Classification of the Modulidae (Caenogastropoda, Cerithioidea), with new genera and new fossil species from the Neogene of tropical America and Indonesia

2014·10.1 MB·English

by  Landau, Bernard

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Classification of the Modulidae (Caenogastropoda, Cerithioidea),
with new genera and new fossil species from the Neogene
of tropical America and Indonesia
Bernard Landau
NaturalisBiodiversityCenter,P.O.Box9517,NL-2300RALeiden,TheNetherlands;CentrodeGeologiadaUniversidadedeLisboa.CampoGrande,
1749-016Lisboa,PortugalandInternationalHealthCentres,Av.InfantedeHenrique7,AreiasSãoJoão,P-8200Albufeira,Portugal;
[email protected][Correspondingauthor]
Geerat J. Vermeij
DepartmentofGeology,UniversityofCaliforniaatDavis,OneShieldsAvenue,Davis,CA95616USA;[email protected]
1
Sonja Reich
NaturalisBiodiversityCenter(GeologyDepartment),P.O.Box9517,NL-2300RLLeiden,TheNetherlands.
ArevisionofthecerithioideangenusModulus,basedonshell exclusivelytropicalAmericangenera,withaMiocene-
charactersandnewfossilmaterialfromtheNeogene Pliocenefossilrecordthatdidnotsurvivetothepresentday.
CaribbeanandIndonesia,showsthegroupnottobemono- IntheIndo-Pacific,Indomodulusgen.nov.firstappearsinthe
generic.Severaldistinctspeciesgroups,withrestrictedgeo- earlyMioceneofIndonesia,afterwhichthereisnofossil
graphicaldistributions,wereidentified.ThegenusModulus recorduntilthePleistocene,andistodayrepresentedbyasin-
Gray,1842,hasageologicalhistorystartinginthelatest gle,widely-distributedIndo-Pacificspecies.Conomodulusnov.
EocenetoearliestOligoceneintropicalAmerica,extendingto gen.isrestrictedtotheMioceneofIndonesia.
thepresentdayCaribbeanandPacificPanamicfaunas,andis Sixnewspeciesweredescribed;fromtheCaribbean:Tro-
foundintheearlyandmiddleMioceneeasternAtlanticand chomodulusparaguanenesisspec.nov.,Laevimoduluscanaespec.
Mediterraneanassemblages,afterwhichthereisagapinits nov.,andLaevimodulusgurabensisspec.nov.;fromIndonesia:
historyuntilthepresentday,whenitisfoundlivingoffWest Conomodulusrenemaispec.nov.,Indomoduluspseudotectum
Africa.Trochomodulusgen.nov.isanexclusivelytropical spec.nov.,andIndomodulusmiocenicumspec.nov.
Americangenus,representedfromthelate-earlyMioceneto
thepresentdayCaribbeanandPacificPanamicfaunas. Keywords:Modulidae,genericrevision,NeogeneCaribbean,Neogene
Laevimodulusgen.nov.andPsammodulusCollins,1934,arealso Indonesia,newgenera,newspecies,palaeobiogeography.
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Introduction Americanfossilspecies,placingspecialattentiontothecol-
1 umellararea.WhilstintheRecentfaunasmodulidsarelo-
TherichanddiverseNeogeneassemblagesfromthenorth- callyfoundinlargenumbers,inthefossiltropicalAmerican
ernDominicanRepubliccollectedaspartofthePanamaPa- andIndonesianassemblagestheyareuncommon.Modulids
leontologyProjectandnowhousedintheNaturhistorisches inhabitrockyandseagrassenvironments.Rockyenviron-
MuseumBasel(Switzerland)andtheBernardLandaucol- mentsareknowntobepoorlyrepresentedinthetropical
lectionshousedintheNaturhistorischesMuseumWien AmericanNeogenerecord(Vermeij,2001a),whereassea-
(Austria)continuetoprovidespeciesasyetunknowntosci- grasscommunitiesarenot.Whymodulidsshouldbesoun-
ence.Althoughallthesespeciesrequireformaldescription, commoninfossilassemblagesisunclear.Nevertheless,this
somearemoreremarkablethanothers.Inthispaperwetake resultsinasmallnumberofspecimensfromalargenumber
theopportunitytodescribesomemodulidsquiteunlikeany ofwidelygeographicallyandstratigraphicallyseparatedde-
ofthoseknowninthefossilorRecentfaunas. posits.Thesmallnumberofspecimensfromeachassem-
Modulidsoccurworldwideintropicalwaters,butare blagemakesintraspecificvariabilitydifficulttoevaluate.
