conilithidae is a proposed taxonomic family of small to medium-sized sea snails , specifically cone snails , marine gastropod mollusks in the superfamily conoidea , the cone snails and their allies .

in 2009 , john k. tucker and manuel j. tenorio proposed elevating the subfamily conilithinae ( previously placed in the family conidae ) to the rank of family .

this was based on a cladistic analysis of anatomical characters , including the radular tooth , the morphology ( i.e. shell characters ) , as well as an analysis of prior molecular phylogeny studies , all of which were used to construct phylogenetic trees .

in their phylogeny , tucker and tenorio noted the close relationship of the various species within the genera in the conilithidae ; this also corresponded to the results of prior molecular studies by puillandre et al. .

tucker and tenorio ’s proposed classification system for the cone shells and their allies ( and the other clades of conoidean gastropods ) is set forth at tucker & tenorio cone snail taxonomy 2009 .

like other species in the superfamily conoidea , these snails are predatory and venomous , able to inject neurotoxins into their prey with their radula .

the species in this family are capable of " stinging " humans , therefore live ones should be handled carefully or not at all .

conilithidae

conilithidae tucker , j . k . & m . j . tenório , 2009 thumbnails
the families conilithidae and conidae - the conus of the eastern pacific . - a conchological iconography # 18
five new species of jaspidiconus petuch , 2004 ( conilithidae : conilithinae ) from the caribbean mollus . . .
biodiversity and habitats of reef molluscs of families conidae and conilithidae ( neogastropoda ) off northern roatan island ( honduras ) .
ebscohost | 99118461 | biodiversity and habitats of reef molluscs of families conidae and conilithidae ( neogastropoda ) off northern roatan island ( honduras ) .
kohniconus is a subgenus of sea snails , marine gastropod mollusks in the genus conasprella , family conilithidae , the cone snails and their allies .
home > journals & occasional publications > conchological iconography > the families conilithidae and conidae - the conus of the eastern pacific . - a conchological iconography # 18
kohniconus is a subgenus of sea snails , marine gastropod mollusks in the genus conasprella , family conilithidae , the cone snails and their allies . [ 1 ]
molluscabase ( 2018 ) . conilithidae tucker & tenorio , 2009 . accessed through : world register of marine species at : urltoken ; = 578794 on 2018 - 07 - 09
[ synonyms : conilithidae tucker & tenorio , 2009 , n . syn . taranteconidae tucker & tenorio , 2009 , n . syn . puncticuliinae tucker & tenorio , 2009 , n . syn . ]
six new gastropods , belonging to the families fasciolariidae , conidae , and conilithidae , recently have been discovered within the biogeographical boundaries of the brazilian molluscan province . these include : poremskiconus fonsecai n . sp . and poremskiconus smoesi n . sp . ( both conidae ) from the cearaian subprovince of northern brazil ; jaspidiconus josei n . sp . ( conilithidae ) from the bahian . . . [ show full abstract ]
a conchological iconography the families conilithidae and conidae - the cones of the eastern pacific : urltoken manuel jimenez tenorio , john k . tucker , henry w . chaney , guido t . poppe , klaus groh : 9783939767428 : books
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 .
( of conilithidae tucker & tenorio , 2009 ) tucker j . k . & tenorio m . j . ( 2009 ) systematic classification of recent and fossil conoidean gastropods . hackenheim : conchbooks . 296 pp . page ( s ) : 136 [ details ]
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 .
six new gastropods , belonging to the families fasciolariidae , conidae , and conilithidae , recently have been discovered within the biogeographical boundaries of the brazilian molluscan province . these include : poremskiconus fonsecai n . sp . and poremskiconus smoesi n . sp . ( both conidae ) from the cearaian subprovince of northern brazil ; jaspidiconus josei n . sp . ( conilithidae ) from the bahian subprovince of central brazil ; and fusinus damasoi n . sp . , fusinus mariaodeteae n . sp . ( both fasciolariidae ) , and lamniconus petestimpsoni n . sp . ( conidae ) from the paulinian subprovince of southern brazil .
prior to 2009 , all cone species were placed within the family conidae and were placed in one genus , conus . in 2009 however , j . k . tucker and m . j . tenorio proposed a classification system for the over 600 recognized species that were in the family . their classification proposed 3 distinct families and 82 genera for the living species of cone snails , including the family conilithidae . this classification was based upon shell morphology , radular differences , anatomy , physiology , cladistics , with comparisons to molecular ( dna ) studies . published accounts of genera within the conidae ( or conilithidae ) that include the genus kohniconus include j . k . tucker & m . j . tenorio ( 2009 ) , and bouchet et al . ( 2011 ) .
