lyle 's flying fox ( pteropus lylei ) is a species of flying fox in the family pteropodidae .

it is found in cambodia , thailand , and vietnam .

it can do fairly well in some urban areas .

1 it is killed for bushmeat in parts of its range .

lyle ' s flying fox

lyle’s flying fox ( pteropus lylei ) is a medium - sized flying fox which forms large colonies high up in trees . lyle’s flying fox has a long dark muzzle , large eyes and a ring of orange fur around the neck , and bears some resemblance to a fox , hence the common name ‘flying fox’ ( 5 ) .
bbc two - lyle’s flying fox - thailand : earth ' s tropical paradise , the central heartland - in pictures . . .
farmers also pose a threat as they consider lyle’s flying fox to be a crop pest , resulting in persecution of this species ( 3 ) .
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lyle ' s flying fox is classified as vulnerable ( vu ) on the iucn red list ( 1 ) and is listed on appendix ii of cites ( 4 ) .
the diet of lyle’s flying fox consists mainly of ripe fruit . however , this species will also feed on nectar , pollen and blossoms to ensure it gets enough energy . fruit is very low in protein and sodium , so the salivary gland of lyle’s flying fox has become specially adapted to ensure this species can extract the required nutrients ( 7 ) .
the population of lyle’s flying fox is believed to be in decline , a trend which is expected to continue due to human pressures on its environment ( 1 ) . habitat loss is a major threat to the population ( 1 ) , with deforestation and construction projects irreversibly destroying the forests that lyle’s flying fox relies on for roosting and for food ( 3 ) .
unlike other bat families , fruit bats do not hibernate . instead , lyle’s flying fox produces heat by shivering , which keeps its body temperature between 33 and 37 degrees celsius ( 6 ) .
the wings and back of lyle ' s flying fox are dark brown or black , which strongly contrast against the bright fur around the head and neck . its lower body varies from a deep dark - brown to a brighter yellow - brown . its breast and belly are black - brown , which is similar to the large flying fox ( pteropus vampyrus ) ( 3 ) .
lyle’s flying foxes are the largest bats in the world , as some species can have a wingspan of up to two metres and weigh around 1 . 5 kg .
the lyle ' s flying fox is listed as least concern ( lr / lc ) , lowest risk . does not qualify for a more at risk category . widespread and abundant taxa are included in this category , on the iucn red list of threatened species
lanlua , p . , sricharoenvej , s . , niyomchan , a . and chico , d . e . ( 2007 ) unique cellular structures in the parotid gland of an old world fruit bat , pteropus lylei ( lyle ' s flying fox ) . italian journal of anatomy and embryology , 112 : 179 - 90 .
lyles’s flying fox can be found in some of southeast asia’s most secluded jungles , as well as in the middle of villages or even large urban metropolises such as bangkok . the species usually roosts in temples in the middle of urban areas ( 3 ) .
unlike the majority of animals in the chiroptera order , which are nocturnal , insectivores and orient themselves using ultrasounds , lyle’s flying foxes are crepuscular , feed on fruit , flowers and nectar and use their sight to guide them while flying and to find food .
the habitat of lyle’s flying foxes is limited to thailand , cambodia , viet nam , and the yunnan province of china . even though little is known about populations within china , lyle’s flying foxes have been listed by iucn as vulnerable or one step away from being considered endangered . until recent hunting restrictions were put into place , fruit bats were considered a delicacy and were hunted legally .
hondo , e . , inoue , n . , maeda , k . , rerkamnuaychoke , w . and duengkae , p . ( 2010 ) movement of lyle ' s flying fox ( pteropus lylei ) in central thailand . journal of wildlife in thailand , 17 : 55 - 63 .
lyle’s flying fox is native to southeast asia and is found in thailand , vietnam , cambodia , malaysia , and yunnan in china . this species has been mostly documented in thailand , where at least 11 colonies have been identified , the largest containing around 3 , 000 individuals ( 1 ) .
although it is a nocturnal species , lyle’s flying fox is very sociable and noisy during the day , as this is when females suckle their young ( 5 ) . large noisy colonies are very conspicuous , but they have few natural predators and so can hang safely up in the trees all day ( 6 ) .
lyle’s flying fox’s primary sense when foraging is vision , as it lacks the echolocation abilities of insectivorous bats . it has well developed teeth which are used to chew fruit while spitting out most of the seeds and pulp ( 5 ) . some seeds are ingested , which is important for the local ecosystem as it allows for the dispersal of seeds into other areas ( 2 ) .
lyle’s flying fox , and other species of bat , have been found to harbor large reservoirs of the nipah virus . although the virus does not harm the bat , it can be deadly to humans , and there is substantial data suggesting that recent outbreaks were caused by bat to person transmission of the virus ( 8 ) .
lyle’s flying fox is listed in appendix ii of the convention on international trade in endangered species ( cites ) , so trade in this species should be carefully monitored ( 4 ) . there are no known populations within protected areas in vietnam or cambodia . however , this species is protected by monks in thailand where populations roost in temple roofs ( 1 ) .
lyle’s flying foxes are the largest bats in the world , as some species can have a wingspan of up to two metres and weigh around 1 . 5 kg . they do not have tails , have a claw on the second digit of each wing and have two incisors on each maxilla . their faces are reminiscent of a small fox and , hence , its common name .
hunting still occurs in some places , but habitat degradation and destruction has also contributed to the 30 % population decline over the past 15 years ( three generations ) . in thailand and cambodia , lyle’s flying foxes are considered pests and are not protected except on temple grounds . for this reason , large , noisy colonies of lyle’s flying foxes can be found colonizing the trees over some buddhist temples in cambodia and thailand . up to 11 colonies are known to exist in thailand and the largest contains close to 3000 individuals .
