher , d . s . & nye , i . w . b . , 1980 ) ; 3 : xx + 243 pp . ( geometroidea - fletcher , d . s . , 1979 ) ; 4 : xiv + 192 pp . ( bombycoidea to zygaenoidea - fletcher , d . s . & nye , i . w . b . , 1982 ) ; 5 : xv + 185 pp . ( pyraloidea - fletcher , d . s . & nye , 1984 ) ; 6 : xxix + 368 pp . ( microlepidoptera - nye , i . w . b . & fletcher , d . s . , 1991 ) . british museum ( natural history ) / the natural history museum , london .
rawlins , j . e . 1984 . mycophagy in lepidoptera . pp . 382 - 483 . in : wheeler , q . & blackwell , m . ( eds ) fungus - insect relationships . perspectives in ecology and evolution . 514 pp . , columbia university press .
robinson , g . s . 1999 . hosts - a database of the hostplants of the world ' s lepidoptera . nota lepidopterologica , 22 : 35 - 47 .
robinson , g . s . , ackery , p . r . , kitching , i . j . , beccaloni , g . w . & hernández , l . m . 2001 . hostplants of the moth and butterfly caterpillars of the oriental region . 744 pp . southdene sdn bhd , kuala lumpur .
robinson , g . s . , ackery , p . r . , kitching , i . j . , beccaloni , g . w . & hernández , l . m . 2002 . hostplants of the moth and butterfly caterpillars of america north of mexico . memoirs of the american entomological institute , 69 : 1 - 824 .
scoble , m . j . ( ed . ) 1999 . a taxonomic catalogue to the geometridae of the world ( insecta : lepidoptera ) . 2 vols . csiro publications , melbourne .
royal botanic garden edinburgh dipterocarpaceae database : http : / / 193 . 62 . 154 . 38 / diptero /
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a quite small species , with a wingspan of only around 25mm , this is a diurnal moth , flying freely in warm sunshine , but tending to rest when the sun goes in .
frequenting acid moorland , there are probably two generations in the south , with moths on the wing from may to august . in the north it flies in june and july .
it has a scattered distribution over most of britain , and the larvae feed mainly on heather ( calluna ) .
ukmoths is built , run and maintained by ian kimber , with thanks to the many kind contributors who provide photos and information .
the ukmoths facebook page is a great place to post your identification queries . more often than not you ' ll get a positive id on most photos fairly quickly .
looking for a specific moth species ? enter just part of the name below .
procache : v317 render date : 2018 - 06 - 15 12 : 26 : 11 page render time : 0 . 2731s total w / procache : 0 . 3354s
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life cycle : the moths fly in one or ( usually ) two generations from april to early september . the caterpillar lives from the summer to october also open on the plant during the day , but well camouflaged . the pupa overwinters .
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fw : 10 - 12mm ; apr - aug in south , jun - jul in north , diurnal ; common heather ( calluna vulgaris ) , bell heather ( erica cinerea ) ; common in heath and moor throughout gb .
§1 new forest , hampshire ; 31 / 08 / 2013 ; male ; fw 10 . 6mm §2 duddon moss , cumbria ; 22 / 07 / 2016 ; female ; fw 10 . 0mm ; netted by day all images © chris lewis
recorded in 18 ( 26 % ) of 69 10k squares . first recorded in 1873 . last recorded in 2018 .
underlying maps using digital map data © norfolk online lepidoptera archive - nola™ 2018 . © james wheeler - n o r f o l k m o t h s 2007 - 2018 . data © nola™ 2018


the habitat consists of exclaves of open steppe vegetation on sands or loess and xerophilous grasslands on alkaline substrates .

the larvae bore the roots of artemisia species .

paracossulus thrips

paracossulus thrips - urdu meaning and translation of paracossulus thrips , translation , multiple word search ( seperate words with space ) , english to urdu machine translation of paracossulus thrips and more .
what is there to think - at sineva by yakovlev paracossulus thrips . correct necessary .
paracossulus thrips is a species of moth of the cossidae family . it is found in hungary , romania , ukraine , russia , kazakhstan and turkey .
enter the name or part of a name you wish to search for . the asterisk character * can be used as wildcard . e . g . ' papilio * ' . keep in mind that the search is only based on the full taxon name .
we are still having problems with the search feature . unfortunately we cannot give a timeline when the advanced search will be fixed .
the id resolving service for stable taxon ids is currently under maintenance . meanwhile , please use the name search in order to find the taxon page .
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regions of the russian federation : the volga - don , east caucasus , the european central black earth , central european , western caucasus , krasnoyarsk , lower volga , prealtay , mid - volzhsky , south west siberian , south ural .
[ 10 ] de jong , y . s . d . m . ( ed . ) ( 2011 ) fauna europaea version 2 . 4 ( faunaeur . org )
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also common in the southern urals , in the south of western siberia , kazakhstan and altai .
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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 . .
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urn : lsid : biodiversity . org . au : afd . taxon : 11b67b9b - 1b16 - 46ec - 9863 - d09e2c2b2879
urn : lsid : biodiversity . org . au : afd . taxon : 1a8c0252 - a2e2 - 4e88 - 8c67 - 7d55b13c8247
urn : lsid : biodiversity . org . au : afd . taxon : 1eb21109 - 3907 - 4ac8 - b4ef - 06173f5907f2
urn : lsid : biodiversity . org . au : afd . taxon : 3ca2d5db - 1b99 - 493d - 9c65 - a3958b4f981c
urn : lsid : biodiversity . org . au : afd . taxon : 53e9bf1d - b31c - 48ba - a013 - 1f54326038d5
urn : lsid : biodiversity . org . au : afd . taxon : a3534fc1 - 8af9 - 40b8 - acb4 - 9e5e3015d538
urn : lsid : biodiversity . org . au : afd . taxon : e7715a02 - cad8 - 4407 - 8933 - abf95d8bd0b3
urn : lsid : biodiversity . org . au : afd . taxon : f4c194ce - 984c - 4def - af0b - 7db8a4a30db1
urn : lsid : biodiversity . org . au : afd . taxon : f7cb0cb7 - 5a6f - 4b7c - 8fe8 - 5096f8bf972d
urn : lsid : biodiversity . org . au : afd . name : 244772
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pseudosimnia pyrulina is a species of sea snail , a marine gastropod mollusk in the family ovulidae , the ovulids , cowry allies or false cowries .

pseudosimnia pyrulina

species margovula pyrulina ( a . adams , 1854 ) accepted as pseudosimnia pyrulina ( a . adams , 1855 )
worms - world register of marine species - pseudosimnia pyrulina ( a . adams , 1855 )
( of margovula pyrulina ( a . adams , 1854 ) ) cate c . n . ( 1978 ) . new species of ovulidae and reinstatement of margovula pyrulina ( a . adams , 1854 ) . the nautilus 92 ( 4 ) : 160 - 167 [ details ]
species pseudosimnia pyrifera c . n . cate , 1973 accepted as pseudosimnia vanhyningi ( m . smith , 1940 )
species pseudosimnia sphoni c . n . cate , 1973 accepted as pseudosimnia vanhyningi ( m . smith , 1940 )
to barcode of life to biodiversity heritage library ( 3 publications ) ( from synonym margovula pyrulina ( a . adams , 1854 ) ) to encyclopedia of life
no one has contributed data records for pseudosimnia lacrima yet . learn how to contribute .