prior to 2009 , all cone species were placed within the family conidae and were placed in one genus , conus . in 2009 however , j . k . tucker and m . j . tenorio proposed a classification system for the over 600 recognized species that were in the family . their classification proposed 3 distinct families and 82 genera for the living species of cone snails , including the family conilithidae . this classification was based upon shell morphology , radular differences , anatomy , physiology , cladistics , with comparisons to molecular ( dna ) studies . [ 2 ] published accounts of genera within the conidae ( or conilithidae ) that include the genus kohniconus include j . k . tucker & m . j . tenorio ( 2009 ) , and bouchet et al . ( 2011 ) . [ 4 ]
( of conilithidae tucker & tenorio , 2009 ) 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 . , available online at urltoken [ details ] available for editors [ request ]
this book present a new classification of the cone shells . it is based on radular morphology to define most of the suprageneric taxa . however , other features such as shell morphology , morphology of the periostracum and operculum and dietary habits are also factored in . the authors consider fossil taxa as well as recent taxa . the classification consists of five families , namely conorbiidae , hemiconidae n . fam . , taranteconidae n . fam . , conilithidae n . fam . and conidae . the family conidae is by far the most numerous in terms of genera both extinct and living , and comprises at least two subfamilies ( coninae and puncticuliinae ) and 64 genera , one of them fossil . the next family in number of genera is conilithidae , with two new subfamilies ( conilithinae and californiconinae ) and 19 genera , two of them extinct . the remaining three families are basal to the other two , and much less diverse : hemiconidae has only one fossil genus assigned , whereas taranteconidae has two extant genera , and conorbiidae consists of three genera , one fossil and two extant . thus , the resulting classification comprises 89 genera in total , from which 27 are introduced as new taxa .
five genera and eight species of gastropods of families conidae and conilithidae were observed in their natural habitats on the southernmost portion of the mesoamerican barrier reef , off the northern coast of roatan island , honduras . fifty per cent of species are widespread caribbean–western atlantic species , whereas 50 % are endemic to the nicaraguan biogeographical subprovince and roatan island . multiple sightings during night scuba diving operations revealed that the reef off northern roatan supports a healthy and diverse population of conoidean gastropods . distribution of all recorded species by depth and habitat type revealed a distinct reef partitioning between the 4 most commonly occurring species .
abstract : five genera and eight species of gastropods of families conidae and conilithidae were observed in their natural habitats on the southernmost portion of the mesoamerican barrier reef , off the northern coast of roatan island , honduras . fifty per cent of species are widespread caribbean–western atlantic species , whereas 50 % are endemic to the nicaraguan biogeographical subprovince and roatan island . multiple sightings during night scuba diving operations revealed that the reef off northern roatan supports a healthy and diverse population of conoidean gastropods . distribution of all recorded species by depth and habitat type revealed a distinct reef partitioning between the 4 most commonly occurring species .
