Bulletin of the Natural Histort Museum. Geology series

84 Pages·2001·7.8 MB·English

by  Natural History Museum (London)

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ISSN 0968-0462
Bulletin of
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NATURAL
W»Tt)RYMi>S6UM
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(3EME«Al. LfflRARy
Geology
Series
VOLUME 57 NUMBER 2 29 NOVEMBER 2001
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Geology Series
ISSN 0968-0462 Vol. 57, No. 2, pp. 83-162
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Bull.nat. Hist.Miis.Land. (Geol.)57(2):83-93 Issued29November2001
The Cenozoic Brachiopod Terebratula: its type
species, neotype, and other included species
THE NATURAL
tflSTORY MUSEUM
DAPHNE E. LEE
S OCT 2001
DepartmentofGeology. UniversityofOtago. P.O. Bo.x56. Diinedin. NewZealand
C.H.C. BRUNTON PB£ScWTHD
DepartmentofPalaeontology. TheNaturalHistoryMuseum. CromwellRoad, LondonSW75BD QEN£RAL UBRARY
EMMATADDEIRUGGIERO
DipartimentodiScienzedella Terradella Universita'diNapoli 'FedericoIT, LargoSanMarcellino 10. 80138
Napoli. Italy
MASSIMO CALDARA & ORONZO SIMONE
DipartimentodiGeologiaeGeofi'iica. Campus Universitario, Via Orahona4, 70125Bari, Italy
Synopsis. Terebratulaterebratula(Linnaeus.1758)hasalongandcomplexhistory.Thespecimennowrecognisedasthetype
wasfirstillustratedbyColonnain 1616,andthefirstuseof'Terebratula'isattributedtoLhwyd, 1699.Colonna'sspecimenwas
refiguredbyKlein,1753,andthespecies/Inom/aterebratulawasdescribedbyLinnaeus.1758.withreferencetotheColonnaand
Kleinillustrations.ThegenusTerebratulawasproposedbyMijllerin 1776.andAnomiaterebratulaLinnaeusdesignatedasthe
typespeciesbyLamarckin 1799.althoughitwasnotanoriginallyincludedspecies.Inspiteofthishistory,thetypeofthegenus
wasneverformallyratified,thewhereaboutsofthetypespecimenwasunknown,andtheageandexactpositionofthetypelocality
wasuncertain.ThispapersummarisesthehistoryofTerebratulaterebratula(Linnaeus)from\616.AnomiaterebratulaLinnaeus
isnowacceptedasthetypespeciesofTerebratula(ICZNruling, 2000).Wehavecollectednewmaterialfromalocalitynear
Andria,Italy,fromwhichColonnacollectedspecimensofTerebratula,andselectedaneotypefromtheCalcarenitediGravina
FormationwhichisPlioceneinage.Twoexistingspecies,Terebratulasinuosa(Brocchi)andTcalabraSeguenza,areplacedin
synonymy with T. terebratula. Three species arecurrently recognisedin Terebratula. ranging in age from MiocenetoEarly
Pleistocenewhenthegenusbecameextinct,probablybecauseofoceancoolingintheMediterraneanregion.
INTRODUCTION genusTerebratulaasdesignatedbyLamarckin1799(Lee&Brunton,
1998;RulingoftheCommission, September2000).Theremaining
objecfivesareachievedinthispaper.
ThebrachiopodgenusTerebratulahasalongandcomplexgeologi-
cal andnomenclaturalhistory.Thenominal genus Terebratula was
proposed by Miiller in 1776, and as pointed out by Muir-Wood DERIVATION OFTHENAME
(1955), it 'is the first valid post-Linnean brachiopod genus".
