Differentiating glacio-eustacy and tectonics; a case studyinvolving dinoflagellate cysts from the Eocene±Oligocenetransition of the Pindos Foreland Basin (NW Greece)

F. J. C. Peeters1,2*, R. P. Hoek1, H. Brinkhuis1, M. Wilpshaar1, P. L. de Boer3, W. Krijgsman4 and J. E.Meulenkamp3

1Laboratorium voor Palaeobotanie en Palynologie, Universiteit Utrecht, Budapestlaan 4, 3584 CD Utrecht, The Netherlands,2Faculteit der Aardwetenschappen, Vrije Universiteit, De Boelelaan 1085, 1081 HV, Amsterdam, The Netherlands, 3Instituut voor

Aardwetenschappen, Universiteit Utrecht, Budapestlaan 4, 3584 CD, Utrecht, The Netherlands, 4Palaeomagnetisch Laboratorium,

Universiteit Utrecht, Budapestlaan 17, 3584 CD Utrecht, The Netherlands

Introduction

Changes in relative sea-level caused bytectonics and by glacio-eustacy areoften difficult to distinguish in the se-dimentary record. High-resolutionstratigraphy and detailed informationon the palaeoenvironment are essentialto discriminate between these possiblemechanisms. Traditionally, chronos-tratigraphic assessment and palaeoen-vir- onmental information, includingproxies for, e.g., relative sea-levelchange, bathymetry and sea surfacetemperature (SST), are generatedthrough (quantitative) micropalaeon-tology, notably the study of calcareousmicrofossils and associated stable iso-tope measurements (d18O, d13C). Intectonically active settings, however,this approach is often difficult, as thesignal becomes diluted due to a highsedimentation rate as a result of a highterrigenous influx. Furthermore, in-creased reworking and limited preser-vation notoriously hamper biochro-nostratigraphic interpretations insuch situations.Dinoflagellate cysts are very sensi-

tive to environmental changes, and arebeing successfully used in high-resolu-tion stratigraphy (cf. Wall et al., 1977;

papers in Head and Wrenn, 1992;Stover and Hardenbol, 1993; Brin-khuis, 1994; Versteegh, 1994; Zeven-boom, 1995). Dinoflagellates whichproduce organic-walled cysts, live par-ticularly in neritic environments andconsequently occur in marginal marinedeposits; in more offshore settings,however, the dinoflagellate signal gen-erally consists increasingly of trans-ported elements. This allows detailedcorrelations along the inshore±offshoretransect and estimates of the relativecontribution of marginal marine ele-ments. If also the proportion of terres-trial palynomorphs (sporomorphs) inoffshore settings is considered, a proxyfor relative sea level change can beestablished. In addition, the ratio be-tween typical low-and high-latitudedinoflagellate cysts has become a well-established tool in the reconstruction ofsea-surface temperature trends (e.g.Edwards et al., 1991; Edwards, 1992;De Vernal, 1994; Versteegh, 1994).In this paper we explore the potential

of quantitative palynological analysis,with an emphasis on dinoflagellatecysts as a tool to discriminate betweentectonically and glacio-eustatically dri-ven sea-level change in tectonically ac-tive settings. Prerequisites for suchassessment are (i) an interval in whichglacio-eustacy is accepted to play animportant role, (ii) an area for which arelatively high-resolution well cali-

brated dinoflagellate zonal frameworkis available for that interval, (iii) knowl-edge of the palaeoecology of dinofla-gellates (iv) an area which qualifies as`tectonically active' at that particulartime, and (v) independent calibration ofthe age assessments.These prerequisites are met in the