this book present a new classification of the cone shells . it is based on radular morphology to define most of the suprageneric taxa . however , other features such as shell morphology , morphology of the periostracum and operculum and dietary habits are also factored in . the authors consider fossil taxa as well as recent taxa . the classification consists of five families , namely conorbiidae , hemiconidae n . fam . , taranteconidae n . fam . , conilithidae n . fam . and conidae . the family conidae is by far the most numerous in terms of genera both extinct and living , and comprises at least two subfamilies ( coninae and puncticuliinae ) and 64 genera , one of them fossil . the next family in number of genera is conilithidae , with two new subfamilies ( conilithinae and californiconinae ) and 19 genera , two of them extinct . the remaining three families are basal to the other two , and much less diverse : hemiconidae has only one fossil genus assigned , whereas taranteconidae has two extant genera , and conorbiidae consists of three genera , one fossil and two extant . thus , the resulting classification comprises 89 genera in total , from which 27 are introduced as new taxa . 296 pp . , 17 figs , 15 b / w & 11 col . pls , hc 4 . bookseller inventory # w12444
species sometimes referred to as ‘cone snails’ but allocated to artemidiconus ( conorbidae ; tucker & tenorio , 2009 ; bouchet et al . , 2011 ) , benthofascis ( conorbidae ; tucker & tenorio , 2009 ; bouchet et al . , 2011 ) , genota ( cryptoconidae in tucker & tenorio , 2009 ; or borsoniidae in bouchet et al . , 2011 ) and genotina ( cryptoconidae in tucker & tenorio , 2009 ; or mangeliidae in bouchet et al . , 2011 ) were excluded . within conorbidae , only benthofascis lozoueti has been sequenced and molecular analysis indicates that the family is separate from conidae ( puillandre et al . , 2011 ) . however , b . lozoueti is the only conorbid species that does not resorb the inner shell walls ( tucker , tenorio & stahlschmidt , 2011 ) . it thus cannot be excluded that the other conorbidae species—which resorb them—may in fact not be confamilial . likewise , as indicated above , molecular data place genota in the borsoniidae ( puillandre et al . , 2011 ) and this clade is not further discussed here . fossil taxa ( hemiconus cossmann , 1889 ; cryptoconus koenen , 1867 ; conorbis swainson , 1840 ; conilithes swainson , 1840 ; eoconus tucker & tenorio , 2009 and plagioconus tucker & tenorio , 2009 ) are not discussed either . consequently , only the conidae , conilithidae and taranteconidae ( sensu tucker & tenorio , 2009 ) are discussed below , i . e . the cone snails as defined by bouchet et al . ( 2011 ) .
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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 .
based on a dataset of 57 genera and molecular sequences of fragments of one mitochondrial ( coi ) and three nuclear ( 28s , 18s and h3 ) genes , puillandre et al . ( 2008 ) published the first molecular phylogeny of conoidea . even though ranks differed , most of the lineages defined by taylor et al . ( 1993 ) were also retrieved in this molecular approach . the new molecular phylogeny of puillandre et al . ( 2011 ) is based on a dataset of 102 conoidean genera ( 87 ‘turrids’ , 5 cones and 10 terebrids ) and sequences of three gene fragments ( coi , 12s rrna and 16s rrna ) . because of the high congruency between the classification based on anatomical characters and the molecular tree obtained , we are inclined to think that our understanding of the phylogeny of the conoidea has now reached a stable position , and that a new classification is warranted . the classification presented below ( fig . 1 b ) aims to transform the molecular phylogeny into an operational classification by : ( 1 ) presenting revised morphological diagnoses for the now redefined families of conoidea ; ( 2 ) allocating ( sometimes tentatively ) all recent genera recognized or used in current literature to the corresponding family .
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 .
diagnosis : shell small - to medium - sized ( 5–40 , usually 10–20 mm ) , fusiform to broadly fusiform , subsutural ramp usually unclearly separated . sculpture mostly strong , usually cancellate , subsutural fold lacking . shell surface often microgranular . apertural armature often well developed , in the form of pleats and denticles on both inner and outer lips ; no true columellar pleats . siphonal canal well expressed , moderately long . anal sinus on subsutural ramp , deep , often ( sub ) tubular . protoconch typically multispiral , up to six whorls , generally smooth with keeled last whorls ; when paucispiral , protoconch usually smooth or spirally striated , sometimes with remaining carination in last portion . operculum always absent . radula of hypodermic marginal teeth that usually have relatively small solid basal part ( fig . 5 ) . distinct ligaments not found . at their tip , teeth can have a weak barb .
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 .
diagnosis : shell small to medium size , 3–30 mm , usually 6–12 mm , oval to low - or high - fusiform , usually with comparatively low spire , often with a shoulder angulation . spiral and axial sculpture both well developed . shell surface often bearing microsculpture of spirally aligned granules . subsutural ramp usually not separated sculpturally . anal sinus on subsutural ramp , shallow to rather deep , rarely tubular . aperture normally not constrained with strong outgrowths or callus pads , rarely denticulate . siphon rather short to moderately long . protoconch typically multispiral , of up to five whorls , with axially ribbed protoconch ii ; spiral cords on protoconch ii present or absent . when paucispiral , protoconch usually spirally lirate . operculum present , with terminal nucleus ( oenopotinae ) , or normally absent . radula of marginal teeth of very variable morphology . teeth can be from semi - enrolled ( fig . 8 f , g ) to true hypodermic . frequently side projections around the base are present and there is large irregularly shaped ‘root’ projecting from the base . barbs may be present , from small to very large ( fig . 8 c ) or absent ( fig . 8 b ) . tooth canal opening laterally .
