Development of Cuvierinidae (Mollusca, Euthecosomata, Cavolinioidea) during the Cainozoic: a non-cladistic approach with a re-interpretation of Recent taxa

2005·2.6 MB·English

by  Janssen, Arie W

BASTERIA, 69: 25-72,2005
Development of Cuvierinidae(Mollusca, Euthecosomata, Cavolinioidea) during
the Cainozoic:
anon-cladisticapproach witha re-interpretationofRecenttaxa
Arie+W. Janssen
National MuseumofNatural History (PalaeontologyDepartment),P.O.Box 9517,2300 RALeiden,the
Netherlands;currently: 12,Triqtal'Hamrija,XewkijaVCT110,Gozo,Malta;
[email protected]
Thispaperis dedicated tothe memoryofDennisCurry
Numerous observationsofmore-or-lesspreciselydatedoccurrencesofCainozoicCuvierinidae
allow lineagestobe reconstructed. The early genera Spoelia Janssen, 1990 andJohnjagtiagen.
nov.(LateOligocene/EarlyMiocene),theroots ofwhicharestillunknown,areincluded inthe
Cuvierinidae.
The Late Oligocene Ireneiatenuistriata (Semper, 1861)is the precursor ofvarious Miocene
species ofIreneia Janssen, 1995. The genus became extinct in the Late Miocene. The genus
Cuvierina Boas, 1886developedfromthe IreneialineageduringtheEarlyMiocene.
InMiddle Miocene timesCuvierinas.lat. splitsupintotwobranches, interpretedhere assub-
genera,viz C. (Cuvierina) andC. (Urceolaria)subgen.nov., eachwithanumberofspeciesoccur-
ringfromthe laterMiocene onward.TheRecentC. columnella columnella’(Rang,1827)’and ‘C.
columnella urceolaris (Mörch, 1850)’,asinterpreteduntil now,areconsidered torepresentthe
finalstages ofthese lineages.RecentCuvierina materialfrom alloceanswasrevised,whichhas
ledtoare-interpretationofextanttaxaand tothe introduction oftwonewRecentspecies.
New taxa here introduced are Cuvierina (C.)pacifica spec. nov., C. (Urceolaria) cancapae spec.
nov., C. (U.) curryi spec.nov., Ireneia gracilisspec. nov.,Johnjagtiagen.nov., and Urceolaria sub-
gen.nov.Aneotypeis designatedforC. (C.)columnella.
Twospecies described fromthe Eoceneofthe United Statesdonotfitthe developmentmodel
of the Cuvierinidae advocated here. For them two genera, Praecuvierina gen. nov. and
Texacuvierina gen. nov. are introduced. Togetherthey form the Praecuvierinidae fam. nov.,
whichisconsidered torepresentanearlieroffshootfromthe Creseidae.
Key words:Gastropoda,Euthecosomata,Praecuvierinidae,Cuvierinidae,lineages,Cainozoic,
newspecies, newsubgenus,newgenus,newfamily.
INTRODUCTION
Thestudyoffossil assemblages ofso-calledpteropods (Gastropoda, Heterobranchia,
Euthecosomata) during thelastdecadeshassupplied interestingnew dataonmorpholo-
gy, distributionand biostratigraphy of many groups of holoplanktonic molluscs. The
presentpaperfocuses onthefamilyCuvierinidae.TheCuvierinidaewereconsidereduntil
recently to be a subfamily, Cuvierininae, in theCavoliniidae. Janssen (2003) raised the
26 BASTERIA, Vol. 69,No. 1-3,2005
Cuvierininae, together withCreseinaeand Clioinae,tofamily level.
Among extanteuthecosomatousgastropods, species of thegenus Cuvierina arewell-
characterisedby anumberof shellfeatures,in particular thestraight, cylindrical to more
or less inflatedshell formand the proximal aperture ofreniformor roundedtriangular
shape. In (near-)adult shells a semispherical, more or less obliquely situated septum is
formed, afterwhichtheapical shellpart(larval shell) is shed.
The genus Cuvierina Boas, 1886,as previously interpreted, comprises but a single
Recentspecies, whichisthetypespecies, viz.C. columnella(Rang, 1827)(compareVander
Spoel, 1967, 1970, 1976). Inaddition, quitea numberof fossil species havesubsequently
beenassigned to Cuvierina:
C. astesana (Rang, 1829)- Pliocene
C. globosa Collins, 1934-Pliocene
CC.. ggrraannddiiss d'Alessandro&Robba,1980-Miocene
C. guttaHodgkinson, 1992- Eocene
C. inflata(Bellardi, 1873)-Pliocene
C. jagtiJanssen, 1995- Pliocene
C. intermedia(Bellardi, 1873)- Miocene/Pliocene
C. ludbrooki(Caprotti, 1962)- Pliocene
C. lura Hodgkinson, 1992 -Eocene
C. miyazakiensis Ujihara, 1966- Pliocene
C.paronai Checchia-Rispoli, 1921-Miocene
C.senonica (Blanckenhorn, 1934)-Miocene
C. torpedo (Marshall, 1918)- Miocene
C. tubulataCollins, 1934-Pliocene
In theadult stage theseareall characterisedby a cylindrical or to a higher or lesser
degree inflatedshell,withaseptumclosing theapical sideoftheshellafterthelarvalshell
parts areshed.