TTeerreebbrraattuulliadtaerwehbriacthulaen(cLoinmnpaaesusse)sisltohoepn-abemaer-ibnegarberrafcohritohpeoOdrsdeorf ThenameTerebratulawasfirstusedinprintbyLhwyd(1699)(Little
Devonian-Recentageandincludesmostbrachiopodslivingtoday. etal.. 1973),andistheoldestgenericnameinthePhylumBrachiopoda
The name Terebratula has been widely usedforover200years: incurrentuse. Terebratulais "so-calledfromtheperforatedbeakof
more than 850 specific names were applied to the genus between theventralvalve'(Littleetal.,1973:2265),andisaquasi-diminutive
1800-1850 alone (Sherborn, 1932). In spite ofthe ubiquity ofthe ofLatin terebratus, the past participle ofterebrare 'to bore'. The
nuTneatrmieleb,rraettcheuenltgaleytn,eursbebearenantdullsiatptelrceeimseatsuiodnnieedwdhuainncadnhsimwtaewrnaeysd.boarAissgiicpnoaqilunletysetbdiaoosunetsdnahebaaovruelt,y btfhiregauccrhoeildolpesocptdeiscoinlmsisetonefsdtafhsreoTAmesrWehibmtroanlteueyla'an(iMnMuuLishrew-uyWdmo',osd,Ocxaf1to9ar5l5do:,gu2we)e.orTfehseh'eplnoloasrmlieyn
acenturyago(Buckman, 1907),notonlywasthetypespeciesofthe came into fairly common use in the 18th century, and some ofthe
numerous brachiopods referred to as Terebratula by other pre-
genusunconfirmed,butitstypelocalityandagewereuncertain. Linneanauthorswerementionedby Muir-Wood(1955).
The present study had several aims. The first objective was to
ratifythe typeofthegenus Terebratulaforinclusion intherevised Thespeciesnameterebratulawasfirstusedinavalidbinomialby
BrachiopodTreatise,followingtherecommendationmadebyMuir- LinnaeusinhisdescriptionofAnomiaterebratulainSystemaNatu-
Wood in the 1965 Treatise volume on the Order Terebratulida. rae(1758:703).Linnaeusgavenoillustration,butreferredtofigures
Secondly, we wished to summarise the complex nomenclatural inColonna (1616c), Lister(1678) and Klein (1753).TheColonna,
historyofthegenusandspecies,Terebratulaterebratula.sinceitwas andColonna/Klein, illustrationsarereproducedinFigs 1,2.
firstillustratedanddescribedin 1616.Thethirdaimwastolocatethe
typespecimenand/ortypelocalityofT.terebratula,or,ifthisproved THE COLONNA ILLUSTRATION OF
impossible, select a neotype to act as namebearer for the order.
Finally,wewishedtodescribetheageandrelationshipsofspecies TEREBRATULA (FIG. 1)
currentlyincludedin Terebratula.
Thefirstobjectivewasachievedwithanapplicationtothe Inter- FabioColonna(1567-1650) (FabiusColumna)wasbom in Naples
national Commission on Zoological Nomenclature to validate the and was one ofthe first natural historians to use copper plates for
selection ofAnomia terebratula Linnaeus as type-species of the engravingbotanicalandzoologicalfigures.Hewroteextensivelyon
©TheNaturalHistoryMuseum,2001
84 D.E.LEE,C.H.C.BRUNTON,E.TADDEIRUGGIERO,M.CALDERAANDO,SIMONE
22 Fabij Columns
^
^^W*^ C«ie}» (LMiitt* u«r/tcfre^iH
bS-^^SS)®SX^(SS^»M@S€2'^
gCTS'S©'^^'^iS^©«^«^i^«>^£"^
Fig.1 Reproductionofpage22inPurpura(Coionna. 1616).
fossils, and 'was one ofthe first to place them ... in a primarily pp. 1-29.andDeglossopetrisdissertatio, pp. 31-39,aretwochap-
biologicalcontext...Coionnaalsoappliedthesameprecisenomen- ters), in which he described and figured a numberofshells, some
claturetohisfossilsastohislivinganimals,distinguishingdifferent fossilandsomeliving.Hisillustrationonpage22(Fig. 1herein)was
kinds of related fossils with more accuracy then ever before' a woodcut of five shells. The upper three specimens are double-
(Rudwick, 1985:42). valvedbrachiopods.whilethelowerspecimensareinternalmoldsof
CoionnawasamemberoftheAccademiadeiLincei(Academyof bivalves. The plate is not numbered, and the five specimens are
Lynxes),andin 1606,hepublishedaworkonnaturalhistory:Minus distinguishedbybriefcaptionsplacedaboveeachspecimen.