notspeciose,withonlyaboutadozenRecentspecies Whenevertheshellcharactersbetweenthefossilpopula-
known,althoughsomeofthesemaybesynonyms.Consid- tionsdifferedconsistently,theywereconsidereddistinct
eringthispaucityofspecies,thefamilyiswellrepresented species.Thismightbeconsideredbysometobearather
inthetropicalAmericanfossilandRecentfaunas,but narrowtaxonomicviewpoint,especiallywiththespecies
seemedtobeverypoorlyrepresentedintheIndo-Pacificfos- heredescribedunderthegenusTrochomodulusgen.nov.(i.e.
silrecord.Inthecourseofthepreparationofthiswork,fur- T.paraguanensisspec.nov.,T.vermeiji,T.tamanensis).How-
therrecordsforfossilmodulidsweresoughtforthe ever,inourexperience,atspecieslevel,tropicalAmerican
Indo-Pacific,andfoundwithinthecollectionsoftheNatu- Neogenefaunastendhavearapidturnoverandbehighly
2 ralisBiodiversityCentre,whichhousesasuperbcollection endemic,and“splitting”ratherthan“lumping”turnoutto
ofIndonesianfossilmolluscs. becorrectinmostcases(Landauetal.,2008).
Untilnow,authorshaveincludedallthelivingspeciesin
asingleworldwidegenus.Wearenotawareofanymolecu- Geological setting
larphylogeneticworkonthegenus.However,examination
oftheshellsof“Modulus”species,clearlyindicatesthatthere DominicanRepublic–Thegeologyandstratigraphyofthe
aredistinctlineageswithinthefamily.Inthispaperwede- DominicanRepublichasbeenextensivelystudiedaspartof
finethesemodulidlineagesbasedonshellcharacters,and thePanamaPaleontologyProject,anditsresultspublished
recordfourspeciesfromtheNeogeneofthenorthernDo- bySaundersetal.(1986),andwillnotberepeatedhere.Up-
minicanRepublic,twoofwhicharenew;andanewspecies datedstratigraphicchartsforthetropicalAmericanareacan
fromthelate-earlyMioceneCantaureFormationof befoundinLandauetal.(2012).
Venezuela,andfourspeciesfromtheMioceneofIndonesia, Indonesia–Thematerialinvestigatedinthisstudyde-
2
ofwhichthreearealsonew.Westressthatunlikeworking rivesfromseveraldifferentlocalitiesinJavaandEastKali-
withRecentshells,whenworkingwiththefossilrecorditis mantan,Indonesia.Banyungantiisasamplinglocalityin
notunusualtofindspecieswithcharactersintermediatebe- Yogcacarta,Java,Indonesia,locatedclosetothevillage
tweengenera.Thisreflectstheevolutionaryprocess,andthe Banyunganti.Thestudiedmaterialwascollectedinun-
placementofsomespecies,suchasIndomoduluspseudotectum namedearlyBurdigalian,fossiliferousbedsoverlyingthe
spec.nov.,remaintosomedegreesubjectiveandopentoal- marineJonggranganFormation(Kadar,1986).Geologyand
ternativerepresentations.Thisworkisnotintendedtobea stratigraphyofthelocalityaredescribedindetailbyReich
monographictreatmentofthefamilyatspecieslevel,butwe etal.(2014).TheJavanesematerialofConomodulusprean-
taketheopportunityofillustratingmanyofthetropical gerensisMartin,1905,wascollectedfromunnamedbedsin
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CadasngamparandCitalahab(TjiTalahab).Theageofboth
localitiesislatemiddleMiocene(Johnsonetal.,submitted).
