harpa gracilis , common name the polynesian harp , is a species of sea snail , a marine gastropod mollusk in the family harpidae , the harp snails .

harpa gracilis

harpa striata lamarck , 1816 : synonym of harpa cabriti p . fischer , 1860
harpa ventricosa lamarck 1816 - ventral harp : synonym of harpa cabriti p . fischer , 1860
harpidae » harpa gracilis , id : 228421 , shell detail « shell encyclopedia , conchology , inc .
worms - world register of marine species - harpa gracilis broderip & g . b . sowerby i , 1829
polynesian harp - harpa gracilis broderip & g . b . sowerby i , 1829 - literature - encyclopedia of life
berkhout ( 1992 ) writes : “h . gracilis is the only harpa shell that has a white protoconch . ”
belongs to harpa according to b . landau and c . marques da silva 2010
described in malacologia . the h . kolaceki has the purple protoconch in common with the tuamotou’s endemic harpa gracilis , but the shape is completely different . rare .
rehder ( 1973 ) writes on harpa gracilis broderip & sowerby , 1829 : “nuclear whorls 3 1 / 2 to 4 , smooth , white . . . ”
however , on the conchology inc . auction i read thus : “the h . kolaceki has the purple protoconch in common with the tuamotou’s endemic harpa gracilis . . . ”
so , can anyone verify for me that there are indeed harpa gracilis ( from the tuamotos or elsewhere ) with a purple protoconch ? perhaps a subspecies ? this would certainly shed some light on this complex of smaller harpa . an image would be most appreciated .
bouchet , p . ( 2015 ) . harpa gracilis broderip & g . b . sowerby i , 1829 . in : molluscabase ( 2015 ) . accessed through : world register of marine species at urltoken ; = 403835
harpa gracilis is the only harp with a white protoconch , clearly distinguishing it from all other harps , whose protoconchs are pink to brown . other differentiating characteristics are the shallow umbilicus , slender form and no shoulder on the body whorl . the harps in the links you sent are , therefore , clearly not gracilis . they are amouretta , as stated by others .
poppe , brulet & dance ( 1999 ) also write : “h . gracilis . . . protoconch is white instead of purple . ”
dealers sometimes use the name gracilis for ( or in relation to ) non - gracilis shells for obvious reasons . regarding h . kolaceki , even poppe , on his auction site selling it , mentioned that it is a
doubtful
species . may very well be a form of amouretta .
about h . gracilis i also wrote in 1990 ( let ' s tune our harps ! xenophora no . 50 pp . 10 - 21 ) :
protococh white .
on the conchology inc . auction what is read : “the h . kolaceki has the purple protoconch in common with the tuamotou’s endemic harpa gracilis . . . ” is an error as it does not match the description of h . kolaceki where the colour of its protoconch is not even mentioned and on the colour photos included in the description it is obvious that the protoconch of h . kolaceki is purple .
i agree with rick . while the shells he illustrated may not be h . gracilis , they are certainly unlike any of the h . amouretta i have ( another 30 specimens to that 50 ) .
broderip w . j . & sowerby i g . b . ( 1829 ) . observations on new or intersting mollusca contained , for the most part , in the museum of the zoological society . zoological journal . 4 : 359 - 379 . , available online at urltoken page ( s ) : 373 [ details ]
dance , s . p . & g . t . poppe , 1999 family harpidae . in : a conchological iconography ( conchbooks , ed . ) , 69 p . [ details ]
rehder h . a . ( 1973 ) . the family harpidae of the world . indo - pacific mollusca 3 ( 16 ) : 207 - 274 . , available online at urltoken [ details ]
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 ]
petit , r . e . ( 2009 ) . george brettingham sowerby , i , ii & iii : their conchological publications and molluscan taxa . zootaxa . 2189 : 1–218 . , available online at urltoken [ details ] available for editors [ request ]
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the star system calculates the number of pieces that were handled by conchology , inc . in the last 15 years :
we want to point out that the star system is only very reliable for philippine shells only , as we handle very few foreign shells in general . as time goes , the system will become more and more performant .
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version 43 . 0 went live 11 / 6 / 2018 - i hope that the majority of issues have been fixed . my email address is on the home page if you see anything wrong .
the following bibliography has been generated by bringing together all references provided by our content partners . there may be duplication .
bouchet p . , gofas s . & rosenberg g . ( 2015 ) . worms mollusca : world marine mollusca database ( version feb 2013 ) . in : species 2000 & itis catalogue of life , 26th august 2015 ( roskov y . , abucay l . , orrell t . , nicolson d . , kunze t . , flann c . , bailly n . , kirk p . , bourgoin t . , dewalt r . e . , decock w . , de wever a . , eds ) . digital resource at urltoken . species 2000 : naturalis , leiden , the netherlands . issn 2405 - 8858 .
broderip w . j . & sowerby g . b . ( 1829 ) . observations on new or intersting mollusca contained , for the most part , in the museum of the zoological society . zoological journal , 4 : 359 - 379 , pl . 9
dance , s . p . & g . t . poppe , . 1999 . amily harpidae . in : a conchological iconography ( conchbooks , ed . ) , 69 p .
petit r . e . 2009 . george brettingham sowerby , i , ii & iii : their conchological publications and molluscan taxa . zootaxa 2189 : 1–218 .
rehder h . a . 1973 . the family harpidae of the world . indo - pacific mollusca 3 ( 16 ) : 207 - 274 .
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broderip w . j . & sowerby g . b . ( 1829 ) . observations on new or intersting mollusca contained , for the most part , in the museum of the zoological society . < i > zoological journal < / i > , 4 : 359 - 379 , pl . 9
broderip , w . j . & sowerby , g . b . ( 1829 ) . observations on new or intersting mollusca contained , for the most part , in the . < em > museum of the zoological society . zoological journal < / i . < / em > 4 : 359 - 379 .
dance , s . p . & g . t . poppe , 1999 family harpidae . in : a conchological iconography ( conchbooks , ed . ) , 69 p .
liu j . y . [ ruiyu ] ( ed . ) . ( 2008 ) . checklist of marine biota of china seas . < em > china science press . < / em > 1267 pp .
petit , r . e . ( 2009 ) . george brettingham sowerby , i , ii & iii : their conchological publications and molluscan taxa . < em > zootaxa . < / em > 2189 : 1–218 .
rehder h . a . ( 1973 ) the family harpidae of the world . indo - pacific mollusca 3 ( 16 ) : 207 - 274 .
rehder h . a . ( 1973 ) . the family harpidae of the world . indo - pacific mollusca 3 ( 16 ) : 207 - 274 .
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interesting , i have over 50 specimens of amouretta from around the world and none of them are like these two shells . can someone post images of similar shells ?
by the way , in the description of h . kolaceki the colour of the protoconch is not even mentioned .
for rick ' s shells . i ' ve documented this conversation on lts with rick ' s photos ( and permission ) and added photos of a
from the solomons in my collection . while there are are some overall shape differences , with my shell more closely resembling forma
krauss , 1848 , i cannot see how rick ' s shells can be distinguished as other than a normal , local , regional variation for the species .
i fully agree with your comment “ i cannot see how rick ' s shells can be distinguished as other than a normal , local , regional variation for the species .
( 1973 ) . rehder presented 11 species of living harps . there have been many new taxa proposed since 1973 . all but one are now generally accepted as forms of one or the other of the 11 species addressed by rehder . only
rehder , 1993 is mostly accepted as a valid addition to his 1973 list . whether or not all 11 of the taxa addressed by rehder ( plus
) are truly separate species or if the many localized forms given species / subspecies names actually represent separate species will have to await a comprehensive dna analysis .
( two or one blotch ) to be able to accept his argument . i would also note that the characters described for
( “body whorl … more broadly ovate and rounded , ” “ribs tend to be narrower and more distant” ) . my own opinion is that
species addressed by rehder and the
accepted
species addressed 31 and 42 years later .