cognitarum stirpium aliquot, ac etiam rariorum nostra coelo Linnaeus(1758: 703)madethreeseparatereferencestotheillus-
orientium [ecphrasis] . . De aquatilibus, aliisque animalibus trationsofbrachiopodsonthisplate inhisdiscussionofspeciesof
quibusdampaucislibellus[plantspp.3-340;animalsI-LXXIII].A Anomia. Under Caputserpentis. 200., he gave a briefdescription,
new edition was published in 1616 (Coionna, 1616a-c), including andreferredtoColumn,purp.22.f.2,i.e.thesmoothbrachiopodon
part111,Purpura(ofwhichDepurpura,aliisquetestaceisrarioribus. the upper right. Brunton & Cocks (1967) discussed in detail the
.
THECENOZOICBRACHIOPODTEREBRATULA 85
Fig.2 FiguresofConchaanomia.theholotypeoi'Anomiaterehratula.2a,woodcutfromColonna(1616):2b,woodcutfromMajor(1675):2c,engraving
fromKlein(1753).
ambiguitieswhicharosewhenLinnaeus(1767)changedhisdescrip- onthefollowingpages.Inparticular,several specimensmentioned
tion ofA. caputserpentis from a smooth, fossil brachiopod to a on pages 23-24are not illustrated at all, although they were num-
capillate living species (now Terebratulina retusa), although he beredsequentiallybytheauthor.
retainedthe reference tothe Colonnafigure. An application to the Thus, Lee & Brunton (1998) assumed that the strongly folded
International Commission on Zoological Nomenclature to change figureontheupperleftonpage22wasthatdescribedinthetexton
the type species of Terebratulina d'Orbigny, 1847 from Anomia thefacingpage(Cap.XII)undertheheading 'ConcharariorAnomia
caputserpentis Linnaeus, 1758, toAnomia retusa Linnaeus, 1758, verticerostrato',andaccordinglytheyconcludedthatthisspecimen
disposed of the ambiguity caused by Linnaeus himself when he wasthatcollectedbyColonnafromAndria.Morerecently,wehave
altered his definition ofA. caputserpentisbetween 1758 and 1767 found that this might not be correct. In five original copies of
(Brunton&Cocks(1967: 295);RulingoftheCommission, 1968). Colonna's Purpura (1616c) held by the University of Naples
Under Terehratula. 201., Linnaeus listed three separate illustra- 'Federico IF (two in the Library, and three in the Department of
tions.Thefirst,'Column,purp.22.f.1',referstotheupperleftfigure BiologiaVegetaleandtheBotanicalGarden)andinanotheroriginal
ofasmooth,stronglyfoldedbrachiopod.Thesecondrefersto 'List, copy held in the Botany Library ofthe Natural History Museum,
angl. 240.t. 8. f. 46', which is a non-plicate Jurassic shell from London, the twofigures at the top ofpage 22 are marked with '4'
England (Lister, 1678). The third reference, which confirms that (upperleft),and'F(upperright).Thesesmallnumbersarenotshown
Linnaeusdidindeedintendtheupperleftbrachiopodtobethetype in the two published reproductions ofColonna's plate (Dollfus &
ofterehratula,isunmistakablythesame(redrawn)Colonnaillustra- Dautzenberg, 1932; Muir-Wood, 1955), norin the 1675 edition of
tionreproducedinafigurebyKlein(1753)(Fig.2herein).Buckman Colonna'sPurpuraeditedbyMajor.
