Balanoidea (Crustacea, Cirripedia) from the Upper Marine Molasse (Lower Miocene) of the Bern area,...

Balanoidea(Crustacea,Cirripedia)fromtheUpperMarineMolasse(EarlyMiocene)oftheBernarea,Switzerland

rené-Pierre Carriol &ursula Menkveld-Gfeller

abstraCt Contrib.Nat.Hist.13:3–17.Within the framework of an integral study of burdigalian (late Early Miocene) fau-

nas from the upper Marine Molasse in the bern area (switzerland), the balanoids

housed in the collections of the natural History Museum bern have been studied.

specimens are assigned to the subfamily of archaeobalaninae and to the balanid

subfamilies Concavinae and Megabalaninae. only in a single case has it been pos-

sible to identify material at the specific level, namely in Chesaconcavus gurlarnensis

Carriol & Schneider, 2008.

rÉsuMÉbalanoidea (Crustacea, Cirripedia) de la Molasse marine supérieure (Miocène infé-

rieur) des environs de berne, suisse – Dans le contexte d'une étude intégrale de la

faune burdigalienne de la Molasse marine supérieure des environs de berne (suisse)

les balanoides conservés dans les collections du Musée d'Histoire naturelle de berne

ont été étudiés. les spécimens ont été attribués à la sous-famille des archaeobala-

ninae ainsi qu'à deux sous-familles de balanidae, les Concavinae et les Megabala-

ninae. une seule détermination a été possible au niveau spécifique, celle de Chesa-

concavus gurlarnensis Carriol & Schneider, 2008.

Keywords:Cirripedia,Taxonomy,Molasse,Miocene,Burdigalian,Switzerland

Mots-clefs:Cirripedia,Taxinomie,Molasse,Miocène,Burdigalien,Suisse

Contributions to natural History No.13: 3–17 ∙20April2010

4 René-PierreCarriol&UrsulaMenkveld-Gfeller

introduction

In a number of papers on the Molasse fauna near Bern there are records ofbalanoidcirripedes(Studer1825;vonFischer-Ooster1861;Bachmann1867;Mayer1872;Kissling1890;Rutsch1928).TheUpperMarineMolasseof thestudyarea(Fig.1) inthevicinityofBern(Switzerland)wasdepositedduringtheEarlyMioceneintheperialpineMolasseBasin(Pfister&Wegmüller1994),inatransitionalareabetweenthewesternParatethys(RhôneBasin)andthecentralParatethysintheeast.

StrataofBurdigalianage in thisareaareassigned to theSenseBeds (atthebase),theBelpbergBedsandtheNiedermattBeds(atthetop)(seeGru-ner2001).

TheSenseBedscomprisehomogeneoussandstoneswithonly fewfossili-ferouslayers(Gruner2001).EversincethedaysofRitter(1742),thefossil-richBelpbergBedshaveremainedthefocusofpalaeontologicalandgeologicalwork.Studer(1825),Bachmann(1867)andKissling(1890)describedthefos-silfaunaaroundBern.TheworkofRutsch(1928)wasforalongtimethemostcomprehensive palaeontological and geological study of the Belpberg Beds.New data on sedimentological and stratigraphical aspects were supplied byKeller (1989) and Schoepfer (1989), and more recently, Pfister & Wegmüller(1994, 1998, 1999, 2000, 2001, 2007a, b) recorded the molluscan faunas,while Kroh & Menkveld-Gfeller (2006) documented the echinoids. Until now,balanidsfromthesedimentswerenotstudiedindetail.MostoftheBurdigalianBalanoideastudiedhereinoriginatefromtheBelpbergBeds,whereassomespecimenswerecollectedfromtheunderlyingSenseBeds.

For an overview of lithostratigraphy and palaeogeography of the SwissMolasse,referenceismadetoBerger&al.(2005a,b).Gruner(2001)assignedtheSenseBedstotheLowerBurdigalian, inviewof their lithostratigraphicposition(Schoepfer1989);theBelpbergandNiedermattbedsareassignedaMiddleBurdigalianage.ForadditionaldiscussionontheageoftheBelpbergBeds (and the"Helvetian",anobsolete termused for the time intervalstud-ied),wereferthereadertoHarzhauser&al.(2003).