Nofurtherinformationonthegeologyorsedimentologyof
thelocalitiesisavailable.
AnoverviewoverthegeologyofEastKalimantancanbe
foundinWilsonetal.,1999andinWilson&Evans(2002).
Thehistoricalsamplelocality114issituatedatthe
MenkrawitRiveratthenorthernMangkalihatPeninsula
(EastKalimantan,Indonesia).Theexactlocalityisunknown.
Sedimentsatthesampledlocalityweredescribedasslightly
sandy,fossiliferousgreyclaysinthetransitionbetween
TabellarandMenkrawitlayers(Beets1941).Theageismid-
dleMiocenebasedonlargebenthicforaminifers(Renema,
2007;Lunt&Allen,2004).Samplinglocality391(Mangkali- Fig.1.Aperturalfeaturesofmodulids;Conomodulusprean-
hat,EastKalimantan)isanhistoricallocalitywithlittle gerensis(Martin,1905),holotypeRGM.10764,unnamedbeds
availableinformationintheproximityoflocality114.Itisof fromthelateMioceneofCadasngampar,Java,Indonesia.
middleMioceneage,stratigraphicallyjustabovelocality114 Height11.7mm(specimentiltedposteriortoverticalaxisto
(Johnsonetal.,submitted). showsiphonalstructures;photoF.P.Wesselingh).
Furthermore,thisstudyincludesrecentlycollectedmate-
rialfromnewsamplinglocalities(TF102;TF110,sampledin
2010,andTF505,referringtothesamelocalitysampledin
2011)inBontang(EastKalimantan,Indonesia).Bontangislo- theThroughflowProjectin2010and2011.Onlymaterial 3
catedintheKutaiBasin,thelargestCenozoicsedimentary fromlocalities114and391(MangkalihatPeninsula)isfrom
basinofKalimantan(Moss&Chambers,1999).Bothlocalities acollectionmadebyLeopoldinthe1920saspartofasurvey
representearlyTortonian(lateMiocene),unnamedbedsin fortheDienstvanhetMijnwezeninNederlandsOost-Indie.
marinesiliciclasticsediments.Severalstudiesonthesampling Themolluscanfaunafromlocality114isdescribedbyBeets
localities,theirstratigraphy,andassociatedmolluscfaunas (1941).ThematerialfromJavaincludesaholotypefromthe
haverecentlybeencarriedoutbyRenemaetal.(submitted); Martincollection(Leloux&Wesselingh,2009),asinglespec-
Kusworoetal.(submitted);andReichetal.(submitted). imenfromCitalahab,andmaterialfromtheBanyunganti
faunacollectedin2006byF.P.WesselinghandW.Renema,
anddescribedbyReichetal.(2014).
Material and Methods
SuperfamilyCerithioideaFleming,1822
TheCaribbeanmaterialdescribedhereisfromthePanama FamilyModulidaeP.Fischer,1884
PaleontologyProject(PPP)collection,housedintheNaturhis-
torischesMuseumBasel(NMBcoll.),Switzerland,andthe Thegenericrevisionproposedhereinisbasedonshellchar-
BernardLandaucollection,nowdepositedintheNaturhis- acters.Weplaceparticularimportanceontheaperturalchar-
torischesMuseumWien(NHMWcoll.),Vienna,Austria. acters,especiallythoseofthesiphonalareasituated
AllIndonesianmaterialdescribedinthisstudyishoused immediatelybelowthecolumellartooth.Thesiphoncanbe
intheNaturalisBiodiversityCentre,Leiden,TheNether- markedeitherbyadepressiononthecolumellarcallusorby
lands(indicatedbyRGMnumbers).ThematerialfromEast agrooveonthemedialedgeofthecolumellarcallus,orin
Kalimantan(Indonesia)islargelyfromcollectionsmadeby somemodulidgeneraacombinationofthetwo(Fig1).
