B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 105 Recognition of the marine gastropod Caecum tumidum Carpenter, 1858 (Caenogastropoda, Truncatelloidea) in the Pliocene of the North Sea Basin Dick F. Hoeksema Watertoren28,NL-4336KCMiddelburg,TheNetherlands;[email protected] Jonathan A. Todd DepartmentofEarthSciences,NaturalHistoryMuseum,CromwellRoad, LondonSW75BD,UnitedKingdom;[email protected] tativesoftheCaecinae. 105 In1858Carpenterdescribedthefossil“Caecumtra- InhiscatalogueofshellsfromtheCrag,Wood chea”sensuWood(1842,1848)fromthePliocene (1842:459)listedaspecimenof“Caecumtrachea?” BritishCragMolluscaasnewtoscienceandnamedit foundintheCorallineCrag(Pliocene;Sutton,Suffolk, Caecumtumidum.Alectotypeisselectedtostabilize UK).Thequestion-markindicatesthatWoodatthat thespecies.C.tumidumappearsalsopresentinthe timehaddoubtsaboutthisidentificationthatheex- PlioceneofBelgiumandTheNetherlands.AsC.tu- pressedasfollows,“TherecentBritishshellisregu- midumuntilrecentlyhasbeenmistakenforCaecum larlyannulatedandsmooth,theannuliinmysingle trachea(Montagu,1803)=Caecumimperforatum(Kan- cragspecimenaremoreirregularandrugose.” macher,1798)aswellasforCaecummammillatum InhisMonographoftheCragMollusca,Wood Wood,1848,diagnosticcharactersandphotographs (1848:115-116)omittedthequestion-mark,evidently arepresentedtodistinguishthethreespecies. consideringhisspeciestobeconspecificwiththeRe- centCaecumtrachea(Montagu,1803). Keywords:Gastropoda,Caenogastropoda,Truncatelloidea, TenyearslaterCarpenter(1858:426-427),“having Caecidae,Caecinae,Caecum,Pliocene,NorthSeaBasin,Belgium, subjectedallthespecimensintheBritishMuseum,as TheNetherlands,UnitedKingdom. wellassomesentmebyMr.Wood,toaveryrigidex- aminationunderthemicroscope”,recognizedandde- scribed“C.trachea”sensuWoodasnewtoscience, Introduction namingitCaecumtumidum. Jeffreys(1867:76),discussingC.trachea,wrote,“I ThefamilyCaecidaeconsistsofagroupofinterstitial donotconsideritaBritishfossil;forIbelievethe benthicgastropods,inwhichtheshell,afterthelarval shellsdescribedandfiguredbySearlesWoodfrom stagewhichisregularlycoiled,isformedbyalong, theCorallineCragarenotthisspecies,buthisC.mam- slightlycurvedtube.Duringthegrowth,earlystages millatum.”Jeffreysdidnotrefertothedescriptionby maybeeitherpreserved(subfamiliesStrebloceratinae Carpenter. andPedumicrinae)ordiscardedandtheposteriorend Wood(1872:87)didnotrespondtoJeffreys.He sealedbyaseptum(subfamilyCaecinae)(Pizziniet positivelyrespondedtoCarpenterbyinsertinginhis al.,2013:1).Inthispaperweonlydealwithrepresen- SupplementtotheMonographoftheCragMolluscaa Basteria79(4-6):105-113(2015) B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 106 surveyof“theCragCaecidaeaccordingtoCarpenter’s Material and methods views(thoughIdonotfullyagreewithhim)”,together withafootnoteinwhichWoodremindedreadersofhis Diagnosticcharacters.–Nofronietal.(1997:5)sum- “noteofinterrogationagainstC.trachea”inhiscata- marizedfourvalidlydiagnosticcharactersthatmay logue(Wood,1842:459;seeabove).Despitethisfoot- distinguishCaecumspecies:1.shellsize,2.shapeof note,twoyearslaterin1874,Woodlisted“C.trachea”in septum,3.presence/absenceandtypeof theSynopticalList(page209)andIndex(page224)of (micro-)sculptureand4.presence/absenceofan hisSupplement,merelymentioning“C.tumidum aperturalvarix.Theyadd,“Itisstressedthatthemor- Carp.”inthecolumnof‘Remarks’(page209). phologicalcharactershereemployed(especiallythe