SEPM Society For Sedimentary Geology

Sequence Stratigraphy, Biostratigraphy, and Taphonomy in Shallow Marine Environments
Author(s): Carlton E. Brett

Source: PALAIOS, Vol. 10, No. 6, Tenth Anniversary Theme Issue (Dec., 1995), pp. 597-616
Published by: SEPM Society for Sedimentary Geology
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TENTHANNIVERSARYTHEMEISSUE 597
597
Sequence Stratigraphy,
Biostratigraphy,
and Taphonomyin Shallow
Marine Environments
CARLTON E. BRETT
DepartmentofEarth and EnvironmentalSciences, UniversityofRochester,Rochester,N.Y. 14627
PALAIOS, 1995, V. 10, p. 597-616 marinefloodingsurfacesoccurat bases ofparasequences,

winnowedshell beds at parasequence tops. At a larger
Sequence stratigraphy providesan integratedframework scale, thin,sheet-likeskeletal deposits may formtrans-
withinwhichto examine historicalpatterns of paleon- gressivesystemstracts of thirdordersequences. Fossil
tological phenomena. Depositional sequences are the Lagerstdtten(typicallyinvolvingrapid burialand/oran-
stratigraphicrecordoffluctuationsofsea level and sed- oxia) are characteristicofthe transgressive to earlyhigh-
imentation,environmentalparameters that also exert stand interval.Althoughhigherbackgroundsedimenta-
critical controlson the distributionof shallow marine tionrates typifythe laterhighstand,fossileventbedsare
organisms, and thepreservationand accumulationoftheir commonlyless recognizabledue to dilutionand biotur-
skeletalremains.As such,stratigraphic paleontologyand bation effects.Close integrationofpaleontologicand se-
sequence stratigraphyshare multipleassociations. quence stratigraphicdata should fostera greatlyim-
Biostratigraphy is a criticaltoolforrelative age-dating proved understandingofbiases and relationshipsofbiotic
and correlationof depositionalsequences; in turn,sedi- and abiotic processes in the accumulationof the strati-
mentologicaland depth-relatedvariablesexerta primary graphicrecord.
controlon the occurrenceof zonally significantfossils.
The mostreadilydated portionsof manysequences are
offshorecondensedfacies. Correlationof a hierarchyof INTRODUCTION
discontinuity surfaces,however,permitsextensionofbio-
stratigraphicdatinginto less fossiliferous facies.In turn, Stratigraphicpaleontologyand sequence stratigraphy
minglingof fossils of distinct zones provides a key to sharemanyfundamental goalsand approaches.Bothfields
recognitionofcondensedsections,and truncationofzones seekto establisha detailedchronostratigraphic framework
points to significantunconformitiessuch as sequence withinwhichto interpretprocessesin earthhistory;both
boundaries.The combinationof refinedgraphicbiostra- are concernedwiththe genesisof particulartypesofsed-
tigraphy,cycle-basedecostratigraphy, and sequencestra- imentarydepositsand in theeffects offluctuating sea level
tigraphy,sequence biostratigraphy, will ultimatelyper- and sedimentationpatterns.Fossils are the primarytools
mit extremelyprecise stratigraphiccorrelationand dat- of chronostratigaphy and biostratigraphy enables intere-
ing of marinestrata. gionalcorrelationof depositionalsequences;however,se-
Taphonomicattributesoffossilassemblagesalso relate quence stratigraphy may,in turn,permitmuchmorede-
closely to sequence stratigraphy.Taphofaciesvarypre- tailed resolutionof time withinbiozones. The sequence
dictablyin depositionalsequences because ofthe depen- paradigmmakes a numberof predictionsthat mayaid in
dence offossilpreservationupon rates of burialand en- interpreting particulartypesof fossilaccumulations,in-
vironmentalenergy.A spectrumof preservationshould cludingextraordinary biotas or Lagerstitten.Conversely,
exist fromhighlycorroded,fragmentedremains,typical as sedimentary particles,fossilsmaybe sensitiveindicators
of erosivelowstandor early transgressiveconditions,to of the dynamicsof sedimentationand depositionalenvi-
intact multielementskeletons, characteristicof rapid ronmentsthat may aid in recognitionof sequences and
backgroundsedimentationduringmid-highstand progra- theirboundaries.As such, close integrationof paleonto-
dation;geochemicallystable lags ofcorrodedfossils(e.g., logicand sequence stratigraphic data shouldyieldimpor-
bone beds) typifymanyhighlycondensedsections.Skel- tant insightsintoearthhistory(Kauffmanand Sageman,
etal accumulationsdevelop duringintervalsof lowsedi- 1992;Sageman,1992;Gomezand Fernandez-Lopez,1994).
ment input. Sediment-starvedaccumulationsoverlying The model of sequence stratigraphy, originallydevel-
? 1995,SEPM (SocietyforSedimentary
Copyright Geology) 0883-1351/95/0010-0597/$3.00