(1907: 528) pointed out that Lister's figure did not match the The following section attempts to clarify the problems we have
description given by Linnaeus, and that the Colonna-Klein figure encountered.
'must be taken as the holotype, which, in fact, has been the usual
practice'. 1 ColonnadescribedfourConchaeinthetextonpages23-24and
Linnaeus included the central figure in Colonna's plate in his markedthemwithaRomanorArabicnotation.Ofthese,onlythe
'Hysterita. 203. Mus. Tess. 90. t.5.f.1,2,2. Column, purp. 22. f. 3? first (I) and the fourth (4) are figured on page 22. The four
Trilobos.'Thislarge,stronglyribbedandfoldedfossilrhynchonellid Conchaeare:
doesnotcorrespondtotheMus. Tess.illustradonlistedbyLinnaeus,
whichisan internal moldofSchizophoria(Bruntonetal., 1967). (i) ConcharariorAnomiaverticerostrato. I. Cap. XII ("I"on
ThetwoupperbrachiopodsintheColonnafigurearebothsmooth page 22 and 'F on page 23). The description of 'F corre-
terebratulid brachiopods with large open foramens and strongly spondstofigure 'F(page22,topright).
delineatedgrowthbands.Acrossbothdrawingsisthecaption: 'Con- (ii) minor!('2'attheedgeofthemarginofline29onpage23).
cha anomia vertice rostrato', andbeneaththe left figure, although ThisspecimenisnotfiguredbyColonna.
probablyrelatingtothecentralbrachiopodontheplate,istheword (iii) Altera Neptunia maiorIII. imbricata. Cap. XIII ('III' on
AhAlentaoedmreaida(aCnsoohntelhclehraw)i.rtOahrniaoptrihcAeanlaodbjmeaiackae'n)vt.erpAtaligcteeho2r3ou,sgthirnattPhoue.rrIep.uiCrsaapn,.othXseIcaIclh(eaoprotn'errtahriees (iv) pCpaaoggneech224a2),AaTnnhodimsi'saIpVe'IcVio.mfempnaariggseinno2et4)fu.ingdTuorhseead.bdyeCsaCcpro.ilpotXniInoVan..o(f'4''ToVn
illustration,thebrachiopodontheupperrightisatnaturalsize{icon correspondstofigure '4'(page22,topleft).
magnitudinemaequatonpage23 inPurpura). 2. The first shell, (that is the smooth, unfolded specimen on the
upperrightonpage22)comesfromAndria.Thethirdcomesfrom
Nettuno(50kmsouthofRome).Thesecondandthefourthcome
from the Museum of Ferrante Imperato in Naples and their
CONCHAANOMIAVERTICE ROSTRATO
provenanceisnotgiven.
3. ColonnathoughtthefourConchae{icthefiguredandunfigured
Some ofthe problems which have made it difficult forearlierand specimens described on pp. 23-24) were similar to each other.
present authors to define Terehratula arise from the lackofcorre- Indeed,intheindexonp.41 threeoftheseareincludedundera
spondencebetweenthefiguresonpage22ofColonna,andthetext single name 'Conchaanomiaquae, rariorverticerostratoPlin.
.
86 D.E.LEE,C.H.C.BRUNTON,E.TADDEIRUGGIERO,M.CALDERAANDO.SIMONE
descript.23.icon.22.Alteraimbricata.24.alteramargineimdosa. upperright) certainly came from Pliocene strata nearAndria, and
ibid. &25'.Thetwofiguresonthetopofpage22arejoinedunder that Colonna himself regarded these specimens as similar to one
thesamecaption 'Conchaanomiaverticerostrato' another.