Harmer(1923:848)repeatedJeffreys’opinionon septumandthemicrosculpture)arevalidlydiagnostic thefossil“C.trachea”,butfollowedWood:“Jeffreys whenobservedinfull-grown,wellpreservedspeci- consideredthattheCorallineCragfossildescribedby mens;therangeofvariabilityofmanycharactersin Woodunderthepresentnamecouldnotbeidentified someinstance,forcestoanalyseasmanycharactersas withtherecentC.tracheaofMontagu,inwhichthe possibleineachcase,andaslargesamplesaspossible concentricribsaremoredistinctandcloselyarranged, toavoidmisinterpretations.”Forexamplethemi- preferringtoregarditasavarietyofWood’sC.mam- crosculptureshouldnotbehandledastheonlydiag- millatum.Onthewhole,however,…IadoptWood’s nosticfeature,“Oftenspeciesthatareusuallystriate, view,whichistheonemostgenerallyaccepted.” presentspecimenswithveryweak(orevennearlyab- HarmerdidnotevenmentionCarpenter’sdescription sent)microsculpture(andviceversa).” ofC.tumidum. Terminology.–BasedonFretter&Graham(1978: Since1874thetaxonCaecumtumidumCarpenter, 234),Lightfoot(1992:172-173)andNofronietal.(1997: 1858seemstohavebeenforgotten.Unjustly,aswe 4-5):apicalend–thesmaller,narrower,closedupper willshow. endofthetube;septum–closureoftheshellafterthe InpreparationforPartIIoftheatlasDefossiele previousstageisdiscarded;mucro–appendageoften schelpenvandeNederlandsekustHoeksema&Raad visibleontheseptum;aperture–theroundopeningat (2015)studiedthefamilyCaecidae.Thankstothesup- thelowerendofthetube;aperturalend–thelower portofthemembersoftheWerkgroepGeologieofthe endofthetube;varix–transversethickeningorring 106 KoninklijkZeeuwschGenootschapderWetenschap- precedingtheedgeoftheaperture,whichalwayssig- pen(Middelburg,TheNetherlands)adozensamples nifiesanadultshellandtheendofgrowth;dorsalside offossilCaecumspecimenscollectedfromthebeaches –convexsideofthetube(dorsalaspectofseptumis oftheWesterschelde,provinceofZeeland,The “twelveo’clock”);ventralside–concavesideofthe Netherlands,weremadeavailableforstudy.Thesam- tube;ribs–longitudinal,raisedsculpture(equivalent plescontainundisputedspecimensofCaecumglabrum tothespiralsculptureofthenormallycoiledgas- (Montagu,1803)andC.mammillatumWood,1848. tropods);rings–transverse,raisedsculpture(equiva- ShellsofathirdCaecumspeciesatfirstsightresem- lenttotheaxialsculptureofthenormallycoiled bledC.trachea,butacloserexaminationshowedcon- gastropods);microsculpture–sculpturevisibleathigh sistentdifferences.Consequentlythefirstauthor magnification. studiedthefossilCaecumshellsintheRoyalBelgian Systematics.–Thecurrentsystematicsisbasedon InstituteforNaturalSciences(Brussels,Belgium)and theWorldRegisterofMarineSpecies(Bouchet& thosefromDutchboreholesdepositedintheNaturalis Gofas,2015),consultedinJuly2015. BiodiversityCenter(Leiden,TheNetherlands).TheC. Abbreviationsandacronyms.–AR:coll.A.C. trachea-likespeciesispresentinbothmuseums,all (Riaan)Rijken,Middelburg,TheNetherlands;DH: specimenswereobtainedfromPliocenedeposits, coll.D.F.(Dick)Hoeksema,Middelburg,TheNether- whichindicatesthatthebeach-collectedspecimens lands;coll.:collection;fig(s):figure(s);Fig(s):figure(s) alsocanbeconsideredtobeofPlioceneorigin.Study- inthepresentpaper;FN:coll.F.A.D.(Freddy)van ingthepublicationsontheBritishPlioceneCragMol- Nieulande,Nieuw-enSt.Joosland,TheNetherlands; luscamentionedabove,Carpenter’sdescriptionofC. frm(s):fragment(s);GS:coll.G.F.