The genera and biogeography of Fasciolariinae (Gastropoda, Neogastropoda, Fasciolariidae)

2012·1.4 MB·English

by  Snyder, Martin Avery

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The genera and biogeography of Fasciolariinae
(Gastropoda, Neogastropoda, Fasciolariidae)
Martin Avery Snyder
DepartmentofMalacology,AcademyofNaturalSciencesofPhiladelphia,19thandBenjaminFranklinParkway,Philadelphia,PA19103-1195,USA.
MuséumNationald’HistoireNaturelle,55,rueduBuffon,F-77005Paris,France;
[email protected][correspondingauthor]
Geerat J. Vermeij
DepartmentofGeology,UniversityofCaliforniaatDavis,OneShieldsAvenue,Davis,CA95616;[email protected]
William G. Lyons
4227PorpoiseDrive,SE,St.Petersburg,FL33705,USA;[email protected]
31
TheclassificationofFasciolariinae,agroupoflargepreda- 1870),Recent,SouthAfrica;LiochlamysDall,1889(type
toryfasciolariidneogastropods,isrevisedatthegeneric species:MazzalinabulbosaHeilprin,1886),EarlyPleistocene,
level.Typeselectionsforallgeneraaremade,andalltypes Florida;Lugubrilariagen.nov.(typespecies:Fasciolaria
areillustrated.Onthebasisofshellcharactersofmostfossil lugubrisA.Adams&ReeveinReeve,1847),Pleistoceneand
andalllivingspecies,werecognizefourteengenera:Africo- Recent,SouthAfricaandNamibia;PleuropocaFischer,1884
lariagen.nov.(typespecies:FasciolariarutilaWatson,1882), (typespecies:MurextrapeziumLinnaeus,1758),Late
Recent,AgulhasBank,SouthAfrica;Aurantilariagen.nov. MiocenetoRecent,Indo-WestPacific;Pliculofususgen.nov.
(typespecies:FasciolariaaurantiacaLamarck,1816),Early (typespecies:FasciolariascalarinaHeilprin,1886),Early
MiocenetoRecent,westernAtlanticandEurope;Australaria MiocenetoPleistocene,southernandeasternUnitedStates
gen.nov.(typespecies:PyrulaaustralasiaPerry,1811),Mid- (containingspeciesformerlyincludedinTerebraspira);Tere-
dleMiocenetoRecent,southernandeasternAustralia; braspiraConrad,1862(typeandonlyspecies:Fasciolariaele-
CincturaHollister,1957(typespecies:PyrulahunteriaPerry, gansEmmons,1858),EarlyPliocene,easternNorthAmerica;
1811),PliocenetoRecent,easternNorthAmerica;Fasciolaria andTriplofususOlsson&Harbison,1953(typespecies:Fasci-
Lamarck,1799(typespecies:MurextulipaLinnaeus,1758), olariagiganteaKiener,1840),PliocenetoRecent,westernAt-
EarlyPliocenetoRecent,westernAtlantic;Filifususgen.nov. lanticandeasternPacific.WealsodescribeAustralaria
(typespecies:FususfilamentosusRöding,1798),Recent,Indo- tenuitestaspec.nov.
WestPacific;Granolariagen.nov.(typespecies:Murexsalmo UnusualfeaturesthathaveevolvedintheFasciolariinae
Wood,1828),EarlyMiocenetoRecent,tropicalAmerica;Kil- includeanenvelopedshell(Liochlamys),smallfoldsonthe
burniagen.nov.(typespecies:FasciolariaheynemanniDunker, innersideofthesiphonalcanal(Pleuroploca,Pliculofusus,
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Terebraspira),beadedordiscontinuousridgesontheinner Miocene,Europe;TarantinaeaMonterosato,1917(type
sideoftheouterlip(someAurantilaria,Filifusus,Pliculofusus, species:MurexlignariusLinnaeus,1758),MiocenetoRecent,
Terebraspira),andacorrugatedoperculum(Aurantilaria, Europe;andSaginafususWenz,1943(typespecies:Fusus
someTriplofusus). priceiE.A.Smith,1887),northernAustraliaandIndonesia.
SeveraltaxaassociatedinthepastwithFasciolariaor ViridifususisassignedtoFusininae(Fasciolariidae);
Fasciolariinaearereassignedtoothermajorgroups.Besides TarantinaeaandNeolatirusareassignedtoPeristerniinae;
PleiaFinlay,1930,whichBeu(2011)reassignedtoPeristerni- andSaginafususisassignedtoMelongenidae.
inaeinFasciolariidae,thesetaxainclude:Fususbuxeus
Reeve,1847(typespeciesofViridifususgen.nov.Recent,
CapeVerdeArchipelago);NeolatirusBellardi,1884(type Keywords:Fasciolariidae,Fasciolariinae,Gastropoda,Miocene,
species:FasciolariarecticaudaFuchs,1877),MiddletoLate Pliocene,Recent,shellmorphology,classification,biogeography.
Introduction confinedtoasinglerowattheshoulderangulation.In
Australaria,sculpturalsimplificationandthetendencyto
ThefasciolariidgastropodsubfamilyFasciolariinaeisa loselirae(internalridges)particularlycharacterizespecies
small,morphologicallycompactgroupoflargetogigantic thatoccurincool-temperatewaters,suchasVictorianand
buccinoideanneogastropods.Despitetheirlarge,showy SouthAustralianpopulationsofA.australasia.Akeel-like
shells,manyofwhichwerewellknowntoearlyEuropean entrancefoldtothesiphonalcanalcharacterizesmany(but
naturalists,thegroupremainstaxonomicallyandecologi- notall)speciesofAustralariaaswellasspeciesofFilifusus.
callypoorlyknown.Thissurprisingignoranceiscommon Theentrancefoldneverdevelopsintoakeel-likefeaturein
32 amonglarge-bodiedmolluscsgenerally,includingmany PleuroplocaandAurantilaria,eveninspecieswithafullcom-
generawithinTurbinidae,Turbinellidae,andMelongenidae, plementofspiralrowsofnodes.Ourdiagnosestherefore
amongothers. reflectmaximalexpressionsofcharacters.
