conorbiinae was a subfamily of small to quite large sea snails , marine gastropod mollusks in the family conidae .

this subfamily has also been written by several authors as " conorbinae " .

( and was a long time considered a subfamily of the turridae . bouchet , kantor et al. elevated in 2011 the subfamily conorbiinae to the rank of family conorbidae . this was based on anatomical characters ( radular tooth and shell characters ) and a dataset of molecular sequences of three gene fragments

conorbiinae

vera - peláez . 2004 . genotina genotae new species and new genus and genota nigeriensis new species of the subfamily conorbiinae ( gastropoda , turridae ) . systematic , biogeography , stratigraphy and phylogeny of conorbis , genotina and genota genera . pliocenica , 4 : 95 - 106
vera - peláez . 2004 . genotina genotae new species and new genus and genota nigeriensis new species of the subfamily conorbiinae ( gastropoda , turridae ) . systematic , biogeography , stratigraphy and phylogeny of conorbis , genotina and genota genera . pliocenica , 4 : 95 - 106 [ details ]
( of genota nigeriensis vera - peláez , 2004 ) vera - peláez . 2004 . genotina genotae new species and new genus and genota nigeriensis new species of the subfamily conorbiinae ( gastropoda , turridae ) . systematic , biogeography , stratigraphy and phylogeny of conorbis , genotina and genota genera . pliocenica , 4 : 95 - 106 [ details ]
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appendix 2 : list of all the family and subfamily names available for the conoidea .
a new genus - level classification of the conoidea is presented , based on the molecular phylogeny of puillandre et al . in the accompanying paper . fifteen lineages are recognized and ranked as families to facilitate continuity in the treatment of the names conidae ( for ‘cones’ ) and terebridae in their traditional usage . the hitherto polyphyletic ‘turridae’ is now resolved as 13 monophyletic families , in which the 358 currently recognized genera and subgenera are placed , or tentatively allocated : conorbidae ( 2 ( sub ) genera ) , borsoniidae ( 34 ) , clathurellidae ( 21 ) , mitromorphidae ( 8 ) , mangeliidae ( 60 ) , raphitomidae ( 71 ) , cochlespiridae ( 9 ) , drilliidae ( 34 ) , pseudomelatomidae ( = crassispiridae ) ( 59 ) , clavatulidae ( 14 ) , horaiclavidae new family ( 28 ) , turridae s . s . ( 16 ) and strictispiridae ( 2 ) . a diagnosis with description of the shell and radulae is provided for each of these families .
comparison of the last two conoidean classifications with the new classification proposed in this article . a . taylor et al . ( 1993 ) and bouchet & rocroi ( 2005 ) . 1 elevated to familial rank by bouchet & rocroi ( 2005 ) . 2 absent in taylor et al . ( 1993 ) . b . proposed classification . c . tucker & tenorio ( 2009 ) . † fossil taxa .
genus - group names have been allocated to the newly defined families based on the following critera : our aim was to provide an exhaustive list of conoidean genus - group names based on recent type species or reasonably recognized as recent in the literature . genera present exclusively in the fossil record are not included . in very few cases , obviously erroneous attribution of recent species to entirely fossil genera also excluded these genera from our listing . junior homonyms and nomina nuda are also not included because they are unavailable for nomenclatural purposes . when the status of a nominal genus is uncertain , we adopted a ‘valid until synonymized’ approach ; that is , a name is regarded valid ( irrespective of our opinion ) if it has not been synonymized in literature . the names of synonyms are placed after corresponding valid names ( the synonyms applied to the valid subgenus name are positioned after the subgenus name ) . all the genus and subgenus names are also listed in alphabetic order in appendix 1 .
( 1 ) 224 genera were assigned to a family based on shell characters , and phenetic resemblance to those genera with radula and / or molecular characters available ; those are marked 1 .
( 2 ) 103 genera were classified on the basis of radula morphology ( both our own and published data ) , and congruence between radula and molecular characters for those genera that were sequenced ; these are marked 2 .
( 3 ) 98 genera were classified in a family on the basis of the molecular data ; these are marked 3 .
( 4 ) 63 genera and subgenera do not fall easily into the morphological groups resulting from the preceding steps , but they have tentatively been assigned to a family as a working hypothesis ; these genera are preceded by a question mark .
( 5 ) 173 subgenera and / or synonyms are also listed ; subgenera are in parentheses , synonyms in square brackets .
to illustrate the range of radular types in the molecularly defined clades , the radulae were extracted as far as possible from the specimens used for the molecular analyses or from conspecific specimens ( after rehydration when soft parts had been dried ) , cleaned with diluted bleach , rinsed in distilled water , mounted on stubs , air dried , coated with gold - palladium , and investigated with a jeol jsm 840a scanning electron microscope . some of the previously photographed radulae ( examined by yuik and / or j . d . taylor ) were additionally illustrated .
for radular descriptions we mostly followed the terminology accepted and discussed by kantor & taylor ( 2000 ) . in radular formulae , the parentheses indicate partial or complete fusion of lateral and rachidian teeth ( for more details , see kantor , 2006 ) .
although the names conoidea and toxoglossa are used interchangeably in the taxonomic literature , we have avoided the name toxoglossa because ( i ) it is not typified and cannot be used for a family–group name , and ( ii ) many of the included taxa do not have a toxoglossate radula . within the conoidea , ranking of the clades was determined by a conservative approach , thus retaining the names terebridae and conidae s . s . ( the latter including the cone snails profundiconus , californiconus , conasprella , conus and taranteconus ) in their accustomed usage at family rank . an alternative would have been to recognize only two families , conidae sensu taylor et al . ( 1993 ) and turridae ( including the turridae , terebridae , drilliidae and pseudomelatomidae ) , with the resulting inconvenience that conidae s . s . and terebridae , although monophyletic , would lose their traditional usage and the vast associated literature dealing with these names that largely ignores the taylor et al . ( 1993 ) and bouchet & rocroi ( 2005 ) classifications .