(George)Simons, tumidumappearedstrikinglyapplicabletothisspecies. Middelburg,TheNetherlands;HJ:coll.H.A.A.(Bart) Thesecondauthortrackeddownthespecimensof“C. deJong,Middelburg,TheNetherlands;HR:coll.H.J. trachea”sensuWoodintheNaturalHistoryMuseum (Harry)Raad,Kruiningen,TheNetherlands;LK:coll. (London,UK),thesyntypesonwhichCarpenter(1858: L.(Lex)Kattenwinkel,Goes,TheNetherlands; 426-427)hadbasedhisdescriptionofC.tumidum,and, NHMUK:NaturalHistoryMuseum(formerlyBritish asexpected,theyprovedtobeconspecificwiththe MuseumofNaturalHistory),London,UnitedKing- mentionedfossilC.trachea-likespecimensfromBel- dom;NMR:NatuurhistorischMuseumRotterdam, giumandTheNetherlands. Rotterdam,TheNetherlands;pl(s):plate(s);PM:coll. Basteria79(4-6),2015 B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 107 P.W.(Peter)Moerdijk,Middelburg,TheNetherlands; Pouwer,2013:9-10,fig.1),1spmexcoll.AR, RBINS:RoyalBelgianInstituteforNaturalSciences, RGM.1008223(Fig.7a);2spms,1frmLK;Borssele, Brussels,Belgium;RGM:FossilMolluscacoll.Natu- DeKaloot(provinceofZeeland,TheNetherlands), ralisBiodiversityCenter(formerlyRijksmuseumvan beach,2spms(Figs4-6),2frmsFN;4spms,5frms GeologieenMineralogie),Leiden,TheNetherlands; HJ;2spms,1frmHR;1frmPM;Nieuwvliet,Zwarte spm(s):specimen(s);WoRMS:WorldRegisterofMa- Polder(provinceofZeeland,TheNetherlands), rineSpecies(Bouchet&Gofas,2015). beach,1frmHR;Domburg(provinceofZeeland,The Typematerial.–IntheNaturalHistoryMuseum Netherlands),beach,1spm,RGM,excoll.J.vanDal- (London,UK),notypespecimensselectedbyCarpen- sum(infortunatelylost). terhimselfcouldbefound.Theonlysampleof“C. Specimenslabeledas“C.imperforatum”inRGM trachea”fromtheCorallineCragFormationislot fromPliocenedepositsinboreholesinTheNether- NHMUKPIG2066,donatedtothemuseumbyWood landsallappearedtobelongtoC.tumidum(April (Fig.1).Thewholelotcomprises19specimensad- 2015). heredtoathinblackpaperstripgluedontoathicker InadditiontothespecimensofC.tumidumthe whitepieceofcard.Originallyatleastanotherfour firstauthorstudiedforcomparisonasampleof specimenswerepresent,butatsometimethesehave beachedspecimensoffreshdeadC.trachea(excoll. beenremovedandthepresentwhereaboutsofthese G.J.Geuze,DH;Figs11-14)fromMartinhal,S.Portu- shellsareunknown.Onthecardonecanreadin gal,andsamplesofC.mammillatumwashedashoreat Wood’shandwritinginpencil“Cor.Crag/Caecum thebeachesofBaarland(seeabove;AR,LK;Figs15- trachea?/Sutton”;acrossandgreenstickerindicate 17),Borssele/DeKaloot(FN,HJ,HR),Ritthem(GS), thespecimenthathasbeenfigured(Wood,1848:pl. Vlissingen(HR),Nieuwvliet/ZwartePolder(HJ)and 20fig.5),whichisconfirmedbyacrossonthemu- DeBanjaard(HJ),provinceofZeeland,TheNether- seumlabel(Fig.2).AsCarpenter(1858:427)basedhis lands. descriptionofC.tumidumon“allthespecimensinthe BritishMuseum,aswellassomesentmebyMr. Systematics Wood”andasall19specimensseemconspecific,lot NHMUKPIG2066canbeconsideredatypeseriesof ClassGastropodaCuvier,1795 syntypes.Thespecimenwhichhasbeenfigured SubclassCaenogastropodaCox,1960 107 (Wood,1848:pl.20fig.5)andtowhichCarpenter OrderLittorinimorphaGolikov&Starobogatov,1975 (1858:427)referredinhisoriginaldescription,ishere SuperfamilyTruncatelloideaGray,1840 selectedaslectotypeandrenumberedNHMUKPI FamilyCaecidaeGray,1850 G2066(1)(Fig.3),makingtheother18specimens SubfamilyCaecinaeGray,1850 paralectotypes.Thelectotypeis2.79mmlongand amongthelargestofthe19typespecimens. CaecumFleming,1813 Othermaterialexamined.