Material studied

Withoneexception, thecollectionofBalanoidea fromtheBurdigalianUpperMarineMolasseoftheBernareahousedattheNaturalHistoryMuseumBern(NMBE)doesnotcompriseanycompletespecimens.Allareshells,withoutopercular plates, filled with an indurated sandstone matrix which cannot be

5EarlyMioceneBalanoideafromBern

A

Münsingen

Kiesen

5 km

Upper Freshwater Molasse (OSM)

Upper Marine Molasse (OMM)Belpberg Beds s.l.and Niedermatt Beds

Upper Marine Molasse (OMM)Sense Beds

Plateau Molasse:

basal thrust of the thrusted and folded Plateau Molasse

1-6 Localities

A = Oberbalm

studyarea

N

Fig. 1: simplified geological map of the study area, indicating the different localities which yielded the balanoidea described herein: – 1: Kräjeren near rüschegg; – 2: Hohburg/Hohburggraben (belp-berg); – 3: aarwald (belpberg); – 4: Cheergraben (belpberg); – 5: stockeren near bolligen; – 6: banti-ger near bolligen (modified after Pfister & Wegmüller 1994; Kellerhals & al. 1999).


prepared,precludingobservationof theshell interior.Thematerial iscon-tainedintheO.Hug,H.Haas,B.HostettlerandW.Bühlercollections,and,inpart, stems from excavations initiated by the NMBE at localities at the Belp-berg,attheBantigerandfromKräjerennearRüschegg(Fig.1).

systematic part

SubclassCirripediaBurmeister,1834

SuperorderThoracicaDarwin,1854OrderSessiliaLamarck,1818SuborderBalanomorphaPilsbry,1916

SuperfamilyBalanoideaLeach,1817FamilyArchaeobalanidaeNewman & Ross,1976

SubfamilyArchaeobalaninaeNewman & Ross,1976

Diagnosis(afterBuckeridge1983:p.82)Wallofeight,sixorfourcompartments;parietessolidorwithsingle,uniformrowoftubes; interlaminatefiguressimple;basiscalcareousormembranous,whenmembranous,wallsolid.

GenusActinobalanusMoroni,1967

Typespecies:Balanus (Hesperibalanus?) actinomorphusMoroniRuggieri,1952,byoriginaldesignation.

Plate 1

Fig. 1: Actinobalanus sp., nMbE a3230, belpberg, Cheergraben, belpberg beds, burdigalian, excava-tion nMbE 1983.Fig. 2: Actinobalanus sp., nMbE b8639, belpberg, Hohburggraben, belpberg beds, burdigalian, b. Hostettler Colln.Fig. 3: Notomegabalanus sp., nMbE b8553, belpberg, Hohburggraben, belpberg beds, burdigalian, b. Hostettler Colln.Fig. 4: Notomegabalanus sp., nMbE D2981, aarwald, belpberg, belpberg beds, burdigalian, b. Hostettler Colln.Fig. 5: Notomegabalanus sp., nMbE D741, rüschegg, Kräjeren, belpberg beds, burdigalian, H. Haas Colln.scale bars: 1 cm.


1

2

4

3

5

A

AB


Diagnosis (afterZullo&Perreault1989:p.2;withinbracketsarecharactersnotobservableinthematerialstudied):

Parietesmonolamellar,solid,smoothorcostate;parietalribsnotdenticu-late; radii solid, narrow to broad, disparietal or paraparietal (fine striationspresentonsuturaledgesofA. dolosus);basiscalcareous,tubiferous;[interiorof scutum and tergumusually rugose; scutal adductor ridge present;nocal-lusbetweenscutaladductorandarticularridges;scutaldepressormusclepitabsentorsmallanddeep,withoutcrests;tergalspurnarrowtobroad,short,usuallyacuminate,basallyroundedorsubtruncate,withspurfasciole].

Actinobalanus sp. (Pl.1,Figs.1,2)

Occurrence:Belpberg(CheergrabenandHohburg),CantonofBern,Switzerland.

Material:NMBE A3230: a group of shells on an internal mould of a tapetid bivalve;NMBEB8689:anothergroupofshellsonasimilarsubstrate.Oneoftheshellsretains opercular plates of which only the external side is visible, but poorpreservationprecludesassessmentoftaxonomiccharacters.

Measurements:NMBEB8689(Pl.1,Fig.2),opercularorificediameter(rostro-carinal);specimenA:2.6mm;specimenB:1.8mm.

Description:Wallsmall,lowconical,whitetolightbrown;parietessolid,externallyribbedontheupperpartandcostateonthe lower;orificewide, rhomboidal; radiinarrow;basisround,calcareous,tubiferous.Opercularplates,preservedinasinglespecimen,adhereclosely tomatrixobscuring their internal features,theirexternalsurfacebeingtooerodedtobeofuse.