Ourpurposeinthispaperistooutlinegenus-level Typespeciesareidentifiedforallgeneraandtype-speci-
classificationandbiogeographicrelationshipsoftheFascio- mensarelocatedordesignatedforeachtypespecies;allare
lariinaebasedonourexaminationofshellcharactersofthe illustrated.Wealsoexcludeseveralgeneraandspecies
livingspeciesandofmanyfossiltaxa.Eightnewgeneric usuallyassignedtoFasciolariinaeandsuggestplacementin
namesareintroduced(Africolaria,Aurantilaria,Australaria, othergroups.Anadditionalnewgenericname(Viridifusus)
Filifusus,Granolaria,Kilburnia,Lugubrilaria,Pliculofusus)and isintroducedtoaccommodatesomeofthosespecies.Wedo
anewspeciesisdescribed(Australariatenuitesta).Wehave notcommentonthetaxonomicstatusofmanyofthespecies
chosentonamethesegroupsatthegenuslevelratherthan namesatthistime.Weareawarethatsomenameslistedin
atthelevelofspeciesgroupasataxonomichypothesistobe our“includedspecies”listshavebeensynonymizedby
testedlaterwithmoleculardata.Thegeneraaredistin- others,butfeelthatthestatusofthesetaxashouldbere-
guishedfromeachotherbycombinationsofcharacters examinedinlightofourgenericconclusions.Wenotethat
ratherthanbysingletraits. moderngeneticworkhastendedtosupporttheworkof
Nearlyeverycharacterusedincombinationtodiscrimi- “splitters”asopposedto“lumpers”withrespecttovarious
nateamonggeneravariesinexpressionamongspeciesand otherexaminedgenera.
evenamongpopulations.Forexample,althoughPleuroploca,
Aurantilaria,andAustralariamayallhavemultiplerowsof Abbreviations:AMS,Australian,Museum,SydneySouth,NSW,Aus-
nodesonthelastwhorl,therearespeciesandpopulationsin tralia;ANSP,AcademyofNaturalSciences,Philadelphia,PA,USA;
eachgenusinwhichnodesareeitherentirelyabsentor BMSM,Bailey-MatthewsShellMuseum,Sanibel,FL,USA;CAS,Cali-
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forniaAcademyofSciences,SanFrancisco,CA,USA;ICZN,Interna- theshellexterior.Theouterliplacksanadapicalsinusinall
tionalCommissionofZoologicalNomenclature;IWP,Indo-westPacific; generaexceptsomemembersofAustralaria,afeaturepres-
MHNG,Muséumd’HistoireNaturelleGenève,Geneva,Switzerland; entinallPeristerniinaeexceptsomespeciesofHemipolygona
MNHN,MuséumNationald’HistoireNaturelle,Paris,France; Rovereto,1899(seeVermeij&Snyder,2006).Nofasciolariine
NHMUK,TheNaturalHistoryMuseumoftheUnitedKingdom,Lon- hasdevelopeddenticlesontheinnersideoftheouterlip,
don,England;NSMT,NationalMuseumofNatureandScience,Tokyo, featuresthatarewell-developedinseveralperisterniinegen-
Japan;PRI,PaleontologicalResearchInstitute,Ithaca,NewYork; eraincludingPeristerniaMörchl852,TurrilatirusVermeij&
SBMNH,SantaBarbaraMuseumofNaturalHistory,SantaBarbara,CA, Snyder,2006,andLatirusMontfort,1810,amongothers.The
USA;SL,Shelllength;USNM,NationalMuseumofNaturalHistory, siphonalprocessofFasciolariinaeisalwaysparalleltothe
SmithsonianInstitution,WashingtonDC,USA;TU,TulaneUniversity axisofcoiling,notdeviatedeitherdorsallyortothesideas
(prefixforpaleontologicallocalitiesandassociatedcollections;speci- insomePeristerniinae.AlthoughtheFasciolariinaeand
mensnowatUF);UF,FloridaMuseumofNaturalHistory,Gainesville, mostPeristerniinaearecharacterizedbythepresenceof
FL,USA;WFIS,WagnerFreeInstituteofScience,Philadelphia,PA, abapicalcolumellarfolds,thereareimportantdifferencesin
USA;WGL,CollectionofWilliamG.Lyons,St.Petersburg,FL,USA. themagnitudeandorientationofthefolds.ThefoldsinFas-
ciolariinaeareelongateandveryobliqueandthischaracter
Systematic part unitesthespeciesofthesubfamily.