Landau, B., Vermeij, G.J., Reich, S.–ClassificationoftheModulidae
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Tropical American and eastern Atlantic nodosusMacsotay&Campos,2001.Pacific;M.cerodes(A.
modulid genera Adams,1851)(Figs12-16),M.disculus(Philippi,1846)(Figs17-
19).RecenteasternAtlantic,WestAfrica;M.guerneiDautzen-
ModulusGray,1842 ber,1900,M.turbinoides(Locard,1897).
FossiltropicalAmerican:M.liveoakensisMansfield,1937
Typespecies.—TrochusmodulusLinnaeus,1758,bysubse- (latestEocene-earliestOligocene:SuwanneeLimestone;for
quentdesignation(J.E.Gray,1847).Recent,Caribbean. ageofassemblageseeHerbert&Portell,2002);M.turbinatus
Reviseddescription.—Shellsmedium-sized,solid, (Heilprin,1887)(lateOligocene:Chattian;Silexbedsofthe
turbiniform.Spiredepressedinmostspecies.Sutureim- TampaLimestoneofFlorida);M.biconicusGardner,1947
pressedtonarrowlycanaliculated.Sculptureofbroadaxial (Figs20-22),M.compactusDall,1892,andM.imitatus
ribsorfoldsonspirewhorlsandadapicalhalfofwhorl,de- Schmelz,1991(Figs23-25)(earlyMioceneBurdigalian,
velopedintotuberclesatshoulderinmostspecies;spiral ChipolaFormationofFlorida);M.woodringiMansfield,1930,
cordsnarrow,rounded,elevated,stronglydevelopedon M.lindaePetuch,1987,M.caloosahatchiensisPetuch,1994,M.
base,wheretheyareweaklytubercularinsomespecies. bermontianusPetuch,1994(Figs9-11)(Plio/Pleistoceneof
Aperturesmall,circular;outerlipsharp,liratewithin.Col- Florida).
umellarcallushardlydeveloped,narrow,withasharp FossilEurope:M.basterotiBenoist,1874(early-middle
lamella-likecolumellartoothabapically.Siphonaldepres- Miocene).
sionnarrowtoabsent;siphonalgroove,deep.Smallumbili- Discussion.—Modulusisthemostspecies-richgenusof
calchinkpresent. itsfamilyinthelivingfauna.Houbrick(1980)interpreted
Speciesincluded.—RecenttropicalAmerican:Atlantic;M. thetaxonModulusmodulusbroadlytoincludepopulations
modulus(Linnaeus,1758)(Figs2-8),M.lindaePetuch,1987,M. withdirectdevelopment,suchastheonehestudiedonthe
kaicheraePetuch,1987,ModuluspaceiPetuch,1987,M.bermon- eastcoastofFlorida,aswellaspopulationsinwhichlarvae 5
tianusPetuch,1994(Figs9-11),ModulusbayeriPetuch,2001,M. hatchasfree-swimmingveligers,suchasthoseinColombia
andBermuda.Thespeciesiscertainlyhighlyvariablein
shapeandintheheightofthespire(Figs2-8).Contraryto
Figs2-19.Modulusspecies(allphotosB.Landau).2-8,Modulusmodulus Houbrick(1980),wesuspectthatM.moduluscomprisessev-
(Linnaeus,1758).2-4,NHMW2013/0314/0022(exBLcoll.),height13.8 eralspecies,whicharegeographicallydistinct.Forexample,
mm,LantanaRoad,PalmBeachCounty,Florida,USA,FortThompson wenotethatspecimensfromCuraçao,Aruba,andtheAt-