–BergenopZoom (provinceofNoord-Brabant,TheNetherlands),bore- CaecumtumidumCarpenter,1858(Figs3-10) hole2,RvD9283,90-91mbelowsurface,Oosterhout Formation–mollusczoneMolD1,Pliocene(Piacenz- Caecumtrachea?-Wood,1842:459(question-markbyWood). ian),1spm,RGM.937740;Dongen(provinceof NotC.tracheaMontagu,1803. Noord-Brabant,TheNetherlands),borehole Caecumtrachea–Wood,1848:115-116,pl.20fig.5;1872:87; B44G.0043,175-176mbelowsurface,OosterhoutFor- 1874:209,224.Harmer,1923:847-848,pl.64fig.32.Trausel mation–mollusczoneMolD1,Pliocene(Piacenzian), &Slieker,2015:NMR26235.NotC.tracheaMontagu,1803. 1spm,RGM.937737.a(Figs7b,8-10);Kallo(province Caecum(Anellum)tumidum–Carpenter,1858:426-427. ofOost-Vlaanderen,Belgium),Verrebroekdok,Lillo Caecummammillatum–Jeffreys,1867:76.Marquet,1997:18,28- Formation,OorderenMember,Atrinalevel,Pliocene 29fig.3;1998:70fig. (Piacenzian),1spm,RBINSno.IRScNBIST5897 Caecumimperforatum-VanRegterenAltena,Bloklander&Poud- (Marquet,1997:18,28-29;Marquet,1998:70fig.;asC. eroyen,1955:29,pl.6fig.58;1965:17,pl.6fig.58. mammillatum);Kallo,Deurganckdok,LilloFormation, LuchtbalorOorderenMember,Pliocene(LateZan- Originaldescription.–Caecum(Anellum)tumidum cleantoPiacenzian),4spms,LK;Baarland(province Carpenter,1858. ofZeeland,TheNetherlands),beachnearCamping Testatereti,sublaeviseuannulisobsoletisornata;aper- Scheldeoord,insanddredgedfromtheWester- turahaudcontracta,annulotumidocincta;septomamil- scheldeatthe‘OverloopvanHansweert’and/orthe lato,plusminusvetumente;marginelateraliconvexo, ‘DrempelvanHansweert’intheWesterscheldeestu- dorsumversussubangulato;operculum?-. ary,derivedfromPliocenedeposits(Moerdijk& Long.0.11,lat.0.018-0.024. Hoeksema, D.F. & Todd, J.A.–CaecumtumiduminthePlioceneoftheNorthSeaBasin B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 108 108 Figs1-10.Caecumtumidum,Carpenter,1858.1,typeseriesfromtheCorallineCragFormation,Pliocene(Zanclean),Sutton,Suffolk,UK, NHMUKPIG2066;2,museumlabelofNHMUK,withacrossindicatingthatthecrossonthecardofthetypeseriesindentifiesthefig- uredspecimen;3,lectotype,length2.79mm,NHMUKPIG2066(1)=thespmpicturedbyWood(1848:pl20fig5);4-6,2spms,length2.7 and2.9mm,washedashoreatBorssele,DeKaloot,FN;4,scale500micrometer;5,aperturalendFig4a,showingavarix,scale100mi- crometer;6,detaillowerparttubeFig4a,showingmicroscopicpitting,scale50micrometer;7-10,2spms;7,scale500micrometer;7a,1 spm,length2.5mm,Baarlandbeach,provinceofZeeland,TheNetherlands,Pliocene,excoll.AR,RGM.1008223;7b,1spm,length2.8 mm,Dongen(provinceofNoord-Brabant,TheNetherlands),boreholeB44G.0043,175-176mbelowsurface,OosterhoutFormation– mollusczoneMolD1,Pliocene(Piacenzian),RGM.937737.a;8,aperturalendFig7b,scale100micrometer;9,apicalend,scale100mi- crometer;10,detaillowerparttubeFig7b,showingmicroscopicpitting,scale50micrometer. Basteria79(4-6),2015 B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 109 Hab.Sutton,Cor.Crag.Mus.Brit.&c. oftenshowsathickeningoranelevatedsmoothring =C.trachea,SearlesWood,Crag.Moll.p.115,pl.20,f. precedingtheedgeoftheaperture(Figs4-5,7-8;com- 5(?pars),nonMont.etauct. pareVanRegterenAltenaetal.,1955&1965:pl.6fig. [Shellslender,round,almostsmoothorobsoletely 58,asC.imperforatum).Dimensionsofthelongest annulated;aperturenotconstricted,withswollen specimenmeasured(Fig.4b):length2.93mm,diame- ring;septummamillate,moreorlessswollen;lateral ter0.49mm(apicalend)-0.62mm(aperture);meas- sides(i.e.hacspeciedorsali(Carpenter,1858:420)=ob- uredlikeL,dandDinfig.2ofPortaetal.