Eocene/Oligocene (E/O) transition inthe Pindos Foreland Basin (PFB) inEpirus (NW Greece). Glacio-eustati-cally induced sea-level changes are gen-erally acknowledged to occur in thisinterval (cf. Saunders et al., 1984;Milleret al., 1987; Kennett and Barker, 1990)while during the late Eocene and earlyOligocene, the Ionian Zone became thesite of a foreland basin developing infront of the Pindos thrust (Fig. 1). Thiscaused a pronounced shift in sedimen-tation from carbonates into a thickterrigenous-clastic-dominated succes-sion (Aubouin, 1959, 1965; IGSR,1966;Underhill, 1989). A detailed dino-flagellate zonal scheme of the UpperEocene to Lower Miocene for the cen-tral Mediterranean was recently estab-lished by Wilpshaar et al. (1996). Forthis scheme multiple sections from cen-tral, NE and NW Italy were used andcalibrated against calcareous planktonand palaeomagnetic stratigraphies on afirst-order basis. For the present study amagnetostratigraphic analysis was per-formed to cocalibrate the chronostrati-graphic interpretations.

*C 1998 Blackwell Science Ltd 245

ABSTRACTIn an attempt to discriminate between tectonically induced sea-level changes and glacio-eustacy, the Ekklissia and Arakthossections (Epirus, NW Greece) are examined, applying (dinocyst)palynology, sedimentology and magnetostratigraphy. Thesections, located in the Pindos Foreland Basin, both comprisethe transition from pelagic limestones to hemipelagic silty claysand turbidite sandstones, reflecting the onset of flyschsedimentation as a result of the Pindos thrust activity. Despitean overall tectonic overprint, relative changes of sea level can bereconstructed, using (i) continental/marine palynomorph ratios,

(ii) relative abundance of inshore and offshore dinoflagellatecysts, and (iii) taxa indicative of relatively cold and warm sea-surface temperature, that can be calibrated against the GlobalPolarity Time Scale (GPTS). Increased fluxes of marginal marineand continental palynomorphs coincide with colder periods on a`third-order' scale, which thus appear to be related to glacio-eustatic trends in sea-level. The larger scale is attributed to theincreasing effect of tectonics and acts on a `second-order scale'.

Terra Nova, 10, 245±249, 1998

AhedBhedChedDhedRef markerFig markerTable markerRef endRef start

*Correspondence: Tel: +31/(0)20-4447419;

Fax: +31/(0)20-6462457;

E-mail:peef@geo.vu.nl

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Materials and methods

The Epirus region in northwesternGreece (Fig. 1) is part of the NW±SE-trending Hellenides-Dinarid orogenicbelt and a classic area for the study offoreland basin development (cf. Au-bouin, 1959; Underhill, 1989). In theIonianZone (Fig. 1) of the PFB, pelagiccarbonates are overlain by a sequenceof terrigenous clastic deposits, oftenreferred to as `flysch' (Aubouin, 1959;Piper et al., 1978; Fleury, 1980). Twosections, the Arakthos and Ekklissiasection (Fig. 1, see also IGSR-IFP,1966), incorporating this conspicuouschange in lithology were selected forthis study (Fig. 2). The Pindos Zoneserved as themain source area of clasticsediments of the Ionian Zone. TheArakthos section is more proximal tothe thrust front than the one near Ekk-lissia. Both sections were sampled forpalynological and palaeomagnetic ana-lysis. The Ekklissia section used in thisstudy slightly differs from the one men-tioned in IGSR (1966). The section weused is located just North of the villageof Ekklissia and reflects the completerecord from limestone to flysch. Sam-ple processing and the quantitativeanalysis of palynological slides, followthe methods discussed in Brinkhuis(1994). Taxonomy of dinoflagellatecysts corresponds to that in Lentinand Williams (1993), Brinkhuis andBiffi (1993) andBrinkhuis (1994). Stan-

dard palaeomagnetic cores were drilledwith an electric drill. From the lime-stones of the lower part of the sectionsorientatedhandsampleswere taken.Allsamples were progressively demagne-tized, either by applying alternatingfields or stepwise thermal demagnetiza-tion. The natural remanent magnetiza-tion (NRM) was measured on ahigh-sensitivity 2G Enterprises cryo-genic magnetometer, using DC squids.