598 BRETT
oped fromseismicrecordsofcontinentalshelfsedimentary tributionand preservationof ancientorganisms.Manyof

deposits,has foundbroad applicabilityin manymarine theseaspectsremainincompletely exploredto thepresent
and nonmarinesedimentarysettings(Sloss, 1963; Vail et timebutthesequencestratigraphic modelprovidesa pow-
al., 1977a,b, 1991;Haq et al., 1987;Wilguset al., 1988and erfulheuristictoolforinvestigating patternin lifehistory.
papers therein;Galloway,1989; Haq, 1991; Macdonald, Simultaneouswith the developmentof the sequence
1991). Sequence stratigraphyprovides an integrated stratigraphic paradigm,paleontologistshave begunto de-
framework withinwhichto examineaspects of local and velop a numberof integrativemodels that relateto bio-
globalsea levelchange,tectonicsand subsidence,and pat- stratigraphic resolution(Shaw, 1964;Sadler, 1981;Schin-
ternsof sedimentation.Moreover,the modelis hierarchi- del, 1982; Edwards, 1984, 1989) and taphonomicbiases
cal, encompassinga numberof scales of processas rep- and processes (Behrensmeyerand Kidwell, 1985, 1993;
resentedbysmallscale shallowing-upward parasequences, Norris,1986; Brett and Baird, 1986; Speyer and Brett,
sets of parasequencesand largeand smallscale unconfor- 1986,1988,1991;Allisonand Briggs,1991;Kidwell1991a,b;
mity-bounded bundlesor depositionalsequences. Einsele et al., 1991). Many of these conceptsprovidepo-
Sequences are the stratigraphicrecordof major (typi- tentiallystronglinkageswith sequence stratigraphy, al-
cally0.5 to 5 millionyear;Vail et al., 1991) fallsand rises though,the relationshipsamong these disciplineshave
of relativesea level; theirlocal developmentreflectsthe generallynot been exploredin depth.
interplayof eustasy,tectonicsubsidence,and sedimen- In the presentpaper,I will reviewthese linkages,with
tation.Ideally,sequences are divisibleinto threephases, respectto biostratigraphy, taphonomy, and thestratinomy
the lowstand(or shelfmargin),transgressive, and high- of skeletalaccumulations.Examples are taken fromcon-
stand systemstracts,each delimitedbelow by a discon- tinentalshelf,forelandbasin, and epicontinentalsea de-
tinuity, thesequence boundary,transgressive surface,and posits,whichdominatethe onshorestratigraphic record
maximumflooding(also termedmaximumstarvationor of shallowmarineenvironments. The focusof thispaper
downlap) surface,respectively (Van Wagoneret al., 1988; is primarilyon 1) the controlthat episodic stratigraphic
Vail et al., 1991; Fig. 1, herein).Systemstracts,in turn, accumulationexertson patternsofpreservation ofmarine
are comprisedof smallerscale parasequences(ca. 20,000 fossilsand 2) the uses of paleontologicaldata forthe in-
to 400,000year) shallowingupwardcycles. terpretation ofdepositionalsequences.Space does notper-
In practice,forshallow shelfsettings,lowstand(shelf mit fulldiscussionof the ecologicaland evolutionary re-
margin)systemstractsare typicallyrestrictedor absent, sponsesoforganismsto sequenceforming processes;these
and depositionalsequences consistof geneticallyrelated topicswill be exploredin a forthcoming paper.