(1993:3). servedfromthedorsalside)convex,dorsalsidein- Locustypicus.–Sutton,Suffolk,UnitedKingdom clinedtobeingsomewhatangular;operculum Stratumtypicum.–RamsholtorSudbourneMem- unknown.Length2.79mm,diameter0.46-0.61mm]. ber,CorallineCragFormation,Pliocene(LateZan- Remarks:Carpenter’smeasurementsareininches, clean). 1.0inch=25.4mm.AccordingtoSclater(1893:440), Distribution.–C.tumidumiswidelydistributedin pages413-432ofCarpenter’spaperwerepublishedin thePlioceneoftheNorthSeaBasin,geographically 1858,andpages433-448in1859.AsC.tumidumwas fromUnitedKingdom(Sutton,Suffolk)toBelgium describedonpages426-427,theyearofpublicationof (provinceofOost-Vlaanderen)andTheNetherlands thisspeciesis1858. (provincesofZeelandandNoord-Brabant)andstrati- Diagnosis.–AfossilCaecumwithanentirelyor graphicallyfromLateZanclean(CorallineCragFor- partiallytransverselyrugosetube;theseptumele- mation,UnitedKingdomandpossiblyLuchtbal vated,hemispherical,obliquelytruncatedwithakeel Member,Belgium)toPiacenzian(OorderenMember, andtheaperturalendoftenwithavarix.Sometimes BelgiumandOosterhoutFormation–mollusczone microscopicpits,largestbetweentherugae.Length: MolD1,TheNetherlands). 2.9mm. Reviseddescription.–Theadultshellisaminute, Discussion brittle,regularlycurvedtube.Thesurfaceoftheshell usuallybears,entirelyorpartially,irregular,rugose, InthestudyofC.tumidumthreeothertaxaplayan ring-shaped,transversethickenings(Figs3-10).The importantrole,viz.C.imperforatum(Kanmacher, tubeandseptumshowanirregularpatternofmicro- 1798),C.trachea(Montagu,1803)andC.mammillatum scopicpits,largestbetweentherugae(Marquet,1997: Wood,1848.AccordingtoWoRMS(consultedinJuly 109 18,28-29fig.3,1998:70fig.;Figs5-6,8-10).Theapical 2015)C.imperforatumhastobeconsideredadubious endissealedbyanelevated,moreorlesshemispheri- synonymofC.trachea. calseptum,oftensomewhatobliquelytruncatedat Caecumtrachea(Figs11-14)occursfromthe theventralsideandcylindricalwithakeelatthedor- Mioceneuntilthepresentdayandisstillacon- salside(thekeelisusuallypositionedat12o’clockto stituentoftheRecentEuropeanandNWAfrican 3o’clock,sometimesalsoweaklypresentaroundthe fauna.Thetubeisentirelyorpartiallycoveredwith restoftheseptum;Figs4,7,9).Theaperturalend rings,theapicalendwithasomewhatsunkenseptum C.tumidum C.trachea C.mammillatum Figs3-10 Figs11-14 Figs15-17 entirelyorpartiallywithtrans- entirely,sometimespartially, entirelysmoothorwithsomefainttrans- macrosculpturetube verselyrugose,ring-shaped coveredwithrings versering-shapedthickenings thickenings microsculpturetube irregularpits oftenlongitudinalstriae oftenlongitudinalstriae elevated,hemispherical, somewhatsunk,witharounded sunk,withanarrowpapilliform septum obliquelytruncated,withakeel mucro roundedmucro sometimeswithathickeningoranele- oftenwithathickeningoranele- sometimeswithathickening vatedsmoothringprecedingtheedgeof aperturalend vatedsmoothringprecedingthe precedingtheedgeoftheaper- theaperture,and/oranarrowengraved edgeoftheaperture ture ringatsomedistancefromtheaperture maximumlength 2.9mm 3.8mm 3.4mm Table1.C.tumidumshowsconsistentdifferencesfromC.tracheaandC.mammillatum. Hoeksema, D.F. & Todd, J.A.