( of amphiperas pyrulina a . adams , 1855 ) lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
pseudosimnia albomarginata cate , c . n . , 1978 : solomons ; japan ( ? )
species pseudosimnia filia azuma , 1974 accepted as primovula rosewateri ( c . n . cate , 1973 )
species pseudosimnia whitworthi c . n . cate , 1973 accepted as diminovula whitworthi c . n . cate , 1973
species pseudosimnia coroniola c . n . cate , 1973 accepted as diminovula coroniola ( c . n . cate , 1973 )
species pseudosimnia culmen c . n . cate , 1973 accepted as diminovula culmen ( c . n . cate , 1973 )
species pseudosimnia emilyreidae c . n . cate , 1973 accepted as diminovula kosugei ( c . n . cate , 1973 )
species pseudosimnia perilla c . n . cate , 1973 accepted as diminovula dautzenbergi ( f . a . schilder , 1931 )
species pseudosimnia striola c . n . cate , 1973 accepted as diminovula dautzenbergi ( f . a . schilder , 1931 )
species pseudosimnia translineata c . n . cate , 1973 accepted as margovula translineata ( c . n . cate , 1973 )
species pseudosimnia carnea sensu f . a . schilder , 1941 not poiret , 1789 accepted as sandalia triticea ( lamarck , 1810 )
( of amphiperas pyrulina a . adams , 1855 ) adams a . ( 1855 [
1854
] ) . descriptions of thirty - nine new species of shells , from the collection of hugh cuming , esq . proceedings of the zoological society of london . 22 : 130 - 138 , pl . 28 . , available online at urltoken [ details ]
species pseudosimnia incisa azuma & c . n . cate , 1971 accepted as diminovula incisa azuma & c . n . cate , 1971
species pseudosimnia marginata g . b . sowerby i , 1828 accepted as margovula marginata ( g . b . sowerby i , 1828 )
species pseudosimnia pyriformis sensu kuroda , 1958 not g . b . sowerby i , 1828 accepted as diminovula alabaster ( reeve , 1865 )
hans - martin braun added the english common name
dwarf red simnia
to
pseudosimnia carnea ( poiret , 1789 )
.
» species pseudosimnia ( diminovula ) fruticum ( reeve , 1865 ) accepted as prionovolva brevis ( g . b . sowerby i , 1828 )
hans - martin braun added the english common name
dwarf red egg cowrie
to
pseudosimnia carnea ( poiret , 1789 )
.
hans - martin braun added the english common name
blood - stained simnia
to
pseudosimnia carnea ( poiret , 1789 )
.
species pseudosimnia fulguris azuma & c . n . cate , 1971 accepted as primovula fulguris ( azuma & c . n . cate , 1971 )
species pseudosimnia kandai c . n . cate & azuma in c . n . cate , 1973 accepted as testudovolva nipponensis ( pilsbry , 1913 )
species pseudosimnia nubila c . n . cate & azuma in c . n . cate , 1973 accepted as diminovula whitworthi c . n . cate , 1973
species pseudosimnia pyriformis sensu f . a . schilder , 1941 not g . b . sowerby i , 1828 accepted as diminovula alabaster ( reeve , 1865 )
species pseudosimnia aurantiomacula c . n . cate & azuma , 1973 accepted as diminovula aurantiomacula ( c . n . cate & azuma , 1973 ) ( original combination )
( of aperiovula aurora omi , 2003 ) omi y . 2003 . a nnew species of aperiovula ( gastropoda : ovulidae ) collected near the boso peninsula , japan . venus 62 ( 1 - 2 ) : 11 - 18 [ details ]
lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
( of aperiovula aurora omi , 2003 ) lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
( of aperiovula albomarginata c . n . cate , 1978 ) cate c . n . ( 1973 ) . a systematic revision of the recent cypraeid family ovulidae . the veliger . 15 ( supplement ) : 1 - 117 . , available online at urltoken [ details ]
( of aperiovula albomarginata c . n . cate , 1978 ) lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
( of aperiovula yukitai azuma , 1974 ) cate c . n . ( 1973 ) . a systematic revision of the recent cypraeid family ovulidae . the veliger . 15 ( supplement ) : 1 - 117 . , available online at urltoken [ details ]
( of aperiovula yukitai azuma , 1974 ) lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
( of prionovolva aureomarginata shikama , 1973 ) cate c . n . ( 1973 ) . a systematic revision of the recent cypraeid family ovulidae . the veliger . 15 ( supplement ) : 1 - 117 . , available online at urltoken [ details ]
( of prionovolva aureomarginata shikama , 1973 ) lorenz f . & fehse d . ( 2009 ) the living ovulidae . a manual of the families of allied cowries : ovulidae , pediculariidae and eocypraeidae . hackenheim : conchbooks . [ details ]
( of prionovolva aureomarginata shikama , 1973 ) liu j . y . [ ruiyu ] ( ed . ) . ( 2008 ) . checklist of marine biota of china seas . china science press . 1267 pp . ( look up in imis ) [ details ] available for editors [ request ]
( of aperiovula c . n . cate , 1973 ) cate c . n . ( 1973 ) . a systematic revision of the recent cypraeid family ovulidae . the veliger . 15 ( supplement ) : 1 - 117 . , available online at urltoken page ( s ) : 36 [ details ]
gofas , s . ; le renard , j . ; bouchet , p . ( 2001 ) . mollusca . in : costello , m . j . et al . ( eds ) , european register of marine species : a check - list of the marine species in europe and a bibliography of guides to their identification . patrimoines naturels . 50 : 180 - 213 . ( look up in imis ) [ details ]
worms - world register of marine species - margovula c . n . cate , 1973
species margovula aboriginea c . n . cate , 1973 accepted as diminovula aboriginea ( c . n . cate , 1973 )
species margovula changi ma , 1997 accepted as diminovula margarita ( g . b . sowerby i , 1828 )
species margovula kosugei c . n . cate , 1973 accepted as diminovula kosugei ( c . n . cate , 1973 )
iredale t . ( 1935 ) . australian cowries . the australian zoologist . 8 ( 2 ) : 96 - 135 , pls 8 - 9 . , available online at urltoken page ( s ) : 103 ; note : not available : no description [ details ]
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osedax mucofloris is a species of bathypelagic polychaetes that is reported to sustain itself on the bones of dead whales .

translated from the mixed greek and latin used in scientific names , " osedax mucofloris " literally means " snot-flower bone-eater " , though the less-accurate " bone-eating snot-flower worm " seems to be the form actually used .

the species is found in north east atlantic where it is abundant .