. indo - pacific mollusca , volume 3 , no . 16 . delaware museum of natural history .
are quite variable within populations and across geographic range . so variable that they occur readily across taxa , especially those that share common geographic distributions and influences , and most often cannot be relied upon to distinguish between species when confronted with a particular specimen . in my presentations i have limited the features discussed to those that should be relied upon to distinguish among taxa . i have presented the protoconchs for all , but did not find this feature to be helpful ( too variable , too often missing or incomplete , and too similar ) in distinguishing among taxa , except in a few cases (
) . i did not find color to be helpful for taxa that are otherwise close and from the same locales . i did not find reliance on “blotching” to be more than partially helpful without linkage to other confirming features . i have addressed the distribution of parietal glazing and found it to be quite helpful and distinctive for many taxa ( especially when linked to other features ) , but not decidedly so for the most problematic taxa (
) . i do not present a general description of each taxa , which is available many places elsewhere ( see rehder 1973 ) . rather , for some i present some background information and then start my descriptions with the parietal glazing and follow with those features that i found can best be used to distinguish among taxa .
the following table presents the features i found best allow identifying and distinguishing the 12 taxa . the green cells describe key features that should be identified first ( and in some taxa , alone or linked to other “greens , ” are sufficient to identify a taxon ) . the pink cells describe features very helpful in narrowing the possibilities for otherwise similar taxa . the yellow cells describe features that i found very consistently separate taxa within the geographic range of
. i apologize for the tiny text in the table , but i wanted to get it all on one page .
, and from a dozen to several dozen for the others , limited examination of shells displayed at shell shows by exhibitors and dealers , some images from the web ( usually too poor to be helpful ) , and images in literature ( also usually also too poor to be helpful ) . obviously , my sample is limited in terms of actual material for close examination , and my conclusions should be tempered accordingly . i would be happy to hear from those with specimens in any of these taxa that question or confirm my observations (
) . however , i would also hope that you can provide good quality photos or would be willing to loan the specimens for a photo session . i will continue to add to these presentations with comments or more photos / material contributed by other fans of
. three terms ( subsutural plateau , shoulder and t ) should be understood . normally , “shoulder” refers to the area from the suture to an inflection point ( a pronounced downward angle ) or , when absent , the periphery . all
, the shoulder would normally be the area from suture to the inflection point ( where spines are located ) . i have defined the area from the suture to the inflection point as the subsutural plateau . and , when i refer to the shoulder , i am referring narrowly to the spiral line representing the inflection where the subsutural plateau turns downward ( and is where the rib spines normally occur ) . i have observed that all mature
has three . as a matter of shorthand i may use t1 , t2 , t3 or t4 to refer to the teleoconch whorls . so ,
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do not be surprised to find cod or mackerel in paris in the late middle ages ( 14th - 15th c . ) . . . the sea did not come that far , but trade did .
in compliance with laws and intellectual property practices in the scientific world , unpublished data under five years ( from excavation reports , analyses , but also academia ) is never communicated .
the dots represent the sites for which a study ( or a simple recording ) of fauna and / or flora was performed ; these sites were recorded in the database zooarchaeological and archaeobotanical inventories of france ( i2af ) . the lack of dots can mean both a lack of data and / or lack of recording .
the dots remain gray when the selected species is absent from the site . symbols of different colour mark the presence of the species in different periods .
to select / deselect one or more periods or to remove the dots corresponding to the listed sites , check / uncheck in the right side .
hovering a dot reveals in the left corner at the bottom of the map , the name of the department or municipality . one click enables the list of sites present on the municipality .
the selection of a site opens a new page that displays general information relating thereto ( other name ( s ) known , location . . . ) , the bibliographical reference ( s ) connected to the species , and a summary of plant and animal species identified per large time period .
a timeline summarizes all known information on the species . the relevant periods or sub - periods when the detail is known , are highlighted in red . tool tip : passing over a period ( palaeolithic for example ) displays the date range of the period .
access to more detailed information is accessible only by convention , after login , for scientists affiliated to groups or institutional programs ( national center of scientific research , national institute of preventive archaeological research , for example ) or associations ( international council for archaeozoology , for example ) .
warning : the data available reflects the progression status of knowledge or the availability of the inventories . it should never be considered as comprehensive .
national inventory of natural heritage , website : https : / / inpn . mnhn . fr .
full reference : f . e . eames . 1952 . a contribution to the study of the eocene in western pakistan and western india : c . the description of the scaphopoda and gastropoda from standard sections in the rakhi nala and zindar pir areas of the western punjab and in the kohat district . philosophical transactions of the royal society of london series b 236 : 1 - 168
type specimen : bmnh g . 68305 , a shell . its type locality is fossil bed f . 2556 , zinda pir , which is in a lutetian marine shale in the domanda formation of pakistan .
holotype nhmw 2009z0076 / 0001 . upper miocene . lopez section , rio yaque del norte , upstream of the mouth of angostura gorge , dominican republic .
buccinum testa costis aequilibus longitudinalibus , distinctis mucronatis , columella laevigata .
. recent . benghala . refers to buonanni , 1681 : plate 185 ( a ) ; lister , 1688 : pl . 992 [ fig . 55 ] ( b ) ; petiver , 1702 : pl . 48 fig . 13 ( c ) ; rumphius , 1705 : pl . 32 figs . k , l , m ( d ) ; gualtieri , 1742 : pl . 29 figs . c . d , e , g ( e ) ; argenville , 1742 : pl . 20 fig . d ( f ) ; regenfuss , 1758 : pl . 2 fig . 14 ( g ) .
rv 5167 . philippines , cebu province , olango island ; by diver , depth 20 m ; june 2012 .
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new records of diplomys labilis ( bangs , 1901 ) ( mammalia , rodentia , . . .
first record of boana maculateralis ( caminer & ron , 2014 ) . . .
first record of the bignose unicornfish , naso vlamingii ( perciformes , . . .
ambidexter symmetricus manning & chace , 1971 ( decapoda , processidae ) : . . .
check list is a peer - reviewed , open access , on - line journal devoted to publishing annotated list of species , notes on geographic distribution of one or a few species , and distribution summary of a taxonomic group . these data are essential for studies on biogeography and provide a baseline for the conservation of biodiversity as a whole . the first step to undertaking effective conservation action is to understand species’ geographic distribution . check list was established to cater to this need by publishing papers on the geographic distribution of species and higher taxonomic groups .
doaj , scopus , zoological abstracts , ebsco host , and index copernicus . member journal of the brazilian association of science editors ( abec ) and of the committee on publication ethics ( cope ) .
the shell has an ovate - oblong shape . it is more or less inflated , generally pretty thin , enamelled , and provided with parallel , longitudinal , inclined and acute ribs . ; the body whorl is much larger than all the others together . the spire is slightly elevated . the aperture is large , oval , dilated , strongly emarginated inferiorly , and without siphonal canal . the outer lip is bordered by the last rib . the columella is smooth , simple , nearly straight and pointed at the base .
the animal has a flattened head , which supports a pair of pretty long , thick , and conical tentacles , with a small protuberance at their base , internally , where the eyes are situated . the mouth is simple , surrounded by a muscular margin , and furnished with a small , slender and pointed trunk . the organ of excitement is elongated , cylindrical , situated on the right side . the locomotive organ is very large , very broad at the anterior part , which is ear - shaped , and distinguished by a deep emargination upon each side . the posterior extremity is caducous , and destitute of an operculum .
the fleshy part of this mollusk is very strong , and very large . its foot is enormous , thick , and extended considerably out of the shell . it cannot be wholly contained within the aperture , before which , by contracting itself , it forms a margin .
the foot is as if divided into two portions . the anterior broader , arcuated , ear - shaped , with a marginal furrow , and joined to the posterior part by a kind of neck . this latter , more extended , is somewhat oval , pointed , and slightly inflated above , without any appearance of operculum . when the animal is violently disturbed , it breaks off the posterior extremity of its foot , in order to withdraw itself more completely within its shell . in consequence of this an operculum would be useless to it , for it would be liable to be carried away by the rupture of the foot . therefore , it is not possessed .
all the external parts of the animal are strongly colored with spots and plates of a brownish red , intermingled with other yellowish spots . the middle portion is frequently crossed by a brown band .
this marine species is circumtropical , except the western atlantic ocean . it also occurs off australia ( northern territory , queensland , western australia ) .