–CaecumtumiduminthePlioceneoftheNorthSeaBasin B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 110 110 Figs11-14.Caecumtrachea(Montagu,1803),1spm,length2.8mm,Martinhalbeach,Algarve,S.Portugal,Recent,excoll.G.J.Geuze,DH; 11,scale500micrometer;12,aperturalend,scale100micrometer;13,apicalend,scale100micrometer;14,detailtube,showingmicro- scopiclongitudinalstriae,scale50micrometer. withaprotruding,eccentric(12-1o’clock),rounded fromPliocenedepositsintheNorthSeaBasin.The mucro,theaperturalendsometimeswithathickening tubehasasmoothsurface,sometimeswithsomefaint precedingtheedgeoftheaperture.Oftenami- rings,theapicalendwithasunkenseptumwithanar- crosculptureoflongitudinalstriae.Length:3.8mm. row,(oftenvery)protruding,eccentric(12-1o’clock), (VanAartsen,1977:15figs22-23,17;Chirli,2006:66- papilliform,roundedmucro,theaperturalendsome- 68,pl.28figs7-10;Chirli&Linse,2011:89,pl.26fig. timeswithathickeningoranelevatedsmoothring 3a;Fretter&Graham,1978:233-234;Jeffreys,1867:75- precedingtheedgeoftheaperture,and/oranarrow 77;Landauetal.,2004:pl.14fig.1,61-62;VanderLin- engravedringatsomedistancefromtheaperture den&Moolenbeek,2000:89-91figs27-28;Panetta, (compareVanRegterenAltenaetal.,1955&1965:pl.6 1980:288-289,294-295figs4-6;Portaetal.,1993:2-5 fig.56).Oftenamicrosculptureoflongitudinalstriae. figs1-9). Length:3.4mm.(Wood,1848:116,pl.20fig.4; Caecummammillatum(Figs15-17)isafossilCaecum Harmer,1923:847,pl.64fig.31;VanRegterenAltena Basteria79(4-6),2015 B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 111 Figs15-17.CaecummammillatumWood,1848,1spm,length3.4mm,showingmicroscopiclongitudinalstriae,Baarlandbeach,province 111 ofZeeland,TheNetherlands,Pliocene,excoll.AR,RGM.1008222;15,scale500micrometer;16,aperturalend,showingavarix,scale100 micrometer;17,apicalend,scale100micrometer. etal.,1955:29,1965:17,pl.6fig.56;Hoeksema& C.trachea.Mostlikelyalsofig.9ofHoeksema&Raad Raad,2015:28fig.10).Theseptumlookssomewhat (2015:28;asC.trachea)canbereferredtosuchaspeci- likethatoftheRecentCaecumvitreumCarpenter,1859 menofC.mammillatum(herefiguredinFig.18).So (Nofronietal.,1997:9fig.16). Wood’sfig.5amayrepresentC.tracheaorperhaps ThemicrosculptureonthetubeofaCaecumshell, C.mammillatum,butnotC.tumidum. ifpresent,isusuallyonlyvisibleonwellpreserved StudyingsurveysoftheEuropeanandNorth specimens.ItcanbeobservedonC.tumidum,C.tra- AfricanCaecidae,neitherthepresentdayfauna(Van cheaandC.mammillatumusingobliquelightingatmi- Aartsen,1977;Panetta,1980;Nofronietal.,1997;Van croscopemagnificationsgreaterthanrespectively derLinden&Moolenbeek,2000)norotherfossilfau- 40x,40xand10x. nas(Portaetal.,1993;Landauetal.,2004;Chirli,2006; ConsideringthespecimensofC.tumidumnow Chirli&Linse,2011)containaCaecumspeciesmatch- presentinthetypeseries,thedescriptionof“C.tra- ingC.tumidum.Theseptumofthepresentspecies, chea”sensuWood(1848:115-116,pl.20fig.5)isnot moreorlesshemisphericalwithakeel,mostclosely accurateinallrespects.Theobservations“length1/8 resemblesthatofCaecumarmoricumDeFolin,1869, ofaninch”(=about3.18mm)and“theclosedtermi- butthetubeofthatRecentspeciesissmooth,witha nationpreciselyresemblingtherecentspeciesin delicatelongitudinalmicrosculptureandwithoutan beingobliquelytruncatedandobtuselypointed”are aperturalvarix(Hoeksema&Segers,1993:81,82figs notcorrect.Thefigs5(withlongitudinalstriae,clear- 1A-E2A-D,85,87fig.7A-B).Theirregular,rugose, estvisiblebetweentherings)and5a(aseptumwith ring-liketransversethickeningsarereminiscentof roundedmucro)arereminiscentofC.trachea.In CaecumgougerotiMoroni&Ruggiere,1985,butthe Carpenter’s(1858:427;1859:435)opiniontheseptum transversethickeningsonthesurfaceofthisMiocene- offig.5aprobablyfitswithintherangeofvariationof Pliocenespeciesaremainlyconcentratedontheapi- C.mammillatum,someformslikefig.5aapproaching calendofthetubeandtheseptumofC.gougeroti Hoeksema, D.F. & Todd, J.A.