Secondly, six original copies ofColonna (1616c) that we have
The figure in the centre ofpage 22, described on pages 24-25 as consultedhave the numbers:'r, "4'and T' written beside the three
'Concha altera Anomia striata [trilobos] rarior. I. Cap. XV.', is brachiopodsfiguredonpage22.Thesenumbersarenotincludedin
consideredasanothershellanditsnumerationstartsagainfrom T". Major'sedition.Linnaeus(1758, 1767)andBrocchi(1814),ifaware
Muchoftheconfusionovertheidentityofthetwofiguresonthe of these numbers, used different, modern numbering (i.e. f.1 for
topofpage22stemsfromthefactthatColonnadescribedtheshell Colonna'sspecimen4ontheupperleft),andMajor(1675)andLee
placedontheupperrightfirst,andonafollowingpagediscussedthe &Brunton(1998)consideredthatColonna'sdescriptiononpage23
shellportrayedontheupperleft,usingadifferentpracticefromthat referredtothefigureontheupperleft.
which became well established in the following centuries. The Thirdly,evenifthespecimenofColonna's(1616c; 22)upperleft
confusion increased further when later authors assigned to these figure is from an unknown locality, it was filled with white, loose
figures two numbers that Colonna had never employed. Thus sediment(Colonna, 1616c:24),anditispossiblethatitwascollected
Linnaeus (1758), following modern convention, designated the fromthesamePliocenecalcarenitesatAndria.
stronglyfoldedspecimenontheupperleftasf. 1 (ie.,figure 1)and FromacloseexaminationoftheColonnawoodcut,itseemslikely
thatontheupperrightasf2(ie.,figure2).Brocchi(1814),andthe thathisspecimenwassomewhatdeformed.Ithasalarge,openfora-
presentauthors,unfillately,havedonethesame. men,andtwostronglydevelopedplicae/foldsonthedorsalvalvethat
The problem ofdeducing which shell description accompanied beginatanearlystageofgrowth,andwouldhaveresultedinastrong
whichillustrationwascompoundedinasecondeditionofColonna's sulciplication(;;;a/-g;HCM«(yo5a)oftheanteriorcommissure.Theartist
work, with the associated text from the earlier edition subdivided (?Colonna himself) may have exaggerated the depth ofthe folds,
into numbered paragraphs, which was reprinted posthumously in althoughthedepictionoftheotherbrachiopodsontheplateseemtobe
1675byJ.D.Major.Inthisedition,theredrawnfigureofthestrongly faithfultoreality.Noundeformedspecimenscollectedbytheauthors
foldedspecimenfromtheupperleftofpage 22 in the 1616edition havefoldsasstronglydevelopedasthosedepictedinthewoodcut.
(i.e.AnomiaterebratulaofLinnaeus)appearedonpage32withinthe FromthemanyspecimensofTerebratulacollectedbytheauthors
textreferringtotheAndrialocality. Similarly, in the Dictionarium from Colonna's Andria locality and from elsewhere in Italy, it is
this figure was noted by Major as that described in Cap. 12, of apparent that the brachiopods in any fossil assemblage/population
Colonna(1616c: 23). varyconsiderablyinthedegreeoffoldingandmayberectimarginate
Acomparison ofthe three figures (Fig. 2 herein) shows that the tobiplicateorsulciplicate.Thus,bothspecimenslabelledbyColonna
KleinfigurelistedbyLinnaeus(Klein, 1753;Tab.NostraXI.n.74), asConchaanomiaverticerostratoarespeciesofTerebratula(sensu
wasredrawnfromthislateredition(Major, 1675)(Fab.Columnaede lato). and given the wide variation in populations of Neogene
Purp. Cap. XII$.3. pag. 32 [Klein 1753,p. 171-2]). Terebratula, might be conspecific. Certainly, the specimens of
Dollfus & Dautzenberg (1932) correctly interpreted the refer- Terebratula terebratula figured by us in this paper (Figs 6-9) fall
ences toMajor'sfigures, but addedtotheconfusion, bygivingthe somewhere in the middle of the two short-looped brachiopods
samecitationforboththeupperrightandupperleftfigures:ie.'p.22, illustrated in Colonna's woodcut. The specimen of Pliocene
fig.l (2e)'. These authors also assumed that the caption 'Altera' TerebratulaterebratulafromMonte Mario selectedandfiguredby
pertainedtotheupperleftfigure,whereasitundoubtedlyreferstothe Buckman(1907)andillustratedinthe 1965 Treatise,iscrushedand
figureinthecenter('Altera[trilobos]').becausethecaptions inall deformed in a similar manner. In the Natural History Museum,
sevenfiguresinColonna'sPurpura(pp. 13,16,20,22,27,30,33)are London,thereareseveralcollectionsofTerebratulafromthissame
placedover,andneverunder,therelevantfigures. horizon at Monte Mario, near Rome. These specimens, which are
almostcertainlyconspecific, varyfromsmallrectimarginate(juve-
nile) specimens (labelled T. depressa) to large rectimarginate
DISCUSSIONAND COMMENTS individuals (named T. grandis), to examples with moderate
sulciplication(labelledT. ampullaorT terebratula).