procaris is a genus of shrimp in the family procarididae .

it contains the following species : procaris ascensionis chace & manning , 1972 procaris chacei hart & manning , 1986 procaris hawaiana holthuis , 1973 procaris mexicana von sternberg & schotte , 2004 procaris noelensis bruce & davie , 2006

procaris

phylum arthropoda subphylum crustacea class eumalacostraca order decapoda family procarididae procaris sp . kensley , 1988
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u . s . fish and wildlife service . 2003 . candidate assessment and listing priority assignment form - procaris hawaiana . u . s . fish and wildlife service , pacific islands office . 6 pp .
bruce , a . j . ; davie , p . j . f . ( 2006 ) . a new anchialine shrimp of the genus procaris from christmas island : the first occurrence of the procarididae in the indian ocean ( crustacea : decapoda : caridea ) . zootaxa . 1238 : 223 - 252 . [ details ] available for editors [ request ]
two of the known pools containing procaris hawaiana lie within a state natural area reserve . the rarity of this shrimp contributed to the current protection received by the maui anchialine pools ( holthuis 1973 ) . no conservation agreements between federal , state , or private landowners have been drafted or initiated and , aside from placement of some pools / pool systems within reserves , virtually no conservation activities have been conducted .
ty - jour ti - a new anchialine shrimp of the genus procaris ( crustacea : decapoda : procarididae ) from the yucatan peninsula t2 - proceedings of the biological society of washington . vl - 117 ur - urltoken pb - biological society of washington cy - washington , py - 2004 sp - 514 ep - 522 sn - 0006 - 324x au - sternberg , richard v au - schotte , marilyn er -
@ article { bhlpart49127 , title = { a new anchialine shrimp of the genus procaris ( crustacea : decapoda : procarididae ) from the yucatan peninsula } , journal = { proceedings of the biological society of washington . } , volume = { 117 } , copyright = { in copyright . digitized with the permission of the rights holder . } , url = urltoken publisher = { washington , biological society of washington } , author = { sternberg , richard v and schotte , marilyn } , year = { 2004 } , pages = { 514 - - 522 } , }
main threat = alien species . greatest potential threat is cited as the introduction of non - native fish into the pools ( u . s . fish and wildlife service 1998 ; 2003 ) . anchialine pools have been used to discard or hold bait - fish and / or aquarium fish . these fish either directly consume the native shrimp or , as with introduced tilapia ( oreochromis mossambica ) , out - compete the native herbivorous species of shrimp which typically serve as the prey - base for the rarer , predatory species of shrimp . habitat modification is also a threat ; up to 90 % of the anchialine pools on the island of hawaii have been either destroyed or altered . on the island of hawaii , dr . r . brock ( pers . comm . , 1998 ) estimates that up to 90 percent of the anchialine pools have been destroyed or altered by human activities . although the two known maui pools , which are contain procaris hawaiana , occur within a protected state reserve , habitat modifications by early hawaiians and later inhabitants have occurred in the area .
chace , f . a . jr . & r . b . manning , 1972 . two new caridean shrimps , one representing a new family , from marine pools on ascension island ( crustacea : decapoda : natantia ) . — smithsonian contributions to zoology 131 : 1 - 18 . [ details ]
de grave , s . & c . h . j . m . fransen . ( 2011 ) . carideorum catalogus : the recent species of the dendrobranchiate , stenopodidean , procarididean and caridean shrimps ( crustacea : decapoda ) . zool . med . leiden . 85 ( 9 ) : 30 . ix . 2011 : 195 - 589 figs 1 - 59 . ( look up in imis ) [ details ] available for editors [ request ]
bouchet , p . ; fontaine , b . ( 2009 ) . list of new marine species described between 2002 - 2006 . census of marine life . [ details ]
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phyllobranchiate gills ; maxillipeds and pereiopods with strong exopods ; none of pereiopods chelate or subchelate ; rostrum small and unarmed ( kensley , 1988 ) .
are quite similar , showing few differentiating characters ( hart & manning , 1986 ) .
abele and felgenhauer ( 1985 ) have studied the ecology of the ascension species . they report that smaller individuals spend most of their time in crevices , while larger individuals are observed swimming in open water . examination of gut contents reveals that they feed on both plant matter and crustaceans including amphipods and atyid shrimp .
( felgenhauer et al . , 1988 ) . although more than 1 , 000 specimens of
were observed in the field , no ovigerous females were seen . a single female maintained in the laboratory bore approximately 60 bright orange eggs on the endopods of pleopods . the large size of these eggs ( 0 . 83 - 0 . 93 mm ) suggests the existence of a zoeal larval stage ( felgenhauer et al . , 1988 ) .
hart and manning ( 1986 : 416 ) note that the similarities between species and their highly anomalous distribution in marine caves indicate an extremely slow rate of evolution . they suggest that ,
, or its predecessors , may , at one time , have been widely distributed throughout the oceans , surviving today only in cryptic habitats removed from some of the environmental pressures necessitating change .
felgenhauer , b . e . , l . g . abele and w . kim . 1988 . reproductive morphology of the anchialine shrimp
hart , c . w . and r . b . manning . 1986 . two new shrimps ( procaridae and agostocarididae , new family ) from marine caves of the western north atlantic .
kensley , b . 1988 . new species and records of cave shrimps from the yucatan peninsula ( decapoda : agostocarididae and hippolytidae ) .
schram , f . r . 1986 . crustacea . oxford university press , oxford , xii + 606 pp .
this shrimp was discovered in february 1972 by j . a . maciolek in anchialine lava pools at cape kinau , maui and also in anchialine pools at lua o palehemo , south point on the big island of hawaii . only three populations of this rare shrimp are known .
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u . s . fish and wildlife service ( usfws ) . 1989a . endangered and threatened wildlife and plants ; animal notice of review . federal register , department of the interior 54 ( 4 ) : 554 - 579 .
apparently endemic to the hawaiian islands where it has only been recorded from three anchialine pools .
due to latency between updates made in state , provincial or other natureserve network databases and when they appear on natureserve explorer , for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data . please refer to
known from the hawaiian islands of hawaii ( 1 site ) and maui ( 2 sites ) ( holthuis , 1973 ; maciolek 1983 ; u . s . fish and wildlife service 1998 ; 2003 ) . current range considered to be relictual of an early global distribution .
known from two sites on maui and one site on hawaii ( usfws , 1998 ) .
known from one site on hawaii and two sites on maui ( holthuis , 1973 ; u . s . fish and wildlife service 1998 ; 2003 ) .
it was estimated to occur in the thousands at the hawaii site ( kensely and williams 1986 ) . due to their rarity within these locations since the previous estimate , population estimates have not been attempted .
it was estimated to occur in the thousands at the hawaii site ( kensely and williams 1986 ) .
in hawaii , there are estimated to be over 650 anchialine pools , with an estimated 90 % of these occurring on hawaii . approximately 90 % of the pools on that island have been destroyed or otherwise impacted by development or other human uses ( richard brock , university of hawaii , personal communication , 1998 in usfws , 1998 ) .
in the state of hawaii , there are estimated to be over 650 anchialine pools , with an estimated 90 % of these occurring on the island of hawaii . unfortunately , approximately 90 percent of the pools on that island have been destroyed or otherwise impacted by development or other human uses ( usfws , 2003 ) .