Besidestheextremelylargesizeofseveralfasciolariines,
SubfamilyFasciolariinaeGray,1853 especiallyinTriplofusus,butalsoinFasciolariaLamarck,
1799,PleuroplocaP.Fischer,1884andtheextinctnewgenus
Description.—Shellfusiform,medium-sizedtoextremely Pliculofusus(Terebraspiraauctt.,nonConrad,1862),theFasci-
large;outerlipusuallymediallyconvex,withoutadapical olariinaehaveevolvedthreecharacterstatesthatareother-
sinus;outer-lipedgeusuallywithpairedcrenulations;inner wiseunknownintheFasciolariidae.Theseare(1)an 33
lipadherent,abapicallybearingtwocolumellarfoldsanda externallyglazedshell(intheextinctgenusLiochlamysDall,
moreprominentfoldatentranceofsiphonalcanal;adaxial 1889),indicatingenvelopmentbythemantleorfoot;(2)the
marginofsiphonalcanaloftenerect;siphonalprocessparal- extremedevelopmentoftheentrancefoldtothesiphonal
leltoaxisofcoiling. canalintoakeel-likefeature,asseeninthegeneraAus-
Remarks.—TheFasciolariinaearemorphologicallysimi- tralariaandFilifusus;and(3)thepresenceofdistinctfolds
lartothefasciolariidsubfamilyPeristerniinaeTryon,1880. alongtheadaxialmarginofthesiphonalcanalabapicalto
Bothgroupsarecharacterizedbythepresenceofabapical theentrancefold,asseeninadultsofPleuroploca,Pliculofusus
columellarfolds,anentrancefoldtothesiphonalcanalsitu- andTerebraspira.
atedimmediatelyanteriortothecolumellarfolds,anda Forgenerathattypicallycontainspecieswithnodular
(usually)crenulated,simpleouterlipthatisusuallylirateon axialribs,ourdescriptionsmayonlymentionthatcondition.
itsinner(adaxial)side.Theseliraearepresentinfully However,mostspeciesthattypicallyhaveshellswithnodu-
formedshells.However,matureindividualsinTriplofusus laraxialribsalsooccur,notuncommonly,inanodoseforms.
Olsson&Harbison,1953,areoftensmoothontheinnerside Individualsorevenlocalpopulationswithanodoseshells
oftheouterlip,andallAfricolarialacksuchlirae.Internal occuramongspeciesofTriplofusus,Pleuroploca,Filifusus,
liraearecreatedlateintheshelldepositionalprocess,anda Granolaria,Aurantilaria,andAustralaria,andseveralofthese
shellwithanapparentlycompletelip(asevidencedbya anodoseformshavereceivedseparatenames.Smoothforms
relativelythickedge)maynotyethavedepositeditsfull ofspecieswithnormallynodoseshellsmayresultfrom
complementoflirae.Nevertheless,theFasciolariinaeare ecophenotypy(Aurantilariaaurantiaca),localgeographic
relativelyhomogeneousinshellfeatures.AllFasciolariinae isolation(Filifususfilamentosusaltimastus),regionalvariation
haveanadherentinnerlipthatmergesimperceptiblyinto (Pleuroplocatrapeziumaudouini;P.t.lischkeana)orforreasons
Snyder,M.A.etal.–GeneraandbiogeographyofFasciolariinae
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notyetunderstood(Triplofususgiganteusreevei;Granolaria (1970:99,100,textfig.5)andBandel(1984:144,textfig.260,
valenciennesii;Pleuroplocat.trapezium;Australariaaustralasia pl.17figs9,10).
coronata;Lugubrilarialugubris).Andofcoursesomegenera
havespecieswithtypicallynodoseshellsandotherswith Includedspecies:
typicallysmooth(anodose)shells.Ifthetypespecieshas FasciolariabullisiLyons,1972:Recent,offwesternFloridaandYucatán
nodoseshells,theotherconditionsmaynotbementioned. Platform,Mexico;
Taxonomically,mostmembersoftheFasciolariinaehave FasciolariacalusaPetuch,1994:EarlyPleistocene(CaloosahatcheeForma-
beenassignedtotwogenera:FasciolariaandPleuroploca, tion),Florida;
thoughsomeAmericanspecieshavebeenplacedin FasciolariahollisteriWeisbord,1962:PleistocenetoRecent,western
TriplofususandtheextinctTerebraspiraandLiochlamys.These Venezuela,CaribbeanColombia,andAruba;
generahavenotbeencriticallycomparedorcarefully Fasciolaria(distansvar.)monocingulataDall,1890:EarlyPleistocene
defined,andinthecaseofFasciolariaandPleuroplocaascon- (CaloosahatcheeFormation),Florida;
ventionallyused,theyareheterogeneous,comprising FasciolariaokeechobeensisTucker&Wilson,1932:MiddlePleistocene
severaldistinctclustersthatweprefertotreatasseparategen- (BermontFormation),Florida;
era.Webelievetheproposalofformalgenericnamesforthese FasciolariseminolePetuch,1994:EarlyPleistocene(CaloosahatcheeFor-
clusterswillfacilitatefuturemolecularandphylogenetic mation),Florida;
studiesofthissurprisinglyoverlookedgroupofgastropods. FasciolariasemistriataG.B.SowerbyI,1850:EarlyPliocene(GuraboFor-
mation),DominicanRepublic;
FasciolariaLamarck,1799 FasciolariasemistriataleuraWoodring,1928:Plio-Pleistocene(Bowden
Typespecies:MurextulipaLinnaeus,1758,bymonotypy. Formation),Jamaica;
Lectotype:specimenfiguredbyd’Argenville,1842:pl.13fig.K,selected FasciolariasemistriatamareanaWeisbord,1962:Plio-Pleistocene(Lower
34 herein(Fig.1). MareFormation),Venezuela;
FasciolariatephrinadeSouza,2002:Recent,offHonduras,Nicaraguaand
Description.—Shelllarge(2largestlivingspeciesto275 northernColombianislands,westernCaribbean;
and278mmSL),broadlyfusiform,usuallyratherthin;pro- MurextulipaLinnaeus,1758:LatePleistocenetoRecent,NorthCarolina
toconchwithdistinctorfaintaxialribletsonfinalwhorl; toFlorida,theGulfofMexicoandCaribbeanSeatoSurinameand
teleoconchsculptureusuallypredominantlyoflowtoobso- Maranhão,andAmapá,Brazil.