Thejuvenile shell(seefig. 14foranexample) isregularly conical, and hasanelongate
protoconch withasingle, near-apical swelling andaroundedtip,closely resembling cer-
tainspeciesofthe family Creseidae(especially Styliola, inwhich,however,thetipis point-
ed). TheshedlarvalshellofCuvierinaspecies differsstronglyfromthefully grownspeci-
mens, and was, at least twice, described as a separate species (Creseis caliciformis
Meisenheimer,1905;Styliola sinecosta Wells, 1974).
Towardstheaperture theadult shellbecomes dorso-ventrally flattenedand frequently
evenslightlyconcaveontheventralsideclosetotheaperture,whichthereforehasareniform
orroundedtriangular shape in adapical view,withaslightlythickenedrim asareinforce-
mentontheinsideof theapertural margin. The dorsalapertural margin is slightly higher
thantheventralone.Theshellsurfacemayormaynothaveamicro-ornamentation,observ-
ableata magnificationof 12 *,ofnumerous close-setlongitudinal lines,usually more clear-
ly developed in the lowertwo-thirds oftheadultshell. Thismicro-ornament,together with
thegrowth lines, may give theshellabeautifulreticulate pattern, especially well visibleat
thoseplaceswherelightreflectsontheshell'ssurface.Intheillustrationsgivenherethisorna-
mentcouldonly be drawnschematically, andinreality ismuchfinerthanseenin thedraw-
ings.
Recent Cuvierina populations are widely distributed in tropical and subtropical
oceans (seealsoVanderSpoel, 1967;fig. 352). They are virtuallyabsent, however,in the
Mediterraneanand theRedSea.
Basedonshellcharacterstwoformas ofCuvierinacolumnellausedtobe distinguished
Janssen: Development ofCuvierinidae during the Cainozoic
27
besidesthetypical form,viz. f.urceolaris(Morch, 1850)andf.atlanticaVander Spoel, 1970.
Later authorsconsideredtheseformastobesubspecies: C. columnellacolumnella,C. colum-
nellaurceolarisand C. columnellaatlantica, although atleastC. c. urceolaris and C. c. colum-
nellawere consideredtobe sympatric.
Belowit willbe demonstratedthat‘C. c. columnella’and ‘C. c. urceolaris’are distinct
species, which developed along separate lineages and are thereforeplaced in different
subgenera. Additionally, thestudy ofmanyRecent Cuvierinasamples has leadtothedis-
criminationofnofewerthanfiveseparatespecies, differingin shellmorphology,biomet-
riccharacteristicsandgeographic distribution.
ABBREVIATIONS
MNHN Museum nationale d'Histoirenaturelle,Paris, France.
RGM NationalMuseumofNaturalHistory (Palaeontology Department), Leiden,
theNetherlands(formerlyRijksmuseum vanGeologieen Mineralogie).
RMNH NationalMuseumofNaturalHistory (Malacology Department), Leiden,
theNetherlands(formerlyRijksmuseum vanNatuurlijke Historie).
SMF Senckenberg Museum,FrankfurtamMain,Germany.
USNM NationalMuseumofNaturalHistory, SmithsonianInstitute,Washington, U.S.A.
ZMUC Zoologisk Museum,Kobenhavn,Denmark.
H shellheight
St. Station
W shellwidth
Allmeasurements arein mm.
MATERIALANDMETHODS
Apart fromthemanyfossilsamples housedin theRGM-collections aswellas insev-
eral private collections, alargenumberof Recent samples wereexamined:predominant-
ly bottom-samples from the CANCAP-expeditions, kept in theRMNH-collections, and
mainly plankton-net samples, especially thosefrom the DANA expeditions, housed in
ZMUC. Somesamples collectedduringtheMETEORexpeditions, housedin theSMF-col-
lectionswere also available.Finally, arestrictednumberof samples fromMNHN,Paris,
was beforeme.
Measured parameters, indicatedin fig. 28, are: shell height, shell width,aperture
width,diameterofseptum,position ofstrongest inflation.
All measurements wereexecuted witha WildM5 stereo binocular microscope at a
magnification of 12 *, using a 120 unitsmicrometerruler, in whicheachunit represents
0.083mm.Theunitswerelaterconvertedtomm. Ofthe saidparametersonly theposition
ofstrongestinflationsometimesis difficulttomeasure, especially inweaklyinflatedspec-
imens,inwhich thesidelinescanbealmostparallel. In such casesthelineperpendicular
tothemeasuring rulerin theWild devicewasveryhelpful.Still, theresults showawider
spreading forthisparameterthanforallothers.