osedax mucofloris

olingo added text to
bone - eating snot - flower worm ( osedax mucofloris )
on
osedax mucofloris glover , kallstrom , smith and dahlgren , 2005
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osedax mucofloris extends out of the whale bone into the seawater . the major features of this portion are :
high symbiont diversity in the bone - eating worm osedax mucofloris from shallow whale - falls in the north atlantic .
based on available data ( which are clearly very limited ) , osedax mucofloris is associated with shallower depths than other known osedax species , having been collected at 30 and 125 meters .
a detailed description of the morphology of osedax mucofloris has been published in the scientific journal proceedings of the royal society of london .
high symbiont diversity in the bone - eating worm osedax mucofloris from shallow whale - falls in the north atlantic . - pubmed - ncbi
molecular analysis of osedax mucofloris using coi and 18s rrna sequences confirmed genetic seperation from morphologically similar o . rubiplumus and o . frankpressi ( glover et al . , 2005 ) .
many osedax species have numerous pinnules on their palps that give the crown a feathery appearance . of the known osedax species , only o . mucofloris and o . frankpressi have pinnules oriented all inward ( the pinnules in the other species being turned outward , both inward and outward , or absent ) . most known osedax species have red palps ( the palps of o . frankpressi are red with two white lateral stripes in living worms ) , but the palps of osedax mucofloris are white to pink . ( rouse et al . 2004 ; vrijenhoek et al . 2009 )
osedax worms are marine annelids closely related to deep sea vent / seep - associated . . .
citation : huusgaard rs , vismann b , kühl m , macnaugton m , colmander v , rouse gw , et al . ( 2012 ) the potent respiratory system of osedax mucofloris ( siboglinidae , annelida ) - a prerequisite for the origin of bone - eating osedax ? plos one 7 ( 4 ) : e35975 . urltoken
schander , c . , rapp , h . t . , & dahlgren , t . ( 2010 ) . osedax mucofloris ( polychaeta , siboglinidae ) , a bone - eating marine worm new to norway . fauna norvegica , 30 , 5 - 8 . urltoken
detailed examination of at least 50 mature specimens indicated an absence of the dwarfed males reported for northeast pacific osedax species . however , o . mucofloris appeared able to reproduce and grow to maturity within one month on defaunated bones placed into aquaria . ( glover et al . 2005 )
the bone - eating siboglinid polychaete osedax mucofloris glover , källström , smith & dahlgren , 2005 is reported from norwegian waters for the first time . dense growth was found on bovine bones deposited at 118 meters depth off western norway . dwarf males were observed for the first time . the two specimens sequenced were identical to haplotypes previously found at a swedish whale fall . the possibility of finding additional species of osedax is discussed .
as with the trunk of o . mucofloris , the anterior vestimentum of vestimentifera lacks branching vessels between the ventral and the muscularized dorsal vessel [ 33 ] . the resemblance between the blood vascular systems adds new evidence to the hypothesis that the trunk of osedax and the vestimentum of vestimentifera are homologous regions [ 21 ] .
the measured o 2 depleted bone environment as well as ferrous sulphide precipitations ( figure 8e ) supports the suggested exposure of osedax to sulphide . furthermore the large root surface in the related lamellibrachia is highly efficient in hydrogen sulphide uptake [ 17 ] . since the main trophic source of osedax compared to other siboglinids ( including lamellibrachia ) seems to be heterotrophic bacteria rather than sulphide - detoxifying chemoautotrophic bacteria [ 8 ] , osedax may instead possess physiological adaptations to detoxify sulphide .
in the present study we investigated whether osedax mucofloris is exposed to an inhospitable hypoxic or anoxic microenvironment by measuring the o 2 distribution surrounding the embedded root system . we assessed possible morphological and / or physiological adaptations to the environmental conditions through detailed morphological studies of the respiratory surfaces and the blood vascular system as well as respiratory measurements of o 2 consumption . the results are discussed in relation to the unique environment , endosymbionts , and embedded root structure of osedax as well as compared with studies of related annelids .
it is important to understand osedax mucofloris reproduction , as scientists want to know how these animals are able to disperse between isolated whale - fall habitats . when the first species of osedax was described , it was noted that the species seemed to exhibit very strong sexual dimorphism . the males did not develop into mature adults . instead , they appeared to remain as larval - sized individuals attached to the side of females , fertilising eggs as they were released into the water column . dwarf males have been described for :
osedax mucifloris appeared to be able to reproduce and grow to maturity within one month on defaunated bones placed in a aquaria ( glover et al . , 2005 ) .
to date , osedax mucofloris has only been recorded from 2 whale carcasses in the swedish fjord kosterfjord ( 58° 53 . 1 n , 11° 06 . 4 e ) . it may be widespread in the atlantic , however . the problem is finding whale remains on the sea floor - even if there are a lot of them , they are small relative to the vast expanses of the ocean .
our data show that o . mucofloris inhabiting bone in cuvettes ( purple and yellow dots , figure 9 ) have a lower mo 2 than o . mucofloris inhabiting sectioned cow bone ( green , blue , red dots , figure 9 ) . along with high standard deviations , this highlights the difficulties of measuring o 2 consumption on these embedded worms . the variations are most likely caused by the difference in blind respiration measurements and the large biological activity present on decaying bone , which was difficult to quantify . future studies should thus be carried out to more precisely determine the mo 2 of o . mucofloris and the contribution of other o 2 - consuming surfaces .
the circular trunk musculature in o . mucofloris is weakly developed , as found in several other annelids [ 26 ] , [ 27 ] . the diagonal musculature revealed in the present study , may compensate for the weak circular musculature by having a similar supportive function . diagonal musculature has most likely been mistaken for circular musculature in several annelids [ 26 ] , [ 27 ] , including previous studies of osedax [ 15 ] , [ 21 ] .
beggiatoa live in the restricted interface between hydrogen sulphide presence and oxygenated water [ 51 ] . o . mucofloris must therefore be in contact with toxic sulphide concentrations . photographer : helena wiklund , department of zoology , göteborg university , sweden .
the shallow water species osedax mucofloris glover , källström , smith , dahlgren , 2005 , has been found at 30 m and 125 m water depth off the coast of tjärnö , sweden [ 12 ] , [ 13 ] , and at 120 m depth in bjørnafjord , norway [ 24 ] . currently these are the only records of osedax from the atlantic , though other species are found in shallow waters in both the east and west pacific [ 4 ] , [ 25 ] . the 125 m deep locality near tjärnö is characterized by stable oceanic salinity ( 34–35‰ ) , temperature ( 5–7°c ) and dissolved o 2 concentration ranging from 4 . 6 to 6 . 3 ml o 2 l −1 [ 13 ] .
previous studies of other close relatives of osedax such as vestimentifera and sabellidae have shown anterior branchial structures to be of vital importance when tube - bound tissue is not ventilated [ 29 ] – [ 31 ] . likewise , the anterior branchial structures of o . mucofloris seem morphologically adapted to facilitate efficient o 2 uptake form the surrounding water . as with the branchial plume of vestimentifera [ 31 ] , [ 32 ] , the anterior palps and pinnules of o . mucofloris have a large surface area to volume ratio and short diffusion distances . furthermore , o 2 uptake is optimized by the two ventilating ciliary bands on each palp , a feature also seen in other siboglinidae and in sabellida in general [ 20 ] , [ 31 ] , [ 33 ] .