transgressiveand highstanddeposits, bounded by ero-
sional lowstandunconformities (Baum and Vail, 1988). SEQUENCE STRATIGRAPHY
Sequence boundariesare thusmostcommonly unitedwith AND BIOSTRATIGRAPHY
transgressivesurfacesand overlainby thin lag deposits
reworkedduringinitialtransgression(see Holland, 1993; The mostdirectrelationshipbetweensequence stratig-
Elder et al., 1994). Transgressivesystemstractsare typi- raphyand paleontologylies withinthe generalrealmof
callythinto stratigraphically condensed,retrogradational biostratigraphy (Haq, 1991;Loutitet al., 1988,1991;Ver-
(upward deepening) successions that onlap sequence teuil and Norris,1992; Posamentierand Goodman,1992;
boundaries.They are bounded at the top by a thirdim- Armentrout, 1992). The connectionis obvious;in orderto
portantdiscontinuitysurface,the surfaceof maximum testthenotionthatsequencesare globaland synchronous,
flooding, or surfaceofmaximumstarvation(Van Wagoner independentdata on age relationshipsmust be utilized.
et al., 1988; Baum and Vail, 1988). Highlycondensedde- Fossil zonationhas been recognizedas the keyto relative
posits both above and below this surfacereflectminimal age datingand correlationsince the early1800s and this
sedimentaccumulationduringthe intervalwhen rate of continuesto be trueto the presentday,althoughbiostra-
sea level rise is at a maximum.In manysequences it is tigraphyhas undergonesomethingof a revolutionin its
convenientto subdividethe highstandinto early(aggra- ownright.Fossil rangedata, and firstand last appearance
dational)and late (progradational)systemstracts;as noted datums(FADs and LADs, respectively), particularly when
in this paper, a fourthtype of discontinuity, the mid- coupled into the more sophisticatedgraphiccorrelation
highstanderosionsurfaceand associated condensedbed techniquesthathave been developedwithinthe last three
("precursorbed") mayprovidean unambiguousboundary decades, providean increasingly refinedbiostratigraphic
betweenthese phases (Brettand Baird, in pressa). framework forsedimentaryrocks (Shaw, 1964; Edwards,
The sequence stratigraphic paradigmmakes a number 1984, 1989). Most claims forthe correlatability of sedi-
of predictionsabout stratigraphicpatternand its rela- mentarysequencesultimatelyhingeuponbiostratigraphic
tionshipto sea level,subsidence,and sedimentationpro- correlations(Vail et al., 1977a, b; Haq et al., 1987; Ross
cesses. On the basis of sequence stratigraphy furtherde- and Ross, 1988; see Hallam, 1992,forsummaries).There
ductionscan be made about the distribution ofbothlith- are cases of discrepancieswhichbringforthcriticalques-
ological and paleontologicalaspects of strata.Sequences tionsabouttheefficacy ofsequencestratigraphy as a global
recordfluctuationsof a numberof parameters,such as correlationmethod,as wellas some ofthe assumptionsof
relativesea level and sedimentationrates,whichare po- biostratigraphy (Hubbard,1988; Hallam, 1989,1992) and
tentiallyof criticalimportancein governing the local dis- Miall (1991) has notedthatbiostratigraphic zonationmay