Sea-level and sea surface temperaturereconstructions

Traditionally the fraction of terrestrialpalynomorphs (sporomorphs) relativeto the total amountof palynomorphs, isused as a proxy for continental influ-ence. Some authors (e.g. Traverse andGinsburg, 1966) also use this ratio as anindication of relative sea-level change.In this paper the continental±marineproxy (CM) is expressed as the percen-tage of pollen and spores in the totalpalynomorph assemblage:

CM � C

C�M�1�

in which C=continental palyno-morphs (pollen and spores) andM=marine palynomorphs (dinofla-gellate cysts). Brinkhuis (1994) pro-posed a model for the distribution ofdinoflagellate cyst groups across theTrento (NE Italy) shelf during late Eo-cene to earlyOligocene times. Themod-el is based on known (Recent) andempirically established (palaeo)ecologi-cal information and distributionpatterns of quantitatively important di-noflagellate cyst groups. In order tointerpret the dinoflagellate cyst associa-tion, division into four ecologicalgroups was made, namely (i) taxa re-presenting restricted marine to innerneritic watermasses (RM), (ii) innerneritic taxa (IN), (iii) outer neritic taxa(ON), and (iv) oceanic taxa (OC) (Table1). Both the CM ratio and fluctuationsin the dinoflagellate cyst ecologicalgroups may be regarded to primarilyreflect the distance to the coast, whichmay be interpreted in terms of relativechanges in sea level. The reconstructionof trends in SST is based on relativeabundance of the oceanic genera Impa-gidinium and Nematosphaeropsis. Hereall Impagidinium spp. except for Impa-gidinium pallidum and Impagidinium ve-

lorum are considered to represent low-latitude, warm waters (cf. Brinkhuisand Biffi, 1993). The latter two taxaare regarded to represent the influencefrom colder watermasses. This assess-ment is based largely on present-daydistributions, and, in the case of Impa-gidinium velorum, on fossil evidence (cf.Bujak, 1984). All the taxa used for theSST-proxy range through the investi-gated interval. The following equationrepresenting relative SST (Zevenboom,1995), is used:

Results and discussion

Dinoflagellate cysts

All samples contain rich and relativelywell-preserved palynological associa-tions except for a few palynologicallybarren samples in the lower part of theEkklissia section. Most dinoflagellateassociations are well diversified andcharacterized by high percentages ofSpiniferites/Achomosphaera spp., Op-erculodinium spp., and Systematophoraspp. Furthermore, Impagidinium spp.and Nematosphaeropsis spp. are com-mon in the lower and middle parts ofthe sections. Deflandrea spp., System-atophora spp. and Homotryblium spp.,show higher percentages in the upperparts. Occasionally, Areosphaeridiumspp., Thalassiphora pelagica, and Tec-tatodinium spp. are abundant as well.The qualitative dinoflagellate cyst

distribution allows the recognition ofthe Lower Oligocene Adi, Rac and CinMediterranean dinocyst Biozones ofBrinkhuis and Biffi (1993), spanningparts of Chron C13r±C12r (34.0±32.5Ma, Berggren et al., 1995) in both sec-tions. At Arakthos, the uppermost Eo-cene Aal and lowermost Oligocene GseBiozone are represented also (Fig. 2).

Palaeomagnetism

The ChRMdirections show on averagea 488 clockwise tectonic rotation for theArakthos section, which is in goodagreementwith the results fromHornerand Freeman (1983) andKissel and Laj(1988) who report rotations of 45±508.This large rotation of the ChRM en-ables us to distinguish between a pri-mary normal direction and a normaloverprint. The presence of both normaland reversed directions confirms theprimary origin of the magnetization.Unfortunately, some samples show

246 *C 1998 Blackwell Science Ltd

Differentiating glacio-eustacy and tectonics . F.J.C. Peeters et al. Terra Nova, Vol 10, No. 5, 245±249. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Ekklissia

Arakthos

ALBANIA

GREECE

?CORFU

Pindos Zone

Pre-Apulian ZoneIonian Zone (PFB)Gavrovo Zone

40 km.0

studied section

20oE 21oE40oN

39oN

exte

rnal

midd

leint

erna

l

Pindos thrust

?