o . mucofloris has only recently been discovered and it is not yet known if it is under threat . one possibility is that intensive whaling over the last 300 years has reduced the available habitat for this polychaete worm , a previously unknown consequence of the whaling industry .
the quantity and complexity of capillaries in the root structure of o . mucofloris reflects the high o 2 demand of osedax , presumably for the metabolism of its heterotrophic endosymbionts and production and development of eggs . similar capillaries are visible on the exposed ovisac of o . frankpressi ( figure 2f in [ 1 ] ) and o . roseus ( figure 4e in [ 21 ] ) , while the present study shows the presence of capillaries supplying the more distally placed root tissue and bacteriocytes . however , the extent of capillaries does not match the extensive capillary network of the trophosome in other siboglinidae [ 20 ] , [ 33 ] . this is in accordance with the original description [ 1 ] mentioning the lack of a discrete trophosome , the osedax trophosome instead being diffuse and beneath the epidermis of the embedded tissue .
when corrected for background respiration , the measured o 2 consumption ( mo 2 ) of o . mucofloris ranged almost across a factor of ten from 220±93 µg o 2 g −1 h −1 to 2053±1950 µg o 2 g −1 h −1 ( table 1 ) , depending on the approach used for measurement . mo 2 measured on o . mucofloris inhabiting sectioned bone pieces ( b1–b3 ) vs . mo 2 measured o . mucofloris inhabiting bones in cuvettes ( c1 , c2 ) , resulted in two distinctly different ranges of mo 2 . the mo 2 of c1 and c2 was 220±93 µg o 2 g −1 h −1 and 238±29 µg o 2 g −1 h −1 , respectively , while the mo 2 of b1–b3 range from 976±201 µg o 2 g −1 h −1 to 2053±1950 µg o 2 g −1 h −1 . methods and ranges are commented further in the discussion .
g . w . rouse , s . k . goffredi , and r . c . vrijenhoek ( 2004 ) .
osedax : bone - eating marine worms with dwarf males
. science 305 : 668–671 . doi : 10 . 1126 / science . 1098650 . pmid 15286372 .
specimens were carefully dissected from the bone and fixed for immunohistochemistry , benzidine staining and histology . prior to dissection , osedax mucofloris were anesthetized for 5–10 minutes in a 1∶1 solution of seawater and mgcl 2 ( isotonic to seawater ) . anaesthetized animals were gently dissected with scalpels and fixed at 4°c over night in 4 % paraformaldehyde in 0 . 15 m phosphate - buffered saline ( pbs ) with 5 % sucrose , ph 7 . 4 . subsequently , the animals were rinsed 4–6 times for 30 min in pbs with 5 % sucrose and were then stored at 4°c in pbs with 0 . 05 % sodium azide ( nan 3 ) .
an extended root surface ( and high area to volume ratio ) is likewise found in both lamellibrachia ( e . g . , [ 16 ] ) and osedax ( yet branched ) . however , the main function has so far been interpreted as very different . whereas the osedax root surface is suggested to mainly facilitate uptake of organic compounds ( with less focus on the possible congruent uptake of sulphide ) [ 9 ] , the root of lamellibrachia is found to be the main respiratory surface of hydrogen sulphide necessary for the chemosynthesis of the symbionts [ 16 ] , [ 17 ] , [ 28 ] .
the estimated diffusion distance in o . mucofloris ( pinnule epidermis 1–2 µm , palp epidermis ∼30 µm ) is furthermore comparable to what has been found for r . pachyptila ( pinnules ∼2 . 00 µm ; branchial filaments ∼25 µm ) and ridgeia piscesae ( pinnules ∼1 . 00 µm ; branchial filaments ∼17 µm ) [ 31 ] , [ 32 ] .
world - wide whale worms ? a new species of osedax from the shallow north atlantic
. proc . biol . sci . ( national center for biotechnology information ) 272 ( 1581 ) : 2587–92 . 22 december 2005 . doi : 10 . 1098 / rspb . 2005 . 3275 . pmc 1559975 . pmid 16321780 .
though beyond the scope of the present paper , microanalytical approaches such as microsensors [ 40 ] , [ 41 ] and functional imaging techniques [ 42 ] could yield a more complete mapping of the chemical microenvironment of osedax , including the exact levels of sulphide exposure and spatio - temporal dynamics of o 2 in the root system . it would also be interesting to know whether osedax have sulphide - binding properties of their haemoglobin as found in vestimentifera [ 43 ] , [ 44 ] , as they might use them to transport toxic sulphides from the area surrounding the root structure to e . g . , the palps , somehow releasing the toxic compounds to oxygenated seawater or detoxifying them .
two nerves , originating at the anterior part of the brain , innervate each palp . one nerve runs abfrontally , between the two lateral ciliary bands and the other nerve runs laterally underneath one of the ciliary bands ( figure 5a ) . further details on the female osedax nervous system will be described elsewhere ( worsaae & rouse , unpublished ) .
the unique bone - eating organism , osedax ( siboglinidae , annelida ) was first described in 2004 from a whale fall located at 2891 m depth in monterey bay , pacific ocean [ 1 ] . since its first discovery it has been found on multiple whale falls in the pacific and atlantic oceans , artificially deployed cow bones [ 2 ] as well as on other vertebrate bones such as those of teleost [ 3 ] . there are five formally described species , with at least a further 12 species known from genetic evidence [ 4 ] , [ 5 ] . in addition , convincing fossil traces of osedax have been found in oligocene and pliocene mammal bones [ 6 ] , [ 7 ] .
osedax worms are marine annelids closely related to the deep sea vent / seep - associated vestimentiferan worms . the sessile ( i . e . , fixed in one place ) females bore into the bones of whale carcasses - - and possibly bones of other vertebrates ( rouse et al . 2004 ; glover et al . 2005 ; vrijenhoek et al . 2008 ) .
we do not yet know the full answer . one theory is that it overcomes the problem of males and females finding each other if they are unable to move ( sessile ) , as osedax females are . the males are able to move around the females , fertilising their eggs . it is easier to do this if you are small and motile , or dwarfed .
micro sensor measurements of o 2 concentrations in proximity to the bone interface were conducted , through agar - filled holes , on osedax - colonized bone fragments in cuvettes . the obtained profiles of o 2 concentration from the aerated seawater , and inwards showed steep o 2 gradients towards both the bone and tissue surface ( figure 8a , b ) . anoxic conditions or very low o 2 levels were found at the bone surface , within the bone , and in proximity to the embedded root tissue of o . mucofloris ( table 2 ) . the average o 2 flux at the bone and tissue interface was 0 . 028±0 . 0024 nmol o 2 cm −2 s −1 ( n = 5 ) and 0 . 029±0 . 0040 nmol o 2 cm −2 s −1 ( n = 3 ) , respectively .