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 ) .
remarks : the genera cruziturricula and fusiturricula form an unsupported group that is sister to drilliidae . in the studied cruziturricula arcuata ( reeve , 1843 ) , the very characteristic radula differs from the drilliidae : 1 - 0 - 0 - 0 - 1 . marginal teeth are loosely enrolled with little overlap of the edges , with two barbs on the tip and a tongue - shape extension at the base ( fig . 13 e ) . the type species of fusiturricula , turris fusinella dall , 1908 , is different from what is currently conceived as belonging to that genus ( e . g . williams , 2006 ) , but those species are similar to cruziturricula sensu auctt . the only data on the radula was provided by powell ( 1966 : 31 ) , who stated ( without mentioning the species ) : “radula of marginals only , wishbone - type , but long and narrow” . although cruziturricula and fusiturricula definitely do not belong in drilliidae and may represent a new family , they are here provisionally placed in drilliidae for lack of a better alternative .
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 .
diagnosis : shell medium - sized to rather large ( usually 15–30 mm , up to 85 mm high ) , broad - fusiform to turreted - fusiform , with high spire and usually moderately long siphonal canal . subsutural ramp usually well developed , with very shallow to rather deep anal sinus situated on its lower part or shifted abapically , with apex at almost a peripheral position . sculpture variously developed , from almost smooth shell surface to well - developed axial ribs and spiral cords . protoconch only known as paucispiral , up to c . 2 . 5 smooth whorls . operculum with medio - lateral nucleus . radular formula 1 - ( 1 - r - 1 ) - 1 . the central formation is composed of very thin , broad , plate - like lateral teeth and a small , but sometimes strong , narrow rachidian . the central formation is variously developed , sometimes appearing as a pronounced tooth ( fig . 18 b ) , sometimes clearly consisting of three elements ( clionella sinuata , see taylor et al . , 1993 ; fig . 18 a , b ) , to its nearly complete reduction ( fig . 18 c ) . marginal teeth usually duplex , with sharp - edged major limb and a deep socket where an accessory limb is inserted , often with angulation distal to the socket ( arrow in fig . 18 e ) . in toxiclionella the marginal teeth are hypodermic , loosely enrolled , attached along their length to radular membrane , having two barbs at the tip and a tooth canal opening subterminally ( fig . 18 d ) .
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 .
diagnosis : shell generally small , 5–25 mm , usually 7–15 mm high , shortly claviform , with relatively low spire and a short , truncated , poorly differentiated siphonal canal . subsutural ramp usually poorly differentiated . axial sculpture almost always present , usually as strong sinuate ribs . spiral sculpture normally weak or obsolete , often with glossy shell surface . anal sinus on subsutural slope , weak to moderately deep , often constrained by callus . protoconch of up to 3 . 5 medially carinate but otherwise smooth whorls when multispiral , but usually paucispiral and smooth . operculum with terminal nucleus . radular formula : 1 - 0 - 0 - 0 - 1 . marginal teeth duplex , with lanceolate major limb and usually narrow accessory limb , which is inserted in a shallow socket . major limb often with angulation lateral to the place of accessory limb insertion . rarely ( some paradrillia , cf . kilburn , 1988 , fig . 17 ; inkinga , fig . 19 b ) the additional limb is of similar size to the major limb and the teeth become trough - shaped in transverse section , with a collar near the base ( arrows in fig . 19 b ) . in several species of horaiclavus the radular apparatus is absent .
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 .
diagnosis : shell of medium to large size ( usually 20–30 mm , up to 110 mm high ) , short - to high - fusiform , usually with a high spire and a long ( rarely short and truncated ) siphonal canal . axial sculpture weak or absent . anal sinus on whorl periphery . protoconch typically multispiral , up to six whorls , protoconch i smooth , protoconch ii with arcuate axial riblets ; reduced paucispiral protoconch usually smooth , may have arcuate axial riblets . operculum fully developed , with terminal nucleus . radular formula typically 1 - ( 1 : r : 1 ) - 1 . small and narrow rachidian and plate - like laterals are fused together , together constituting a central formation of different development ( kantor , 2006 ) , varying from a well - defined broad central tooth ( fig . 21 b ) to a tooth clearly formed of three elements ( fig . 21 a ) through a gradual reduction of rachidian and / or laterals to complete absence ( fig . 21 d ) . marginals duplex , of variable morphology , from broadly oval and flattened ( fig . 21 e , f ) with nearly equally developed limbs to awl - shaped and divided only in basal part ( fig . 21 a ) . in most cases , the major limb is large and knife - shaped , while the accessory limb is dorsal and more weakly developed ( fig . 21 ) . in iotyrris , marginal teeth have equally developed limbs that form a shallow broad trough ( fig . 21 d ) .