fourteen clades of rank equivalent to conidae s . s . and terebridae are recognizable from the molecular phylogenetic tree ( puillandre et al . , 2011 ) . forty - three family–group names within conoidea are nomenclaturally available ( appendix 2 ) , of which five are based on a genus with a fossiltype species ( andoniinae vera - pelaez , 2002 ; cryptoconinae cossmann , 1896 ; hemiconidae tucker & tenorio , 2009 ; johnwyattidae serna , 1979 ; siphopsinae le renard , 1995 ) . these five cannot be applied to a molecular clade and will not be used in our classification . names were applied to clades based on the position of their type genus in the tree . if more than one family–group name was applicable , the valid name was determined by priority . if the type genus of a nominal family name was not sequenced , application of the name was determined by reference to the morphologically most similar genus used in the analysis .
because the molecular taxon sampling is still too patchy for such levels , we have abstained from extending the classification below family level ( i . e . subfamilies , tribes ) , even when some molecular clades obviously match previously recognized ‘subfamilies’ ( e . g . californiconinae , oenopotinae , zemaciinae , zonulispirinae ) . they are all included in a family ( appendix 2 ) , without precluding their usefulness and potential taxonomic validity .
each family ( except the conidae and terebridae , already extensively covered and illustrated in other recent works , e . g . röckel , korn & kohn , 1995 ; terryn , 2007 ; tucker & tenorio , 2009 ) is illustrated by one or several shells , radulae and protoconchs , covering the morphological variability of the group . as far as possible , specimens used for the molecular analyses were used for illustration . however , since a substantial part of the morphological variability was not covered by our dataset , shells and radulae of other specimens are also illustrated .
diagnosis : shell medium - sized to large or very large , normally 20–50 mm , up to 170 mm high , conical or biconical , with narrow aperture and short siphonal canal . shell with considerable internal remodelling due to inner wall resorption . spiral sculpture usually developed , axial sculpture absent or in form of shoulder tuberculation . anal sinus on subsutural ramp , shallow to moderately deep . operculum present , small , with terminal nucleus . radula of marginal hypodermic teeth , generally harpoon - shaped , barbed at tip , often with complex inner structure of folds and serration , base small and swollen , tooth canal opening ( sub ) terminally , rarely laterally . subradular membrane vestigial . teeth can be attached to the membrane by long or very long flexible ligament . tooth wall forms several overlapping layers .
remarks : in conidae and the other families that are included in this major clade ( family conidae sensu taylor et al . , 1993 ) , the radula consists only of marginal teeth that are usually enrolled with completely overlapping edges ( hypodermic ) . teeth at their formation in the radular sac are already enrolled and they are attached to the radular membrane only by their bases , sometimes through a long flexible ligament . in the ‘turrids’ with enrolled teeth , these are attached to the membrane along most of its length ( kantor & taylor , 2000 ) . a molecular phylogeny of the conidae is currently in preparation ( c . meyer , personal communication ) .
diagnosis ( from tucker & tenorio , 2009 ) : “radular tooth : anterior fold is usually present ; basal spur is directed toward the apex of the tooth or parallel with the tooth base ; the waist , base and c - fold are absent ; terminating cusps , serratins and accessory process are also absent . shell characters : the interior of the shell is extensively remodelled including the columellar region ; nodules are absent but cords may be present . shells can be squatly conical to elongated or biconical . ”
radulae . borsoniidae . a . genota mitriformis ( wood , 1828 ) * * , mnhn im200742293 ( shell : fig . 2 m ) . b . borsonia sp . * * , mnhn im200717932 ( shell : fig . 2 d ) . c . bathytoma neocaledonica puillandre et al . , 2010 * * , mnhn im200717857 ( shell : fig . 2 e ) . d . borsoniidae gen . 1 * * , mnhn im200717911 ( shell : fig . 2 b ) . symbol : * * , sequenced specimen .
diagnosis : shell small to large ( 5–80 mm ) , fusiform to biconic , sometimes with strong to obsolete columellar pleats . sculpture usually well developed , axial ribs sometimes obsolete to absent . siphonal canal short to moderately long . anal sinus on subsutural ramp , deep . protoconch when multispiral with up to five whorls , initially smooth and then with arcuate axial riblets , when paucispiral up to two smooth whorls . operculum with terminal nucleus , fully developed to missing . radula of hypodermic marginal teeth that usually have a weakly developed solid basal part , often attached to the ligament ( marked by an arrow on fig . 3 c ) . tooth canal opening ( sub ) terminally or , sometimes , laterally . at their tip teeth can have weak to rather strong barb ( s ) ( genota , fig . 3 a ) . overlapping of the tooth edges is weak ( fig . 3 c ) . in zemacies , the radula is completely absent .
remarks : this is a rather heterogeneous group . obviously , it is not fully resolved as it is based on molecular data and comprises rather conchologically different clades . this could be explained by the fact that many taxa of this group are among most ancient of conoideans , known since the palaeocene ( zemacies , borsonia , tomopleura ) or eocene ( bathytoma , genota , microdrillia ) . the loss of apomorphies by mutation could be more important for old taxa ( puillandre et al . , 2011 ) .
the names borsoniinae bellardi , 1875 , and pseudotominae bellardi , 1875 , were established simultaneously . as first revisers , under art . 24 of the iczn code , we here give precedence to the former over the latter .
shells . a–f . clathurellidae . a . nannodiella ravella ( hedley , 1922 ) * , mnhn im200742350 , philippines , panglao 2004 , st . t9 , 09°33 . 5′n , 123°49 . 5′e , 97–120 m , sl 5 . 8 mm ( radula : fig . 5 a ) . b . comarmondia gracilis ( montagu , 1803 ) , mnhn , le brusc , cap sicié , provence , france , 40–100 m , sl 23 mm . c . etrema cf . tenera ( hedley , 1899 ) * * , mnhn im200717869 , philippines , panglao 2004 , st . s21 , 09°41 . 7′n , 123°50 . 9′e , 4–12 m , sl unknown ( broken ) . d . glyphostoma rostrata sysoev & bouchet , 2001 , mnhn , new caledonia , bathus 4 , st . dw896 , 20°16′s , 163°52′e , 315–350 m , sl 21 . 5 mm . e . lienardia nigrotincta ( montrouzier , 1872 ) * , mnhn , touho , new caledonia , 20°45 . 2′s , 165°16 . 3′e , intertidal , sl 6 . 9 mm ( radula : fig . 5 b ) . f . strombinoturris crockeri hertlein & strong , 1951 , lacm 747–37 , off isabel island , mexico , 18–33 m , sl 47 . 5 mm . g–i : mitromorphidae . g . lovellona atramentosa ( reeve , 1849 ) * , mnhn , philippines , panglao 2004 , st . m2 , 09°32 . 8′n , 123°45 . 9′e , 0–2 m , sl 9 . 0 mm ( radula : fig . 6 a ) . h . mitromorpha metula ( hinds , 1843 ) * , mnhn im200742339 , philippines , panglao 2004 , st . b8 , 09°37 . 1′n , 123°46 . 1′e , 3 m , sl 3 . 1 mm ( radula : fig . 6 b ) . i . anarithma sp . * , mnhn , philippines , panglao 2004 , st . s5 , 09°37 . 1′n , 123°46 . 1′e , 2–4 m , sl 6 . 9 mm ( radula : fig . 6 c ) . abbreviation and symbols : sl , shell length ; * , sequenced species ; * * , sequenced specimen . photo credits : b . buge ( c ) , m . g . harasewych and d . tippett ( f ) , p . maestrati ( d , e ) .