Formation,latePleistocene;5-6,NHMW2013/0314/0045(exBLcoll.), lanticcoastofCostaRica(Vermeijcollection)differfrom
height14.1mm,ElCastillo,LaIsabella,northernDominicanRepublic, moretypicalM.modulusinhavingobsoletenodesanda
LaIsabellaFormation,latePleistocene.7-8,NHMW2013/0314/0024(ex highspire;thesespecimenstypicallylackliraeontheinner
BLcoll.),height17.5mm,BocaChica,MargaritaIsland,NuevaEsparta sideoftheouterlip.SeveralspecieswithintheM.modulus
State,Venezuela,TortugaFormation,latePleistocene.9-11,Modulus speciesgroupwereerectedbyPetuch(1987,1994,2001)
bermontianusPetuch,1994,NHMW2013/0314/0025(exBLcoll.),height basedonsmalldifferencesofshellmorphology.Arevision
9.7mm,CapelettiBros.Pit,DadeCounty,Florida,USA,BermontFor- ofthesespeciesisbeyondthescopeofthiswork,butatleast
mation,latePleistocene.12-16,Moduluscerodes(A.Adams,1851).12-14, someofthemarelikelytobevalid.IntheeasternPacific,the
NHMW2013/0314/0026(exBLcoll.),height11.6mm.15-16,NHMW verystronglynodoseM.cerodesusuallylackslirae(Figs15-
2013/0314/0027(exBLcoll.),height10.8mm,SantaInesBaynear 16),althoughtheyarepresentinsomespecimens(Figs12-
Mulegé,BajaCaliforniaSur,Mexico,latePleistocene.17-19,Modulus 14).WhetherM.cerodesistrulydistinctfromthesympatric
disculus(Philippi,1846),“bluehouse”,BuricaPeninsula,Panama,Ar- M.disculus(Figs17-19)isdoubtful.Molecularstudieswillbe
muellesFormation,CharcoAzulGroup,latePleistocene. neededtoclarifythetaxonomyoflivingwesternAtlantic
andeasternPacificspeciesofModulus.
Landau, B., Vermeij, G.J., Reich, S.–ClassificationoftheModulidae
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Figs20-31.Modulusspecies(allphotosB.Landau).20-22,ModulusbiconicusGardner,1947,NHMW2013/0314/0028(exBLcoll.),height8.7mm,Tenmile
Creek,CalhounCounty,Florida,USA,ChipolaFormation,upperBurdigalian,earlyMiocene.23-25,ModulusimitatusSchmelz,1991,NHMW
2013/0314/0029(exBLcoll.),height16.4mm,FarleyRiver,CalhounCounty,Florida,USA,ChipolaFormation,upperBurdigalian,earlyMiocene.26-31,
Modulusbasteroti(Benoist,1873).Figs26-28:NHMW2013/0314/0030(exBLcoll.),height20.2mm,MoulindeGamachot,Uzeste.29-31,NHMW
2013/0314/0031(exBLcoll.),height12.0mm,Lariey,Saucats,Gironde,France,lowerAquitanian,earlyMiocene.
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Trochomodulusgen.nov. Modulusmodulusbasileus(Guppy).Woodring;1928(inpart):343,pl.25
fig.17,pl.26figs1-2(notfigs3-4=Modulussp.);Perrilliat,1972:46,
Typespecies.—ModuluscatenulatusPhilippi,1849,desig- pl.14figs9-14.NotModulusbasileus(Guppy,1873).Petuch,1994:
natedherein.Recent,PanamicPacific. 64,pl.11fig.E.