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


parcoblatta fulvescens , the fulvous wood cockroach , is a species of cockroach endemic to the united states and possibly canada that measures around 13 mm ( 0.5 in ) long .

parcoblatta fulvescens

might be virginica or even fulvescens cause some of the males in the pics u posted look similar to fulvescens males .
moved from zebra wood cockroach . this one actually appears to be p . fulvescens , which is probably the most common species of parcoblatta in ok
i guess this female is a parcoblatta uhleriana . they are the most common here .
1 . female parcoblalta fulvescens feed selectively on diets differing in nutrient content when given a choice during the reproductive cycle .
hello . i caught some parcoblatta and i cannot find pictures of many of the species . i might have four species of parcoblatta right now . can somebody post some pictures of p . zebra , p . lata , p . fulvescens , p . caudellia and p . notha ? if you can post pictures of any of the other species of parcoblatta i would love to see them too !
ps . male p . lata has black abdomen by the way . fulvescens look similar to p . lata but doesn ' t have black abdomen .
life history and habitat associations of the broad wood cockroach , parcoblatta lata ( blattaria : blattellidae ) and other native cockroaches in the coastal plain of south carolina .
journal article : life history and habitat associations of the broad wood cockroach , parcoblatta lata ( blattaria : blattellidae ) and other native cockroaches in the coastal plain of south carolina .
life history and habitat associations of the broad wood cockroach , parcoblatta lata ( blattaria : blattellidae ) and other native cockroaches in the coastal plain of south carolina . ( journal article ) | osti . gov
horn , scott , & hanula , james , l . life history and habitat associations of the broad wood cockroach , parcoblatta lata ( blattaria : blattellidae ) and other native cockroaches in the coastal plain of south carolina . . united states . doi : 10 . 1603 / 0013 - 8746 ( 2002 ) 095 [ 0665 : lhahao ] 2 . 0 . co ; 2 .
horn , scott , and hanula , james , l . tue .
life history and habitat associations of the broad wood cockroach , parcoblatta lata ( blattaria : blattellidae ) and other native cockroaches in the coastal plain of south carolina .
. united states . doi : 10 . 1603 / 0013 - 8746 ( 2002 ) 095 [ 0665 : lhahao ] 2 . 0 . co ; 2 . urltoken
horn , scott , and hanula , james , l . life history and habitat associations of the broad wood cockroach , parcoblatta lata ( blattaria : blattellidae ) and other native cockroaches in the coastal plain of south carolina . . united states : n . p . , 2002 . web . doi : 10 . 1603 / 0013 - 8746 ( 2002 ) 095 [ 0665 : lhahao ] 2 . 0 . co ; 2 .
wood cockroaches are an important prey of the red - cockaded woodpecker , picoides borealis , an endangered species inhabiting pine forests in the southern united states . these woodpeckers forage on the boles of live pine trees , but their prey consists of a high proportion of wood cockroaches , parcoblatta spp . , that are more commonly associated with dead plant material . cockroach population density samples were conducted on live pine trees , dead snags and coarse woody debris on the ground . the studies showed that snags and logs are also important habitats of wood cockroaches in pine forests .
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sometimes taxonomists create new names for groups that already have a name . they may do this because they are unaware of the original name , or they may think the organism before them belongs to a different group when in fact it does not . if two or more names are found to apply to the same group , they are considered synonyms . in most cases , the first name takes priority and is considered to be the valid or accepted name . however , there can be exceptions , and it ' s not always easy to determine which of a series of synonyms should be considered valid or accepted . here we list the synonyms provided to eol by our classification partners . we also include other versions of the name that most likely refer to the same group , for example , misspellings in the literature or different variations of the authorship associated with the name .
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upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington
contributed by bryan e . reynolds on 17 july , 2016 - 11 : 28am last updated 21 july , 2016 - 8 : 33am
thanks for the clarification , alan , and sorry for jumping the gun . these buggers sure look similar . next time i ' ll post into the family level .
a lot of the species in this genus look alike so it ' s hard to distinguish them from one another , even for the experts .