Only adult,full-grown specimens weremeasured.Inmanycases remnantsofthelar-
28 BASTERIA, Vol. 69,No. 1-3,2005
val shellare still adhering to the teleoconch,below the septum. Frequently the shell is
transparantenough to measure thepositionof theseptum,butin some cases partsof the
remaining larval shellwallhadtoberemoved.
THEFOSSIL RECORD OF CUVIERINIDAE
Early taxa
Thequestion whichfossil generashouldadditionally beincludedin theCuvierinidae
is subject of discussion.From the Eoceneof northernAmericaHodgkinson et al.(1992)
included the genera Bucanoides Hodgkinson, 1992, Loxobidens Hodgkinson, 1992, and
TibiellaMeyer, 1884 inthisfamily(at thattime stillCuvierininae).
Loxobidens,however,exclusively knownby its typespecies L.aduncus (Hodgkinson,
1992)ofwhichonly apertural fragments areknown,has acertainresemblancetothecre-
seid genusCamptoceratops, dueto thedenticlesontheapertural flange(Hodgkinson etal.,
1992: pi. 12, figs 1-5).
Bucanoides(with typespecies B.tenuisHodgkinson, 1992andtwoadditionalspecies),
is characterisedby a conicalshell form, and thepresence of aseptum, afterthejuvenile
shellparts areshed.Thisclosely resemblesthecreseid genusEuchilotheca,and thereforeI
alsoexcludeBucanoidesfromtheCuvierinidae,infavourofplacement in theCreseidae.In
all these species the shell surface lacks longitudinal micro-ornament. The apertural
flanges present in mostof thespecies of thesegeneraalso supportanassignment to the
Creseidae,as such structures are not foundin Cuvierinidaeto date.
Thetypespecies ofTibiella(T. marshiMeyer, 1884)is characterisedby thepresenceof
aseptum, similar tothatseen in Euchilothecaand Bucanoides,whereasitsapertural struc-
tures closelyresemblethoseof Thecopsella Cossmann,1988.Thus,Tibiellais includedhere
in theCreseidaeas well.
The question whether thetype species of Eocene Thecopsella (= T.fischeri Cossmann,
1888)belongs in the Euthecosomataor in the Caecidae(compare Tembrock, 1965)is not
problematic, as itsprotoconch is straight,notspirallywound,andtypically creseid, unlike
some species with a spiral protoconch included in this genus by Tembrock [1965:
Thecopsella lituus (Cossmann, 1899); T. undulataTembrock, 1965; T. latdorfense Tembrock,
1965]and R. Janssen (1978: T. tenuiannulataR. Janssen, 1978) thatindeedbelong in the
Caecidae.
Cuvierina is closely related toand,asindicated by fossil occurrences, afurther devel-
opmentof therecently introducedgenusIreneia, theoldestknownspecies ofwhichis the
NWEuropean Late Oligocene (Chattian) I. tenuistriata(Semper, 1861). Species of Ireneia
resemble Cuvierinain shellshape, morphology of larvalshelland aperture, andpresence
of longitudinal micro-ornamentation ontheadult shell,but they differin thatthelarval
shellis retained, not shed.
The first genuine species of Cuvierina originated from the Ireneia-root during the
Oligocene-Miocene transition.Apparently thenewly developed principle oflarval shell-
shedding was advantageous, although necessitating drasticchanges in theanimal's vital
functions,suchas thecapabilityto generateaclosingseptumandre-attachmentofthecol-
umellarmuscleatahigher position ontheinner shellwall.Consequently, Ireneiabecame
extinct during thefinal Miocene,whereasCuvierinasurvives tothepresent day.
The development of Cuvierina from Ireneiaas advocated here disregards two early
taxa, introducedfrom the Eocene of the United States, i.e., C. lura Hodgkinson, 1992
(Lutetian)and C. guttaHodgkinson, 1992 (Bartonian). Thetypematerialof thesespecies
]anssen:Development ofCuvierinidaeduring theCainozoic 29
(seenSeptember 1999)is similartotrue cuvierinids.However,several differences,such as
the surprisingly small dimensionsand the complete absence of ornament,should be
noted.As thewell-documenteddevelopmentof Cuvierinafrom Ireneiastartedlater,in the
Early Miocene, the two Eocene species are here consideredto represent anearly, and
unsuccessful, offshootofCreseidae.In thesecircumstancesthetwo taxa cannot bemain-
tainedin Cuvierina,andbelowI introduceforthemnew genera.
After the onsetof theCuvierina side-branch thegenus Ireneiastill develops several
newspecies duringtheMiocenewhichretain thetypical featuresofthegenus,viz. I.nieu-
landiaJanssen,1995,I.testudinaria(Michelotti, 1847),I.calandrellii(Michelotti, 1847)[inclu-
sive of its probable synonym I. urenuiensis (Suter, 1917)], and I. gracilis spec. nov. The
youngestspecies inthislineage is I. marqueti Janssen,1995, fromtheLateMioceneof the
NorthSeaBasin, characterisedbythepresenceof transverse annulations.