Fig. 1 Map of the Epirus area (NWGreece) and location of the investigatedsections in the Ionian Zone. During thelate Eocene to early Oligocene the IonianZone became the site of a foreland basin(Pindos ForelandBasin)which developedas a result of the Pindos thrust activity.

Paper 198 Disc

such a low intensity that no reliabledirections could be determined (e.g. atEkklissia below and above a normalpolarity interval). The ChRM direc-tions of the Arakthos section recordallow the recognition of a well-definednormal polarity interval, comprisingthe upper part of the Gse Zone, thewhole Adi Zone and the lower part ofthe Rac Zone (Fig. 2). Despite someuncertainties, the results from the Ekk-lissia section indicate a reversed polar-ity for the Cin Zone and a normalpolarity for the lower part of the RacZone (Fig. 2). Comparison of the bios-tratigraphic and magnetostratigraphicresults with the scheme of Wilpshaar etal. (1996), indicates that the normalpolarity interval in both sections repre-sents Chron C13n. Palaeomagneticdata thus confirm that both sections

are largely time-equivalent, and, more-over, they indicate that the transitionfrom limestones to terrigenous clasticdeposits is older at Arakthos than atEkklissia (Fig. 2).

Sea-surface temperature and sea-levelreconstructions

Trends in dinoflagellate cyst ratios andecological groups can be interpreted asfluctuations in continental influence,relative distance to the coast and sea-surface temperatures (Fig. 2). The low-er parts of the sections are dominatedby outer neritic and oceanic taxa, in-dented by small excursions of restrictedmarine and inner neritic ecologicalgroups, reflecting relatively outer neri-tic to open marine conditions for thelower part of both sections. An increas-

ing CMproxy, in both sections, mimicsthe shift from carbonates to terrigenousclastic sedimentation. Similarly, an up-ward increase of restricted marine andinner neritic dinoflagellate cysts indi-cates increasing influence of marginalmarine settings. On average, the per-centages of sporomorphs and marginalmarine dinoflagellate groups is higherat Arakthos than at Ekklissia, reflect-ing the more distal setting of the latter.At Arakthos, where the uppermost Eo-cene is identified, three distinct inter-vals are recorded in the CM curve; thefirst associated with the Aal and Gsezones (5 20% continental palyno-morphs), the second within the Adiand Rac and lower part of the CinZones (20±40% continental palyno-morphs), and the third within the CinZone (4 40% continental palyno-morphs). The distribution pattern ofthe oceanic ecological group shows asimilar three fold division.

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Terra Nova, Vol 10, No. 5, 245±249 Differentiating glacio-eustacy and tectonics . F.J.C. Peeters et al.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

SST �2� �I:velorum� I:pallidum� � � Nematospaeropsis spp:

�Impagidinium spp:� � Nematospaeropsis spp:�2�

Paleom

. sam

p.

Paly. s

amp.

12R

3N ?

c/(c+m)100% 100%

ecological groups

Adi

Rac

Cin

nodata

PM iPM r

DC zLit

holog

y

dinocyst SSTcoldwarm

mfs

SB

SB

Cin

Paly. s

amp.

ecological groupsPaleom

. sam

p.

NormalReversed

No diagnosticsignal

paleomagnetic data

restricted marine

outer neritic

inner neritic

oceanic

ecological groups

SMW

SMW

SMW

HST

HST

TST

Ekklissia(distal)

c/(c+m)100% 100%

30

35

40

45

50

5

10

15

20

25

13N

13R

12R

nodata

Arakthos(proximal)

Adi

Rac

Gse

Aal

PM i

PM rDC z

Thickn

ess [

m.]