Description.—Shellsmedium-sized,relativelysolid,
trochiform.Spireofmediumheighttomoderatelydepressed, Materialanddimensions.—Maximumheight11.6mm,NHMW
straight-sidedtocyrtoconoidinprofile,whorlsstraight-sided 2013/0314/0013/1(exBLcoll.)(Figs45-47),NHMW2013/0314/0014/1
toconcave.Sutureimpressed.Sculptureofnarrowspiral (Figs48-51),NHMW2013/0314/0015/3,LopezSection,RíoYaquedel
cords,withasinglemorestronglydevelopedcordatsuture, Norte,BaitoaFormation,early-middleMiocene.
whichdevelopsintostrongkeel-likeperipheralcordonlast
whorl,cordsmoreprominentonbase.Aperturesmall,circu- Discussion.—Trochomodulusbasileus(Guppy,1873)issome-
lar;outerlipsharp,liratewithin.Columellarcallusweakly whatvariableinrespecttothestrengthofthespiralsculp-
thickenedandhardlyexpandedoverventer,withasharp tureandthecarinaiscoarselytubercularinsomespecimens,
lamella-likecolumellartoothabapically.Siphonaldepression asshownbythespecimensillustratedherefromtheDo-
narrowtoabsent;siphonalgroove,deep.Smallumbilical minicanRepublic(Figs45-51),thosefromthelatePliocene
chinkmaybepresentorabsent. PiacenzianAgueguexquiteFormationofMexico(Perrilliat,
Speciesincluded.—RecenttropicalAmerican:Atlantic;M. 1973,pl.14figs9-14),andthosefromthePlioceneBowden
carchedonius(Lamack,1822)(Fig.33),M.calusaPetuch,1988 FormationofJamaica(Woodring,1928,pl.25fig.17,pl.26
(Figs34-36).Pacific;ModuluscatenulatusPhilippi,1849(Fig.32). figs1-2).However,someofthespecimensfiguredby
FossiltropicalAmerican:M.willcoxiDall,1892(Figs37- Woodring(1928,pl.26figs3-4)withaxialsculpturearenot
39)(earlyMioceneBurdigalian,ChipolaFormationof thisspecies,butaspeciesofModulus.Theshellillustratedby
Florida);M.tamanensisMaury,1925(Figs40-41)(BrassoFor- Petuch(1994,pl.11fig.E)asModulusbasileusfromtheearly 7
mation,middleMioceneofTrinidad);M.vermeijiLandau& PleistoceneCaloosahatcheeFormationofFloridahasvery
Silva,2010(Figs42-44)(CaujaraoFormation,lateMiocene; weakspiralsculptureonthespirewhorlsandabovethekeel
andArayaFormation,earlyPlioceneofVenezuela),Mon- onthelastwhorl,andinouropinionismorelikelytorepre-
odontabasileaGuppy,1873(Figs45-51)(BaitoaFormation, sentTrochomoduluscalusa(Petuch,1988).
earlyMioceneoftheDominicanRepublic;BowdenForma- Trochomoduluswillcoxi(Dall,1892)fromtheearly
tion,PlioceneofJamaica;andCaloosahatcheeFormation, MioceneBurdigalianChipolaFormationofFlorida,T.tame-
PleistoceneofFlorida). nensis(Maury,1925)fromthemiddleMioceneBrassoFor-
Etymology.—Namereflectingthetrochiformshell mationofTrinidadandT.vermeiji(Landau&Silva,2010)
shapeofthisgroupofmodulids.Gendermasculine. fromthelateMioceneCaujaraoFormationandearly
Discussion.—SpeciesofTrochomodulusaredistin- PlioceneArayaFormationsofVenezuelaalldifferfromT.
guishedfromthoseofModulusmostimportantlybytheir basileusinhavingnoumbilicuspresent.
trochiformratherthanturbiniformshellshape,andbytheir Distribution.—Early-middleMiocene:BaitoaForma-
sculpture,whichconsistsofstrongspiralcords,withakeel- tion,DominicanRepublic(thispaper);Pliocene,Bowden
likeperipheryandnoaxialsculpture.Thecharacterofthe Formation,Jamaica(Woodring,1928);latePliocene:
siphonaldepressionandgrooveissimilarinbothgenera. AgueguexquiteFormation,Mexico(Perrilliat,1973).