SEQUENCE STRATIGRAPHYIN PALEONTOLOGY 599
not have sufficient

resolutionto test forsynchroneity of ficationofotherwisecrypticsequenceboundariesorpermit
somekeysurfaces.Nonetheless, in general,physically
based determinationof theirrelativemagnitude.For example,
sequence and eventstratigraphy have matchedquite fa- Sagemanand Johnson(1985) wereable to identify a major
vorablywithtraditionalbiostratigraphic zonation. sequence boundarywithinoffshoremudrocksby docu-
mentingprogressiveshorewardtruncationof ammonite
Sequence Biostratigraphy biozonesin the Cretaceousof the WesternInterior.Here
erosionwas demonstrablymore severein uprampareas.
Processes controllingbiostratigraphic range data and Sageman et al. (1995) used the widespreaddistribution of
thosethatproducesedimentary sequencesare notentirely thisunconformity to argueforeustatic,as opposedto local
independent.Sequence boundingunconformities and un- tectoniccontrolon the sea level lowstand.
conformities associatedwithmaximumfloodingmay im- Obviously,apparentrangesoffossilsmayvarygeograph-
pose artificialtruncationson the rangesof variousfossil icallywithina sequence and should not be taken at face
taxa. In particular,MacLeod (1991) and Holland (1992, value withoutfurther scrutiny. Nonetheless,it is also quite
1994, 1995) argue that some of the apparent blocks of likelythatmanyoriginations and extinctionsoforganisms
faunal stability(such as those used to recognizestages) are relatedin real biologicalwaysto eventsthatproduced
may be an artifactof episodic stratigraphy. The coinci- sequence boundariesor otherunconformities withinse-
dence of firstand last appearance datumswithsequence quences (e.g.,Dockery,1986). Major dropsand risesin sea
boundingunconformities in some instancescertainlycan level or concurrent climaticor oxygenation effectsmaybe
be attributedto incompletenessof the recordassociated criticalcontrolsin determining the patternoflifehistory.
witherosionand nondepositionat thesesurfaces.Possibly, Major risesin sea levelmayconnectformally isolatedbio-
a staggering offirstand last appearancesis maskedin this geographicareas permitting therapidspreadofplanktonic
way,makingthe fossilrecordappear moreepisodicthan marinelarvae acrosswide areas encompassingseveralba-
was the truepatternof evolutionaryhistory.Such an ar- sins.For example,Hallam (1983) demonstrated thatEarly
tifactis particularlyto be suspectedin cases whereinthe Jurassictransgressions permittedcommunication between
boundingunconformities of sequences representa large faunas of Europe and westernNorthAmerica.Likewise,
fractionor even moretimethan that enclosedwithinthe the stressassociated withrapid sea level fall or rise and
preservedsedimentaryrocks (Ross and Ross, 1994; Hol- deoxygenation ofmajorportionsofthewatercolumnmay
land, 1995). produce local extinctionsand thus producethe typesof
In a landmarkstudy,Holland (1995) used computer changesthatareassociatedwithzonalorstagelevelbound-
simulationofFADs and LADs to demonstratethatstrati- aries(see forexample,Hallam,1992,p. 186;Dockery,1986).
graphicrangesof fossilspecies may be highlybiased rel- Moreover,certainportionsofsedimentary sequencesare
ativeto theoriginalpatterns,as a resultofsamplingerrors, predictablymoreimportantthanothersin biostratigraph-
facies changes,species' ecologicalattributesand episodic ic study.Notably,condensedsections,typicallythin,fos-
accumulationof sediments(Fig. 1). In particular,species siliferouslimestonesand organic-rich darkshalesthatform
rangeswithindepositionalsequencesare predictedto dis- in offshoreareas of sequences duringtimes of relatively
play a stronginfluenceof gaps and abruptfaciesshifts. rapid sea level rise and marineflooding,have provenex-
Althoughspecies originationsand extinctionswere pro- tremelycriticalin thedevelopmentofbiostratigraphic cor-
grammedto occurrandomlythroughtime,Holland's sim- relations(Loutitet al., 1988,1991;Posamentierand Good-
ulations showed a strongclusteringof FADs and LADs man, 1992). These condensedbeds are typicallyenriched
above sequence boundariesand major floodingsurfaces in biostratigraphically importantfossilsand may permit
(Fig. 1). Both themigrationoffaciesin sedimentary cycles datingof corresponding less condensedand less fossilif-
(parasequences)and the absence of a stratigraphic record erous continentalshelfor shallowmarinesediments.For
at sequence boundariesand maximumfloodingsurfaces example,condensed,black"core shales" in mid-continent
mayseverelytruncatetheobservedrangesofspecies.Hol- Pennsylvaniancyclothemsyield biostratigraphically di-
land suggeststhatmanyapparentlyabruptor punctuated agnosticconodontassemblagescriticalforcorrelationof
patternsof species distributionmay be artificially pro- the cyclothems(Heckel,1977,1986). In turn,commingling
duced, or at least sharpened,by the discontinuousrecord of fossilsof different biozones providescriticalevidence
of facies.This bias can be reducedby studyingfossildis- forlong-termor biostratigraphic condensation(Fiirsich,
tributionpatternson a regionalbasisto minimizetheprob- 1971, 1978; Fiirsichand Aberhan,1990; Kidwell,1991a;
lem oflocal faciescontrols.True pulses oforigination and Gomez and Fernandez-Lopez,1994).
extinctionmaybe discriminated fromartifactual"spikes" Detailed sequence stratigraphy offersthe potentialof
iftheydo notoccurimmediately above sequencebounding refining correlations,at leastwithinlocal basins,byseveral
unconformities or maximumfloodingsurfaces,or if they ordersofmagnitudeoverthosewhichhavebeenpreviously
occursynchronously in manyfacies.Even wherethespikes establishedon the basis of biostratigraphy; the combina-
are located at sequence discontinuitiestheymay be rec- tion of detailed physicalsequence stratigraphy and bio-
ognizable as real if theygreatlyexceed modelledvalues stratigraphy has been termedsequence biostratigraphy
forclusteringof FADs or LADs assumingrandomorigi- (Loutitet al., 1991;Posamentierand Goodman,1992).The
nationand extinction,as in Holland's models. recognitionof widespread floodingsurfaces bounding
Conversely,biozone truncationsmay aid in the identi- parasequencespermitsa levelofcorrelational precisionon