radulae . clathurellidae . a . nannodiella ravella ( hedley , 1922 ) * , mnhn im200742350 ( shell : fig . 4 a ) . b . lienardia nigrotincta ( montrouzier , 1872 ) * , mnhn ( shell : fig . 4 e ) . c . lienardia jousseaumei ( hervier , 1896 ) , mnhn , philippines , panglao 2004 , st . b7 , 9°35 . 9′n , 123°51 . 8′e , 4–30 m . symbol : * , sequenced species .
radulae . mitromorphidae . a . lovellona atramentosa ( reeve , 1849 ) * , mnhn ( shell : fig . 4 g ) . b . mitromorpha metula ( hinds , 1843 ) , mnhn im200742339 ( shell : fig . 4 h ) . c . anarithma sp . * , mnhn ( shell : fig . 4 i ) . symbol : * , sequenced species .
diagnosis : shell small - to medium - sized , 3–30 mm , usually 5–10 mm high , biconic , of mitriform shape . sculpture rather smooth , with dominant spiral elements . aperture narrow , with or without 1–3 columellar pleats , sometimes with denticles within . siphonal canal short or indistinct . anal sinus from indistinct to rather shallow indentation on weakly pronounced subsutural ramp . protoconch multispiral or paucispiral , up to 4 . 5 smooth whorls . no operculum . radula of hypodermic , marginal , relatively short , awl - shaped teeth with large swollen solid basal part ( fig . 6 ) . distinct ligaments present , short . tooth canal opening subterminally or laterally . at their tip , teeth can have a weak barb ( fig . 6 c ) .
remarks : mitrolumninae was established as a substitute name for diptychomitrinae . mitromorphidae and mitrolumninae were published the same year but mitromorphidae ( 19 may 1904 ) has priority over mitrolumninae ( 31 august 1904 ) .
radulae . mangeliidae . a . benthomangelia trophonoidea ( schepman , 1913 ) , mnhn im200717835 ( shell : fig . 7 c ) . b . toxicochlespira pagoda sysoev & kantor , 1990 * * , mnhn im200717925 ( shell : fig . 7 o ) . c . mangeliinae gen . 2 . mnhn im200910331 , philippines , panglao 2004 , st . s26 , 9°41 . 50′n , 123°51 . 00′e , 21 m , sl unknown ( broken ) . d . eucithara cf . coronata ( hinds , 1843 ) * * , mnhn im200717900 ( shell : fig . 7 f ) . e . anticlinura sp . * * , mnhn im200742513 ( shell : fig . 7 e ) . f . mangeliidae gen . sp . , mnhn , philippines , panglao 2004 , st . t26 , 9°43 . 3′n , 123°48 . 8′e , 123–135 m . g . mangelia powisiana ( dautzenberg , 1887 ) . plymouth , england , after taylor et al . ( 1993 ) . symbol : * * , sequenced specimen .
radulae . raphitomidae . a . buccinaria pendula bouchet & sysoev , 1997 , mnhn , new caledonia , bathus 4 , st . cp948 , 20°33′s , 164°57′e , 533–610 m . b . daphnella pulvisculus chino , 2006 , mnhn , new caledonia , bathus 4 , st . dw927 , 18°56′s , 163°22′e , 444–452 m . c . gymnobela yoshidai kuroda & habe in habe , 1962 , mnhn , norfolk ridge , norfolk 2 , st . dw2058 , 24°40′s , 168°40′e , 591–1032 m . d . kermia irretita ( hedley , 1899 ) , mnhn , new caledonia , lifou 2000 , st . 1419 , 20°55 . 6′s , 167°04 . 5′e , 0–5 m . e . daphnella cladara sysoev & bouchet , 2001 , mnhn , norfolk ridge , lithist , st . cp09 , 24°53′s , 168°22′e , 518–540 m . f . daphnella mitrellaformis ( nomura , 1940 ) , mnhn , new caledonia , lifou 2000 , st . dw1649 , 20°54′s , 167°01′e , 150–200 m . g . miowateria sp . , mnhn , fidji , musorstom 10 , st . cp1354 , 17°43′s , 178°55′e , 959–963 m . h . spergo fusiformis ( kuroda & habe in habe , 1962 ) , mnhn , tongatapu , tonga , bordau 2 , st . cp1566 , 21°02′s , 175°18′w , 530–531 m .
remarks : this is the largest and most variable taxon in the conoidea , as concerns the number of species , with the largest vertical range ( intertidal to hadal depths ) .
radulae . cochlespiridae . a . cochlespira radiata ( dall , 1889 ) , mnhn , se brazil , after kantor & taylor , 2000 . b–c . sibogasyrinx sp . * * , mnhn im200717701 ( shell : fig . 11 b ) . symbol : * * , sequenced specimen .
diagnosis : shell of moderate size , about 20–30 mm , up to 100 mm high , high - pagodiform to fusiform , with a tall spire and usually a long siphonal canal . axial sculpture poorly developed or absent , subsutural ramp usually smooth . anal sinus deep , on subsutural ramp . protoconch paucispiral , smooth . operculum with terminal nucleus . radula 1–0 - r - 0–1 . rachidian broad , subrectangular or arched , with a single rather large cusp ( fig . 12 ) , rarely absent ( some aforia ) . marginal teeth duplex , with well developed accessory limb .
remarks : this is a clade with poor congruence between molecular and shell characters . the contents of most genera need revision and the family limits remain uncertain .