glover , a . g . ; kallstrom , b . ; smith , c . r . ; dahlgren , t . g . 2005 . world - wide whale worms ? a new species of osedax from the shallow north atlantic . proceedings of the royal society b - biological sciences 272 ( 1581 ) : 2587 - 2592 page ( s ) : 2 589 [ details ]
o 2 consumption was measured on o . mucofloris inhabiting three sectioned cow bone pieces ( b1 , b2 and b3 ) and bones in two cuvettes ( c1 and c2 ) . measurements were either initiated directly after the annelids protracted subsequent to the disturbance of being moved ( c1 , c2 , b1 ) or after one night of acclimatization ( b2 , b3 ) . all measurements were corrected for background respiration . for b2 and b3 the background respiration was measured using the same water and bones ( after dissection and 48 hours of acclimation to restore biological activity ) . for c1 , c2 and b1 the background respiration was the mean value of measurements using new water , bones in two cuvettes and three sectioned bone pieces without o . mucofloris . different respiration chambers were used for sliced bone pieces ( volume : 186 ml ) and cuvettes ( volume : 51 ml ) .
the present study shows that osedax mucofloris has a higher weight specific o 2 consumption ( mo 2 ) than other resting annelids ( figure 9 ) [ 18 ] . this may reflect an elevated demand of the embedded tissue due to presence of heterotrophic aerobic endosymbionts , which have a higher metabolism than regular tissue . the measured mo 2 actually corresponds to that found for riftia pachyptila [ 22 ] , [ 23 ] ( figure 9 ) , possessing a vast amount of chemoautotrophic bacteria in their trophosome . furthermore , a high mo 2 may also reflect oxidative sulphide detoxification . the high mo 2 corresponds well with the large sbsa and elaborate branchial structures , which are both usually associated with animals exhibiting a high o 2 demand . this correlation is also found in arenicola marina with smaller sbsa and lower mo 2 ( red square , figure 9 ) .
a : sketch of the path of the longitudinal trunk vessels into the root structure , drawn from the light microscope with a camera lucida of a benzidine stained o . mucofloris female . trunk twisted in midsection . b : dic light micrograph of a benzidine stained o . mucofloris female . lateral view , trunk twisted in midsection . ventral and dorsal blood vessels continues , folded , into the anterior part of the ovisac / root system . c : close up of blood vessels near ovisac . d : close up of blood vessels supplying more distally placed capillaries . e : capillaries supplying tissue and endosymbionts . abbreviations : blood traces ( b ) , blood vessel ( bv ) , capillaries ( cap ) , dorsal blood vessel ( dbv ) , egg cluster ( ec ) , oviduct ( od ) , ovisac ( os ) , palp ( p ) , root structure ( r ) , trunk ( t ) , ventral blood vessel ( vbv ) .
glover , a . g ; kallstrom , b . ; smith , c . r ; dahlgren , t . g ( 2005 ) .
world - wide whale worms ? a new species of osedax from the shallow north atlantic
. proceedings of the royal society b : biological sciences 272 ( 1581 ) : 2587–2592 . doi : 10 . 1098 / rspb . 2005 . 3275 . issn 0962 - 8452 . pmc 1559975 . pmid 16321780 .
the trunk of osedax mucofloris encloses two major longitudinal blood vessels ( figures 2e , 6a , b ) . the dorsal vessel is highly muscularized with circular musculature throughout its length ( figure 2d , e ) . no lateral connecting vessels between the major longitudinal vessels or epidermal capillaries were found in the trunk . from the trunk , the two blood vessels continue posteriorly into the root structure and increase in diameter ( figure 6a , b ) . when studied under the light microscope , the muscularized dorsal vessel is visible as a defined tube along the trunk , and continues into the anterior root structure , curling up in the centre anterior to the ovisac . the curling configuration is most likely caused by contraction of the basal part of the trunk . the exact further path of the vessel was difficult to determine . the confocal laser scanning microscope ( clsm ) studies also showed the continuation of the dorsal blood vessel around the ovisac , revealing cylindrical musculature at the ovisac , corresponding in diameter to the musculature of the dorsal blood vessel of the trunk ( figure 4b ) .
a : osedax mucofloris extend its palps and pinnules , with large respiratory surfaces , into the overlying o 2 - rich water in order to uptake o 2 . o 2 is then distributed to the buried root system through the extensive blood vascular system also supplying the heterotrophic endosymbionts . the o 2 distribution to the root system is crucial as local uptake is not possible in the anoxic bone environment . the anoxic environment is partly produced by intense bacterial processes ( green arrows ) utilizing o 2 at the bone surface . hydrogen sulphide is produced by anoxic bacterial processes within the bone matrix during decomposition of organic content using sulphate . b : schematic illustration of assumed blood flow in palp and pinnules , longitudinal section . blue vessels carrying venous blood through afferent vessels , red vessels carrying arterial blood through efferent vessels . c : schematic illustration of assumed blood flow in palp and pinnules , transverse section . likewise blue vessels carries venous blood through afferent vessels , red vessels carries arterial blood through efferent vessels . note that the palp blood vessels are created by an invagination of the basement membrane . green indicates musculature .
a rough estimate was made of the surface area of the anterior crown , calculated from palp length given by glover et al . [ 12 ] , and the dimensions of palps and pinnules of o . mucofloris found in the present study . this results in a weight specific branchial surface area ( sbsa ) of ∼22 cm 2 g −1 fixed mass , which is similar to the sbsa of riftia pachyptila and higher than the sbsa of fish , crabs and other annelids [ 31 ] . this is especially obvious when compared to the sbsa of e . g . , arenicola marina ( 4 . 00 cm 2 g −1 ) [ 34 ] , although this species can also take up o 2 across its general epidermis .
interestingly , the present study showed that each pinnule is equipped with a distal sensory cell and external sensory cilia . these structures may sense disturbance in the water to avoid predators , as osedax need not sense food items or reproductive indicators in the water [ 1 ] , [ 8 ] , [ 35 ] . additionally , sensing of currents may also be beneficial in order to orientate the palps e . g . , for optimal uptake of dissolved o 2 . riftia pachyptila does not have sensory structures on the pinnules , but does have long , separate sensory filaments that lack ciliary bands and pinnules . these structures , with unknown function , are placed between pinnulated filaments [ 33 ] .
a : single z - stack image of the ‘trunk - root system’ connection , note the bundles of longitudinal muscles . b : musculature located by the ovisac , assumed to be the posterior end of the longitudinal dorsal blood vessel . c : single z - stack image of pinnules , circular musculature encircling the pinnular loop , note the distal perikaryon and sensory cilia . d : single z - stack image showing a longitudinal section of the pinnule in c , note the internal nerve . e : depth coded z - stack , pinnule nerves in osedax ‘yellow - collar’ . abbreviations : cilia ( ci ) , circular muscles ( cm ) , lateral ciliary band ( lcb ) , longitudinal muscles ( lm ) , nerve ( n ) , perikaryon ( pe ) .