( 250 - 1000 square km ( about 100 - 400 square miles ) ) known from the hawaiian islands of hawaii ( 1 site ) and maui ( 2 sites ) ( holthuis , 1973 ; maciolek 1983 ; u . s . fish and wildlife service 1998 ; 2003 ) . current range considered to be relictual of an early global distribution .
has pink to light - red pigmentation that is darkest along the midline with the dorsal thorax being white to yellow . black pigments are associated with the eyss . conspicuous chelapeds ( claws ) are lacking . locomotion is accomplished by swimming with the swimmerets ( pareopods and uropods ) and occurs just above the substate to mid - water ( usfws , 1998 ; 2003 ) .
found in anchialine pools with salinities generally ranging from 19 - 25 ppt . anchialine pools are tidally influenced saline pools that occur along the coast but are not openly connected to the ocean . this includes water in natural depressions , fissures , quarries , and craters . the site on hawaii is an anchialine pool that is connected to deeper waters leading to a submerged lava tube ( kensley and williams 1986 ) .
use the generic guidelines for the application of occurrence ranks ( 2008 ) . the key for ranking species occurrences using the generic approach provides a step - wise process for implementing this method .
zoological data developed by natureserve and its network of natural heritage programs ( see local programs ) and other contributors and cooperators ( see sources ) .
holthuis , l . b . 1973 . caridean shrimps found in land - locked saltwater pools at four indo - west pacific localities ( sinai peninsula , funafuti atoll , maui and hawaii islands ) , with the description of one new genus and four new species . zoollogische verhandelingen 128 : 3 - 55 .
kensley , b . and d . williams . 1986 . new shrimps ( families procarididae and atyidae ) from a submerged lave tube on hawaii . journal of crustacean biology 6 ( 3 ) : 417 - 437 .
maciolek , j . a . 1983 . distribution and biology of indo - pacific insular hypogeal shrimps . bulletin of marine science 33 : 606 - 618 .
maciolek , j . a . 1986 . environmental features and biota of anchialine pools on cape kinau , maui , hawaii . stygologia 2 ( 1 / 2 ) : 119 - 129 .
u . s . fish and wildlife service ( usfws ) . 1998 . category and listing priority forms .
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ridgely , r . s . , t . f . allnutt , t . brooks , d . k . mcnicol , d . w . mehlman , b . e . young , and j . r . zook . 2003 . digital distribution maps of the birds of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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patterson , b . d . , g . ceballos , w . sechrest , m . f . tognelli , t . brooks , l . luna , p . ortega , i . salazar , and b . e . young . 2003 . digital distribution maps of the mammals of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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this procarid shrimp has a thin , fragile integument ( hard outer covering of the body ) . the carapace and short , triangular rostrum are unarmed , and the eyestalks split into two blunt lobes . all five pairs of walking limbs ( pereiopods ) are similar , each with large , bristle - like structures ( 2 ) .
recorded only from green bay cave in hamilton parish , bermuda ( 2 ) .
classified as critically endangered ( cr ) on the iucn red list 2006 ( 1 ) .
this information is awaiting authentication by a species expert , and will be updated as soon as possible . if you are able to help please contact : arkive @ urltoken
anchialine coastal bodies of standing waters that have no surface connections to the ocean but display both tidal fluctuations and salinity ranges characteristic of fresh and brackish waters , indicating the presence of subsurface connections to the watertable and ocean . carapace the hard shell covering the upper surface of part of the body of a crustacean . rostrum central , forward - projecting and occasionally long spine between the eyes of crustaceans .
dr . thomas m . iliffe department of marine biology texas a & m ; university at galveston 5007 ave . u galveston tx 77551 united states of america iliffe @ urltoken http : / / www . urltoken
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mclaughlin et al . 2005 . common and scientific names of aquatic invertebrates from the united states and canada : crustaceans . american fisheries society special publication 31
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .


stictea is a genus of moths belonging to the subfamily olethreutinae of the family tortricidae .

delimitation of this genus versus strepsicrates is not fully resolved , and some species now listed here were formerly placed there .

stictea

stictea mygindiana ( = olethreutes mygindiana ) cowberry marble - norfolk micro moths - the micro moths of norfolk .
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by t . m . gilligan 1 , j . baixeras 2 , j . w . brown 3 , and k . r . tuck 4
urltoken is pleased to offer the complete world catalogue of the tortricidae ( t @ rts ) ! this is a complete list of all world species , utilizing the world catalogue published in 2005 as the foundation for the database . version 3 . 0 of the online catalogue contains 15 , 099 records representing 10 , 883 species . more than 1 , 600 records have been updated from ver 2 . 0 ( jul , 2012 ) , and more than 3 , 000 records have been updated from the original catalogue . the database is completely searchable and contains photos of over 1 , 200 type specimens .
t @ rts will be updated regularly both with corrections from the original world catalogue and with additions since its publication . as such , these pages will serve as the most up to date information on current tortricid nomenclature . if you find any errors in the data presented here or have any questions / comments , please use the contact form to send the authors an email .
we are indebted to all of the original authors of the world catalogue ( j . w . brown , j . baixeras , r . brown , m . horak , f . komai , e . metzler , j . razowski , and k . tuck ) for providing the basis for this project . we would also like to thank the dozens of individuals who have provided corrections or updates to the database since it was first placed online in 2007 .
gilligan , t . m . , j . baixeras , j . w . brown & k . r . tuck . 2014 . t @ rts : online world catalogue of the tortricidae ( ver . 3 . 0 ) . urltoken
1 colorado state university , bioagricultural sciences and pest management , 1177 campus delivery , fort collins , co 80523 , usa 2 institut cavanilles de biodiversitat i biologia evolutiva , universitat de valencia , apartat oficial 2085 , 46071 valencia , spain 3 systematic entomology laboratory - usda [ retired ] , smithsonian institution , p . o . box 37012 , national museum of natural history , washington , dc 20013 , usa 4 curator - microlepidoptera [ retired ] , entomology department ( dc2 - 2n ) , natural history museum , cromwell road , london sw7 5bd , uk
unless noted , all images on these pages are copyright © 2003 - 14 by todd gilligan . please do not download , copy , print , or otherwise distribute any images from these pages without the permission of the author . contact form .
a species of heathland and moorland , this attractive moth flies during the afternoon and early evening .
it can be found on the wing during may and june . it is a species of northern britain , occurring from staffordshire northwards to scotland and into ireland .
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 .
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at cawston ? in barrett ' s 1873 norfolk list . barrett notes
i have had no opportunity of identifying this species
.
recorded in 1 ( 1 % ) of 69 10k squares . first recorded in 1874 . last recorded in 1874 .
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
confused by a class within a class or an order within an order ? please see our brief essay .
to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
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museum für naturkunde leibniz - institut für evolutions - und biodiversitätsforschung invalidenstr . 43 10115 berlin germany e - mail : fauna - eu ( at ) mfn - berlin . de website : urltoken
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notes : nationally scarce ( nb ) on heathland and moorland throughout much of northern england , scotland and western ireland . not recorded in hampshire or on the isle of wight to date . wingspan 15 - 20 mm . differs from o . arbutella by its larger size and the characteristic purplish brown coloration of the forewing , which in o . arbutella is purplish red , and by the generally more broadly fasciate markings ; in o . mygindiana the head is dark grey but in o . arbutella it is purplish red or ferruginous ; the male may be readily distinguished from both sexes of o . arbutella by its light grey hindwing [ bradley ] . larva feeds on cowberry , bearberry and bog - myrtle , living between shoots spun together with silk .
please note that the nbn gateway map service has been terminated as of 1 april 2017 .
as soon as a replacement map service is available , distribution maps will hopefully appear here again .
in the meantime , you can get some idea of distribution from the nbn atlas website .
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gogoselachus ( " shark from the gogo formation " ) is an extinct genus of cartilaginous fish known from the late devonian of australia .

it is one of the earliest well-preserved devonian chondrichthyans ( 20 million years older than cladoselache ) , as much more of the fish than just teeth and scales were preserved .

this rare preservation reveals some unique discoveries about the evolution of the cartilage that was inside later cartilaginous fish such as sharks , rays , and chimaeras .