sibogasyrinx was proposed as a subgenus of leucosyrinx for two species characterized by a low position of the peripheral angle . during our examination of radulae of species provisionally attributed to leucosyrinx , two distinct radulae types were found , differing in the presence of the rachidian , without apparent congruence with shell morphology . in sibogasyrinx pyramidalis , the type species of sibogasyrinx , as well as in the sequenced specimen ( fig . 11 b ) , the radula is characterized by the presence of a well - developed rachidian . although the sequenced specimen is conchologically closer to typical leucosyrinx , the molecular sequence suggests affinities to cochlespira .
radulae . drilliidae . a–b . splendrillia sp . * * , mnhn im200717847 ( shell : fig . 11 l ) . c . clavus exasperatus ( reeve , 1843 ) , mnhn , new caledonia , lifou 2000 , st . 1420 , 20°47 . 7′s , 167°09 . 35′e , 4–5 m . d . imaclava pilsbryi ( bartsch , 1950 ) , after kantor & taylor , 2000 . e . cruziturricula arcuata ( reeve , 1843 ) * * , nhmuk moea 20100541 , gulf of panama , jtd - 00–34 , 08°26 . 24′n , 79°09 . 14′w , 66–68 m ( shell : fig . 11 i ) . symbol : * * , sequenced specimen .
diagnosis : shell small - to medium - sized , usually 15–25 mm , up to 50 mm high , with a rather high spire and usually truncated base . spiral sculpture often obsolete . anal sinus on subsutural ramp , deep , ( sub ) symmetrical , sometimes tubular . protoconch usually paucispiral ( up to two whorls ) , smooth or abapically carinate . operculum with terminal nucleus . radular formula 1 - 1 - r - 1 - 1 , rarely 1 - 1 - 0 - 1 - 1 , central tooth small , from narrow unicuspid ( fig . 13 b ) to subrectangular with additional cusps ( fig . 13 c ) , rarely reduced to completely absent . lateral teeth broad , pectinate , and arched , marginal teeth from simple flat and sharply pointed ( fig . 13 a ) to duplex with slightly thickened edges and to loosely enrolled with the small barb near the tip ( imaclava , fig . 13 d ) .
radulae . pseudomelatomidae . a . tiariturris spectabilis berry , 1958 * * , nhmuk moea 20100540 , gulf of panama , jtd - 00 - 34 , 08°26 . 24′n , 79 09 . 14′w , 66–68 m ( shell : fig . 14 q ) . b . comitas onokeana vivens dell , 1956 , mnhn , new caledonia , montrouzier , st . 1269 , after kantor & taylor ( 2000 ) . c , d . comitas sp . * * , mnhn im200717918 ( shell : fig . 14 a ) . e , f . crassiclava turricula ( sowerby , 1834 ) , off nacascola , west side of bahia culebra , costa rica , after kantor et al . ( 1997 ) . symbol : * * , sequenced specimen .
radulae . pseudomelatomidae . a . leucosyrinx sp . * * , mnhn im200717846 ( shell : fig . 14 g ) . b . zonulispira sp . * * , nhmuk moea 20100536 , gulf of panama , jtd - 00 - 18 , 08°19 . 50′n , 78°47 . 71′w , 25–32 m ( shell : fig . 14 v ) . c . carinodrillia dichroa pilsbry & lowe , 1932 * * , nhmuk moea 20100530 , gulf of panama , jtd - 00 - 18 , 08°19 . 50′n , 78°47 . 71′w , 25–32 m ( shell : fig . 1 j ) . d . ptychobela suturalis ( gray , 1838 ) * * , det . j . a . todd , yk , nhmuk moea 20100560 , off southern hong kong , sta . 71 . e . cheungbeia robusta ( hinds , 1839 ) * , nhmuk moea 20100557 , coll . b . morton , off southern hong kong , sta . 70 ( shell : fig . 14 k ) . f . inquisitor sp . * * , mnhn im200717851 ( shell : fig . 14 e ) . symbols : * , sequenced species ; * * , sequenced specimen .
diagnosis : shell small to rather large , 15–100 mm high , claviform to fusiform . spiral and axial sculpture generally well developed , often strong . subsutural fold often present . anal sinus on subsutural ramp , usually moderately deep to very deep , often constrained by callus rendering anal sinus subtubular . protoconch usually paucispiral , sometimes multispiral , with up to three whorls , smooth or sometimes axially or spirally sculptured on later whorls . operculum with terminal nucleus .
1 - ( 1 - r - 1 ) - 1 – comitas type ( includes also knefastia and antiplanes ) . the central formation is rather variable in degree of development of the rachidian tooth and fusion of three teeth . in some comitas ( fig . 15 c , d ) , the central formation looks like single well - defined tooth [ as in comitas murrawolga ( garrard , 1961 ) ] , while in comitas onokeana vivens dell , 1956 , knefastia and antiplanes , the rachidian is totally reduced and the formation appears as two poorly developed paired plates ( reduced laterals ) ( fig . 15 b ) . marginal teeth in comitas and knefastia flat , broadly oval , with thickened edges and teeth tips and without pronounced accessory limb . in antiplanes marginal teeth narrowly elongate , with well - developed accessory limb .
1 - 1 - 0 - 1 - 1 – crassiclava type . differs from the comitas type by the better defined laterals ( fig . 15 e , f ) that are low , arcuate and sharply curved towards the midline of the ribbon .
1 - 0 - r - 0 - 1 – pseudomelatoma type ( also includes hormospira and tiariturris , fig . 15 a ) . rachidian with strong cusp and subrectangular base ( contrary to the other types with a central formation formed by fused lateral and rachidian ) . despite neither ontogeny nor folding of the radula have been examined , we tentatively treat this structure as a true rachidian . marginal teeth simple , solid and strongly curved , attached to the membrane by rather a narrow base and free along most of their length .