Thus, our selection ofa neotype from a locality described and
Thisdiscovery,asourmanuscriptwasalmostreadyforsubmission collected by Colonnaclosely follows the recommendations ofthe
and after the ICZN had approved the selection of a neotype for Code.
Terebratula terebratula (Linnaeus), raises some issueswhichneed ItshouldbenotedthatMuir-Wood's(1955:fig.2)captionforher
furtherdiscussion.Ifindeedtheprovenanceofthespecimenselected reproductionoftheoriginalColonnafigureismisleading.Thecaption
by Linnaeus asthetypeoiAnomiaterebratulaisunknownandthe reads 'Reproduction ofearly drawings of Brachiopods figured as
specimenislost,thenthebasisforthespecies,andconsequentlythe 'Conchaanomia',andtakenfromFabioColonna'sfirsteditionofde
genus,familyandsuperfamilywouldremain uncertain. Purpura, 1616,p.22.ThefirstfigureisofaspecimenfromMte.Mario,
However,sinceLee&Brunton(1998)havealreadynominateda nearRome,andisprobablyofTertiary(?Pliocene)age;theothertwo
neotype from Colonna's locality nearAndria, the neotype locality figuresmayrepresentJurassicforms'. However, asshownhere,the
now becomes the type locality for the species, regardless of the firstfigureisfromanunknownlocality,whilethesecondspecimenis
locality ofthe original Colonnabrachiopod(International Code of ofPlioceneageandcamefromnearAndria,notRome.
Zoological NomenclatureArticle 75f).
Is it possible then to determine the provenance of the original
specimen ofColonna selected by Linnaeus as the type ofAnomia TYPE LOCALITY OF TEREBRATULA
terebratula! TEREBRATULA
Firstly, it is obvious that Colonna's caption: 'Concha anomia
vertice rostrato' applies to both ofthe brachiopods figured on the Colonna(1616c: 23)describedthelocalityfromwhichhecollected
upperrightandupperleftofpage22,thefirstofwhich(thatonthe his specimen of Terebratula shown on the top right ofhis figure
THECENOZOICBRACHIOPODTEREBRATULA 87
Fig.3 ReproductionofPacichelli'slate 17"'centurymapofAndria.Northisatthebottomofthefigure,andthechurchofSantaMariadeiMiracoli(2in
thekey)iswestofthecityneartherighthandedgeofthemap.
(Colonna, 1616:22)asfollows: 'Wefoundthisshellfullofthewhite oysters,scatteredpectinids,echinoidsandbrachiopods(Fig.5).The
sediment[~tophaceaconcretione']onwhichthatwholeslopingarea upper2mthickbedisawell-cemented,veryfossiliferouscalcarenite
orhillismade.Thisisconstitutednotsomuchofloosesediments,as withadiscontinuousoysterbednearitsbase.FossilsincludeOstrea,
offragmentsofvariousshellsandunbrokenshellstoo.Wecollected Chlaniys,internalmouldsofbivalvessuchasVeneridae,echinoids,
this one and others in the small valley or ditch a little below the calcareousalgaeandbrachiopods.