gogoselachus

what gogoselachus might have looked like , as restored from the scant fossil remains .
gogoselachus lynnbeazleyae : the first recorded shark from the late devonian gogo formation of western australia .
the fossil , named gogoselachus lynnbeazleyae , is the first shark to be found in this area .
right scapulocoracoid wam 09 . 6 . 145–007 of gogoselachus lynbeazleyae , . . . | download scientific diagram
sciency thoughts : gogoselachus lynnbeazleyae : the first recorded shark from the late devonian gogo formation of western australia .
teeth of gogoselachus are distinctive with many small cusps . the image far right is a ct - scanned tooth showing internal structure .
gogoselachus was clearly a fast - swimming predator that hunted other fishes using its jagged teeth to snare prey . gogoselachus lived on an ancient reef that teemed with many kinds of large predatory placoderm fishes , so had to hold its own in this piscine rat race .
gogoselachus lynnbeazleyae discovered by professor john long at the gogo fossil site , australia . testing shows that the three - dimensional remnant s… | pinteres…
however , when the authors recently examined the microstructure of the cartilage of gogoselachus using micro ct scanning , they discovered the matrix holding the cartilage together in the jaw contained bone cells .
structure of gogoselachus lynbeazleyae endoskeleton . ( a–c ) gogoselachus calcified cartilage ( a ) sem showing tessellate layout ( b , c ) horizontal section through tissue ( d ) transverse ct scan of right meckel ' s cartilage . scale bar is 0 . 1 mm in ( c ) . abbreviations : cp , cell processes ; l , lacunae . long et al . ( 2015 ) .
publishing their findings in the journal plos one , scientists said the fossil of gogoselachus lynbeazleyae dates to the late devonian period , around 380 million years ago , and shows the evolutionary transition from bones to cartilage .
concluding , the authors said the shark is an
exceptionally preserved
fossil that
suggests gogoselachus represents a transitional step toward the tessellated prismatic calcified cartilage which is today recognised as the main diagnostic character of the chondrichthyes
.
but this idea has just been challenged due to the discovery , announced today in the journal plos one , of a 380 - million - year - old fossil shark from western australia named gogoselachus lynbeazleyae that shows remnant bone cells present in its cartilaginous skeleton .
the specimen described by long et al . comprises a set of disarticulated but clearly associated remains comprising both left and right meckel ' s cartilages , nasal cartilage , ceratohyal , hyomandibula , basibranchial cartilage , both scapulocoracoids , and associated teeth and scales . this is described as gogoselachus lynbeazleyae , where ‘ gogoselachus ’ means gogo - shark and ‘ lynbeazleyae ’ honours lyn beazley of the university of western australia for her ‘contribution to scientific progress in western australia’ . the specimen is thought to be the first known acid - prepared mineralized devonian shark remains .
gogoselachus lynbeazleyae is an exceptionally preserved acid - prepared fossil chondrichthyan from the late gogo formation devonian of australia . its lower jaws have an expanded cotylus and small mandibular knob , and the slender scapulocoracoid has just 2 facets for radial articulations . the teeth have well - developed labial cusplets and their structure is intermediate between the ctenacanthiform and symmoriiform condition . the distinctive calcified cartilage forming the endoskeleton has multiple layers of nonprismatic subpolygonal tesserae separated by a cellular matrix . this suggests gogoselachus represents a transitional step toward the tessellated prismatic calcified cartilage which is today recognized as the main diagnostic character of the chondrichthyes .
gogoselachus cartilage showing the separate units called tesserae making the up the lower jaw ( left ) , and a thin section showing bone cells ( red line ) inside the matrix which binds the tesserae together ( right ) . image on the left is 0 . 5mm across , image on right is about 0 . 1mm across .
gogoselachus lynbeazelyae gen . et sp . nov . long , burrow , ginter , maisey , trinajstic , coates , young , senden urn : lsid : zoobank . org : act : ef04ad3e - 865d - 4fa5 - a6ff - f4ed6359fd26 ; urn : lsid : zoobank . org : act : 2ed167f8 - 9506 - 4d43 - b08e - 237291937b11
preparation of gogoselachus lynbeazleyae from the gogo formation , western australia . ( a ) meckel ' s cartilage as exposed on collection , before acetic acid etching . ( b ) specimen during early acid preparation . ( c ) meckel ' s cartilages after full preparation , shown articulated in dorsal view . long et al . ( 2015 ) .
( a–c ) gogoselachus calcified cartilage ( a ) sem wam 09 . 6 . 145–031 showing tessellate layout ( b , c ) horizontal section through tissue wam 09 . 6 . 145–032 ( d ) transverse ct scan of right meckel ' s cartilage . scale bar = 0 . 1 mm in ( c ) . abbreviations : cp , cell processes ; l , lacunae .
fig 4 . right scapulocoracoid wam 09 . 6 . 145–007 of gogoselachus lynbeazleyae , gogo formation , western australia . ( a ) lateral ( b ) posterior and ( c ) medial views , with close up of articulatory area for pectoral fin in e , f . d , articulation area for pectoral fin articulation on left scapulocoracoid for comparison . abbreviations as for fig 3 .
the picture above shows the specimen ( b ) during early acid bath preparation , whilst ( c ) shows the meckelian cartilage once it has been removed from the surrounding rock . the species has been named gogoselachus lynbeazleyae . the genus nomenclature comes from the name of the geological formation and the greek “selachos” which means shark . the trivial name honours neuroscientist professor lyn beazley ( university of western australia ) , in acknowledgement of her role as an ambassador for science .
here we present new data from the first well - preserved chondrichthyan fossil from the early late devonian ( ca . 380–384 mya ) gogo formation lägerstatte of western australia . the specimen is the first devonian shark body fossil to be acid - prepared , revealing the endoskeletal elements as three - dimensional undistorted units : meckel’s cartilages , nasal , ceratohyal , basibranchial and possible epibranchial cartilages , plus left and right scapulocoracoids , as well as teeth and scales . this unique specimen is assigned to gogoselachus lynnbeazleyae n . gen . n . sp .
gogoselachus lynbeazleyae is represented by a set of associated elements which were probably held together by ligaments and muscular tissue when the concretion formed around them . they include both left and right meckel ' s cartilages , nasal cartilage , ceratohyal , hyomandibula , basibranchial cartilage , both scapulocoracoids , as well as associated teeth and scales . fig 1 shows the specimen as it was found in the field ( fig 1a ) , after a week in acetic acid ( fig 1b ) , and after the lower jaws had been freed and reassembled into life position ( fig 1c ) .
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i thought you might be interested in this item at urltoken title : plos one publisher : san francisco , ca : public library of science . isbn / issn : 1932 - 6203 oclc : 969745500
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long ja , burrow cj , ginter m , maisey jg , trinajstic km , et al . ( 2015 )
correction : first shark from the late devonian ( frasnian ) gogo formation , western australia sheds new light on the development of tessellated calcified cartilage . plos one 10 ( 6 ) : e0131502 .
the meckel’s cartilages show a jaw articulation surface dominated by an expansive cotylus , and a small mandibular knob , an unusual condition for chondrichthyans . the scapulocoracoid of the new specimen shows evidence of two pectoral fin basal articulation facets , differing from the standard condition for early gnathostomes which have either one or three articulations . the tooth structure is intermediate between the ‘primitive’ ctenacanthiform and symmoriiform condition , and more derived forms with a euselachian - type base . of special interest is the highly distinctive type of calcified cartilage forming the endoskeleton , comprising multiple layers of nonprismatic subpolygonal tesserae separated by a cellular matrix , interpreted as a transitional step toward the tessellated prismatic calcified cartilage that is recognized as the main diagnostic character of the chondrichthyans .
citation : long ja , burrow cj , ginter m , maisey jg , trinajstic km , coates mi , et al . ( 2015 ) first shark from the late devonian ( frasnian ) gogo formation , western australia sheds new light on the development of tessellated calcified cartilage . plos one 10 ( 5 ) : e0126066 . urltoken
academic editor : andrew a . farke , raymond m . alf museum of paleontology , united states
copyright : © 2015 long et al . this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited
funding : this work was supported by grants from the australian research council dp 0772138 awarded to gcy and jal for travel and scanning of specimens at the anu ; and dp1092870 awarded to kmt for travel to collaborate . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
the fossils occur inside limestone concretions which formed around whole fish or parts of carcasses that fell into the deep inter - reef basins between the reef fronts . rapid calcitic concretion formation prevented compaction or distortion from tectonic or gravity - induced sedimentary loading , thus retaining the original shape of the bones and delicate perichondrally calcified cartilages [ 34 ] . geochemical studies indicate that carcasses fell through a euxinic layer within the basin , enabling rare rapid preservation of soft tissue in anoxic benthic conditions [ 43 ] .