1 - 0 - 0 - 0 - 1 – most genera of the family . marginal teeth elongated , narrow , flat , with thickened edge ( e . g . funa , carinodrillia , fig . 16 c ) , or trough - shaped in transverse section , sometimes with small barb near the tip ( cheungbeia , fig . 16 e ) , which may become semi - enrolled ( e . g . pyrgospira , pilsbryspira , zonulispira , fig . 16 b ) , to nearly hollow , where limbs overlap at significant length of the teeth ( ptychobela , fig . 16 d ) .
remarks : anatomically , pseudomelatomidae is the most variable family of conoidea . most genera were formerly included in the subfamily crassispirinae , but the nomenclaturally valid name for this clade is pseudomelatomidae . its constituents includes several taxa that were previously recognized as separate ( sub ) families : zonulispirinae , characterized by semi - enrolled marginal radular teeth ( a character found in several branches of the clade ) , and pseudomelatominae , defined on the basis of the very characteristic solid marginal teeth and strongly developed rachidian .
the genus leucosyrinx has long been a convenient genus for placement of turreted - fusiform species , mostly from deep water of the atlantic and indo - pacific . currently , it is a mixture of species of probably different taxonomic position , and the membership of the genus needs revision . the type species from the northern atlantic , leucosyrinx verrilli ( dall , 1881 ) , has a radula consisting of only duplex , rather robust marginal teeth ( powell , 1966 : text fig . b12 ) , a radula type also found by us in indo - pacific species ( fig . 16 a ) . a second type of radula is found in sibogasyrinx , originally described as a subgenus of leucosyrinx , and which clusters in the molecular tree with the cochlespiridae ( see under that family ) . in our study , true leucosyrinx appears to be sister to the pseudomelatomidae , but this relationship has poor support . as a working hypothesis , we tentatively include leucosyrinx in the pseudomelatomidae . clavatulidae gray , 1853
radulae . clavatulidae . a , b . turricula nelliae ( e . a . smith , 1877 ) * * , nhmuk moea 20100551 , danang , vietnam . c . pusionella compacta strebel , 1914 * * , mnhn im200717830 ( shell : fig . 17 b ) . d . toxiclionella tumida ( sowerby , 1870 ) , south africa , after kantor & taylor ( 2000 ) . e . clavatula xanteni nolf & verstraeten , 2006 * * , mnhn im200717829 ( shell : fig . 17 c ) . f . gemmuloborsonia colorata ( sysoev & bouchet , 2001 ) * * , mnhn im200717849 ( shell : fig . 17 a ) . symbol : * * , sequenced specimen . arrow , see text .
remarks : the genus gemmuloborsonia represents a sister group to the ( clavatulidae + horaiclavidae ) clade , but this node is not well supported . because it resembles much more the clavatulidae than the horaiclavidae in terms of shell and radular characters ( fig . 18 f ) , gemmuloborsonia is provisionally included in the former family . if further studies support that hypothesis , then the diagnosis of the clavatulidae should be amended to account for the presence of weak columellar pleats and multispiral turridae - type protoconch present in gemmuloborsonia . horaiclavidae new family
radulae . horaiclavidae . a . paradrillia sp . * * , mnhn im200742475 ( shell : fig . 17 n ) . b . inkinga sp . , mnhn ( shell : fig . 17 j ) . c . ceritoturris pupiformis ( e . a . smith , 1884 ) * * , mnhn im200717888 ( shell : fig . 17 i ) . d . horaiclavus splendidus ( a . adams , 1867 ) * * , mnhn im200717840 ( shell : fig . 17 h ) . symbol : * * , sequenced specimen . arrows , see text .
remarks : this family shares many characters with pseudomelatomidae , conchologically differing by a small stout shell with short siphonal canal and usually poorly developed spiral sculpture .
radulae also are rather similar to many representatives of pseudomelatomidae and no clear cut distinctions were found . genera currently included in horaiclavidae have been usually included in the crassispiridae ( = pseudomelatomidae ) , and the clear molecular - based division between the two clades seems to be not so clearly reflected in shell - based distinction . therefore , the generic composition of the family is somewhat provisional and needs confirmation by further molecular data and / or a detailed analysis of conchological and radular characters .
shells . turridae . a . xenuroturris legitima iredale , 1929 * * , mnhn im200717684 , vanuatu , santo 2006 , st . dr087 , 15°38 . 5′s , 167°15 . 1′e , 13 m , sl 57 . 0 mm ( radula : fig . 21 c ) . b . iotyrris cingulifera ( lamarck , 1822 ) * * , mnhn im200717685 , vanuatu , santo 2006 , st . fs84 , 15°33 . 6′s , 167°16 . 6′e , 8–9 m , sl 15 . 5 mm ( radula : fig . 21 d ) . c . decollidrillia nigra habe & ito , 1965 , zmmu uncatalogued , southern kurile islands , sl 12 . 8 mm . d . lophiotoma acuta ( perry , 1811 ) * * , mnhn im200717860 , philippines , panglao 2004 , st . r44 , 09°33 . 3′n , 123°43 . 9′e , 2 m , sl 44 . 0 mm . e . turridrupa acutigemmata ( e . a . smith , 1877 ) , new caledonia , 46 m , sl 26 . 5 mm . f . turris babylonia ( linnaeus , 1758 ) * * , mnhn im200717754 , philippines , panglao 2004 , st . r42 , 09°37 . 1′n , 123°52 . 6′e , 8–22 m , sl 79 . 4 mm . g . gemmula rarimaculata ( kuroda & oyama , 1971 ) * * , mnhn im200717838 , coral sea , ebisco , st . dw2533 , 22°18′s , 159°28′e , 360–370 m , sl 13 . 7 mm . h . ptychosyrinx chilensis berry , 1968 , usnm 870005 , s of coquimbo , chile , 31 . 1° s , 71 . 8° w , 179–187 m , sl 21 . 1 mm . i . lucerapex cf . casearia ( hedley & petterd , 1906 ) * * , mnhn im200742448 , philippines , panglao 2005 , st . cp2363 , 09°06 . 0′n , 123°25 . 0′e , 437–439 m , sl 21 . 0 mm ( radula : fig . 21 e ) . j . cryptogemma corneus ( okutani , 1966 ) , zin 58809 / 1 , off shikotan i . , kurile islands , 1450–1530 m , 12 . 3 mm . abbreviation and symbol : sl , shell length ; * * , sequenced specimen .
radulae . turridae . a . turridrupa cf . armillata ( reeve , 1845 ) , mnhn im200740773 , coral sea , ebisco , st . dw2607 , 19°33′s , 158°40′e , 400–413 m . b . gemmula kieneri ( doumet , 1840 ) , mnhn , vanuatu , musorstom 8 , st . cp1123 , 15°07′s , 166°55′e , 262–352 m . c . xenuroturris legitima iredale , 1929 , mnhn im200717684 ( shell : fig . 20 a ) . d . iotyrris cingulifera ( lamarck , 1822 ) * * , mnhn im200717685 ( shell : fig . 20 b ) . e . lucerapex cf . casearia ( hedley & petterd , 1906 ) * * , mnhn im200742448 ( shell : fig . 20 i ) . f . turris crispa ( lamarck , 1816 ) , mnhn , ile ouen - baie du prony , st . 80 , 22°31s , 166°28e , 33 m , ss 39 mm . symbols : * * , sequenced specimen .