FamilyBalanidaeLeach,1817

SubfamilyConcavinaeZullo,1992

Diagnosis (after Pitombo 2004, p. 264; within brackets are characters notobservableinthematerialstudied):

Wallofsixplates,generally large, tubiferous,withonerowof tubeswithor without transverse septa. Sutural edges of radii with prominent trans-verseteethbearingdenticlesonlowersidesonly;innerfacewithlongitudinalabutment near the margin of sheath; alae sometimes cleft; lateral marginof sheath extending over the adjacent ala. Basis tubiferous, multilayered(vesicular)atleastmarginally.[Scutumsometimeswithoutradialstriae,withaconspicuousadductorridge.Tergumwithspurshowinganabruptchangeinthedirectionofgrowthlineswithfurrowmarginsinfolded,basalmarginwithweakdepressormusclecrestslimitedtoborder.Secondmaxillawithanteriormarginofdistallobebearingsmoothacuminatesetae].

GenusChesaconcavusZullo,1992

Diagnosis(afterCarriol&Schneider2008,p.5;withinbracketsarecharactersnotobservableinthematerialstudied):

Alarsuturaledgebifid;[scutaexternallystriate,withtergalsegmentnarrow,andlateraldepressorridgeseparatedfromadductorridge;tergawithsimpleapex].

Typespecies:Balanus concavus chesapeakensisPilsbry,1916,byoriginaldesignation.

Chesaconcavus gurlarnensisCarriol & Schneider,2008 (Pl.2,Figs.2–4)

Chesaconcavus gurlarnensis Carriol&Schneider(2008):p.5,pls.3,4

Additionalsynonymy:Balanus miserLamarck, 1818andBalanus pustularisLamarck, 1818;Wine-

berger(1851):p.87(listofspecies).Balanus concavus Bronn, 1831; De Alessandri (1910): p. 121 (in part; not

smallform), pl.48,figs.2–7(non8).


Occurrence:Burdigalian: Gurlarn (type locality), near Passau (Lower Bavaria, southernGermany);Eggenburgregion(LowerAustria);Stockeren,Bantigerand?Belp-berg,CantonofBern(Switzerland).

Material:NMBEA1801:oneshell;NMBED2972:agroupofthreeshells;NMBEA7986:agroupofshells;NMBED2973: twoshells;NMBED2974: twoshells;NMBED2975:agroupofshells.

Measurements:NMBED2972(Pl.2,Fig.3),rostralheight:specimenA:48.9mm.NMBED2974(Pl.2,Fig.2),carinalheight:specimenA:65.9mm.

Description:Shellhighconical,moderatelylarge,parietesbeige,externallysmooth;orificelarge,subtrigonal;radiisolid,withveryobliquesummits;alaewithsuturaledgesbifidandwithlateralmarginsofsheathextendingoverthealae;sheathwithclearlyvisiblegrowthridges,andoccupyingupperone-thirdtoone-halfofinteriorofshellwall;parietaltubesbearingcloselyspacedtransverseseptain upper third; longitudinal primary parietal septa denticulate at their base;basisbearingasinglerowofseptatetubes,multilayeredmarginally.Scutaandterganotpreserved.

Discussion:IntheabsenceofopercularplatesthespecificattributionoftheseshellsofChesa-concavus isbasedontheresemblancewiththoseofC. gurlarnensis fromtheUpperBurdigalianUpperMarineMolasseofLowerBavaria,southernGermany.

Plate 2

Fig. 1: Megabalanus sp., nMbE a1800, stockeren, sense beds, burdigalian.Fig. 2: Chesaconcavus gurlarnensis, nMbE D2974, stockeren, sense beds, burdigalian.Fig. 3: Chesaconcavus gurlarnensis, nMbE D2972, bantiger, belpberg beds, burdigalian.Fig. 4: Chesaconcavus gurlarnensis, nMbE a1801, stockeren, sense beds, burdigalian.scale bar: 1 cm.


1

2

4

3

A

A

A


SubfamilyMegabalaninaeNewman,1979

Diagnosis (after Pitombo 2004, p. 265; within brackets are characters notobservableinthematerialstudied):

Wallofsixplates,generallylarge,wrinkledorlightlyribbed,tubiferous,withone major row of tubes lacking transverse septa in adults, except for Mega-balanusvinaceus (Darwin,1854)andFosterellaBuckeridge,1983whichhave minor tubes on inner and outer laminae, respectively. Radii tubiferous,sutural edges with prominent transverse septa bearing denticles on upperandlowerside,oronlowersideonly;innerfacewithalongitudinalabutmentnearthesheathmargin.Alanotcleft;lateralmarginofsheathextendingovertheadjacentala.Basis tubiferous,commonlymultilayered(vesicular). [Scu-tum normally witha conspicuous adductor ridge. Tergumwithspur showinganabruptchangeinthedirectionofgrowthlineswithfurrowmarginsinfolded,basalmarginwithweakdepressormusclecrests limited toborder.Secondmaxillawithanteriormarginofdistallobebearingsmoothacuminatesetae].