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beccaloni g . w . ( 2007 - ) cockroach species file online . version 5 . 0
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2 . diets high in carbohydrate and protein content are preferred early in the cycle , while a cellulose - containing diet is readily consumed toward the end of the cycle .
3 . the total diet consumed by females given a choice contained about 16 % protein , and they did not excrete uric acid while on this diet .
4 . females on diets high in carbohydrate or cellulose required longer to complete the reproductive cycle than females on high - protein diet or those given a dietary choice .
5 . nitrogen - stressed females will consume urate - containing fecal pellets , but only if they have utilizable carbohydrate in their diet .
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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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atkinson , thomas h . , philip g . koehler , and richard s . patterson
nomina insecta nearctica : a check list of the insects of north america : vol . 4 : non - holometabolous orders
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 .
the largest family of the order , with ~ 35 spp . ( incl . several adventive ) in 14 genera of 4 subfamilies in our area and ~ 2 , 300 spp . in ~ 220 genera of 7 subfamilies worldwide
american insects : a handbook of the insects of america north of mexico ross h . arnett . 2000 . crc press .
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great ! thank you for the pictures ! i took some picture today . here .
i do not know if these orange males are p . uhleriana or p . lata .
i am guessing that this one is a p . virginica male . they are a lot smaller ( just slightly bigger than the p . bolliana males ) than the other males and they have the dark head . they are faster too .
here is a p . bolliana or what i think is a p . bolliana . adult male .
i do not know what these are . most of the big orange males i think came from black nymphs which i thought were i . deropeltiformis .
here are an adult female and adult male of what i think are p . bolliana .
i know some of the big orange ones molted recently so maybe they are not the normal color yet .
this one i am not sure what it is . what is this adult female ? is it p . lata ? it molted over a day ago ( yesterday in the daytime ) .
this male was slightly different in color . a few of the big orange males are smaller than normal and some have different color like this ( did not get a good look so not sure exactly how ) .
i caught this p . virginica in leaves with p . uhleriana . they are tiny ! she was chewing on me while i was taking a picture . . . . . so it was hard to take a pic without shaking so much
i don ' t think u have any i . deropeltiformis in the pics . there might be some pseudomops nymphs in there by the way .
very interesting ! i do not think she is a p . virginica or a p . fulviscens . she is a lot bigger than the p . virginica ( close to p . uhleriana size ) . she has slightly different wings from p . fulviscens . yes , i can sell some p . uhleriana if they start breeding . it seems to be dryer now outside than it used to be a few months ago and because of that i think there are fewer roaches . i am having a hard time finding many roaches , i mean i can easily find some but just not very many . are the p . uhleriana common in your area ?
i don ' t think p . uhleriana is present in my area ( they do occur in my state but i haven ' t seen any yet ) .
how much are you asking for a pair of p . uhleriana adults ? i ' m interested in about two pairs
only two pairs ? i think i can find more than that . i will look today . can you tell what the p . uhleriana adult males look like ? you live in alabama right ? just pay for the shipping and i will send them to you . how much do you think the shipping will be ?
i found another adult female p . uhleriana today . it is harder than i thought . i do not think i can get very many . the places i go to do not have anymore roaches . i need to find other places . i have only one adult female ischnoptera deropeltiformis so i need a male and i hope one of them are i . deropeltiformis lol . i think i found some more of the ones that look like the pseudomops . do you want me to send some now or in the future if they breed ?
i would love to get some adults soon so that i can not only breed them but i would be able to send some to zephyr on june . do you think it will be possible to send me 2 or more pairs of adults ?
ps this one might be the p . uhleriana male . was this specimen black when he was nymph ?
okay . i do not know if that one was black as a nymph ( i got him as an adult ) . i have one sub - adult male nymph that is black and has that little brown spot close to the tip of the abdomen on top . another sub - adult male roach molted yesterday and it was dark rusty brown like this black one on top . it did not have that brown spot and it was less shiny . i found these p . uhleriana like roaches in thick white oak and red oak like leaves .
perhaps you could put one of the specimen in the fridge for about 10 ~ 15 min . and take the pics of median segment under the wing ?
sure i will do that ! but i am going to have a concert soon so maybe i cannot put the picture here this afternoon .