lyle’s flying foxes will colonize trees in large groups , where they nest and suckle their young during the day . at night they depart to forage for food and have been known to travel 40 miles away to find food . rather than using echolocation as their insectivore cousins do , fruit bats rely on well - developed visual and olfactory senses .
in central thailand ' s forests , plains and city streets , nature lives alongside people .
fujita , m . ( 1988 ) flying foxes and economics . bats magazine , 6 : 1 . available at : urltoken
the size of fruit bats can be truly startling when first encountered at night by strangers to tropical climes and has resulted in reports of vampire bats or flying pterodactyls , but unless you’re a fruit or flower , or smell like one , you have little to fear . okay , a little fear is appropriate . over the past 10 years or so several research studies have found that fruit bats , including lyle’s flying foxes , are hosts for emerging viruses that can infect and kill livestock and
) belongs to the large family of mammalian old world fruit bats or flying foxes . the spanish name is zorro valador de lyle . as the common name suggests fruit bats live on nectar , pollen , blossoms , and fruit , and for this reason their habitats are limited to tropical and subtropical regions of the world . old world fruit bats are the largest bats in the world , with wing spans up to 6 feet ( 1 . 8 meters ) and weighing as much as 2 . 2 pounds ( 1 . 5 kg ) . by comparison , the size of lyle’s flying foxes is about mid - sized within the fruit bat family .
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luby , s . p . et al . ( 2009 ) recurrent zoonotic transmission of nipah virus into humans , bangladesh , 2001 - 2007 . emerging infectious diseases , 15 : 1229 - 1235 .
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this species is listed on appendix ii of cites . there are no known populations within protected areas in viet nam ( son nguyen truong pers . comm . ) or cambodia . the species is protected by monks in thailand ( s . bumrungsri pers . comm . ) .
nothing is known from china about the ecology of this species ( smith et al . 2008 ) . in other areas , it is known to form large colonies in trees that can become stripped of leaves by the bats ' activity . it feeds in orchards and is regarded as a serious pest in thailand ( s . bumrungsri pers . comm . ) . it occurs in mangrove forest in viet nam ( son nguyen truong pers . comm . ) . this species travels up to 50 km between colonies ( s . bumrugsri pers . comm . ) .
in thailand the species is threatened through loss of roosting habitat , as existing trees die and are not replaced , it is also subject to hunting ( s . bumrungsri pers . comm . ) . it is also threatened in cambodia by hunting ( p . bates pers . comm . ) .
echolocation detecting objects by reflected sound . used by bats and odontocete cetaceans ( toothed whales , dolphins and porpoises ) for orientation and to detect and locate prey . gland an organ that makes and secretes substances used by the body . hibernation a winter survival strategy in which an animal’s metabolic rate slows down and a state of deep sleep is attained . while hibernating , animals survive on stored reserves of fat that they have accumulated in summer . insectivorous insect - eating . nocturnal active at night .
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hutson , a . m . , racey , p . a . ( chiroptera red list authority ) , chanson , j . & chiozza , f . ( global mammal assessment team )
justification : listed as vulnerable as it is estimated that a decline of over 30 % has occurred in a fifteen year time period ( three generations ) including the past and the present , and this decline is expected to continue due to increasing trade and hunting pressure and ongoing decline in the extent and quality of its habitat .
this species is known from yunnan in china ( smith et al . 2008 ) , extending in to cambodia , thailand , and viet nam .
the largest known colony in thailand is about 3 , 000 individuals and up to 11 colonies have been identified . there are three known colonies in viet nam ( son nguyen truong pers . comm . ) .
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colonies inhabiting the forest roost in the tops of tall canopy trees , which they revisit year after year ( 5 ) ( 6 ) .
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wilson , e . d . ( 1997 ) bats in question : the smithsonian answer book ( second edition ) . smithsonian institution press , washington d . c .
allen , g . m . ( 1980 ) bats : biology , behavior and folklore . harvard university , dover publications inc .
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with an isolated population in yunnan ( china ) and extending through cambodia , thailand and vietnam .
gestation lasts from 140 to 190 days and only one baby is generally born in each birth , occasionally two . the mother takes the baby right when it is born and it remains clinging to her body for several days . after this period , the baby pup remains on tree branches and holes when the mother goes out to feed .
gregarious animals , they form large colonies that come together to rest in the immense trees in the jungle . the largest known colony is in thailand , comprised of some 3000 specimens .
their populations are shrinking considerably in recent years due to direct hunting , deforestation and the progressive deterioration of their habitat . according to the iucn ( international union for the conservation of nature ) , the species has diminished by some 30 % in the last 15 years .
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one specimen was at least 20 . 4 years old when it died in captivity [ 0671 ] .
[ 0730 ] smith et al . ( 2003 ) , body mass of late quaternary mammals
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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