( a ) meckel ' s cartilage as exposed on collection , before acetic acid etching . ( b ) specimen during early acid preparation . ( c ) meckel ' s cartilages wam 09 . 6 . 145–001 ( left ) , wam 09 . 6 . 145–002 ( right ) after full preparation , shown articulated in dorsal view .
the field work was done in australia with permission of the land owners and leaseholders . there are no regulations pertaining to collecting of fossils in this region of australia apart from land owner permission as stated under the lands administration act , as relevant to pastoral leaseholders : see urltoken .
the specimen is registered in the collections of the western australian museum as wam 09 . 6 . 145 , repository at 69 kew st , welshpool , western australia .
the specimen was acid prepared at museum victoria , melbourne by jal using 10 % acetic acid , with cartilage elements strengthened by mowital b30 in ethanol . the specimen and residues were washed in water to neutralize the acidity , then picked under a binocular microscope to retrieve isolated scales and teeth .
scanning electron micrographs of teeth , scales and cartilage structure were taken using a jeol - 6400 at the centre for microscopy , university of queensland ( scales ) , and using other sems at university of warsaw , poland ( teeth images ) , and the los angeles county museum of natural history , california ( cartilage ) . thin section images were taken using an olympus bx - 50 microscope with dp12 imaging system .
digital transverse sections of the cartilages were studied through composite serial reconstruction from 10μ slices by the ultrafine ct scanner in the dept of applied mathematics , anu , and studied using drishti 2 . 4 software developed by that department ( and now publically available as freeware ) .
the electronic edition of this article conforms to the requirements of the amended international code of zoological nomenclature , and hence the new names contained herein are available under that code from the electronic edition of this article . this published work and the nomenclatural acts it contains have been registered in zoobank , the online registration system for the iczn . the zoobank lsids ( life science identifiers ) can be resolved and the associated information viewed through any standard web browser by appending the lsid to the prefix
urltoken
. the lsid for this publication is : urn : lsid : zoobank . org : pub : 3563562e - 41e0 - 4579 - 8645 - 9c13f7880019 . the electronic edition of this work was published in a journal with an issn , and has been archived and is available from the following digital repositories : pubmed central , lockss .
qm , queensland museum , po box 3300 , south brisbane bc , queensland 4101 .
wam , western australian museum fossil collection ; 49 kew st , welshpool , wa , 6106 .
generic name for the gogo formation and greek ‘ selachos ’ , shark . species name acknowledging professor lyn beazley , of the university of western australia , for her contribution to scientific progress in western australia .
? deihim mansureae
dubious specimen aeu 236
ginter et al . ( 2002 ; plate 4 , figures j–k ) [ 45 ] .
shark
long & trinajstic ( 2010 ; p . 263 , figure 2 ) [ 34 ]
circa 60 km se of fitzroy crossing , north western australia , near the stromatoporoid camp locality , collected by jal , july 7th , 2005 .
a chondrichthyan with cladodont type teeth characterised by the following combination of features : a crown with a prominent median cusp rounded in cross - section , two smaller lateral cusps , and , in asymmetrical forms , a single intermediate lateral cusplet situated between the median cusp and the mesial lateral cusp ; a row of slender accessory labial cusplets at the crown - base interface ; a euselachian type base with a deep aboral depression and a vague orolingual hump . meckel’s cartilage with wide , transverse articular cotylus and small mandibular knob ; ceratohyal with a deep anterior - facing pit towards the posterior of the outer surface ; elongate basibranchial with straight edges ; scapulocoracoid with two diazonal nerve foramina and two pectoral fin basal articulation facets .
the meckel’s cartilages ( figs 1a–1c ; 2a , 2b and 2g ) are c . 5 cm long , deep posteriorly , with a well - developed continuous dental sulcus for the tooth rows ( sul ) , bordered laterally by a raised straight rim . the outer face of the cartilage has a ventral ridge ( fig 1a , vr ) defining a depressed lateral surface for the insertion of the adductor mandibulae musculature . the lower jaw articulatory surface has a broad transverse cotylus ( fig 2g , cot ) flanked medially by a small mandibular knob ( m . kb ) . both the cotylus and the mandibular process are on the same transverse plane . posterior to the cotylus is a thin vertical lamina ( fig 2a and 2c , re . fl ) , the sustentaculum sensu gegenbaur [ 46 ] a structure that has been recently referred to as the retroarticular flange [ 22 ] . the anterior region of the meckel’s cartilage has a well - defined oval symphysial attachment area ( fig 2b , sym ) with a separate subrectangular muscle attachment area posteroventral to the symphysis ( ma ) .
( a–c ) left meckel’s cartilage wam 09 . 6 . 145–001 , medial , lateral and dorsal views . ( d , e ) ceratohyal wam 09 . 6 . 145–005 , lateral and dorsal views . ( f ) nasal cartilage wam 09 . 6 . 145–006 , anterior vie w ? . ( g ) ? epibranchial cartilage wam 09 . 6 . 145–004 . ( h ) basibranchial cartilage wam 09 . 6 . 145–003 . abbreviations : cot , cotylus ; fl . ch , flange on ceratohyal ; fo . ch , fossa on ceratohyal ; ma , muscle attachment area ; m . kb , mandibular knob ; re . fl , retroarticular flange ; sul , sulcus ; sym , symphyseal pit ; vr , ventral ridge .
the ceratohyal ( fig 2c and 2d ) conforms to the generalized chondrichthyan shape [ 44 , 46 ] , having an expanded posterior blade and narrow , slightly rotated anterior region . it is about three - quarters the length of the meckel’s cartilage , and has a large fossa ( fo . ch ) facing anteriorly at its posterior extremity , rimmed by an elevated flange ( fl . ch ) . the anterior end of the ceratohyal is slightly expanded , with two separate , slightly rugose areas on the medial surface . it bears no distinctive grooves or foramina .
a delicate , triangular double - lamina of twisted cartilage resembles a nasal cartilage ( fig 2f ) . it correlates closely to the size and shape of other primitive chondrichthyan nasal cartilages or parts of the nasal capsule [ 14 ] . a shorter , tubular cartilage preserved above the meckel’s cartilages is one of the gill arches ( fig 2e ) , most likely epibranchial 1 by comparison with ozarcus [ 21 ] . a single elongate and transversely concave cartilage plate probably represents the mineralized surface of a basibranchial cartilage ( fig 2h ) . it has a straight posterior margin with dorsally projecting corners ; the anterior end of the unit is m - shaped . midway along , its convex surface is rough with numerous small pores .
the scapulocoracoid ( figs 3 and 4 ) is 8 . 0 cm high , with an elongate , flat , dorsally tapering scapular region and a deeply concave coracoid region ventral to a thickened posterior ridge at the lateral inflection . the scapular blade has a weak posterolateral process ( pla ) . muscle attachment scars are interpreted as follows : the supinator muscle anteriorly on the lateral surface of the scapula ( sup ) ; the mediolateral pectoral retractor muscles near the anterolateral inflection ( flr ) ; the coracobranchialis muscles ( cobr ) above the inflection ridge ; the coracohyoideus muscles ( cohy ) ventrally ; and the pectoral depressor muscle sheet ( pdm ) on the posterolateral surface of the coracoid ( fig 3a and 3b ) . a ridge ( ri ) along the posterior angle of the ventrolateral inflection shows remnants of areas with raised borders enclosing surfaces presumed to be articulation facets for the pectoral fin basal cartilages . the small central facet is completely preserved on the left side ( fig 3b and 3d , pf . ar2 ) , but the lateral and medial surfaces of the ridge have broken off . the medial area shows remnants of a rim shared with the central area , but the lateral area is missing .
the left scapulocoracoid is slightly damaged in the area of the fin articulation but clearly shows two facets for fin radial articulations ( fig 3d and 3e ) . this interpretation is supported by features preserved in the right scapulocoracoid which has a broad triangular facet for the mesial pectoral fin radial attachment ( fig 4b and 4e , pf . ar1 ) clearly visible , as well as a slightly smaller lateral pectoral fin radial facet ( pf . ar2 ) . restorations combining data from both sides show that there was only two fin radial articulations ( fig 3e , fig 4f ) .