letespiralcordswhicharestrongestonthebasalconstric-
tionandsuturalrampandmaybeabsentonthecentral Remarks.—OfthreespecimensofMurextulipainthecollec-
sectorofthebodywhorl(however,someshellswithrugose tionoftheLinneanSociety,BurlingtonHouse,London,one
cordsoverentireteleoconch);axialsculpture,whenpresent, ismarked“489”(thenumbergiventhespeciesbyLinnaeus
consistingofweakroundedribsonearlyteleoconchwhorls; inhis10thedition),aspecimenprobablyinLinnaeus’hands
shellsurfaceornamentedwithmanyintactorinterrupted whenhedescribedthespecies,but,asS.P.Danceindicated
spiralbands;outerlipmediallyconvex,itsedgeoftenwith inanotethataccompaniesthematerial“thatshellisina
strong,sometimespairedcrenulationsrepresentingtermini ‘shockingstate’;thespireisbroken,thesiphonischipped,
ofspiralbands,itsinnersidewithfinesmoothlirae;shallow andthebodywhorlisriddledwithwormholes”.Anotheris
notchorsimpleflexure,causedbyflexureinsuturalramp, marked“562”(thecorrespondingspeciesnumberinthe12th
presentnearjunctionofouterlipandpenultimatewhorl; edition)andoneisunmarked.Theunnumberedspecimenis
parietalridgeatadapicalendofapertureabsent;entrance immatureandalsoshowsconsiderablewear,perhapsfrom
foldmoreprominentthancolumellarfolds,rounded;no usebyhermitcrabs.Preferenceasalectotypemightgoto
foldsabapicaltoentrancefold.Radulaoftypespeciesde- theshellmarked“489”.Linnaeus(1758)citedforMurex
scribedandfiguredbyTroschel(1868:62,pl.5fig.12),Wells tulipafiguresbyBuonanni(1684:fig.187),Rumphius(1705:
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pl.49fig.H),Gualtieri(1742:pl.46fig.A),Dezallierd’Ar- thatmanyofthefossiltaxaarevalid,andthatadditionalRe-
genville(1742:pl.13fig.K)andRegenfuss(1758:pl.9fig.35). centspeciesmayberecognizedwithmolecularsequencing.
Ofthese,thefigurebyd’Argenvilleisleaststylizedandtruest TheearliestspeciesofFasciolariathatweareabletover-
totheformoftheadultshell,andweselectit,ratherthanthe ifyisF.semistriata,whichaccordingtoB.M.Landau(per-
decrepitspecimen“489”,aslectotypeofMurextulipa.Inthe sonalcommunicationtoGJV,16November,2009)isfromthe
1757secondeditionfigureKappearsonplate10. basalGuraboFormation(earlyPliocene)oftheDominican
Asrestrictedhere,FasciolariaisawesternAtlanticgenus Republic.Thislateoriginofthegenuslikelyaccountsforthe
withratherthinshellsinwhichthesculptureisreducedrel- absenceofFasciolariaintheeasternPacific,becausetheCen-
ativetothatofmostotherFasciolariinae.Anexceptionin- tralAmericanseawayconnectingtheAtlanticwiththePa-
volvesthescheepmakeri(Kobelt,1875)formofF.tulipa,a cificwasalreadysubstantiallyconstrictedbythattime.Very
Recentshellthatisheavierthanthetypicalformandwhose similartobutmuchyoungerthanF.semistriataisF.semistri-
entiresurfaceiscoveredwithstrongcords;similarheavily ataleura,fromthePlio-PleistoceneBowdenBedsofJamaica.
sculptedshellssometimesoccuramongspecimensofF.okee- Concurrentwithleura,F.semistriatamareanaandF.hollisteri
chobeensis.Thereisvariationintheextentandstrengthof appearedinthePlio-PleistoceneLowerMareFormationof
spiralsculpture,withF.tulipaandF.hollisterihavingsubsu- Venezuela,followedbyF.calusa,F.okeechobeensis,F.
turalcordsthatarelackinginotherlivingspecies.Fasciolaria seminole,andF.tulipaintheEarlytoLatePleistoceneof
bullisiandF.tephrinaareretainedinFasciolariaratherthan Florida.AlthoughF.tulipanowrangesnorthwardtoNorth
Cincturabecause:1)theirshellslackanyindicationofapari- Carolina,allfossilrecordsofFasciolariafromtheCarolinas
etalridge;2)theyexhibitflexureintheirsuturalramps,al- andVirginiaareofspeciesnowclassifiedinCinctura,
beitslight,thatcreatesaslightangleandinterruptsthe Pliculofusus,TriplofususorTerebraspira.Likewise,aspeciesre-
curvatureoftheouterlip;3)theirspiralbandsarenumerous portedasFasciolariasemistriatafromMiocenebedsofMexico
andofteninterrupted,and4)theirshellsarethinnerand (PerrilliatMontoya,1960:22,23,pl.3figs12-15)andlateras- 35
lighterinweightthanthoseofCincturaofcomparablesize. signedtothemiddleMioceneAgueguexquiteFormation
PlacementoftheEarlyPleistoceneFasciolariamonocingu- (Perrilliat,1981:64)isnotF.semistriataandappearstobea
lataisproblematic.Dall(1890)introducedthetaxonasa speciesofCinctura.SeemoreonthisrecordinRemarksfor
subspeciesofF.distans(now=Cincturalilium)andplace- Cinctura.AlthoughseveraloftheRecentspecies(e.g.,F.