–CaecumtumiduminthePlioceneoftheNorthSeaBasin B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 112 den,TheNetherlands)andAuke-FlorianHiemstrafor makingFigs4-17.WethankTomMeijerandRonald Pouwerforinformationonthepalaeontologicalageof theRGMsamples.PeterMoerdijkisacknowledgedfor hissuggestionstoimprovethetranslationoftheorigi- naldescription.FinallywethankArieJanssenand BernardLandauforcriticallyreviewingthemanu- script. References Fig18.Caecumcf.mam- millatumWood,1848.1 Aartsen,J.J.VAN,1977.RevisionoftheEastAtlanticand wornspm,showingaC. MediterraneanCaecidae.—Basteria41(1-4):7-19. trachea-likeseptum, Bouchet,P.&Gofas,S.,editors,2015.WorldRegisterofMarine length2.7mm,Domburg Species(WoRMS),Mollusca.–www.marinespecies.orgon beach,provinceofZee- 2015-07-31. land,TheNetherlands, Bruyne,R.H.de,Leeuwen,S.J.van,GmeligMeyling,A.W.& Pliocene,excoll.M. Daan,R.,(red.),2013.SchelpdierenvanhetNederlandseNo- Donze,RGM.794216. ordzeegebied.Ecologischeatlasvandemarieneweekdieren (Mollusca).UitgeverijTirion,UtrechtenStichtingAnemoon, Lisse. showsatriangular,roundedmucro(Chirli,2006:63- Carpenter,P.P.,1858-1859.FirstStepstowardsaMonographof 64,pl.27figs5-8). theCaecidae,aFamilyofRostriferousGasteropoda.–Pro- Caecumtumidumseems,likeC.mammillatum,tobe ceedingsoftheZoologicalSocietyofLondon26:413-432 restrictedtothePlioceneoftheNorthSeaBasin,the (1858),433-444(1859). latterbeingthecommonestspeciespresent.C.glabrum Chirli,C.,2006.MalacofaunaPliocenicaToscana,Vol.5 isrelativelyrareinthePlioceneoftheNorthBasin CaenogastropodaCox,1959Part1:1-144,pls1-46.Firenze. 112 (Wood,1848:117;Marquet,1997:18-19,1998:71),but Chirli,C.&Linse,U.,2011.ThePleistoceneMarineGastropods remainsaconstituentoftheNEAtlanticandNorth ofRhodesIsland(Greece):1-447.Firenze. Seafauna(Fretter&Graham,1978:234-235;De Fretter,V.&Graham,A.,1978.TheProsobranchMolluscsof Bruyneetal.,2013:205).AfourthPlioceneCaecum- BritainandDenmarkPart4MarineRissoacea.–TheJournal speciesfromtheNorthSeaBasin,CaecumliratumCar- ofMolluscanStudies,Supplement6:153-241. penter,1858,isonlyknownfromtheCorallineCrag Harmer,F.W.,1923.ThePlioceneMolluscaofGreatBritain FormationofSuttonbytheholotypeintheNatural beingsupplementarytoS.V.Wood’sMonographoftheCrag HistoryMuseum(London,UK)andwhichrequires Mollusca,Vol.2,Part3.-MonographofthePalaeontograph- re-study(Carpenter,1858:421).Incontradictiontothe icalSocietyLondon75(352):705-856,pls57-64.. distributiongivenbyChirli(2006:67-68)andChirli& Hoeksema,D.F.&Raad,H.J.,2015.Defossieleschelpenvande Linse(2011:89),weconcludethatthepresenceoffos- NederlandsekustII,deel10.Hydrobiidae,Cochliopidae, silC.tracheaintheNorthSeaBasinstilllacksconvinc- Iravadiidae,TruncatellidaeenCaecidae.–Spirula402:24-29. ingevidence. Hoeksema,D.F.&Segers,W.,1993.Onthesystematicsanddis- tributionofthemarinegastropodCaecumarmoricumDe Acknowledgements Folin,1869(Prosobranchia,Caecidae).