these are the best pictures i can get . i am scared that i am going to kill them holding them like this . are these two pictures good enough to find out what species they are ? this male is limping and his front legs do not work now . my hands a shaky . what are the p . uhleriana supposed to have ?
have a pair of structures on the median segment ( and not the first abdominal segment ) . these structures do not meet in the midline to form a ridge . the wings are markedly broader than the pronotum in
please could you explain what they mean ? like what the ridge looks like . i might be able to look at them quickly without out hurting them and see what they are .
do you see those two dot looking thing below the pronotum ? that ' s the pair of modified structure that we are looking for .
thank you for the clear pictures . here are some more pictures i took .
this one is a strange one . it laid an ootheca that looked like the other p . uhleriana . but she has smaller and more round wings . i think she is the smallest of them all . cariblatta your p . uhleriana should be laying oothecae by now . have they ?
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abed , d . , brossut , r . , and farine , j . - p . 1993a . evidence for sex pheromones produced by males and females in
abed , d . , chevied , p . , farine , j . - p . , bonnard , o . , le quéré , j . l . , and brossut , r . 1993b . calling behaviour of female
atkinson , t . h . , koehler , p . g . , and patterson , r . s . 1991 . catalog and atlas of the cockroaches ( dictyoptera ) of north america north of mexico .
breed , m . d . 1983 . cockroach mating systems , pp . 268 - 284 , in d . t . gwyne and g . k . moris ( eds . ) .
charlton , r . e . , webster , f . x . , zhang , a . , schal , c . , liang , d . , sreng , i . , and roelofs , w . l . 1993 . sex pheromone for the brownbanded cockroach is an unusual dialkyl - substituted α - pyrone .
farine , j . - p . , evaraerst , c . , abed , d . , ntari , m . , and brossut , r . 1996 . pheromonal emission during the mating behavior of
gautier , j . y . , deleporte , p . , and rivault , c . 1988 . relationships between ecology and behavior in cockroaches , pp . 335 - 351 , in c . n . slobodchikoff ( ed . ) .
gemeno , c . and schal , c . 2003 . sex pheromones of cockroaches . in r . t . cardé and j . millar ( eds . ) .
gorton , r . e . , jr . 1980 . a comparative ecological study of the wood cockroaches in northeastern kansas .
gorton , r . e . , jr . 1981 . behavioral and ecological correlations within a cockroach community .
hanula , j . l . and engstrom , r . t . 2000 . comparison of red - cockaded woodpecker (
hebard , m . 1917 . the blattidae of north america north of the mexican boundary .
kurtti , t . j . and brooks , m . a . 1976 . the dissociation of insect embryos for cell culture .
liang , d . and schal , c . 1993a . calling behavior of the female german cockroach ,
liang , d . and schal , c . 1993b . ultrastructure and maturation of a sex pheromone gland in the female german cockroach ,
nishino , c . , tobin , t . r . , and bowers , w . s . 1977 . electroantennogram responses of the american cockroach to germacrene d sex pheromone mimic .
noirot , c . and quennedey , a . 1974 . fine structure of insect epidermal glands .
roth , l . m . and barth , r . h . , jr . 1967 . the sense organs employed by cockroaches in mating behavior .
schal , c . and bell , w . j . 1985 . calling behavior in female cockroaches ( dictyoptera , blattaria ) .
schal , c . , gautier , j . - y . , and bell , w . j . 1984 . behavioural ecology of cockroaches .
schal , c . , liang , d . , hazarika , l . k . , charlton , r . e . , and roelofs , w . l . 1992 . site of pheromone production in female
schal , c . , liang , d . , and blomquist , g . j . 1996 . neural and endocrine control of pheromone production and release in cockroaches , pp . 3 - 20 , in r . t . cardé and a . k . minks ( eds . ) .
smith , a . f . and schal , c . 1990a . corpus allatum control of sex pheromone production and calling in the female brown - banded cockroach ,
smith , a . f . and schal , c . 1990b . the physiological basis for the termination of calling in the female brown - banded cockroach ,
smith , a . f . and schal , c . 1991 . circadian calling behavior in the adult female brown - banded cockroach ,
sreng , l . 1993 . cockroach mating behaviors , sex pheromones , and abdominal glands ( dyctioptera : blaberidae ) .
sreng , l . 1998 . apostosis - inducing brain factors in maturation of an insect sex pheromone gland during differentiation .