mentinCincturaissupportedbyitspossessionofanemer- bullisi,F.hollisteriandF.tephrina)haveratherlimiteddistri-
gentparietalridge.However,theshellalsohasachanneled butions,therangeofF.tulipa(Amapá,BraziltoNorthCar-
suturalramp,whichcreatesadistinctinflectioninthesu- olina)isamongthegreatestofanywesternAtlantic
turalrampandouterlip,afeaturecharacteristicofFascio- fasciolariid,beingexceededonlybythatoftheperisterniine
laria.WehaveexaminedtwospecimensofFasciolariatulipa Leucozonianassa(Gmelin,1791),whichrangesfromNorth
withwell-developedparietalridges,amongmorethan600 CarolinatooceanicislandsoffcentralBrazilandformerly
specimensthatlackthatfeature,suggestingthattheridge extendedtoBermuda(Lyons&Snyder,2008).Fasciolaria
mayberelatedtoageneusuallyrecessiveintheFasciolaria tulipahasalsobeenrecordedatvirtuallyeveryCaribbeanis-
lineagebutwhichmayhavebeenovertlyexpressedin land,regardlessofthedepthsofsurroundingwaters,alto-
F.monocingulata. getheraremarkablefeatofdispersalforaspecieswith
demersaldevelopment.
Wehavenotattemptedtoevaluateeachofthespecies-
leveltaxaassignedtoFasciolaria.Thisappliesinparticularto CincturaHollister,1957
theseveraltaxaofPlioceneandEarlytoMiddlePleistocene Typespecies:PyrulahunteriaPerry,1811,byoriginaldesignation.
fossils.Giventhepaucispiralprotoconchs(andimplicitly Neotype:USNM615769,86.5mmSL,Recent,Charleston,SouthCar-
nonplanktonicdispersal)ofthelivingspecies,itispossible olina,selectedherein(Fig.3).
Snyder,M.A.etal.–GeneraandbiogeographyofFasciolariinae
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Description.—Shellofsmalltomoderatesize(2largestliv- dlePliocene(Yorktown,JacksonBluffandTamiamiFormations),
ingspeciesto131and140mmSL),broadlyfusiform,slen- Virginia,NorthCarolina,SouthCarolinaandFlorida;
dertorelativelyglobose,solid;protoconchwithorwithout Fasciolaria(Cinctura)rucksorumPetuch,1994:EarlyPleistocene(Nashua
axialribletsonfinalwhorl;teleoconchsurfacegenerally Formation),Florida;
smoothexceptforoccasionalfaintspiralincisionsandvery Fasciolaria(Cinctura)sarasotaensisPetuch,1994:LatePliocene(Upper
lowaxialribsonfirst1-3whorlsandtransversespiralcords Pinecrestbeds,TamiamiFormation),Florida;
onbaseandsiphonalprocess;distinct,uninterruptedspiral Fasciolaria(Cinctura)liliumtortuganaHollister,1957:Recent,NorthCar-
bands,usuallyblack,onallwhorls,bandsonbodywhorl olinatoFlorida,Alabama,MississippiandeasternLouisiana.
varyinginnumberfrom4toasmanyas14,dependingon
species;outerlipmediallyconvex,itsedgesometimesbear- Remarks.—Hollister(1957)differentiatedCincturaasasub-
ingthickenednodesrepresentingterminiofspiralbands,its genusofFasciolaria,principallyonthebasisofaprominent
innersidewithfinesmoothlirae;adapicalflexureabsentin parietalridgethatemergesfromwithintheapertureofthe
outerlip;parietalridgeatadapicalendofapertureabsentin shell.SpeciesheassignedtoCincturaalsolacktheinflected
earliest(earlytomid-Pliocene)material,presentandoften suturalramp,sometimeswithsubsuturalspiralcords,the
prominentinalllate-PliocenetoRecentspecies;entrance resultantflexureontheouterlip,andoften-interruptedspi-
foldmoreprominentthancolumellarfolds,rounded;no ralbandsofFasciolariaspeciesandhavespiralsculpturecon-
foldsabapicaltoentrancefold.Radulaoftypespeciesde- finedtotheconcavebaseofthebodywhorlandthe
scribedandfiguredbyHackney(1945:46,48,pl.1figs5,6, siphonalprocess.WehereelevateCincturatogenus-level
asF.distansauctt.,nonLamarck,1822)andWells(1970:99, rankasacladedistinguishedfromFasciolariabyasuiteof
100,textfig.5). charactersequivalenttothoseofmanysimilartaxarecog-
nizedatgenuslevelinotherfamilies(e.g.,Cypraeidae,Mu-
36 Includedspecies: ricidae,Conidae).
FasciolariaapicinaDall,1890:EarlyPleistocene(CaloosahatcheeForma- Theblue-graycolorandrelativelygloboseformofthe
tion),Florida; shelldepictedbyPerry(1811)forPyrulahunteria(Fig.2)are
FasciolariabeaufortensisWard&Blackwelder,1987:EarlyPleistocene featuresmostoftenfoundinestuarineCincturapopulations
(JamesCityFormation),NorthCarolinaandVirginia; ofthesoutheasternUnitedStateseastoftheMississippi
FasciolariadistansbranhamaeRehder&Abbott,1951:Recent,southern River,butshellsofsimilarcolorandshapealsooccurun-
TexasandBayofCampeche,Mexico; commonlyinsomeMexicanpopulations.Featuresthatdis-
Fasciolaria(Cinctura)capelettiPetuch,1994:MiddlePleistocene(Bermont tinguishtheAtlanticandeasternGulfCincturahunteriafrom
Formation),Florida; thewesternGulfC.liliumarenumbersofprimaryspiral
Fasciolaria(Cinctura)evergladesensisPetuch,1991:MiddlePleistocene bandsonthebodywhorl(4-7onC.hunteria,7-11onC.