ChurchofD.MariadeAndria,whichissituatedonemileoutsidethe On macrofossil (Caldara, 1987; Caldara & Gissi, 1993) and
city'(seetheAppendixforatranslationofColonna'spp.23and24). microfossil(TaddeiRuggiero, 1996)evidence,thelowerpartofthe
InthemoderntownofAndriathenamesoffivechurchesinclude CalcarenitediGravinaFormationislocallyMiddletoUpperPliocene,
theword'Maria',andtofindwhichofthemwastheonereferredto orpossiblyUpperPliocenetoLowerPleistoceneinage.Notraceof
byColonnaitwasnecessarytoconsultPacichelli'slateseventeenth acool-temperaturePleistocenemacrofossilfaunawasfoundduring
centurymapofAndria(Fig.3).ThechurchesofSantaMariaNova(4 ourvisit.
onFig.3)andSantaMariadeiMiracoli(2onFig.3)arebothsituated Thus,thetypelocalityfortheneotypeofTerebratulaterebratida
outsidethetowntothewest(westisontherighthandsideofFig.3), (Linnaeus) is adjacent to the church ofSanta Maria dei Miracoli.
butonlythelatterchurchisbuiltdirectlyonthe'tophaceaconcretione' about2kmwestofAndria,Puglia,ItalyintheCalcarenitediGravina
(= Calcarenite di Gravina Formation). On either side of a small Formation,ofUpperPlioceneage.Thespecimenselectedasneotype
natural valley (now dry) adjacenttothe churchofSantaMariadei comes from beneath a small overhang about 200 m north of the
Miracoli l-3mhighcliffsofwhite, well-cementedcalcareniteout- church. The outcrops are difficult to access, and involve crossing
cropsporadically(Fig.4),andspecimensofTerebratidaarescattered privateproperty.
throughoutthecalcareniteshowingthatthislocalityisundoubtedly
theonevisitedbyColonna.Thebrachiopodsarenotuncommon,but
areoftenfragileand/orbroken. SYSTEMATICDESCRIPTIONS
The basement rocks of the region around Andria are Lower
Cretaceous in age. Overlyingthese with angularunconformity are
25 to 30m of coarse-grained highly fossiliferous marine OrderTerebratulidaMoore. 1952
SuperfamilyTerebratuloideaSchuchert, 1913
biocalcarenites and calcirudites ofthe Calcarenite di Gravina For-
FamilyTerebratulidaeGray. 1840
mation which is widespread in this area. In the vicinity of the
Madonna d'Andria church, the sequence consists of 2m of fine,
bioturbated,massivecalcareniteslackingmacrofossils.Abovethisis Diagnosis. Medium to large, ventribiconvex, rectimarginate to
a coarse, bioturbated calcarenite up to 4m thick which includes uniplicate orbiplicate, rarely unisulcate or sulciplicate, sinooth or
D.E.LEE,C.H.C.BRUNTON,E.TADDEIRUGGIERO,M.CALDERAANDO.SIMONE
Fig.4 OutcropsoftheCalcarenitediGravinaFormation,fromwhichtheneotypewascollected,inthesmallvalleynorthofthechurch.
THECENOZOICBRACHIOPODTEREBRATULA
a
>-<c7
Wm
^ ^10
2 19 3 e» 4 <^6 9 ES3ii
Fig.5 Locationmap,geologicalmapandstratigraphiccolumnfortheAndriaregion,Puglia,Italy.A,alluvialdeposits(Holocene);B,terracedalluvial
deposits(Pleistocene);C,terracedmarinedeposits(Pleistocene);D,CalcarenitediGravinaFormation(UpperPliocene);E,CalcarediBariFormation
(LowerCretaceous):F,Colonna'slocality;G,outcropfromwhichtheneotypeofTerebratulawascollected(seestratigraphiccolumn). 1,Pectinidae;2,
Veneridae;3,gastropods;4,brachiopods;5,echinoids;6,calcareousalgae;7,oysters;8,bioturbations;9,mudpebbles; 10,finecalcarenite; 11,coarse
calcarenite.
with fine radial capillae; loop short, triangular; outer hinge plates uncertainties surrounding the identity, age andtype locality ofthe
usuallyconcaveorflat,commonlyattachedtodorsaledgeofcrural Colonnaspecimen.TheColonnaworkisrare,andnotranslationhas
base,innerhingeplatesrarelydeveloped. hithertobeenavailable. Secondly,there wasdoubtastothecorrect
type speciesforthegenus.