tokro , p . g . , brossut , r . , and sreng , l . 1993 . studies on the sex pheromone of female
wendelken , p . and barth , r . h . 1971 . the mating behavior of
( saussure and zehntner ) ( blattaria , blaberoidea , blattellidae , blattellinae ) .
yang , h . - t . , chow , y . - s . , peng , w . - k . , and hsu , e . - l . 1998 . evidence for the site of female sex pheromone production in
gemeno , c . , snook , k . , benda , n . et al . j chem ecol ( 2003 ) 29 : 37 . urltoken
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presence and absence of bats across habitat scales in the upper coastal plain of south carolina .
abstract during 2001 , we used active acoustical sampling ( anabat ii ) to survey foraging habitat relationships of bats on the savannah river site ( srs ) in the upper coastal plain of south carolina . using an a priori information - theoretic approach , we conducted logistic regression analysis to examine presence of individual bat species relative to a suite of microhabitat , stand , and landscape - level features such as forest structural metrics , forest type , proximity to riparian zones and carolina bay wetlands , insect abundance , and weather . there was considerable empirical support to suggest that the majority of the activity of bats across most of the 6 species occurred
effect of habitat and foraging height on bat activity in the coastal plain of south carolina .
a comparison of bat activity levels in the coastal plain of south carolina among 5 habitat types : forested riparian areas , clearcuts , young pine plantations , mature pine plantations and pine savannas , using time expansion radio - microphones and integrated detectors to simultaneously monitor bat activity at three heights in each habitat type .
tree - roost characteristics of subadult and female adult bats ( nyctieius humeralis ) in the upper coastal plain of south carolina .
tree - roost of evening bats were identified by radio tracking of 14 individuals at the srs . bats roosted in longleaf pine cavities under exfoliating bark in snags near beaver ponds . the roosting occurred in open park like stands . no evening bats roosted in the more dense bottomland hardwood stands or mixed pine hardwood stands . none were observed in loblolly stands .
assessment and comparison of richness , abundance and difference of herpetofauna at five small isolated wetlands located within a commercial forest landscape in the south carolina coastal plain . data indicates small isolated wetlands are focal points of herpetofaunal richness and abundance in managed coastal plain forest and contribute more to regional biodiversity than is implied by their small size or ephemeral hydrology .
factors limiting regeneration of quercus alba and cornus florida in formerly cultivated coastal plain sites , south carolina .
[ cite : 595888 ] bg ' standard ' source covering roach & termite species of the world . navigate by classification tree
[ cite : 1286620 ] link all of the nomenclatural , bibliographic , and specimen data accumulated in missouri botanical garden ' s ( mbg ) electronic databases during the past 30 years are publicly available here . this system has nearly 1 . 3 million scientific names and over 4 . 4 million specimen records . great resource for plant distribution beyond us & canada .
[ cite : 1185947 the collection holds more than 3 . 5 million insect specimens and is one of the largest university insect collections in the world . all groups of insects are represented in the collection , and we are recognized for our holdings of leafhoppers ( cicadellidae ) , beetles ( coleoptera ) , and true flies ( diptera ) . the triplehorn insect collection is housed within the museum of biological diversity , located on the west campus of the ohio state university in columbus , ohio . on 29 april , 2005 , the ohio state insect collection was renamed in honor of dr . charles a . triplehorn , professor of entomology and curator at osu between 1962 and 1992 .
keys to the insects of the european part of the ussr . vol . i : apterygota , palaeoptera , hemimetabola
taxonomy outdated ; otherwise , a sound source with lots of useful info . english version : keys to the insects of the european ussr . ( keys to the fauna of the ussr no . 84 ) jerusalem : israel program for scientific translations , 1967 . 1214 pp .
full text hunter , w . d . , f . c . pratt , j . d . mitchell . 1912 . the principal cactus insects of the united states . usda bureau of entomology bulletin 113 : 1 - 71 .
by furniss , r . l . and carolin , v . m . 1977 .
u . s . d . a . forest service misc . publ . 1339 , 1977
shorthouse j . d . , floate k . d . ( eds . ) arthropods of canadian grasslands , vol . 1 : 199 - 225 , 2010
( translated by leigh e . chadwick ) one of the best books on insects ever written , period . lavishly illustrated by the author . a rare example of beautiful book design and typesetting work . written by a swiss naturalist and painter , an authority on chrysididae . many editions available .
. each record tells when . see dataset links for citations & terms of use .