Litho

logy

dinocyst SSTcoldwarm

SEQUENCESTRATIGRAPHIC

INTERPRETATION

34.0

33.4

32.7

32.5

TA

4.5

TA

4.4

TA

4.3

33.05

absoluteage [Ma]

PM i = paleomagnetic interpretation

PM r = paleomagnetic results

DC z = dinocyst zonationPaly. samp.= palynological samplePaleom. samp = paleomagnetic sample

samples and zonation

HST = highstand systems tractSMW= shelf-margin systems tractTST = transgressive systems tract

SB = sequence boundary

mfs = maximum flooding surface

sequence stratigraphic interpretation

TST

mfs

SB

SB

SB

mfs 33.52

limestone

turbidite sandstone

marl

lithology

Fig. 2 The Arakthos and Ekklissia sections, Continental/Marine (C/(C + M)) ratio, dinoflagellate eco-groups, dinocyst sea-surfacetemperature (SST) reconstruction, dinoflagellate-cyst zonal assignments, palaeomagnetic zonal assignments, sequence-stratigraphicinterpretation and correlation to the cycles of Haq et al. (1988).

Paper 198 Disc

Slightly below the Rac/Cin zonalboundary (middle Early Oligocene) thecombined outer neritic and oceanic di-noflagellate cyst group reach maximumvalues of& 80%of the total associationin both sections. Slightly higher in thesections, the composition of the dino-flagellate cyst association changesabruptly and becomes dominated byinner neritic taxa. This sudden changein composition occurs in both sectionsat the base of the Cin zone and isregarded to be isochronous. Remark-ably, this change in the dinoflagellatecyst assemblage seems tobe unrelated tothe terrestrial influx, since the CM ratioremains virtually constant in this inter-val. The upper part of both sections,assigned to the Cin Zone, is character-ized by high percentages of both thecombined restricted/inner neritic groupand sporomorphs, indicating that mar-ginal marine and continental influxesbecame overwhelmingly dominant.

Sequence stratigraphy

Following the sequence stratigraphicmodel of Vail et al. (1977) and applyingthe CM ratio and dinoflagellate datacombined with lithological changes, atentative subdivision into system tractsis proposed. Using also the availabledinoflagellate zonal scheme, threethird-order cycles can be recognized atArakthos (Fig. 2). Theoldest cycle startsat the base of the limestone channel andends just before the Adi±Rac transition.The top of the second cycle is drawn justabove the last pelagic limestone layer, atthe base of a thick succession of marls,silts and eventually, coarser grainedsandy turbidites.

At Ekklissia, the more condensedsection, changes are less clear, notablyin the lower part of the succession. Afirst sequence boundary is placed at thebase of the inception of more marlysedimentation in the Adi zone. Similarto Arakthos, the top of this cycle isdrawn just above the last pelagic lime-stone layer, at the base of the successionof marls, silts and coarser grained san-dy turbidites. At the Eocene/Oligocenetransitional interval, Haq et al. (1988)also recognize three third-order cycles(TA4.3, TA4.4 and TA4.5); that theywere recognized in central and north-east Italy with the use of dinoflagellatepalynology, and the sequence bound-aries occur in the same biozones (Brin-khuis and Biffi, 1993; Brinkhuis, 1994),stresses their inter-regional character.Although the cycles have been tectoni-cally enhanced (TA4.5) or subdued(TA4.3/TA4.4), their position seemsto agree with theHaq et al. (1988) chartfor the late Eocene and earlyOligocene.The concomitant overall change of

both lithology and palynology indi-cates that tectonics play an importantrole. However, it appears that withinthis larger scaled trend, shorter termedcyclicities occur, which, by using thebiochronostratigraphy, can be attribu-ted to the so-called `third order' cyclessensu Haq et al. (1988). The largerscaled trend may thus be attributed tosecond-order cyclicity.

Tectonics vs. glacio-eustacy

Between the sections the alternatingpattern of colder and warmer intervalscanbe reasonably correlated (Fig. 2). Inthe Ekklissia section, two intervals with

relatively cold SSTs are recognized, onein the Adi and one during lower part ofthe Cin biozone. At Arakthos, threeintervals with relatively cold SSTs arerecognized, namely besides the Adi andlower Cin biozones, but also during theAal biozone. Following the Vail et al.(1977) sequence stratigraphic model, itmay be anticipated that if a relationshipbetween third-order cyclicity and gla-cio-eustatic cycles exists, SSTvalues arelow during late HST and (early) LST.This appears to be the case in our studyof the Arakthos and Ekklissia sections.Hence, our records support an under-lying glacio-eustatic mechanism as thecause for sea-level lowering and in-creased transportation of marginalmarine elements, with a cyclicity of& 500 Kyr (third-order). The lowerorder trend does not show a relation-ship with cool/warm cycles, although itclearly has impact on the compositionof the palynological associations. Thislower, second-order trend with a cycli-city of 4 1 Myr, may thus be ascribedto large-scale compressional tectonics.