Trochomodulusbasileus(Guppy,1873)(Figs45-51) Trochomodulussp.(Figs52-53)
MonodontabasileaGuppy,1873:85,pl.1fig.2;Guppy,1874:434,pl.1 Materialanddimensions.—NMBH20204(Figs12-13),onejuvenile
fig.2. specimenheight3.7mm,width3.9mm.
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(oppositepage) Figs52-58.Trochomodulusspecies(allphotosB.Landau).
Figs32-51.Trochomodulusspecies(allphotosB.Landauunlessindi- 52-53,Trochomodulussp.,NMBH20204,NMBlocality17814,eastcoast
cated).32,Trochomoduluscatenulatus(Philippi,1849),NMR44212, ofCayoAgua,justsouthofPuntadeNispero,BocasdelToro,Panama, 9
height16mm,Mexico,Recent(NMRcollection).33,Trochomodulus CayoAguaFormation,Zanclean,EarlyPliocene,height3.7mm.54-58,
carchedonius(Lamarck,1822),NMR44157,height14mm,SanibelIsland, Trochomodulusparaguanensisspec.nov.,holotypeNHMW
Florida,USA,Recent(NMRcollection).34-36,Trochomoduluscalusa 2013/0314/0023,CasaCantaure,ParaguanáPeninsula,FalcónState,
(Petuch,1988),NHMW2013/0314/0032(exBLcoll.),height9.9mm, Venezuela,uppershellbed,CantaureFormation(earlyMiocene;Burdi-
DaviesPit,OkeechobeeCounty,Florida,USA,CaloosahatcheeForma- galian),height15.3mm.
tion,latePleistocene.37-39,Trochomoduluswillcoxi(Dall,1892),NHMW
2013/0314/0033(exBLcoll.),height37.0mm,TenmileCreek,Calhoun
County,Florida,USA,ChipolaFormation,upperBurdigalian,early
Miocene.40-41,Trochomodulustamanensis(Maury,1925).40,Syntype
PRI1069,height17.5mm.41,SyntypePRI1070,height15.0mm,
Machapoorie,Samana,Trinidad,early-middleMiocene(Photoscour-
tesyofPRI).42-44,Trochomodulusvermeiji(Landau&Silva,2010),Holo-
typeNHMW2010/0038/0007(exBLcoll.),height15.5mm,Cañondelas
Calderas,CubaguaIsland,NuevaEspartaState,Venezuela,ArayaFor-
mation,CubaguaGroup,earlyPliocene.45-51,Trochomodulusbasileus
(Guppy,1873),LopezSection,RíoYaquedelNorte,BaitoaFormation,
early-middleMiocene.45-47,NHMW2013/0314/0013(exBLcoll.),
height11.6mm,width10.6mm.48-51,NHMW2013/0314/0014(exBL
coll.),height11.7mm,width10.7mm.
Landau, B., Vermeij, G.J., Reich, S.–ClassificationoftheModulidae
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Locality.—NMBlocality17814,eastcoastofCayoAgua, gledatperipheralcord.Basemoderatelydepressed,bearing
justsouthofPuntadeNispero,BocasdelToro,Panama. tenspiralcords,wideningtowardscentre.Umbilicusalmost
Stratum.—CayoAguaFormation,Zanclean,Early closed.Aperturesubquadrate,outerlipsharp,angledatpe-
Pliocene. ripheralcord,stronglyanddeeplyliratewithin;analcanal
Discussion.—AsinglesmallspecimenofaTrochomodulus notdeveloped;siphonalnotchdeep,narrow;siphonalde-
speciesfromtheEarlyPliocene,ZancleanCayoAguaForma- pressionnotdeveloped.Columellaexcavatedinmid-por-
tionofBocasdelToroispresentintheNMBcollections.It tion,withstrong,sharp,sub-horizontalcolumellartooth.
mayrepresentsastronglykeeledspecimenofT.basileus,but Columellarcallusweaklydeveloped.