remarks : this group is well defined by its usually narrowly fusiform shell with obsolete axial sculpture and peripheral anal sinus . however , the genus lucerapex , although fully conforming conchologically , occupies a position on the tree ( sister group to turridae + terebridae ) that excludes it from the turridae . it is nevertheless tentatively included here in turridae .
radulae . a , b . strictispiridae . strictispira paxillus ( reeve , 1845 ) , british virgina islands , ansp . a . teeth detached from the membrane and showing the large median flange . b . two teeth , showing the large median flange attached to the membrane . photo submit : j . d . taylor ( a . b ) . c . conorbidae . benthofascis lozoueti sysoev & bouchet , 2001 , mnhn im200742331 ( shell : fig . 2 o ) .
diagnosis : shell medium - sized , to 20 mm , claviform . spiral and axial sculpture well developed , shoulder with a marked subsutural fold . anal sinus deep , laterally directed . parietal callus well developed . protoconch paucispiral in all species examined , smooth . operculum leaf - shaped , with terminal nucleus . radula consisting of a pair of solid awl - shaped marginal teeth that have a prominent flange , located above the base of the tooth and firmly attached to the radular membrane .
remarks : we did not obtain any material of strictispiridae for molecular analysis and the position of the family group remains unclear . the shell resembles that of pseudomelatomidae and the shape of the radular marginal teeth is also somewhat similar to pseudomelatoma , hormospira and tiariturris , but it differs in the absence of the rachidian . among the examined species of strictispira , s . paxillus is characterised by rather unusual characters , such as the absence of the venom gland together with very large and powerful odontophore ( kantor & taylor , 1994 ) . until molecular data are available , we conservatively treat this family as valid , following taylor et al . ( 1993 ) .
diagnosis : shell medium - sized to large , 8–270 mm , usually 30–100 mm high , auger - shaped , with high to very high multiwhorled spire and flattened shell profile , aperture relatively small . siphonal canal short , anal sinus not pronounced . protoconch with up to 5 smooth whorls when multispiral . radular formula 1–0 - 0 - 0 - 1 but radular apparatus absent in many species . marginal teeth range from solid and curved to hypodermic , with or without small barb at the tip . hypodermic teeth without solid bases . in some species ( e . g . impages hectica ) the walls of the teeth are penetrated by numerous holes .
the authors thank b . a . marshall , m . g . harasewych , r . n . kilburn , m . j . tenorio , j . k . tucker and d . tippett for constructive comments and help in completing the lists of genera and subgenera . a . e . fedosov , j . d . taylor , j . mclean , m . g . harasewych and d . l . tippett supplied shell and radula pictures . j . a . todd provided specimens for dissection and photography . barbara buge curated and photographed the voucher specimens in mnhn .
turridae ( mollusca : gastropoda ) of southern africa and mozambique . part 1 . subfamily turrinae
turridae ( mollusca : gastropoda ) of southern africa and mozambique . part 2 . subfamily clavatulinae
turridae ( mollusca : gastropoda ) of southern africa and mozambique . part 3 . subfamily borsoniinae
turridae ( mollusca : gastropoda ) of southern africa and mozambique . part 4 . subfamily drilliinae , crassispirinae and strictispirinae
turridae ( mollusca : gastropoda ) of southern africa and mozambique . part 5 . subfamily taraninae
turridae ( mollusca : gastropoda ) of southern africa and mozambique . part 6 . subfamily mangeliinae , section 1
turridae ( mollusca : gastropoda ) of southern africa and mozambique . part 6 . subfamily mangeliinae , section 2
turridae [ s . l . ] ( mollusca : gastropoda ) of southern africa and mozambique . part 7 . subfamily crassispirinae , section 2
turridae ( s . l . ) of southern africa and mozambique ( mollusca : gastropoda : conoidea ) part 8 . conidae : subfamily mangeliinae , section 3
the new zealand recent and fossil mollusca of the family turridae . with general notes on turrid nomenclature and systematics
the molluscan families speightiidae and turridae . an evaluation of the valid taxa , both recent and fossil , with lists of characteristic species
( see bouchet and rocroi , 2005 for details ) . the status of names including only fossil genera is not discussed
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molluscabase ( 2018 ) . conorbidae de gregorio , 1880 . accessed through : world register of marine species at : urltoken ; = 153962 on 2018 - 07 - 09
bouchet , p . ; kantor , y . i . ; sysoev , a . ; puillandre , n . ( 2011 ) . a new operational classification of the conoidea ( gastropoda ) . journal of molluscan studies . 77 ( 3 ) : 273 - 308 . , available online at urltoken [ details ]
bouchet p . & rocroi j . - p . ( 2005 ) . classification and nomenclator of gastropod families . malacologia . 47 ( 1 - 2 ) : 1 - 397 isbn 3 - 925919 - 72 - 4 . [ details ]
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( of conorbis adamii bozzetti , 1994 ) bozzetti , l . 1994 . nuova specie dalle filippine . world shells 9 : 60 - 62 , 4 figs . [ english title also given as
a new species from philippines
; serial also referred to as : mondo di conchiglie . ] [ details ]
knudsen , j . ( 1952 ) marine prosobranchs of tropical west africa collected by the atlantide expedition , 1945–46 . videnskabelige meddelelser fra dansk naturhistorisk forening i kjobenhavn , 114 , 129–185 , 3 pls . page ( s ) : 176 , pl . 1 fig . 12 [ details ]
( of genota marchandi pin , 1996 ) pin , m . ( 1996 ) . il genere genota h . & a . adams 1853 ( conacea : turridae ) delle coste occidentali africane , con descrizione di genota marchandi n . sp . la conchiglia . 28 ( 279 ) , 53 - 56 . [ details ]
ryall p . , horro j . & rolán e . 2013 . a revision of the genus genota h . & a . adams , 1853 ( gastropoda : conoidea : borsonidae ) from west africa . iberus , 31 ( 2 ) : 1 - 17 . [ details ]
gofas , s . ; afonso , j . p . ; brandào , m . ( ed . ) . ( s . a . ) . conchas e moluscos de angola = coquillages et mollusques d ' angola . [ shells and molluscs of angola ] . universidade agostinho / elf aquitaine angola : angola . 140 pp . ( look up in imis ) [ details ]
( of genota marchandi pin , 1996 ) ryall p . , horro j . & rolán e . 2013 . a revision of the genus genota h . & a . adams , 1853 ( gastropoda : conoidea : borsonidae ) from west africa . iberus , 31 ( 2 ) : 1 - 17 . [ details ]
( of genota marchadi [ sic ] ) ardovini , r . ; cossignani , t . ( 2004 ) . west african seashells ( including azores , madeira and canary is . ) = conchiglie dell ' africa occidentale ( incluse azzorre , madeira e canarie ) . english - italian edition . l ' informatore piceno : ancona , italy . isbn 88 - 86070 - 11 - x . 319 pp . ( look up in imis ) page ( s ) : 37 , fig . 222 [ details ]
( of genota nigeriensis vera - peláez , 2004 ) ryall p . , horro j . & rolán e . 2013 . a revision of the genus genota h . & a . adams , 1853 ( gastropoda : conoidea : borsonidae ) from west africa . iberus , 31 ( 2 ) : 1 - 17 . [ details ]
bouchet , p . ; fontaine , b . ( 2009 ) . list of new marine species described between 2002 - 2006 . census of marine life . [ details ]
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bouchet , p . ; fontaine , b . ( 2009 ) . list of new marine species described between 2002 - 2006 . census of marine life .