Conclusions

The transition from limestone sedimen-tation into siliciclastic sedimentation(flysch) in the Arakthos and Ekklissiasections from the Pindos Foreland Ba-sin occurred at 33.6±32.5Ma, followingthe timescale of Berggren et al. (1995),using dinoflagellate cyst zonation andmagnetostratigraphy. The lower part ofthe transition is characterized by rela-tively high abundances of outer neriticand oceanic taxa, whereas the upperpart is dominated by restricted and in-ner neritic marine taxa. In addition,relative percentages of terrestrial paly-nomorphs increases significantly upsec-tion. In both sections increased fluxes oftransported material, taken as indica-tive for sea-level fall coincide with rela-tively cold dinoflagellate-basedSSTs ona third-order scale sensu Haq et al.(1988). In both sections there seems tobenoobvious relationship betweenSSTtrends and the larger scale (second-or-der) trend as evidenced by the sporo-morph and inshore/offshore dinoflagel-late ratios. It is considered that thislarger scale trend is the expression ofthe increasing effects of tectonics (i.e.approach of the Pindos thrust front),while the smaller scale cyclicity corre-sponds to the third-order cycles of Haqet al. (1988). The chronostratigraphical

248 *C 1998 Blackwell Science Ltd

Differentiating glacio-eustacy and tectonics . F.J.C. Peeters et al. Terra Nova, Vol 10, No. 5, 245±249. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Table 1 Based on empirically established (palaeo) ecological information (Brinkhuis,1994), the dinoflagellate cyst association is divided into four ecological groups. Thesubdivision is on genus level, except for Spiniferites and Hystrychokolpoma whichwere subdivided up to species level dinoflagellate cyst ecological groups

Restricted Inner neritic

Eocladopyxis spp. Areoligera spp. Hystrichokolpoma spp. (pp.)

Homotryblium spp. Cordosphaeridium spp. Operculodinium spp.

Phthanoperidinium spp. Deflandrea spp. Schematophora spp.

Polysphaeridium spp. Glaphyrocysta spp. Spiniferites spp. (pp.)

Wetzeliella cpx. Heteraulacacysta spp. Systematophora spp.

oceanic outer neritic

Impagidinium spp. Ennadeacysta spp. Spiniferites spp. (pp.)

Nematosphaeropsis spp. Hemiplacophora spp. Stoveracysta spp.

Hystrichokolpoma spp. (pp.) Tectatodinium spp.

Reticulatosphaera spp. Thalassiphora spp.

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position of the sequences matches thosereported by these authors, and may beascribed to glacio-eustacy. Therefore, itis concluded that the trends and fluctua-tions in inshore/offshore ratios in thesections mirror the interference of cli-matically controlled (third-order) andtectonically induced (second-order)sea-level fluctuations. This study de-monstrates the potential of quan-titative dinoflagellate studies as a toolfor high-resolution stratigraphic andpalaeoenvironmental analysis.

Acknowledgements

Sponsoring of the AASP initiated `EPIRUSproject' by Phillips (USA), Norsk Hydro(Norway), Statoil (Norway), Unocal(USA), Elf-Aquitaine (France) and Amoco(USA) is gratefully acknowledged, as isadditional funding by the LPP Foundation(The Netherlands). This is publication no.990109 of the Netherlands Research Schoolof Sedimentary Geology. We thank H.Hooghiemstra and an anonymous reviewerfor their comments.

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Received 8 June 1998; revised versionaccepted 19 January 1999

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