in january 2009 , 3 replicate experimental sampling devices , using cow and whale bone , for recruitment of female o . mucofloris ( figure 10 ) were placed at 125 m depth off the coast of tjärnö , sweden ( 58°52 . 976n ; 11°05 . 715e ) in close vicinity to a minke whale carcass sunk in october 2003 [ 13 ] . one device for morphological and reproductive studies was successfully retrieved in may 2009 . a second device was retrieved in november 2009 for in vivo studies ( respirometry and microsensor analysis ) and additional morphological studies . additionally , a piece of cetacean bone deposited at 123 m in the same area in may 2008 was retrieved in january 2009 and used for preliminary investigations and for designing experimental setups . in january 2009 , the bottom water had a salinity of 34 . 6‰ , a temperature of 8 . 4°c , and an o 2 content of 78 . 7 % atmospheric saturation .
pinnules ( up to 100 µm wide ) project perpendicularly from the frontal palp surface between the lateral ciliary bands , with a density of approximately eight pinnules across the palp per 50 µm palp length ( figures 3a , 5f , g ) . at the proximal end of the palps , pinnules are less developed than at the distal end , seemingly growing in length ( up to 170 µm ) synchronously with the growth of the palp ( figure 3a ) . a sensory cell extends through the centre of the pinnule with a distal perikaryon and a few external , presumably sensory , cilia ( figure 4c , d ) . its axon seems to connect with one of the two major longitudinal palp nerves , possibly the one running more laterally beneath the ciliary band . this is supported by similar findings in osedax “yellow - collar
( figure 4e ) .
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read , g . ; fauchald , k . ( ed . ) ( 2018 ) . world polychaeta database .
glover , kallstrom , smith & dahlgren , 2005 . accessed through : world register of marine species at : urltoken ; = 265980 on 2018 - 07 - 09
van der land , j . ( ed ) . ( 2008 ) . unesco - ioc register of marine organisms ( urmo ) . , available online at urltoken [ details ]
: i think it ' s brilliant . describes exactly what it does and what it looks like and piques one ' s interest .
that is the most awful name for a species that i ' ve ever seen . wow .
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feathered plumes , which are the gills , or branchiae , that bring oxygen down to the root structure embedded in the whale bone .
a cylinder - shaped column that extends furthest into the water column . scientists think this structure , an oviduct , shoots fertilised eggs into the water column , perhaps helping the animal to disperse tiny larvae to be carried off by ocean currents .
live specimens are visible as four white to pink palps that emerge from the surface of the whale bone . the palps are 5 - 6 mm in length , and are surrounded at their base by a thin mucous tube . the oviduct is white , and extends to one - third of the length of the uncontracted palps . the palps are of equal length , measuring 0 . 8 mm when contracted , with numerous pinnules 0 . 1 mm in length . at the base of the pinnules there is a ciliary band that runs the entire length of each palp . the pinnules are 0 . 01 mm wide . they are densely packed and coloured white to pink in live specimens , with microvilli 0 . 05 mm in length . the trunk region is 6 - 8 mm in length and 0 . 5 mm wide , and is partially embedded within the bone matrix . the trunk is principally composed of bands of longitudinal muscles , galnds and major dorsal and ventral blood vessels . the mouth and gut are absent . there are ventral plaques on the collar ( peristomial region ) of the trunk .
on dissected specimens , an oviduct is visble running into an ovisac and the root structure , and contains numerous eggs in the trunk region . there is a vascularized root system of 2 - 10 mm in length , which burrows in a shallow depression to depths of 2 - 3 mm into the whale bone matrix in a bracnhed , mycelial form . numerous eggs ( greater than 100 in number ) were relaesed from the ovisac on disturbance . the eggs ranged in diameter from 85 - 90 um . the chaetae and opisthosomal region were not observed . epibiotic rod - shaped bacteria ( length 1 - 1 . 5 um , width 300 nm ) were present over the surface of the trunks , palps and pinnules , but absent from the roots .
found on the bones of an experimentally implanted minke whale carcass at 125 m depth , kosterfjord , sweden ( 58°53 . 1 ` n , 11°06 . 4 ` e ) ( glover et al . , 2005 ) .
depth range based on 1 specimen in 1 taxon . environmental ranges depth range ( m ) : 125 - 125 note : this information has not been validated . check this * note * . your feedback is most welcome .
glover et al . ( 2005 ) reported that disturbance caused the worms to retract completely into the bone . however , when placed in aquaria , with clean , chilled seawater , they would emerge from the bone and be clearly visible to the naked eye . over a period of several minutes , the worms would first extend the palps , and then the oviduct ( see video here ) . any disturbance to the aquarium tank would result in the animals immediately withdrawing into the bone .
means
snot - flower bone - eater
, though the less - accurate
bone - eating snot - flower worm
seems to be the form actually used .
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verna c 1 , ramette a , wiklund h , dahlgren tg , glover ag , gaill f , dubilier n .
max planck institute for marine microbiology , celsiusstr . 1 , 28359 bremen , germany .
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copyright : © 2012 huusgaard 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 : the work was supported by the danish research council ( urltoken ; grant # 272 - 06 - 0260 to k . worsaae ) and the swedish research council ( urltoken ; grant # 2006 - 2768 to t . dahlgren ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
competing interests : one of the authors is currently employed as a senior researcher , phd by uni research , a non profit research institute controlled by the university of bergen . uni research has no competing interests to this work in terms of employment , consultancy , patents or products . this does also not alter the authors′ adherence to all the plos one policies on sharing data and materials .
ciliary bands and sensory structures ( anti acetylated α - tubulin staining and histology ) .
in the description below , we use a dorsal - ventral definition opposite to the one used by rouse et al . [ 1 ] , [ 21 ] , based on a new interpretation ( rouse & worsaae , unpublished ) .
we found two previously unreported broad longitudinal ciliary bands dorso - laterally on each side of the oviduct on the anterior part of the trunk ( figure 2a , b , e ) . the ciliary bundles of the bands consist of multi ciliated cells , appearing pillow - like with numerous , conspicuously short cilia ( < 10 µm long ) projecting outwards from the centre of the elliptical bundles ( figure 2c ) . the bundles are organized in an anterior - dorsal diagonal pattern within the longitudinal bands ( figure 2b ) . the ciliary bands narrow toward the posterior part of the trunk and are replaced by single tufts of cilia scattered basally across the trunk surface ( figure 2a ) . the short cilia appeared immotile , and with several longitudinal nerves running beneath , ( worsaae & rouse , unpublished ) their function may be sensory rather than ventilatory .