itdoesnothavetheumbilicusasstronglydeveloped.The Discussion.—Trochomodulusparaguanensisspec.nov.is
specimenistoojuveniletodeterminewithcertainty. representedbyonlytwospecimens,butitisdistinctive
enoughtowarrantdescription.Itclearlybelongstothe
Trochomodulusparaguanensisspec.nov.(Figs54-58) genusTrochomodulusgen.nov.,characterisedbyitsrather
biconicshell,predominantlyspiralsculpture,muchreduced
Typematerialanddimensions.—HolotypeNHMW2013/0314/0023, axialsculpture,andbyitsverysharp,keel-likeperiphery.
height15.3mm,width14.7mm(Figs54-58);paratype1NMBH20231, TrochomodulusparaguanensisismostsimilartoTrochomod-
height14.0mm,width13.1mm,NMBlocality17516. ulustamanensis(Maury,1925)fromtheMiddleMiocene
Othermaterial.—Knownonlyfromtypematerial. BrassoFormationofTrinidad.SixspecimensofT.tamanensis
intheBaselcollections(NMBlot1o434[sic]fromCaparo
Typelocality.—1kmsouthwestofCasaCantaure,about10 River)differfromtheCantaurespecimeninhavingamore
kmwestofPuebloNuevo,Falcón,Venezuela(=localityGS- elevatedshellandalessangularperiphery.Moreover,the
6-PGNAofGibson-Smith&Gibson-Smith,1979). spirewhorlshavefivespiralcordsasopposedtosixinT.
10 Typestratum.—Uppershellbed,CantaureFormation paraguanensis.Trochomodulusvermeiji(Landau&Silva,2010)
(earlyMiocene;Burdigalian). fromthelateMioceneCaujaraoFormationandearly
Etymology.—Fromgeographicareainwhichitisfound MioceneArayaFormationofVenezuelaisalsosimilar,but
–theParaguanáPeninsula,FalcónState,Venezuela.Gender hasanarrowershell,withalessdepressedbase,andwider
masculine. andfewerspiralcords.Trochomodulusbasileus(Guppy,1873)
Diagnosis.—ATrochomodulusspecieswithamedium- (discussedabove)differsinthesamecharacters,ofanar-
sizedshell,withsixnarrowspiralcordsonspirewhorls,the rowershellwithfewerspiralcords,butalsohasawideum-
abapicalcordslightlymorestronglydeveloped,anarrowly bilicus,whichisalmostabsentinT.paraguanensis.Thisnew
canaliculatedsuture,axialsculptureofcollabralgrowth speciesdiffersfromtheRecenttropicalAmericanAtlanticT.
linesonearlyadultwhorls,basemoderatelydepressed, carchedonius(Lamack,1822)andPacificT.catenulatus
bearingtenspiralcords,andadeepsiphonalgroovewithno Philippi,1849inhavingamuchnarrowerandsharperpe-
siphonaldepression. ripheralkeelandspiralcords.
Description.—Shellmedium-sized,trochiform.Teleo- Distribution.—EarlyMiocene:CantaureFormation,Fal-
conchofsixweaklyconcavewhorls.Sutureimpressed,very cónState,Venezuela(thispaper).
narrowlycanaliculate.Protoconchandearlyteleoconch
whorlssomewhatabraded.Fourthtolastwhorlwithsix Laevimodulusnov.gen.
narrowspiralcords,abapicalcordslightlystrongerand
moredistantfromothers,formingperiphery.Strongly Typespecies.—Laevimoduluscanaespec.nov.,designated
prosoclinecollabralgrowthlinespresentonearlywhorls, herein.LateMiocene,DominicanRepublic.
givingthemfinelyreticulateappearance,growthlinesobso- Description.—Shellssmall,ofmedium-thickness,low-
lete,oralmostso,fromfourthwhorl.Lastwhorlacutelyan- turbiniform.Spiredepressed,whorlsconvex.Suturedeeply
Basteria78(1-3)