–GloriaMaris31(6): 79-88. WearegratefultoGerardGeuze,BartdeJong,Lex Jeffreys,G.W.,1867.BritishConchology,oranaccountofthe Kattenwinkel,PeterMoerdijk,FreddyvanNieulande, MolluscawhichnowinhabittheBritishIslesandthesur- HarryRaad,RiaanRijkenandGeorgeSimons(Werk- roundingseas.Vol.4:1-486,pls1-8.VanVoorst,London. groepGeologie,KoninklijkZeeuwschGenootschap Landau,B.,Marquet,R.&Grigis,M.,2004.TheEarlyPliocene derWetenschappen,Middelburg,TheNetherlands) Gastropoda(Mollusca)ofEstepona,SouthernSpain.Part2: fortheloanordonationofmaterialandallowingusto Orthogastropoda,Neotaenioglossa.–Palaeontos4:1-108. publishtheirrecords.WethankAnneliseFolieand Lightfoot,J.,1992.CaecidaeoftheWesternAtlantic.Part1.–Of RobertMarquetformakingavailabletheCaecumma- SeaandShore14(4):171-185. terialintheRoyalBelgianInstituteforNaturalSci- Linden,J.vander&Moolenbeek,R.G.,2000.Caecidae(Mol- ences(Brussels,Belgium).WethankFrankWesselingh lusca:Gastropoda)fromMauritania.–VitaMarina47(3):81- andRonaldPouwerforallowingustostudytheCae- 92. cumsamplesintheNaturalisBiodiversityCenter(Lei- Marquet,R.,1997.PliocenegastropodfaunasfromKallo(Oost- Basteria79(4-6),2015 B2015-15:Basteria-2015 11/20/2015 3:12 PM Page 113 Vlaanderen,Belgium).Part2.Caenogastropoda:Potamidi- 1955.DefossieleschelpenvandeNederlandsestrandenen daetoTornidae.–ContributionstoTertiaryandQuaternary zeegaten,2.–Basteria19(2-3):27-34,pls5-8. Geology34(1-2):9-29. RegterenAltena,C.O.van,Bloklander,A.&Pouderoyen,L.P., Marquet,R.,1998.DepliocenegastropodenfaunavanKallo 1965.DefossieleschelpenvandeNederlandsestrandenen (Oost-Vlaanderen,België):1-246.Publicatieno.17,Belgis- zeegaten.Eersteserie,tweededruk:1-55,pls1-22.Neder- cheVerenigingvoorPaleontologie,Antwerpen. landseMalacologischeVereniging,Leiden. Moerdijk,P.W.&Pouwer,R.,2013.Laevastartequiespacisnov. Sclater,W.L.,1893.ListoftheDatesofDeliveryoftheSheetsof spec.(Mollusca,Bivalvia,Astartidae)fromthePlioceneof the‘Proceedings’oftheZoologicalSocietyofLondon,from TheNetherlands.–CainozoicResearch10(1-2):9-13. thecommencementin1830to1859.–ProceedingsoftheZo- Nofroni,I.,Pizzini,M.&Oliverio,M.,1997.Contributiontothe ologicalSocietyofLondon61:435-440. knowledgeofthefamilyCaecidae.3.RevisionoftheCaeci- Trausel,J.&Slieker,F.,2015.NMR26235.-www.nmr-pics.nl> daeoftheCanaryIslands(Caenogastropoda:Rissooidea).– Caecidae>PLEISTOCENECaecumtrachea(Montagu, Argonauta10(7-12):3-32. 1803)on2015-07-31. Panetta,P.,1980.LafamigliaCaecidaenelMediterraneo.–Bol- Wood,S.V.,1842.ACatalogueofShellsfromtheCrag.–The lettinoMalacologico16(7-8):277-300. AnnalsandMagazineofNaturalHistory9:455-462,pl.5. Pizzini,M.,Raines,B&Vannozzi,A.,2013.ThefamilyCaecidae Wood,S.V.,1848.AMonographoftheCragMollusca:Vol.1 intheSouth-WestPacific(Gastropoda:Rissooidea).–Bollet- Univalves-MonographofthePalaeontographicalSociety tinoMalacologico49,Suppl.10:1-78. London1(1):i-xii,1-208,pls1-21. Porta,J.de,Martinell,J.&GonzálezDelgado,J.A.,1993.Caeci- Wood,S.V.,1872.SupplementtotheCragMollusca,Vol.3Uni- dae(Gastropoda,Mesogastropoda)delNeógenoyCuater- valvesandBivalves,Part1–MonographofthePalaeonto- narioMarinosdelMediterráneoNoroccidentalydela graphicalSocietyLondon25(113):i-xxxi,1-99,pls1-7. PenínsulaIbérica.–RevistaEspañoladePaleontología8(1): Wood,S.V.,1874.SupplementtotheCragMollusca,Vol.3Uni- 1-13. valvesandBivalves,Part2-MonographofthePalaeonto- RegterenAltena,C.O.van,Bloklander,A.&Pouderoyen,L.P., graphicalSocietyLondon27(123):100-231,pls8-11+add.pl. 113 Hoeksema, D.F. & Todd, J.A.–CaecumtumiduminthePlioceneoftheNorthSeaBasin