InadditiontoIreneiatwoothergenera,occurring duringtheLateOligoceneandEarly
Miocene, areincludedin theCuvierinidae.TheydifferfrombothIreneiaandCuvierinaby
showing reinforcementstructures intheshell.InmonotypicSpoelia Janssen,1995theshell
hassquarish lateralcarinaeinthemedianpartoftheshellandareinforcedaperturalmar-
gin. S.torquayensis Janssen,1990was describedfromtheLate Oligocene ofSAustraliaand
SE France, but in the meantime it was also recognised in the Chattian 'Sternberger
Gestein'intheNorthSeaBasin (RGMcollections). Quiterecently, itwas alsofound tobe
present in thelowermost phosphorite-bearing level in the Maltese archipelago (Gozo,
Wardija;Janssen,2004), belowtheC1mainphosphorite level,andthereforeofAquitanian
age (Rehfeld &Janssen, 1995).Theoriginof Spoelia is stillenigmatic; it appearsunlikely
thatyoungertaxa developedfromthisspecies. Themorphology ofitsprotoconch, andthe
presenceofmicro-ornament,however,show ittobe adistinctcuvierinid.
Thesameis true for‘Cleodora(Creseis)’ moulinsiiBenoist, 1873,a species as yetexclu-
sivelyknownfromtheEarly Miocene(Burdigalian) oftheAquitaine Basin(France), char-
acterisedby anIreneia-likeshell with a distinctmicro-ornament,butwithalongitudinal
dorso-lateralfoldhalfwaytheshellheight,andareinforcedaperturalmargin asin Spoelia.
Itdiffersmarkedlyfromtheothergeneradiscussedhereandbelowanew genusisintro-
duced for it.The morphological characteristics indicate itssystematic position to be in
between Spoelia andIreneia.
Thefirst typical Cuvierina species is C. torpedo (Marshall, 1918), described fromthe
EarlyMiocene ('LateOtaian'=Aquitanian) ofNew Zealand.In thisspecies theshedding
ofthe larvalshelloccursclose totheprotoconch, the septum, therefore, is smalland the
shell retains anelongate spindle-shape with a slenderbasal part, strongly resembling
species ofIreneia, inclusiveof thepresence of thelongitudinal micro-ornament (Janssen,
1995:55).
Although its morphology andapproximate agemakeit likely thatC. torpedo indeed
is thefirstrealCuvierinaspecies splittingofffromtheIreneiaroot,itisdifficulttointerpret
theNewZealandlocalityintermsofpalaeobiogeography. Thesupposed ancestorspecies,
Ireneia tenuistriata,is foundexactly attheopposite side ofthe globe, viz. in the NorthSea
Basin. Most probably this must be explained in terms of as yet incompletely known
palaeogeographical distributions.
Thefurther development of Ireneia and Cuvierina couldvery wellbe studied in the
MediterraneanNeogene, asdiscussedinthenext section.
Cuvierinid developments inthe Mediterraneanarea during the Miocene
In theMediterraneanarea thefirsttypical Cuvierinais foundin theMaltesearchipel-
30 BASTERIA, Vol. 69, No. 1-3,2005
ago. Here relatively deep-water, andthereforepteropod-rich deposits arefoundwithin
theso-calledGlobigerinaLimestoneFormation(Aquitanian toLanghian), andin theBlue
ClayFormation (Serravallian). In theGlobigerina Limestone they occurpredominantly as
phosphatic internal moulds, concentrated in two main phosphorite levels (Rehfeld &
Janssen, 1995), as well as in several subordinatephosphorite concentrations,or, in the
uppermostpart, as scattered occurrences in the sediment. In the Blue Clay Formation
pteropods, andmostotherfossils aswell, are foundas limoniticinternalmoulds.Inboth
cases the shells, and thereforealso apossible micro-ornamentof theshells, have disap-
peared.
In theMalteseMiocene series Cuvierina paronaiChecchia-Rispoli, 1921 is theearliest
species, found in restricted numbers in phosphorite level C 2, dated as
'Burdigalian/Langhian transition'and it continues through all pteropod bearing levels
intotheBlueClayFormation,whichisofSerravallianage(KieneletaL, 1995).
Cuvierinaparonai was originally described, also in aphosphatised state ofpreserva-
tion,fromstrata atGargano (Puglia, Italy) that,basedontheirmicrofauna(d'Alessandro
&Robba, 1980),belong toBlowZoneN17,whichaccording toBerggren etal.(1995)isof
Late Tortonian to Messinian age. The Gargano pteropod assemblages, however, do not
containspecies indicating anageyoungerthanLanghian, andthuswereconsideredtobe
mostprobably reworked.