(BermontFormation),Florida; lilium)andabsence(C.hunteria)orpresence(C.lilium)ofdis-
Fasciolaria(Cinctura)holeylandicaPetuch,1994:MiddlePleistocene tinctribletsontheprotoconch.Perry’sfiguredshellhas7
(lowerBermontFormation),Florida; primarybands,itsprotoconchistoosmallforinspection,
PyrulahunteriaPerry,1811:LatePleistocene(Ft.ThompsonFormation) anditstypeisnotknowntobeextant(Petit,2003:17),soits
toRecent,NorthCarolinatoFlorida,Alabama,Mississippiand statusasan“eastern”or“western”shellisambiguous.Be-
easternLouisiana causeofthisambiguitywehaveelectednottodesignatea
FasciolarialiliumFischervonWaldheim,1807:Recent,Cancun,Mexico lectotypebaseduponthisillustration.
tosoutheasternTexas; Inwhatwasthenacontroversialaction(seeBurch,1957:
Fasciolaria(Cinctura)lindaePetuch,1994:EarlyPleistocene(Caloosa- 9;Petit,2003:15),Hollister(1957)invokedthelawofpriority
hatcheeFormation),Florida; toreplacetheuniversallyrecognizedFasciolariadistans
FasciolariarhomboideaW.B.Rogers&H.D.Rogers,1839:EarlyandMid- Lamarck,1822,withF.liliumFischervonWaldheim,1807
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(calledtheCampecheformbyHollister)forwesternGulf theyarenotMiocene.AnoteappendedtotheTulanemate-
bandedtulipsandPyrulahunteriaPerry,1811(calledthe rialbyDr.EmilyVokesadds:“up.Miocene[reallyL.
Floridaform)foreasternpopulations.Hollisteralsodesig- Pliocene]correlateswithBrightonBeds=uppermost
natedthelatterastypespeciesofhisnewgenus-leveltaxon Pinecrest,”i.e.LatePlioceneorEarlyPleistocene.
Cinctura.PresumingthatPerry’stypeofhunteriawaslost FossilandRecentCincturaareconfinedtowaterscon-
andrecognizingtheneedfornomenclaturalstability,Hollis- tiguoustothesoutheasternUnitedStatesandtheGulfof
ter(1957:75,83)triedtoclarifyandfixtheidentityofhis Mexico,andthereisnoevidencethatthegenushaseveroc-
typespeciesbydesignatingtwospecimenscollectedbyEd- curredintheCaribbeanregionorelsewhere.Wehavetraced
mundRavenelnearCharleston,SouthCarolina(USNM anddisprovedseveralCaribbeanlistingsofRecentspecies
615769)asneosyntypesofPyrulahunteriatherebyfixing nowassignedtoCinctura,andweregardthoseremainingas
Perry’snametotheeasternform.Areasonablereadingof erroneous.TheCincturaspeciesgroupisacontinentalline-
Hollister’srationaleandtheinformationheconsideredre- agemuchlikethemelongenidBusyconclade(e.g.,Busycon,
vealsthathisactionwasjustifiedandhesatisfiedeachofthe Busycotypus,Pyruella)thatevolvedineasternNorthAmerica
severalrequirementssetforthbytheInternationalCodeof andhasnotextendeditsrangebeyondtheYucatanPlatform
ZoologicalNomenclaturefordesignationofaneotype.Re- ofMexicoandintotheCaribbeanBasin.
grettably,though,theCodemakesnoprovisionforneosyn-
types,soHollister’sdesignationcannotstand.Inaccord LiochlamysDall,1889
withHollister’sintent,weselectthe86.5-mmspecimenfrom Typespecies:MazzalinabulbosaHeilprin,1886,byoriginaldesignation.
thisUSNM“neosyntypelot”asneotype. Holotype:WFIS909,59.2mmSL,bysubsequentdesignation,Spamer&
Cincturarhomboidea,anEarlyandMiddlePliocene Forster(1988:48)(Fig.4).
speciesknownfromVirginiatoFlorida,istheoldestknown
representativeofthegenus.Itssmoothshellandlackofan Description.—Shellrathersmallforsubfamily(largest 37
inflectedsuturalramparesimilartofeaturesofitslatercon- speciesto120mmSL),broadlyfusiform,polished,exter-
geners,butearliestexamplesofthespecieslackaparietal nallyenameled,devoidofsculptureexceptforweakthreads
ridge.TheparietalridgeappearsintheupperPinecrestbeds onadapicalhalfofsiphonalprocess;outerlipstronglycon-
oftheTamiamiFormationinFlorida,whereshellswithand vex,crenulated,itsinnersidewithstrongsmoothlirae;junc-
withouttheridge,allstillcalledC.rhomboidea,occurwithin tionofouterlipwithpenultimatewhorlmarkedbydeep
thesamestrata.ShellsoftheLatePlioceneC.sarasotaensis, adapicalnotch;twocolumellarfolds,theabapicaloneofap-
EarlyPleistoceneC.apicina,C.beaufortensis,C.lindaeandC. proximatelyequalprominenceastheroundedentrancefold
rucksorum,MiddlePleistoceneC.capelettii,C.evergladesensis tothesiphonalcanal;foldsabapicaltoentrancefoldon
andC.holeylandica,andtheRecentC.hunteria,C.lilium,and innersideofsiphonalcanalabsent;parietalridgeabsent.