a : dorso - lateral view of a complete specimen , lateral ciliary bands occupies half the length of the trunk . b : dorsal view of the anterior part of the trunk , elliptical shaped cilia bundles are directed anteriorly from the lateral part of the trunk . c : depth coded z - stack , the elliptical shaped cilia bundles constituting the lateral ciliary band are formed by ciliary tufts . d : close - up of transverse section of a trunk . e : transverse section of a trunk , note the muscularized dorsal blood vessel . f : single z - stack image of the trunk musculature , longitudinal muscles beneath circular and diagonal muscles . abbreviations : ciliary tufts ( ct ) , circular muscles ( cm ) , diagonal muscles ( dm ) , elliptical ciliary bundles ( ecb ) , lateral ciliary band ( lcb ) , longitudinal muscles ( lm ) , muscular gap ( mg ) , palp ( p ) , root structure ( r ) , torn ovisac ( to ) , trunk ( t ) , dorsal blood vessel ( dbv ) .
two dense ciliary bands are found on each palp along their epidermal longitudinal lobes on each lateral side , as previously reported [ 12 ] ( figures 3a , b , e , 4c , 5a , f , g ) . the bands consist of multi ciliated cells with 70–90 µm long cilia ( figure 3e ) . the ciliary bands extend from near the basal part of the palps to the distal tip , beating in metachronal waves .
a : lateral view of palps , pinnules increasing in length and development along the palp . b : abfrontal view of the midsection of a palp , ciliary bands on each side . c : close - up of palp musculature . d : lateral view of the muscular palp - trunk connection . e : close - up of the lateral ciliary band of a palp . abbreviations : circular muscles ( cm ) , lateral ciliary band ( lcb ) , longitudinal muscles ( lm ) , muscular bundles ( bu ) , muscular gap ( mg ) , oviduct ( od ) , pinnules ( pin ) .
diagram of a transverse section of the basal part of palps ( a ) , dic light micrographs of benzidine stained palps ( b–d ) and transverse 1 . 2 µm sections of
a : diagram of a transverse section at the basal part of the palp region . circular musculature ( continued green lines ) encircles the longitudinal musculature ( green broken lines ) in a cylinder formation . two gaps separate the longitudinal muscle bands . black lines illustrate the motile lateral ciliary bands and two main palp nerves run along each palp as shown ( blue dots ) . b : pinnular loop filled with blood . c : pinnular loops , broken lines and arrows indicate the assumed direction of blood flow . d : midsection of palp , longitudinal blood vessels and pinnular loops visible . e : transverse section of a pinnule , the pinnular loop enclosed by a membrane fusing in the centre . f : transverse section of the distal part of a palp , arrow tips shows circular musculature . g : transverse section of the distal part of a palp , the two longitudinal blood vessels obvious . abbreviations : circular muscles ( cm ) , epidermis ( ep ) , lateral ciliary band ( lcb ) , left dorsal palp nerve ( ldpn ) , left ventral palp nerve ( lvpn ) , membrane fusion ( mf ) , musculature ( m ) , palp blood vessel ( pbv ) , pinnule ( pin ) , pinnular loop ( pl ) , right dorsal palp nerve ( rdpn ) , right ventral palp nerve ( rvpn ) .
the longitudinal muscles run along the entire length of the trunk ( figure 2a ) , originating posteriorly at the trunk basis , and inserting anteriorly at the base of the four palps ( figure 3d ) . in one individual , it was possible to detect clustering of longitudinal muscles into 14–16 bundles of > 20 muscle strands in the posterior part of the trunk , anterior to the root structure ( figure 4a ) . along , and around the entire trunk , the muscles are distributed in a dense cylindrical formation with an average of six strands per 50 µm ( figures 2d , e , 3d ) . the musculature is slightly separated internally to the oviduct ( possibly by the nerve cords ) , as well as randomly along the trunk , creating minor gaps most likely for mucus gland exits or nerves ( figures 2a , 3d ) . anteriorly , a gap in the musculature is found ventrally at the position of the brain , as well as at each of the four insertion points of the palps . at each of these four points , the longitudinal musculature divides into two bundles ( of each 20–30 strands ) , encircling the insertion point of the palp ( figure 3d ) . the longitudinal palp muscles originate at the base and run along the entire length of the palps to their tips , separated by a smaller frontal and a larger abfrontal gap ( figures 3c , d , 5a ) . no longitudinal musculature was detected in the pinnules .
around the ovisac and anterior root structure the longitudinal musculature divides , and together with the circular muscles , creates a mesh - like structure ( figure 4a ) . the musculature extends posteriorly along the roots in a cylindrical formation , supporting the tissue penetrating the bone .
thin circular muscles are found peripheral to the longitudinal muscles along the entire trunk , with 24 strands per 50 µm ( figure 2d ) . the circular muscles are most dominant in the posterior part of the trunk and continue into the root structure ( figure 4a ) . notably , much thicker diagonal muscle strands were found beneath the circular musculature , but peripheral to the longitudinal muscles ( figure 2f ) . attached at the mid - dorsal line , the strands run diagonally around the trunk in an anterior direction and attach at the mid - ventral line . the diagonal muscles are distributed along the entire length of the trunk , lying further apart ( 35–70 µm ) in the posterior end than along the rest of the trunk ( 5–10 µm ) .
the circular musculature of the palps encircles the longitudinal musculature as a cylinder with ∼18 muscle strands per 50 µm and is evenly distributed along the entire palp ( figure 3c ) . fine circular muscles enclose each vessel of the pinnular loop ( figure 4c , d ) ; they were likewise visible in the semi - thin sections on the outside of the pinnular loop and along the midline of each pinnule ( figure 5f , arrow tips ) , with a spacing corresponding to those shown with phalloidin staining ( figure 4c , d ) .
the ventral blood vessel also continues into the root structure , but as a narrower and less defined vessel , the path of which was even more difficult to determine , than that of the dorsal vessel . blood vessels of different sizes , as well as multiple obvious capillaries within the root tissue were observed ( figure 6b–e ) . larger vessels extend out from the area of the ovisac and divide into thinner vessels ( diameter : 7–30 µm ) . capillaries were detected in the periphery of the root tissue ( figure 6e ) , with distances between the detected capillaries ranging from 65–220 µm .
anteriorly , each palp encloses a pair of blood vessels created by invaginations of the inner lamina of the basement membrane of the epidermis ( figures 5f , g , 7c ) . imaging of live specimens confirmed the presence of two blood vessels , running along the entire length of the palps .
the pinnules are largely filled by a blood cavity lined with a membrane , which fuses in between the two blood cavities along most of the pinnule length , thereby creating the pinnular loop ( figures 5b–g , 7c ) . the palps of the histological sections were ∼300 µm in diameter at the base and the palp epidermis was ∼50–75 µm thick . the palp diameter and the thickness of the epidermis were both found to decrease towards the distal end of the palp .
pinnules vary in diameter along their length and along the palp , with a median diameter of ∼40×100 µm for the pinnules and ∼20–40 µm for each blood vessel . the diffusion distance across the pinnule epithelium was measured on the semi - thin sections to be 1–2 µm , and for the epidermis of the distal part of the palps , the diffusion distance was measured to be ∼30 µm .