( a ) posterior ( b ) medial and ( c ) lateral views . d , close up of posterior face showing articulation area for pectoral fin , e . , interpretation of same area . f , close up of central lateral surface showing diazonal foramen ( df ) . abbreviations : af , articulation facets ; artc , articular crest ; br , break in bone ; cobr , coracobrachialis muscle attachment area ; cohy , coracohyoideus muscle attachment area ; cp , coracoid plate ; df , diazonal foramina ; muscle attachment areas ; flr , mediolateral pectoral retractor muscle attachment area ; pdm , pectoral depressor muscle attachment area ; pla , posterolateral process ; pf . ar1 , 2 , pectoral fin articulation areas 1 and 2 ; pla , posterolateral angle ; pva , posteroventral angle ; ri , ridge ; sup , supinator attachment area .
( a ) lateral ( b ) posterior and ( c ) medial views , with close up of articulatory area for pectoral fin in e , f . d , articulation area for pectoral fin articulation on left scapulocoracoid for comparison . abbreviations as for fig 3 .
the dentition is represented by some 82 individual cladodont - type teeth which show limited morphological variation ; more than half are relatively large ( width at the base–crown junction 1 . 5–2 mm ; figs 5a–5e ; 6a–6f and 6j–6t ) with the others smaller ( all about 1 mm wide ; fig 6g–6i ) . two types of crowns , symmetrical and asymmetrical , were observed among the larger teeth . symmetrical crowns ( fig 6d–6f and 6n–6t ) have a prominent median cusp , standing upright ( in labial view ) or with a slight distal inclination , and only two divergent lateral cusps whose height usually exceeds one - third of the height of the median cusp . no intermediate cusplets are present . in asymmetrical crowns ( figs 5a–5d ; 6a–6c , 6j–6m ) the median cusp is considerably inclined distally ( up to 20 degrees ) , the distal lateral cusp is inclined distally at 45 degrees and the mesial lateral cusp is only slightly inclined mesially . between the median and mesial cusps there is a small intermediate cusplet . the asymmetrical teeth are wider ( mesio - distally ) than the symmetrical ones . the median cusp is up to 2 . 5 mm in height in symmetrical teeth ; in asymmetrical teeth it is relatively shorter and thicker . in all types of teeth it is rounded in cross section , gently curved lingually , but non - sigmoidal . the labial face is ornamented with two to three coarse cristae almost reaching the tip ; the cristae may bifurcate at the basal part . the lingual face is almost smooth , only one or two indistinct cristae extend along the lateral margins . in asymmetrical specimens they are visible near the mesial margin ( fig 6c and 6j ) . the lingual and labial faces are separated with a lateral carina . the shape and ornamentation of the lateral cusps is similar to that of the median cusp .
large tooth wam 09 . 6 . 145–009 in ( a ) lingual , ( b ) labial , ( c ) oral ( d ) aboral views , in natural light . ( e ) sem of tooth wam 09 . 6 . 145–010 , naturally broken near the median surface . ( f , g ) ct - scans of tooth wam 09 . 6 . 145–011 , showing layout of vascular canals , in basal and lingual - lateral views . abbreviations : l - lcl , labio - lingual canals ; ctcl , central canal ; blcl , basolabial canals ; ccl , coronal canals .
( a–c ) wam 09 . 6 . 145–012 in oral , labial , and lingual views . ( d–f ) wam 09 . 6 . 145–013 in labial , basal , and oblique lateral views . ( g–h ) small tooth , wam 09 . 6 . 145–014 , in lingual , and oral views . ( i ) tooth of intermediate size , wam 09 . 6 . 145–015 , in labial view . ( j–m ) wam 09 . 6 . 145–016 in lingual , labial , oral and distal views . ( n–p ) wam 09 . 6 . 145–017 in oral , labial , and lingual views . ( q–t ) wam 09 . 6 . 145–018 in labial , lingual , oral , and lateral views .
at the crown–base interface , on the labial side of all the teeth , is a row of several accessory cusplets . they display various sizes and directions , but generally are slender , sharp and directed orally . in the asymmetrical specimens the largest may exceed the size of the intermediate cusplet . the labial accessory cusplets are vulnerable to abrasion and in most cases they are broken , leaving only the basal parts preserved .
tooth bases of larger teeth are deep , elongated mesio - distally , from elliptical to broadly hexagonal , to trapezoidal in outline , with a lingual extension . a deep depression in the central / labial area of the aboral side is framed by the well developed , labially convex basolabial rim . no clear interlocking devices ( buttons or basolabial projections ) are present , save for a slight orolingual hump , observed in certain , especially small , specimens . the aboral depression and the basolabial rim might have helped in strengthening the connection between the teeth in a tooth file . the base surface , save for the immediate vicinity of the crown on the oral side and the aboral - lingual area , is perforated by numerous canal openings . the largest foramina occur on the orolingual rim and in the aboral depression .
the ct - scans and broken surfaces reveal a network of basal nutritive canals ( fig 5e–5g ) . in the lingual extension of the base there are mainly labio - lingual canals which , in most cases , extend from the orolingual rim to the lingual wall of the aboral depression . these major canals are interconnected by narrower tubes . in the area of the depression ( i . e . below the crown ; called here central canals ) , the directions change to mainly mesio - distal and vertical . and finally , the canals in the basolabial rim are mainly vertical . from the network of central canals , single vascular canals ( coronal canals ) extend into the basal parts of the median and lateral cusps .
the smaller teeth ( fig 6g–6i ) differ from the larger ones not only by the overall size , but also in proportions between the base and crown—in the smaller teeth , in most cases , the base is relatively broader ( see especially fig 6i ) . in contrast to the excellent preservation of many of the larger teeth , most of the recovered smaller teeth have broken cusps . as the teeth were not found in situ , but dispersed around the jaw cartilages , we can only speculate in reconstructing the dentition . it is most probable that the narrow , symmetrical teeth were on the anterior part of the jaw , and the broader , asymmetrical forms were in the lateral tooth families . the smaller teeth are probably juvenile ones , replaced by larger forms , but preserved on the outer side of jaw , as is common among the ctenacanthiforms [ 47 – 49 ] . the occurrence of both symmetrical and asymmetrical forms among the smaller teeth , as well as their broken ( possibly some worn ) cusps , support this view .
tooth bases resemble the “euselachian - type” ( sensu ginter 2005 ) [ 50 ] in having a spongiose structure and numerous labio - lingual canals . the absence of buttons and basolabial projections is a potential synapomorphy with chondrichthyans possessing
euselachian - type
teeth , although the presence of an orolingual hump and basolabial depression suggests that the bases of successive teeth overlapped in similar fashion to those of cladodont chondrichthyans in which buttons and projections are present [ 47 , 49 – 52 ] . the presence of one or two medially situated larger canals in bases of symmetrical teeth ( fig 6p and 6r ) , instead of evenly distributed canals of virtually the same size , possibly represents a stage between the layout in ctenacanthiforms and symmoriiforms , which have median basal canal openings ( lingual and aboral ) which are usually much larger than any other foramina , and
euselachian - type
teeth with evenly distributed canals .