C.branhamaealldisplaytheparietalridgethatcharacterizes
membersofCinctura. Includedspecies:
Asnotedabove,shellsfiguredasFasciolariasemistriataby MazzalinabulbosaHeilprin,1886:EarlyPleistocene(CaloosahatcheeFor-
Perrilliat-Montoya(1960;1981)fromtheMioceneAgueguex- mation),Florida;
quiteFormationnearCoatzacoalcos,Veracruz,Mexicoseem LiochlamysgriffiniPetuch,1994:EarlyPleistocene(CaloosahatcheeFor-
torepresentaspeciesofCinctura.SeveralimmatureCinctura mation),Florida.
specimensexaminedbyoneofus(WGL)fromthe
AgueguexquiteFormationatloc.TU638nearCoatzacoal- Remarks.—ThefossilgenusLiochlamysisuniquewithin
cos,Veracruz,Mexico(E.H.Vokes,1993:149)maybecon- Fasciolariidaeinhavingapolished,enameledexterior,indi-
specific,althoughtheyaretooimmaturetobecertain.These catingenvelopmentbythemantleorfootduringlife(Ver-
seemtobetheonlyrecordsoffossilCincturainMexico,but meij,2005).WecannotacceptPetuch’s(1994)suggestionthat
Snyder,M.A.etal.–GeneraandbiogeographyofFasciolariinae
B76-Snyder_etal:Basteria-2010 12/07/2012 16:50 Page 38
LiochlamysisderivedfromaspeciessuchasFasciolariacalusa Remarks.—ThegenericnameTerebraspirahasbeenwidely
Petuch,1994,alsofromtheCaloosahatcheeFormationof usedforOligocenetoPleistocenefasciolariinesfromthe
Florida.ShellsofLiochlamysaremoreglobosewithrelatively southeasternUnitedStates,butourexaminationofthetype
shorterspires,lackanysubstantialsuturalconstrictionbe- species,FasciolariaelegansEmmons,1858,revealsthespecies
tweenwhorls,andhaveasiphonalprocessrelativelyshorter tobesodistinctivethatitcannotbegroupedwithanytaxa
thanthatofanyotherFasciolariinae.TheshellshapeofF. thatothershaveassignedtoTerebraspira.Asdiscussed
calusaisverymuchlikethatofF.tulipa,withwell-rounded below,thoseotherspeciesarehereassignedtothenew
whorls,distinctsutures,awell-developedsiphonalprocess, genusPliculofusus.
andaprominentprotoconch,butitsshellsurfaceisgener- Thenow-restrictedmonotypicgenusTerebraspirahasa
allysmoother,sculptureonitssuturalrampisfainter,and distinctivecombinationofcharacters.Theabsenceofaxial
protoconchsofthetwospeciesarequitedifferent. sculptureonallbutthefirsttwoteleoconchwhorlsrecalls
Fasciolaria,Cinctura,andmorphotypesofseveralothergen-
TerebraspiraConrad,1862 era,butthespiralsculptureofhigh,wide,flat-toppedcords
Typespecies:FasciolariaelegansEmmons,1858,bymonotypy. isunique.InT.elegans,thereare21suchcordsonthelast
Lectotype:specimenfiguredbyEmmons,1858:fig.114,designated whorl,becomingsmallerandmorecloselyspacedabapi-
herein(Fig.5). cally,and7onthepenultimatewhorl.Thesubsuturalramp
oflaterwhorlsresemblesthespirallycordedrampinFascio-
Description.—Shellofmediumsizeforsubfamily,lengthof lariabutismuchbroader.Theprotrudingparietalridgeof
onespecimenexamined149.8mm,withveryhighspireand TerebraspiraisseenalsoinCinctura,whoseshellis,however,
veryshortsiphonalprocess;teleoconchwithnineweakly mostlysmooth.ThespireofTerebraspiraisexceptionally
shoulderedwhorlsseparatedbydeeplychanneledsuture; high,beingonlyslightlylessthanhalftheheightoftheaper-
38 lastwhorlwithconspicuoussubsuturalramp,separated turepluscanal.Acomparablyhighspireisseenonlyin
frommainpartofwhorlbydeepgroove;axialsculpture somespeciesofthenewgeneraAfricolariaandAustralaria
presentonlyonfirsttwoteleoconchwhorls;otherwhorls (seebelow).Thehigh,roundedsiphonalfascioleisunusual
withhigh,wide,flat-toppedspiralcordsseparatedbydeep forthesubfamily;itissharedonlywithPliculofusus.The
interspaces;aperturebroad,outerlipsomewhatbrokenbut adapicallyextendedapertureintoachannel-likefeatureis
probablyplanar,itsedgewithpairedcrenulations,itsinner exceptionallywellmarkedinTerebraspiraanddiffersfrom
sidewiththickspiralribsrepresentinginterspacesbetween theconditionseeninothermembersofthesubfamily.Ex-
externalcords;internalliraeabsent;adapicalendofaperture plicitcomparisonsbetweenTerebraspiraandthegenera
extendedaschannel;entrancefoldtosiphonalcanal PliculofususandTriplofususaregivenunderPliculofusus.
rounded,moreprominentthantwocolumellarfolds;four Emmons’originalillustrationservestoidentifyT.elegans
veryweakfoldsabapicaltoentrancefoldalongadaxialside andweselectthespecimendepictedinthatfigureaslecto-
ofsiphonalcanal;parietalridgethinbutdistinct,extending type.L.D.Campbellinformsus(e-mailtoGJV,3May,2011)
outofapertureasathickened,polishedribnotcoincident thatEmmons’specimenislost.Oneofthefewknownspeci-
withspiralsculpture;innerlippolished,extendingasglaze mensofT.elegansisintheUSNMcollection,no.429895(Fig.
overpartofventralsideoflastwhorl;siphonalfasciole 6).AccordingtoCampbell,L.W.Wardcollectedthatspeci-
prominent,rounded,withfivespiralthreads. menintheLumberRiversectionatLumberton,NorthCar-
olina,inadepositthatCampbellconsidersastratigraphic
Includedspecies: equivalentoftheRaysorFormationofEarlyPlioceneage
FasciolariaelegansEmmons,1858:EarlyPliocene(RaysorFormation), (3.8Ma).Wehaveexaminedandillustratethisspecimen,
NorthCarolina. andourdescriptionofTerebraspiraisbaseduponit.