a : depth profile of o 2 towards bone surface , blue : agar , red : cuvette wall , green : bone surface . b : depth profil of o 2 towards tissue surface , blue : agar , red : cuvette wall , green : tissue surface . c : schematic drawing of the micro sensor measuring path through the cuvette wall . d : placement of measuring site on wb1 , note the blackened areas indicating presence of ferrous sulphide . e : close - up of measuring sites on wb1 .
micro sensor measurements of o 2 distribution in one mucus tube showed a ∼50 % decrease in the o 2 concentration in the centre of the mucus tube wall as compared to outside the tube ( table 3 ) . direct measurements within the tube were not possible due to disturbance by the worm . measurements of the o 2 microenvironment surrounding the palps showed a strong decrease in o 2 concentrations , when a palp approached the microelectrode measuring tip . at the base and middle of the palp , the o 2 levels were almost zero showing the o 2 uptake to be high in these areas ( table 3 ) . at the distal end of the palp the o 2 concentration was only reduced to approximately 50 % atmospheric saturation , possibly due to decaying palp tips .
the presence of highly vasculated palps and pinnules , the former densely ciliated showed that the anterior crown is the main site for o 2 uptake ( figure 7a–c ) . uptake of o 2 over the trunk surface is possible , but a thicker epidermis , short and seemingly immotile ciliary bands and no obvious respiratory structures suggest that the trunk is a minor site of o 2 uptake . oxygen does , to some extent , diffuse from the surrounding water into the mucus tube , thereby supplying the dwarf males .
the well - developed longitudinal musculature of the trunk serves to retract the trunk and palps into the tube and bone , presumably as protection from predators . as no regular retraction patterns of trunk musculature were detected , retraction into the tube is not considered a significant mode of ventilation for required o 2 . furthermore , the tube only surrounds the trunk , tightly fitting to the base of the trunk and the surface of the bone , preventing any water exchange to the ovisac and roots from the bone surface . the longitudinal musculature is moreover able to stretch the extensive branchial structures ( palps ) into the aerated water in order to increase the o 2 uptake .
the main blood flow within the palp vessels may be generated by the circular body wall musculature of the palp . however , the thin circular musculature of the pinnules surrounding each branch of the looped blood vessel ( figure 4c , d ) most likely assists local blood flow , as suggested for the tentacular vessels of riftia pachyptila [ 22 ] . musculature in anterior appendages has been found in several vestimentifera , in the form of sphincter muscles located in the branchial lamella and filament vessels [ 33 ] .
graph modified from cammen [ 18 ] , the regression line ( log r = −1 . 682 + 0 . 850 * log w ) calculated from measurements of resting nonventilating annelids only . dots : b1 ( red ) , b2 ( blue ) , b3 ( green ) , c1 ( purple ) , c2 ( yellow ) . triangles : previous measured o 2 consumption of r . pachyptila . no sulphide present in water when measuring : red , blue [ 22 ] and green [ 23 ] ; sulphide present in water during measurement : purple [ 23 ] . red square : o 2 consumption of resting arenicola marina [ 52 ] .
the experimental sampling devices were placed at 125 m depth off the coast of tjärnö , sweden ( 58°52 . 976n ; 11°05 . 715e ) in close vicinity to a minke whale carcass sunk in october 2003 [ 13 ] .
four specimens were stained following the protocol of worsaae & rouse [ 45 ] . first staining included the primary antibodies monoclonal mouse anti - acetylated α - tubulin ( sigma t6793 , 1∶200 ) & polyclonal rabbit anti - serotonin ( sigma : s5545 ; 1∶100 / 1∶400 ) or monoclonal mouse anti - acetylated α - tubulin & anti - fmrfamide ( immunostar : 20091 , 1∶100 ) . this was complimented by secondary antibodies ; anti - mouse cy5 ( jackson immunoresearch : 115 - 175 - 062 , 1∶400 ) and anti - rabbit tritc ( sigma t5268 , 1∶200 / 1∶400 ) . hereafter specimens were incubated for 60 min in phalloidin conjugated with fitc or alexa flour 488 ( sigma f5282 or invitrogen a12379 , 0 . 17 or 0 . 33 µmol l −1 phalloidin in pbs ) . specimens were mounted in 100 % vectashield® containing dapi ( vector laboratories inc . , california , usa ) and stored at −18°c . the specificity of primary antibody binding versus e . g . , autoflourescence was tested by omitting one of the primary antibodies , but otherwise treating specimens as described .
specimens were studied using a leica tcs sp5 confocal laser scanning microscope ( clsm ) ( university of copenhagen , faculty of health science , courtesy of m . givskov and t . bjarnsholt ) . leica lasaf computer software or imaris® x64 6 . 0 . 0 ( bitplane ag , zurich , switzerland ) was used to produce projections of z - stacks of clsm images , while further analyses of z - stack series were performed with imaris® x64 6 . 0 . 0 . computed 2d images of muscles , nerve and cilia with relation to respiration and palp morphology were further optimized with adobe photoshop cs3 and adobe illustrator ( adobe system incorporated ) for presentation .
one specimen was embedded in epon and used for histological analysis . semi - thick 1 . 2 µm sections were cut on a microtome ( em uc6 , leica , wetzlar germany ) with a diamond knife ( diatome ; biel , switzerland ) . a small amount of pattex contact adhesive ( pattex compact ; henkel kgaa , düsseldorf , germany ) was diluted with a few drops of xylene in an eppendorf tube and applied to the side of the epon block to make serial sectioning of ribbons possible following the protocol of henry [ 46 ] and ruthensteiner [ 47 ] . bands of ∼20 sections were stained with toluidine blue and mounted in entellan® ( electron microscopy sciences , pennsylvania , usa ) . sections were studied and photographed using light microscopy ( bx50 microscope ; dp71 camera ; cell f software ; olympus , japan ) .
using a modified version of the benzidine staining method by knox [ 48 ] , haemoglobin was stained in four fixed specimens . a 100 % saturated benzidine solution was prepared by adding benzidine to distilled water . the solution was stirred for two hours . fixed specimens were rinsed in running tap water in the same time period . specimens were subsequently incubated in the filtered benzidine solution for 1 hour , also under stirring . next , 3 % hydrogen peroxide was added drop by drop until blood vessels turned dark blue . specimens were either mounted in glycerol directly or dehydrated in a series of alcohol acidified with drops of 0 . 1 % acetic acid , where after tissues were cleared in xylene and mounted in d . p . x between two cover slips . specimens were analyzed and photographed under a light microscope ( bx50 microscope ; dp71 camera ; cell f software ; olympus , japan ) .
o 2 consumption was measured in seawater kept at ∼100 % atmospheric saturation using intermittent respirometry in accordance with vismann and hagerman [ 49 ] . the experimental setup was placed in a constant temperature room at 6°c . each experiment encompassed 3–8 measuring sequences consisting of a flushing period of 10–30 min and a measuring period of 30–45 min . the set - up had a chamber flushing rate of 30 ml min −1 and a shunt water flow of 8 ml min −1 past the o 2 electrode ( e5046 , radiometer medical aps , brønshøj , denmark ) .