In Cuvierinaparonaishedding ofthelarvalshelloccurs atalarger diameteroftheshell
andconsequently theclosing septumis considerably widerthaninitssupposed ancestor,
C. torpedo. TheItalianand Maltesespecimens of thisspecies areall preserved as internal
moulds, but in some specimens in perfect shell preservation fromthe Badenianof the
CentralParatethys (Zorn, 1991)itcouldbeobservedthatthelongitudinal micro-ornament
is clearlypresentin thisspecies as well. The shellformisalmostcylindrical withamore
conicalbasal part. Usually, as in mostcuvierinids, the shelldemonstratesa slight dorso-
ventral flattening, especially towardstheaperture. Thisspecies apparently is the precur-
sor of two separatelineages, onein which thecylindrical shell-formis maintained,and
anotherinwhich theshelltoa higheror lesser degree isinflated.
Theearliestspecies in the latterlineage is C. intermedia(Bellardi, 1873), whichup to
nowwas only known fromPliocenedeposits.Aquitetypicalspecimen (B.M.Landau col-
lection, Albufeira, Portugal; seeJanssen,1995: 41, asC. ? intermedia) fromtheSerravallian
of CacelaVelha(Algarve, S. Portugal) demonstratesthat C. intermediaalready separated
fromtheC.paronai root in MiddleMiocene times.Alsospecimens fromthelaterMiocene
of S.Italy(Puglia, Salento,Lecce area) are hereincludedin C. intermedia.Theinflationof
theshellin thisspecies is gradual and notvery strong, and thereisnopreapertural con-
striction, indicating its closerelationship withC. paronai.
D'Alessandro & Robba(1980) alsomentionedreworkedoccurrences of C. paronai in
thesamedeposits oftheLeccearea. Theydatedtheseas transitionalbetweenBlowzones
Nl4 and Nls, i.e., LateSerravallian.Thepteropod assemblage fromtheselocalitiescon-
tains,amongst other species, two taxa that donot occurin the Gargano area.These are
both species of the genusCuvierina, indicatedby d'Alessandro & Robba (1980) as ‘C.
columnellaurceolaris(Mörch, 1!850)',here consideredtorepresentanewspecies((C. curryi,
seebelow) and C. grandis d'Alessandro&Robba, 1980.
Interestingly, thesetwo cuvierinidscanbe directlylinkedtoinsituoccurrences in the
Maltese archipelago, where they are restrictedto the SerravallianBlue Clay Formation.
Herewefind typicalspecimens ofC. grandis. Theyresemble C.paronai intheircylindrical
shellform,butthebaseof theshell generally is less conical, andthey differmarkedly in
size,reaching aheight ofapproximately 18mm. They arestillabsent inthelowerpartof
theBlue Clay,but relatively commonin theupperpart.
Janssen: Development ofCuvierinidaeduringthe Cainozoic 31
NexttothisC.grandis and C.paronai yetanother cuvierinid ispresentin theBlueClay.
This species differs fromboth C. grandis and C. paronai by its smallerand moreinflated
shell.They areidenticalwiththespecimens fromSalentoidentifiedasC. columnellaurce-
olarisbyd'Alessandro& Robba.
Furthercuvieriniddevelopment isdemonstratedby apteropod assemblage foundin
the so-calledTellaroFormationof SESicily. This deposit superficially resemblestheBlue
Clay Formation of Malta, as both consist of clayey sediments, and contain fossils in a
limonitisedstate of preservation, with, e.g., the cephalopod Sepia presentin both, and
withmanyfurthersimilaritiesin thebenthicfaunas.Di Geronimoetal.(1981) datedthese
Sicilianclaysas Globorotaliaacostaensis Zone (= BlowZone 17;Tortonian),but erroneously
correlated them with the upperpart of the Maltese Blue Clay Formation.Martini (in
Janssen, 1999b: 116) analysed a nannoplankton sample from the Sicilian locality and
indeedfound ittobelongto theNN10zone(Middle Tortonian).
Theholoplanktonic molluscanassemblage of theSicilianTellaroFormationincludes
threespecies ofCuvierina(Janssen, 1999b).Thecommonest inthismaterialhasashellwith
a strong inflationjust below mid-height and a reniformaperture, apparently a further
development ofthe so-called‘C. columnellaurceolaris’(= C. curryi, describedbelow) from
Salento.The materialcannot specifically bedistinguished from Cuvierinainflata (Bellardi,
1873),describedfromtheEarly PlioceneofN.Italy.Other, lesscommonspecimens in this
materialstill resembleC. paronai; they donotreach thelarge dimensionsof C.grandis. A
thirdgroupofspecimens differsfrombothby thepossession of anelongate, slenderand
cylindricalshellandby atriangular shape oftheaperture.
Thislatter formis also known fromN. Italy (Turin Hills),in a locality namedTetti
Borelli, fromwherePavia & Robba (1979) extensively describedtheholoplanktonic mol-
luscanassemblage (revisedin Janssen, 1995).ThesimilaritybetweenthePoggio Musenna
andTettiBorelliplanktonic molluscanfaunasbeyond any doubtindicatesthatbothare as
good ascoeval (Janssen, 1999b).