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Pliculofususgen.nov. FasciolariascalarinamacgintyiM.Smith,1936:Pleistocene,Florida;
Typespecies:FasciolariascalarinaHeilprin,1886,designatedherein. TerebraspiraseminolePetuch,1994:Pliocene,Florida;
Holotype:WFIS904,158.6mmSL,bysubsequentdesignation,Spamer FasciolariasparrowiEmmons,1858:EarlyPliocene,NorthCarolina.
&Forster(1988:56)(Fig.7).
Etymology.—ThenameisLatin,formedofplicula(small
Description.—Shellmedium-sizedtolarge,to190mmSL fold)andfusus(spindle),referringtothepresenceofsmall
(P.scalarinus),high-spired;suturedeeplyimpressedbutnot foldsontheadaxialsideofthesiphonalcanal;thegenderis
channeled;teleoconchsculptureofbroad,lowaxialribs masculine.
withroundedcross-section;spiralsculptureofhigh,
roundedorΛ-shapedcordsoverentireshell;aperturenar- Remarks.—MostspeciesinPliculofususgen.nov.wereas-
rowtobroad,outerlipmediallyconvex,itsedgewith signedbylatetwentieth-centuryauthorstoTerebraspira(see
pairedcrenulations,itsinnersidewithbeadedordiscontin- e.g.Campbell,1993;Petuch,1994).AsnotedunderTere-
uouslirae,especiallyonanterior(abapical)sector;adapical braspira,however,allthesespeciesdifferinsignificantways
endofouterlipnotformingaperturalextension;entrance fromthetypeandonlydescribedspeciesofTerebraspira,T.
foldtosiphonalcanalrounded,moreprominentthancol- elegans(Emmons,1858).Thiscircumstancenecessitatesthe
umellarfolds;fourdistinctfoldsabapicaltoentrancefoldon proposalofthenewgenusPliculofusus.
adaxialsideofsiphonalcanal;parietalridgethin,distinct, SpeciesofPliculofususgen.nov.aremostsimilarto
notextendingbeyondedgeofaperture;innerlipcallus speciesofTerebraspiraandTriplofusus,butatleasteightfea-
spreadingontoventralsideoflastwhorl,recessedwhereit turesdistinguishthegenera:(1)Pliculofusushasaglazed
bordersexternalsculptureoflastwhorl;siphonalprocess innerlipthatextendsontotheventralsideofthelastwhorl,
shorttomoderatelylong,bearingdistinctroundedsiphonal asinTriplofususandTerebraspira,butincontrasttothosegen-
fasciole. eraitsedgeisrecessedwithrespecttotheexternalsculp- 39
ture,implyingresorption;(2)thereisathinbutdistinct
Includedspecies: parietalridgeinPliculofusus,asinTerebraspira,butthisridge
FasciolariaacutaEmmons,1858:Pliocene(DuplinandTamiamiForma- isnotextendedbeyondtheapertureasitisinTerebraspira;
tions),VirginiatoFlorida; Triplofususlackstheridge;(3)asinTerebraspira,Pliculofusus
TerebraspiracalusaPetuch,1994:Pliocene,Florida; hassmallfoldsontheadaxialsideofthesiphonalcanal,al-
TerebraspiradiegelaePetuch,1994:Pliocene,Florida; thoughtheyarestrongerthaninTerebraspira;thesefoldsare
TerebraspirakissimmeensisPetuch,1994:Pliocene,Florida; absentinTriplofusus;(4)thereisaprominent,rounded
TerebraspiralabelleensisPetuch,1994:Pleistocene,Florida; siphonalfascioleinPliculofususandTerebraspira,butthis
TerebraspiralindaePetuch,1994:Pliocene,Florida;
Terebraspiramaryae,Petuch,1994:Pliocene,Florida; (nextpage)Figs1-6.Fasciolariinaespecies.1,LectotypeofMurextulipa
FasciolarianodulosaEmmons,1858:Pliocene(Duplin,RaysorandTami- Linnaeus,1758,typespeciesofFasciolaria;figureafterd’Argenville,
amiFormations),NorthCarolinatoFlorida; 1742:pl.13fig.K;2,OriginalillustrationofPyrulahunteriaPerry,1811,
TerebraspiraokeechobeensisPetuch,1994:Pliocene,Florida; typespeciesofCinctura;figureafterPerry,1811:pl.50fig.4;3,Neotype
TerebraspiraosceolaiPetuch,1994:Pliocene,Florida; ofPyrulahunteriaPerry,1811,typespeciesofCinctura;USNM615769,
FasciolariapetrosaDall,1915:LateOligoceneorEarlyMiocene(Tampa 86.5mmSL;4,HolotypeofMazzalinabulbosaHeilprin,1886,type
Formation),Florida; speciesofLiochlamys;WFIS909,59.2mmSL;5,LectotypeofFasciolaria
FasciolariaramondiMaury,1902:EarlyMiocene(ChipolaFormation), elegansEmmons,1858,typespeciesofTerebraspira;figureafterEmmons,
Florida; 1858:fig.114;6,Terebraspiraelegans(Emmons,1858),USNM429895,
FasciolariascalarinaHeilprin,1886:EarlyPleistocene(Caloosahatchee 149.8mmSL.
Formation),Florida;
Snyder,M.A.etal.–GeneraandbiogeographyofFasciolariinae
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