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traditionally , all cone shells have been included in the linnean genus conus . tucker & tenorio ( 2009 ) have recently proposed an alternative shell - and radula - based classification that recognizes 4 families and 80 genera of cones . in worms , we currently still recognize a single family conidae ( following puillandre et al . 2011 ) , but tucker & tenorio ' s 80 genera classification is presented as
alternative representation
. [ p . bouchet , 14 aug . 2011 ]
conidae ( also see coninae ) is a taxonomic grouping of predatory sea snails , marine gastropod molluscs in the superfamily conoidea . some classifications of the cone snails , including the 2014 classification , group only cone snails in the family conidae . other classifications have previously grouped the cone snails in a subfamily of this family , the coninae .
until 1993 , with the taxonomic changes proposed by taylor , et al . ,
taxonomic changes as of 2009 and 2011 , based upon molecular phylogeny ( see below ) , have elevated the subfamilies which were previously in the family turridae to the status of families in their own right , leaving the family conidae once again containing the species which were traditionally placed in that family .
the family conidae currently ( march 2015 ) contains over 800 recognized species , while linnaeus knew only 30 valid species .
coninae fleming , 1822 — synonyms : conulinae rafinesque , 1815 ( inv . ) ; textiliinae da motta , 1995 ( n . a . )
clathurellinae h . adams & a . adams , 1858 — synonyms : defranciinae gray , 1853 ( inv . ) ; borsoniinae a . bellardi , 1875 ; pseudotominae a . bellardi , 1888 ; diptychomitrinae l . bellardi , 1888 ; mitrolumnidae sacco , 1904 ; mitromorphinae casey , 1904 ; lorinae thiele , 1925 sensu opinion 666
raphitominae a . bellardi , 1875 — synonyms : daphnellinae casey , 1904 ; taraninae casey , 1904 ; thatcheriidae powell , 1942 ; pleurotomellinae f . nordsieck , 1968 ; andoniinae vera - pelaez , 2002
in 2009 john k . tucker and manuel j . tenorio proposed a classification system for the cone shells and their allies ( which resorb their inner walls during growth ) was based upon a
( which are a distinct large and diverse group ) from the cone snails , and creates a number of new families .
in 2011 bouchet et al . proposed a new classification in which several subfamilies have been raised to the rank of family :
based on mitochondrial dna and nuclear dna testing , and builds on the prior work by j . k . tucker & m . j . tenorio ( 2009 ) , but does not include fossil taxa .
recognized extant genera within the cone snails as per j . k . tucker & m . j . tenorio ( 2009 ) , and bouchet et al . ( 2011 ) , include : .
and the genera recognized by tucker & tenorio 2009 are considered to be
alternate representations
.
. using 329 species , the authors carried out molecular phylogenetic analyses . the results suggested that the authors should place all cone snails in a single family , conidae , containing four genera :
following taylor et al . in 1993 , and prior to 2011 a large number of diverse genera were included within the family conidae , however as a result of molecular phylogeny studies in 2011 many of these genera which were in the family conidae ( commonly known as turrids ) have been moved back to the turridae or placed in new families within the superfamily conoidea . the following list of genera is maintained for historical reasons :
there are around 30 records of humans killed by a cone snail . human victims suffer little pain , because the venom contains an analgesic component . some species can kill a human in under 5 minutes , from where the name
cigarette snail
as one only has time to smoke a cigarette before dying . the molluscs can attack if provoked and can sting through a wetsuit with their harpoon , which resembles a transparent needle .
normally cone snails ( and many species in the superfamily conoidea ) use the venom to immobilize prey . it consists of a mixture of peptides , called conopeptides . their venom is made up of 10 to 30 amino acids , but occasionally as many as 60 . the venom of each cone snail species may contain as many as 200 pharmacologically active components . it is estimated that more than 50 , 000 conopeptides can be found because every species of cone snail is thought to produce its own specific venom .
cone snail venom , in more recent years , has come to interest biotechnologists and pharmacists because of its potential medicinal properties . production of synthetic conopeptides has started , using solid - phase peptide synthesis .
w - conopeptide , from the species conus magus is the basis of the analgesic drug prialt , an approved treatment for pain said to be 1000 times as powerful as morphine and used as a last resort in specific application . conopeptides are also being looked at as anti - epileptic agents and to help stop nerve - cell death after a stroke or head injury . conopeptides also have potential in helping against spasms due to spinal cord injuries , and may be helpful in diagnosing and treating small cell carcinomas in the lung .
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foregut anatomy , feeding mechanisms , relationships and classification of conoidea ( toxoglossa ) ( gastropoda )
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classification and nomenclator of gastropod families
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tucker j . k . & tenorio m . j . ( 2009 ) systematic classification of recent and fossil conoidean gastropods . hackenheim : conchbooks . 296 pp . , at p . 133
p . k . bandyopadhyay , b . j . stevenson , j . p . ownby , m . t . cady , m . watkins , & b . olivera ( 2008 ) , the mitochondrial genome of conus textile , coxi - conii intergenic sequences and conoidean evolution . molecular phylogenetics and evolution 46 : 215 - 223 .