Theslender Cuvierinajagti Janssen, 1995 found at Tetti Borelli, also present in the
Poggio Musenna assemblage, appeared tohavebeenintroducedearlieras a scaphopod,
i.e. Dentaliumludbrooki Caprotti (1962: 96, pi. 16, figs 4-6) from Early Pliocene clays of
CastelTArquato innorthern Italy (Janssen, 1999a). The Tetti Borellispecimens clearly
demonstratethepresenceof alongitudinal micro-ornament.
From allthese dataitmaybe concludedthat theancestral species Cuvierina paronai
developed two lineages, one characterisedby a cylindrical shell and a triangularrather
thanreniformaperture (Cuvierina s.str.), and theotherby aninflatedshellandreniform
aperture [Cuvierina (Urceolaria) subg. n., describedbelow]. Inthe courseof the develop-
mentof Cuvierina s.str. a tendency to decrease thelongitudinal micro-ornamentis seen
fromtheEarly Plioceneonwards.
Cuvierinainthe Pliocene
Thetrendsobserved inthedevelopmentof Miocenecuvierinidspecies continuedur-
ing thePliocene.Thesubgenus Urceolariacontinues untiltheZanclean/Piacenziantransi-
tionwithC. intermediaand C.inflata. The Recent‘Cuvierina columnellaurceolaris’apparent-
ly is theconcluding form in this development, andthereforeC.urceolarisis here consid-
eredadistinctspecies. Thisspecies is well-knownfromtheIndianandPacificoceans,and
wasrecently also collectedfromstratainthePhilippines, thatafterafirstanalysis arepre-
sumably Piacenzianin age(Janssen, 2000 unpublished).
Janssen (1995) synonymised Cuvierina globosa Collins, 1934, from the Pliocene
32 BASTERIA, Vol. 69,No. 1-3,2005
Agueguexquite Formation in Mexico with C. inflata. From the Early Plioceneof Japan
(Blowzones 18-20) Ujihara (1996,figs 6.16-28) illustratedspecimens fromthislineage as
C.intermedia,whichbecause oftheirstronglyinflatedshellsandpreapertural constriction
agreebetter with C. inflata.
The lineage containing the cylindrical species (Cuvierina s. str.) continues in the
MediterraneanEarly Pliocenewith C.ludbrooki. Thisspecies also occurs in the Japanese
Pliocene(Ujihara etal, 1990;Ujihara, 1996;as C. cf.tubulata) andwas recently alsofound
to occur, together with C. intermedia, in Pliocene deposits of the Fiji Islands (Andrew
Grebneffcollection).
Less slender is a common species known from the MediterraneanZanclean and
Piacenzian,C. astesana(Rang, 1829).In thisspecies areductionofthelongitudinal micro-
ornamentis seen,andtheaperturevariesfromreniformtotriangular. The widerange of
variation that couldbe observed in the abundantmaterial available from French and
Italianpopulations of C. astesana includes formsstrongly resembling theLateMiocene-
PlioceneC. ludbrooki, which is here consideredto be its ancestral species. TheMexican
species C. tubulataCollins, 1934 is hereinterpreted to beprobably a synonymof C. aste-
sana.Just asingle, doubtfulspecimen of C.astesanaisknown fromtheNorthSeaBasin.
Another species, described as C. miyazakiensis Ujihara, 1996 (not seen), from the
Pliocene of Japan (Blow zones18-20), seemsto make a perfect connection to Recent ‘C.
columnella’.
RecentCuvierina
RecentCuvierinidaewere extensively studiedbyVanderSpoel (1967,1970,1976).In
his 1967paper thisauthor, asmanybeforehim, stillaccepted just asingle species, with
two formas, viz. Cuvierinacolumnellaf. columnella(Rang, 1827) and f.urceolaris (Morch,
1850). The main distinguishing characteristic was considered to be the position of the
greatestshellwidth (close to caudalclosing septum in f. columnellaand inthe middleof
theshell in f. urceolaris). Specimens withthe greatestwidthin betweenthese two points
wereindicatedintermediates.
In 1970VanderSpoel introduceda thirdforma, f.atlantica, subdividing thef.colum-
nella into asmall Indo-Pacificform(columnella), and alarger, predominantly Atlanticform
(atlantica). Later authors consideredthese formas to be subspecies, although at least f.
columnellaandf.urceolariswereknown tobesympatric.
Since Van der Spoel (1970) explicitly introduced f. atlantica as an infrasubspecific
taxonthenamewasvalidatedas aspecies groupnamebyBé,MacClintock& Currie(1972:
58) (ICZN arts. 15.2and45.5.1), which authorsgaveanextensive description oftheshell
micro-architecture in‘C. columnellaatlantica’.