Gaetani& Sacca(1985),inapaperdealingwith systematicsand
SubfamilyTerebratulinaeGray, 1840 shell structure ofbrachiopods ofMiocene - Pleistocene age from
Genus TEREBRATULA Muller, 1776: 249 southernItaly,commentedontheproblemofrecognisingTrerei'rarw/a.
Theyconcludedthatthereweretwovalidspecies:Terebratulasinuosa
Diagnosis. Medium to large, subpentagonal to broadly oval, (Brocchi)andT.calabraSeguenzawhichwererestrictedtotheUpper
smooth; anterior commissure rectimarginate to uniplicate or Miocene and Pliocene respectively. Cooper (1983) identified large
sulciplicate; beak short, erect; foramen large, symphytium partly sulciplicatespecimensofPlioceneagefromMonteMario,Rome,asT.
visible.Pediclecollarshort;cardinalprocessflatandsemielliptical ampulla. Otherauthors (Taddei Ruggiero, 1986, 1990, 1994, 1996)
to a thickened boss; outer hinge plates narrow or lacking; crural have identified large, shellbed-forming Pleistocene specimens as T.
processes may be long; loop short, broadly triangular; transverse scillae Seguenza. Until variation within large populations of the
bandnarrow,formingalowarch. speciesplacedinTerebratulahasbeenstudied,weareabletorecognise
only three valid species ofTerebratula - T terebratula (Linnaeus,
Typespecies. AnomiaterebratulaLinnaeus, 1758,bythesubse-
1758),7^am/7H//fl(Brocchi, 1814)and7iir/Z/oeSeguenza, 1871.
quentdesignationofLamarck(1799: 89).
Geographicrange. Italy, Sicily, Malta, Spain,Algeria. Terebratula terebratula (Linnaeus, 1758) Figs 6-9
Stratigraphicrange. Miocene-EarlyPleistocene. \15%AnomiaterebratulaLinnaeus: 703.
Remarks. The great majority of the thousand or more specific Synonyms
names attributed to Terebratula have long been accommodated in 1.Anomiasinuosa Brocchi, 1814: 468, is an objective synonym
othergenera.However,althoughT.terebratulaistheoldestavailable because Brocchi gave no figure, but referred to "Column. 22,
name for the medium to large, smooth, short-looped Miocene - f.r,whichistheholotypeofAnomiaterebratula.
Pleistocene terebratulids from Italy and the circum-Mediterranean 2. Terebratulacalabra Seguenza, 1871: 64
region,ithasrarelybeenusedintheliteratureorinidentificationof 3. TerebratulacostaeSeguenza., 1871:67;TaddeiRuggiero, 1994:
specimensforthereasonsoutlinedabove.Thisisduemainlytothe 206.
90 D.E.LEE.C.H.C.BRUNTON.E.TADDEIRUGGIERO.M.CALDERAANDO.SIMONE
Figs6-9 Neotypeand3topotypesofTerebratulaterehiatida(Linnaeus)fromaColonnalocalityatMadonnadeiMiracoli,Andria,Italy.6a-c,dorsal,
lateralandanteriorviewsofneotype.NHMBG152(length55.4mm);7a-c,dorsal,ventralandanteriorviewsoftopotype.NHMBG194(length
50.8mm);8a,b,ventralandanteriorviewsoftopotype,NHMBG195.showingM-shapedanteriorcommissure;9a,b,ventralviewsofloopoftopotype.
NHMBGI96.Allfiguresnaturalsize,exceptFig.9b.