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t . f . duda , jr . & a . j . kohn ( 2005 ) , species - level phylogeography and evolutionary history of the hyperdiverse marine gastropod genus conus , molecular phylogenetics and evolution 34 : 257 - 272 .
t . f . duda , jr . & e . rolan ( 2005 ) , explosive radiation of cape verde conus , a marine species flock , molecular ecology 14 : 267 - 272 .
b . vallejo , jr . ( 2005 ) , inferring the mode of speciation in the indo - west pacific conus ( gastropoda : conidae ) , journal of biogeography 32 : 1429 - 1439 .
n . puillandre , s . samadi , m . boesselier , a . sysoev , y . kantor , c . cruaud , a . couloux , & p . bouchett ( 2008 ) , starting to unravel the toxoglossan knot : molecular phylogeny of the
turrid
( neogastropoda : conoidea ) , molecular phylogenetics and evolution 47 : 1122 - 1134 .
bouchet p . , kantor yu . i . , sysoev a . & puillandre n . ( 2011 ) .
a new operational classification of the conoidea
. journal of molluscan studies 77 : 273 - 308 . doi : 10 . 1093 / mollus / eyr017 .
tucker , j . k . & stahlschmidt , p . ( 2010 ) a second species of pseudoconorbis ( gastropoda : conoidea ) from india . miscellanea malacologica 4 ( 3 ) : 31 - 34 .
watkins , m . , corneli , p . s . , hillyard , d . , & olivera , b . m . ( 2010 ) molecular phylogeny of conus chiangi ( azuma , 1972 ) ( gastropods : conidae ) . the nautilus 124 ( 3 ) : 129 - 136 .
tucker , j . k . , tenorio , m . j . & stahlschmidt , p . ( 2011 ) the genus benthofascis ( gastropoda : conoidea ) : a revision with descriptions of new species . zootaxa 2796 : 1 - 14 .
tucker , j . k . & tenorio , m . j . ( 2011 ) new species of gradiconus and kohniconus from the western atlantic ( gastropoda : conoidea : conidae , conilithidae ) . miscellanea malacologica 5 ( 1 ) : 1 - 16 .
petuch , e . j . & sargent , d . m . ( 2011 ) new species of conidae and conilithidae ( gastropoda ) from the tropical americas and philippines . with notes on some poorly - known floridian species . visaya 3 ( 3 ) : 116 - 137 .
petuch & drolshage ( 2011 ) compendium of florida fossil shells , volume 1 mdm publications , wellington , florida , 432 pp .
urltoken classification : traditionally , all cone shells have been included in the linnean genus conus . tucker & tenorio ( 2009 ) have recently proposed an alternative shell - and radula - based classification that recognizes 4 families and 80 genera of cones . in worms , we currently still recognize a single family conidae ( following puillandre et al . 2011 ) , but tucker & tenorio ' s 80 genera classification is presented as
alternative representation
. [ p . bouchet , 14 aug . 2011 ]
n . puillandre , e . strong , p . bouchet , m . boisselier , v . couloux , & s . samadi ( 2009 ) ,
identifying gastropod spawn from dna barcodes : possible but not yet practicable
, molecular ecology resources 9 : 1311 - 1321 .
tucker j . k . & tenorio m . j . ( 2009 ) , systematic classification of recent and fossil conoidean gastropods , conchbooks , hankenheim , germany , 295 pp .
biggs , j . s . , watkins , m . showers corneli , p . and olivera , b . m . ( 2010 ) . defining a clade by morphological , molecular , and toxinological criteria : distinctive forms related to conus praecellens a . adams , 1854 ( gastropoda : conidae ) . nautilus 124 : 1 - 19 ( naming new species and moving species from kurodaconus to turriconus ) .
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applied microbiology and biotechnology 79 ( 1 ) : 1 - 9 . doi : 10 . 1007 / s00253 - 008 - 1385 - 6 .
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conoserver : updated content , knowledge , and discovery tools in the conopeptide database
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. smithsonian institution press , washington and london .
monteiro a . ( ed . ) ( 2007 ) . the cone collector 1 : 1 - 28 .
taylor , j . d . , kantor yu . i . & sysoev a . v . ( 1993 ) .
foregut anatomy , feeding mechanisms , relationships and classification of conoidea ( toxoglossa ) ( gastropoda )
. bull . nat . hist . mus . ( zool . ) 59 : 125 - 169 .
berschauer d . ( 2010 ) . technology and the fall of the mono - generic family the cone collector 15 : pp . 51–54
puillandre n . , meyer c . p . , bouchet p . , and olivera b . m . ( 2011 ) , genetic divergence and geographical variation in the deep - water conus orbignyi complex ( mollusca : conoidea ) , zoologica scripta 40 ( 4 ) 350 - 363 .
puillandre n . , duda t . f . , meyer c . , olivera b . m . & bouchet p . ( 2015 ) . one , four or 100 genera ? a new classification of the cone snails . journal of molluscan studies . 81 : 1 - 23
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urltoken
conidae is a taxonomic family of minute to quite large sea snails , marine gastropod molluscs in the superfamily conoidea .
current taxonomic changes as of 2009 and 2011 , based upon molecular phylogeny ( see below ) , have elevated the subfamilies which were previously in the family turridae to the status of families in their own right , leaving the family conidae once again containing the species which were traditionally placed in that family .
the family conidae currently still contains well over 600 recognized species , which traditionally have all been placed in the genus conus .
clathurellinae h . adams & a . adams , 1858 — synonyms : defranciinae gray , 1853 ( inv . ) ; borsoniinae a . bellardi , 1875 ; pseudotominae a . bellardi , 1888 ; diptychomitrinae l . bellardi , 1888 ; mitrolumnidae sacco , 1904 ; mitromorphinae casey , 1904 ; lorinae thiele , 1925 sensu opinion 666
recognized extant genera within the cone snails as per j . k . tucker & m . j . tenorio ( 2009 ) , and bouchet et al . ( 2011 ) , include :
note : an asterisk * indicates genera that are considered to be within the family conilithidae by tucker & tenorio 2009 .