If we consider the development of Cuvierina populations during the Neogene, as
describedabove, the conclusionis obvious that theRecent populations that used tobe
identifiedasC. columnellas.lat.arenot monospecific.Whatwasconsideredtobe'thetyp-
ical form' and C. columnellaatlantica, withhardly inflated, cylindrical shells, alack of (at
bestareduced) longitudinal micro-ornamentandtriangularapertureoutlinesarethesuc-
cessors of theparonai-grandis-ludbrooki-astesana-miyazakiensis-line, whereastheinflatedC.
columnellaurceolariswithitsobviousmicro-ornament andreniformapertureconcludesthe
lineage intermedia-curryi-inflata.
Of Recent Cuvierina a large numberofbottom samples fromthe northernAtlantic
Ocean, covering the Madeira, Canary and Cap Verde areas,but also from the Guyana
Coast and the eastern Caribbean, was available.In this materialthe occurrence of C.
Janssen:Development ofCuvierinidae during the Cainozoic 33
atlanticaisconvincingly demonstratedby itsrelatively largeand slendershell(H min.6.7
mm,max. 10.5mm,meanvalue8.3mm,H/Wmin.3.2, max.3.9, meanvalue3.55,n=190),
thevirtualabsence ofmicro-ornament,thetriangularaperture andthevery lowposition
of the point of strongest inflation.Itis mainly foundaround Madeiraand theCanaries
and isalsopresentinthewesternAtlantic, in theCaribbeanandmoretotheNorth.
A largenumberof samples, however,predominantly originatingfromtheCapVerde
area, differstrongly fromthisatlanticatype.Notonly are theshellsclearly lessslender(H
min.7.6mm,max.9.3mm,meanvalue8.44 mm,H/Wmin.2.8,max.3.6, meanvalue3.07,
n= 118),butthey alsohavea distinctmicro-ornament, andareniformrather thantrian-
gularaperture.These characteristicslinkthemtotheIndo-PacificC. urceolaris, fromwhich
they differby larger dimensionsand aless pronounced inflationof theshell, whichalso
is situatedatalowerpoint. Thisuntilnowunrecognised formisalsopresentin somesam-
ples fromthe Guyana Coastand the Caribbean.It is describedbelowas anewspecies,
Cuvierinacancapae.
TheAtlantic materialfromthe DANAexpeditions, housedinZMUC, substantiates
theexistence oftwo taxa in thisarea.From thislatter materialC.atlantica appearsalsoto
bepresentintheS Atlantic,butIhadinsufficientsamples availabletoobtainafairideaof
its distributionthere.It seemsthatthisspecies showsthebipolaritythatis well-knownin
pteropods.
FromtheIndianandPacificOceansIcouldstudy manysamples fromtheZMUC col-
lection. This collectionincludes several hundreds ofalcohol samples, mainly collected
alive during theDANA-expeditions and most of them(re-)identified in the 1970sby S.
VanderSpoel.
Cuvierinaurceolarisinvariably iseasily recognised inthismaterialbyitssmallsizeand
distinctly inflatedshell(H min.5.1 mm,max.6.6mm, meanvalue5.99mm,H/Wmin.2.3,
max. 2.8, mean value 2.55, n = 163), clear micro-ornament and reniform aperture.
Specimens fromtheIndianOceanonaverageareslightly smaller(H meanvalue5.49mm,
n=47)thanthose fromthePacific(Hmeanvalue6.19 mm,n=116).
C. columnellacolumnella,as characterisedby Van der Spoel (1972: 120,fig. 19) alsois
readily identifiedbybeinglargerthan C.urceolaris,moreslender(Hmin. 6.6mm,max. 8.9
mm,meanvalue7.75 mm, H/Wmin.2.9,max. 4.0,meanvalue3.46, n=135)and hardly
inflated, withthe point ofmaximum inflationsituatedratherhigh,and furthermoreby
thecomplete absenceofmicro-ornament and atriangular aperture.
Many samples, however,identifiedas C. columnella(f.) columnellaby VanderSpoelin
the ZMUC collectionIhadto re-identify as C. urceolaris,becauseof their inflatedshells
and distinctmicro-ornament,as aresultofwhichthegeographic distributionof‘C. colum-
nella’s. str. was drasticallyreduced.
The quiteunexpected resultofthisis, that'typical' C. columnellasensuVanderSpoel,
1970is exclusively found tobepresentin thePacific, bothNandSoftheequator,witha
barren zonein between. The N Pacific form(H min. 6.6 mm, max. 8.5 mm, meanvalue
7.59mm,H/W min. 2.9, max. 3.5, meanvalue3.23,n=36) differsinmeasurements from
theS Pacificone(H min.7.1,max. 8.9, meanvalue7.81 mm,H/Wmin. 3.3,max.4.0,mean
value3.54,n=99). Thisleadstotheoddconclusion, thatC.columnellaaspreviously under-
stood, doesnot occur in its type locality, which is 'la mer des Indes', theIndianOcean
(Rang, 1827:323).