( a ) presumed flank scale wam 09 . 6 . 145–019 , anterocrown view . ( b , c ) presumed flank scale wam 09 . 6 . 145–020 , anterocrown and posterior views . ( d ) presumed flank scale wam 09 . 6 . 145–021 , posterobasal view . ( e , f ) presumed flank scale wam 09 . 6 . 145–022 , posterobasal view and magnification of undersurface of crown denticulation . ( g , h ) very small scale wam 09 . 6 . 145–023 , broken , showing crown view of one half and basal view of the other half . ( i ) abraded umbellate scale wam 09 . 6 . 145–024 , crown view . ( j , k ) pultschuppe or squamae proniae scale wam 09 . 6 . 145–025 , crown and posterior views . ( l ) stellate tessera wam 09 . 6 . 145–026 , crown view . ( m ) probable branchial denticle plate wam 09 . 6 . 145–027 , lateral view . ( n ) vertical transverse section through presumed flank scale wam 09 . 6 . 145–028 . ( o ) oblique vertical section through umbellate scale wam 09 . 6 . 145–029 . ( p , q ) horizontal section through the crown of ? head tessera wam 09 . 6 . 145–030 , whole section and magnification to show spiralfasern structure . scale bars = 0 . 1 mm in all figures except ( p ) , where scale bar = 1 mm .
thin sections show that the scales grew by appositional addition of odontodes laterally and posteriorly ( fig 7n and 7o ) ; shorter , more upright odontodes are added anteriorly . the thin bone bases show several inner growth zones having separate bases , with bases underlying the whole scale formed only in the later growth periods ( fig 7n and 7o ) . each odontode has a relatively wide pulp canal , surrounded and partly infilled by ‘spiralfasern’ , and interconnected by radial and circular canals . the ‘spiralfasern’ structure ( sensu gross ) [ 54 ] , rather than being fibrous , appears to be formed by spiralling rings of atubular dentine which developed centrifugally to partly fill the wide pulp canals . fine branching dentine tubules permeate the outer solid areas of the odontodes . innermost ( oldest ) odontodes have separate ? acellular bone bases , but the youngest growth zones have bases which underlie new and old growth zones . sharpey’s fibres extended through the full - width zones , but not the individual odontode bases . horizontal section of a pultschuppe scale shows centrifugal growth zones ( fig 7p and 7q ) .
the endoskeletal elements are formed of mineralized tessellated cartilage ( fig 8 ) that varies in thickness , corresponding to the number of layers of tesserae . the meckel’s cartilages , scapulocoracoids and the median ventral section of the basibranchial unit have two layers , whereas the gill - arch cartilages and the articulatory block on the scapulocoracoid are finer , with mostly one but rarely two layers . thin sections of loose cartilage fragments ( fig 8b and 8c ) show a structure similar to typical shark tessellated calcified cartilage , with subpolygonal tesserae showing rings and waves of liesegang [ 55 ] . however , rather than having fibrous connections between the tesserae as found in modern sharks [ 56 ] , the tesserae are mostly surrounded by a matrix incorporating elongate lacunae , some with fine processes extending from them ( fig 8c ) . ct scans ( fig 9h ) show the varying thickness of the cartilage in different areas of the endoskeletal elements ( fig 8d ) .
the morphology of the teeth is unique among the devonian chondrichthyes , showing a mix of features found in several shark groups . the crown is somewhat similar , mostly in the proportions of the cusps , to that of ctenacanthus concinnus [ 64 ] and cladodoides wildungensis [ 65 , 66 ] , but the median cusps of the latter two are labially flattened whereas in gogoselache lynbeazleyae they are biconvex . the asymmetry in devonian ctenacanthids , albeit common , is never advanced to a point of occurrence of an intermediate cusplet only on one side . moreover , the tooth base in ctenacanthids is provided with a distinct orolingual button and a straight , shelf - like basolabial projection , which are missing in g . lynbeazleyae teeth .
( a–c ) arduodens flammeus hairapetian & ginter , 2009 , from chahriseh , aeu 610 , in lingual , aboral , and lateral views . ( d–g ) deihim mansureae ginter , hairapetian and klug , 2002 . ( d , e ) holotype , igpuw / ps / 5 / 1 , from hutk , in oral and lateral views . ( f , g ) putative anterior tooth , aeu 239 , from hodjedk , in labial and oral views . scale bar = 0 . 5 mm .
based on isolated teeth from the famennian of iran , ginter et al . [ 45 ] proposed a strong heterodonty within the dentition of d . mansureae . most of the specimens attributed to that species are various forms of crushing teeth . only two specimens can be considered as clutching teeth , and these were tentatively included in d . mansureae because of the structure of the base and the presence of labial accessory cusplets . it was suggested [ 45 ] that the teeth were from the anterior tooth families and the whole dentition functioned in a way similar to modern heterodontus , with a few minute anterior clutching teeth and a large lateral and posterolateral crushing apparatus ( [ 68 ] fig24c ) . however , the reconstructed whole dentitions of g . lynbeazleyae and d . mansureae were completely different , with a total absence of crushing teeth associated with wam 09 . 6 . 145 . therefore , we consider g . lynbeazleyae and d . mansureae to belong to separate , albeit probably related , genera .
a single tooth from the upper devonian of the canning basin , western australia , emerikodus ektrapelus [ 69 ] bears some similarity to the new form . the preservation of the tooth is not very good , but the form of the main parts of the crown and base are quite similar to those of g . lynbeazleyae . however emerikodus displays one very unusual feature : the row of accessory cusplets is apparently situated on the lingual , and not labial , side of the crown .
the presumed normal body scales of gogoselache lynbeazleyae appear intermediate between the ‘cladodont’ type sensu gross [ 53 ] and the ctenacanthid type sensu reif [ 70 ] . the ‘cladodont’ scale category was based on isolated scales from the middle devonian ohio bonebeds [ 54 , 71 ] and the upper devonian of iowa , rhineland and harz [ 54 ] . the scales are characterized by having a base formed of lamellar bone layers penetrated by sharpey ' s fibres , and a crown plate comprising circular or semicircular zones of odontodes ; growth was by addition to the margin of the scale crown and a new ( cellular or acellular ) bone layer under the whole base .
several occurrences of ctenacanthid and cladodont type scales have been reported from eastern and western australian middle to late devonian microvertebrate assemblages [ 72 – 74 ] . other rich assemblages from the ? upper givetian aztec siltstone at mt crean , antarctica include ctenacanthid - type scales similar to those of gogoselache , that are most likely from antarctilamna prisca [ 75 ] . the acanthodian and chondrichthyan microremains in the mt crean assemblage [ 76 ] show a marked similarity to early frasnian faunas from iran [ 77 ] .
comparing gogoselache scales with those of older taxa , the flank scales with their concave bases and polyodontode crowns resemble those of leonodus mader from the lochkovian of northern spain [ 78 ] . doliodus problematicus from the early emsian of new brunswick , canada [ 15 ] , and antarctilamna prisca young from the givetian of antarctica [ 75 ] . unlike gogoselache , all of these taxa have diplodont teeth , indicating that the scale form is probably ancestral to both shark lineages determined by tooth morphology .
the structure of the calcified cartilage in the endoskeleton of gogoselache is of interest as an apparent transitional form between gcc and pcc . ct scans and thin sections show a calcified tissue forming the outer shell of endoskeletal elements , more derived than the simple calcified cartilage comprising separate globules found in osteostracan agnathans [ 55 ] , acanthodians [ 79 ] and some stem chondrichthyans ( fig 9a and 9b ) . placoderms differ from other gnathostomes in having perichondral bone plus globular ( gcc ) and / or uncalcified cartilage forming their endoskeleton ( fig 9c and 9d ) in instances where such elements are preserved at all [ 53 , 73 , 80 ] .
we would like to thank field team members of the 2005 gogo expedition , l . hatcher , m . nossal , b . choo and m . karkeek ; and r . f . miller ( nbmg ) for image of doliodus calcified cartilage . we thank giar - ann hung for assistance with the sem photography at la county museum of natural history . dr adam summers , university of washington , provided helpful discussion on the cartilage histology . this work is a contribution to igcp 596 .
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sharks are one of the oldest and least changed of all the living back - boned jawed creatures . but because their skeletons are made of cartilage much of their early fossil record is poor .
cartilage is a rubbery tissue that forms the framework for bones to ossify ( harden ) upon . it’s why babies have rubbery legs when they begin to walk , as the bones haven’t fully ossified around the cartilage cores . our ears and noses have cartilage frameworks too , which lack bone , but still support the soft structures we hear and smell with .