GenusMegabalanusHoek,1913

Diagnosis(emendedafterCarriol2005,p.123;withinbracketsarecharactersnotobservableinthematerialstudied):

Parieteswithonerowoftubes;primarydenticlesofsuturaledgesofradiiwithregularsecondarydenticlesonbothupperandlowersides;basisgener-allytubiferous.[Tergumusuallywithoutbeak].

Typespecies:Lepas tintinnabulumLinné,1758,byoriginaldesignation.

Megabalanussp.(Pl.2,Fig.1)

Occurrence:Stockeren,CantonofBern,Switzerland.

Material:NMBEA1800:agroupofshells.

Measurements:NMBEA1800(Pl.2,Fig.1),rostralheight:specimenA:29.2mm.


Description:Walltubuloconical,whitetocreamcoloured,withsmoothsurface;radiiwidewith horizontal summits; sutural edges of radii with well-developed primarydenticlesbearingsecondarydenticlesonupperandlowersides;parietespene-tratedby longitudinalpores;basiswithradiatingtubes.Scutaandterganotpreserved.

GenusNotomegabalanusNewman,1979

Diagnosis(afterCarriol2002,p.2;withinbracketsarecharactersnotobserv-ableinthematerialstudied):

Parieteswithonerowof tubes,suturaledgesof radiiwith irregularsec-ondarydenticlesonlowersideonly. [Tergumwithoutbeakandspurfurrowopen].

Typespecies:BalanusdecorusDarwin,1854,byoriginaldesignation.

Notomegabalanussp.(Pl.1,Figs.3–5)

Occurrence:?Belpberg, Belpberg (Aarwald, Hohburg, Hohburggraben, Cheergraben) andRüschegg(Kräjeren),CantonofBern,Switzerland.

Material:NMBE A2978: two shells; NMBE A2979: one shell; NMBE A3181: a groupof shells; NMBEA 3182: one shell; NMBE A3183: fragment of shell; NMBEB7913:agroupofshells;NMBEB8552:agroupofthreeshells;NMBEB8553:one shell; NMBE B8554: two shells; NMBE B8555: one shell on the internalmouldofagastropod;NMBED741:agroupofshells;NMBED2976:agroupofshells;NMBED2977:oneshellattachedtoanoystershell;NMBED2978:agroupofshells;NMBED2979:oneshell;NMBED2980:agroupofthreeshellsonaninternalmouldofagastropod;NMBED2981:oneshell;NMBED2982:a group of shells; NMBE D2983: one shell; NMBE D2984: one shell; NMBED2985:agroupofshells.

Measurements:NMBED741 (Pl.1, Fig. 5),rostralheight:specimenA:13.9mm;opercularori-


ficediameter(rostro-carinal):10.0mm.NMBEB8553,rostralheight:14.7mm;carinalheight:12.0mm;opercularorificediameter(rostro-carinal):10.0mm.

Description:Wall high conical, with smooth surface, red brown to light brown, parieteswith longitudinaldarkerstripes,penetratedby longitudinalpores;orificesmall, subdiamond-shaped; radii and alae rather wide, with horizontal sum-mits; sutural edges of radii with well-developed primary denticles bearingsecondarydenticlesonlowersideonly;basiswithradiatingtubes.Scutaandterganotpreserved.

acknowledgements

We wish to thank P. Loubry (Muséum national d'Histoire naturelle, Paris)for takingphotographs,M.Hohn(NaturhistorischesMuseumderBurgerge-meindeBern)fordrawingFigure1andourreferee,J.W.M.Jagt(Natuurhisto-rischMuseumMaastricht),forhiscarefulreviewofthemanuscript.

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addresses of the authors:

René-PierreCarriolDépartementHistoiredelaTerreMuséumnationald'HistoirenaturelleCasePostale3857rueCuvierF–75231ParisCedex05

E-mail:[email protected]

Dr.UrsulaMenkveld-GfellerNaturhistorischesMuseumderBurgergemeindeBernBernastrasse15CH–3005Bern.

E-mail:[email protected]

Balanoidea (Crustacea, Cirripedia) from the Upper Marine Molasse (Lower Miocene) of the Bern area,...

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