This 'problem', however,israpidly solved, as apartfromtypical C. urceolaris, anoth-
er as yet unrecognised formoccurs, intheIndianOceanaswellasin theSPacific, which
was also identifiedC. columnella (f.) columnellaby Van der Spoel, but differs by larger
dimensionsand aclearmicro-ornament.InitiallyIconsideredthesespecimens tobeiden-
ticalwith C. cancapae, buttheir measurements and shape of aperture resemble closely
34 BASTERIA, Vol. 69,No. 1-3,2005
SSuubbggeennuuss ssppeecciieess DDiissttrriibbuuttiioonn sshhaappee aappeerrttuurree oorrnnaammeenntt nnaammeeiinnVVaannddeerrSSppooeell,,11997700
CCuuvviieerriinnaa aattllaannttiiccaa NN&&SSAAttllaannttiicc ccyylliinnddrriiccaall ttrriiaanngguullaarr ±±aabbsseenntt aattllaannttiiccaa((ppaarrttiimm))
CCuuvviieerriinnaa ccoolluummnneelllala IInnddiiaann,,SS..PPaacciiffiicc ccyylliinnddrriiccaall ttrriiaanngguullaarr pprreesseenntt ccoolluummnneellllaa,,aattllaannttiiccaa((ppaarrttiimm))
CCuuvviieerriinnaa ppaacciiffiiccaa NN&&SSPPaacciiffiicc ±± ccyylliinnddrriiccaall ttrriiaanngguullaarr aabbsseenntt ccoolluummnneelllala((ppaarrttiimm))
UUrrcceeoollaarriiaa ccaannccaappaaee CC&&SS AAttllaannttiicc lleessssiinnffllaatteedd rreenniiffoorrmm pprreesseenntt aattllaannttiiccaa((ppaarrttiimm))
UUrrcceeoollaarriiaa uurrcceeoollaarriiss IInnddiiaann,,WWPPaacciiffiicc iinnffllaatteedd rreenniiffoorrmm pprreesseenntt uurrcceeoollaarriiss,,ccoolluummnneellllaa ((ppaarrttiimm))
Table 1.MaincharacteristicsanddistributionofRecentCuvierinaspecies.
thoseof C. atlantica, fromwhich they in fact differpredominantly by the presenceof a
clearmicro-ornament. Thisformis commonly presentinthesouthernIndianOcean,and
no doubtit isthis formthat was described asC. columnellabyRang, 1827.
Unfortunately it is impossibleto check this onthetype material,of whichonly the
labelsurvives (Van derSpoel, 1976: 191).Rang's illustrationshows anextremely slender
shell, with H/W-ratio = approximately 5.1, avalue thatexceeds by far the highest value
(H/W =4.0) found in allmy measurements,and therefore Iconsider the drawing to be
incorrect.Theheight of the specimen illustratedbyRang is 11 mm and thatagreesper-
fectlywithmy findings (seeMeasurements,fig.29).
In thesecircumstances Ibelieveitnecessary to designate a neotypeforC. columnella,
differing fromwhatis currently understoodunderthatname, and which thereforeis in
need ofanewname.In thesystematic partbelowIproposethenameC. pacifica forwhat
used tobe considered'typical C. columnellafromthePacific. Theneotypeof C. columnel-
lais designated in materialfromtheMNHN collections, where thetypeof C. columnella
Rang, 1827,used tobe housed.
In summary, I conclude that nofewer thanfive species shouldbe distinguished in
RecentCuvierina, inpart differentfromprevious interpretations ofthesenames,andwith
almost completely separate geographic distributions. Asummary of these is given in
Table1.
In C.pacifica aswellasin C. urceolaris,toalesser degreealsoin C.columnella,themeas-
urements indicatethatpopulations fromdifferentareas differsignificantlyin sizeand/or
proportions; thesemighteventually beinterpreted as geographic subspecies.
However, themain characteristicsof someof theseextant taxa unfortunately donot
fullycorrespond totheoverallpicture asseeninthetwolineagesforthefossilspecies. The
newinterpretation of C.columnella,thetypespecies ofCuvierina, indicatesthatthelossof
longitudinal micro-ornament apparently doesnotoccur in all Recent speciesof Cuvierina
s. str.,although areductionofthemicro-ornamentisonly found in thissubgenus. Thusit
isratherthecylindrical shellformandtriangularaperturethatcharacteriseCuvierinas. str.
Still,in theNPacificformofC.pacificaspecimens withmoreor less inflatedshellsarealso
present.
C. urceolaris, the type species of Urceolaria, characterises the lineage with inflated
shells, but C.cancapae, also includedin thatsubgenus, has only a weakly inflatedshell,
whereasitsothermorphology clearly suggests Urceolaria.
Thisapparentmixture offeaturesintheRecentpopulations mightsuggestthatweare
dealing with hybridisation, e.g. cancapae= atlantica x urceolaris, or columnella=pacifica x
urceolaris,butthisidea isby nomeanssupported by thevarious distributionpatterns.
Anotherpossible explanation forthese 'hybrids' might be that thegenusCuvierinais
in thecourse ofspeciation. Itmustbeborneinmind,thatofthefossiltaxaweusuallyhave
only a very poor knowledge of their horizontaldistributionand thereforeof their geo-