600 BRETT
FirstOccurrences WaterDepth(m) LastOccurrences

1
FIGURE -Relationship between numbers of simulated
first
and lastappearance datums (FAD,LAD)andrelative waterdepthintwoidealized
sequences.Notedivision
depositional ofthesequencesintosmallscale shallowing upwardcycles(parasequences)and systemstracts;LST
= lowstandsystemstract;TST = transgressive systemstract;HST = highstand systemstract;sb = sequenceboundary; ts = transgressive
surface;mfs= maximum floodingsurface(orsurfaceofmaximum absenceofstippling
starvation); withinwaterdepthcurveindicateslackof
depositionduringthelowstand.InthismodelbyS. M. Holland(1995)each of 1000 taxa had an equal probability oforiginatingor becoming
within
extinct each ofeighty 50-thousand yeartimesteps.Dashedvertical linesrepresentthemaximum number offirstorlastapearancethat
wouldbe expectedat 0.001 probability ifall FADs and LADswererandomly distributed.Notethatsignificantly
non-random spikesof FADs
occurabovesequenceboundaries andTST boundaries; and non-random spikesofLADsoccurbelowsequenceboundaries andTST flooding
surfaces.These spikesare strictly offaciesdiscontinuities
artifacts andtimegaps intherecordofsequences.Bracketedintervals, markedat
margins bydash patternindicatepositionsofcondensedsectionsthatare enriched inzonallyimportantfossils.Figureslightlymodifiedfrom
Holland(1995); reprinted
bypermission ofPaleontologicalSociety.
theorderofa fewhundredsofthousandsofyears(Goodwin The most detailed use of ecostratigraphy as a tool for

and Anderson,1985;Loutitet al., 1991).Whereevensmall- identifying and correlatingsmall scale sea level fluctua-
erscale sedimentary rhythms displaya nestedhierarchical tionsis that of Cisne and Rabe (1978). Using supposedly
patternwithinparasequences (e.g., fivedecimeterscale isochronoustransectsprovided by K-bentonitesin the
carbonate-shalerhythmsper parasequence), Milanko- Middle Ordovicianof New York State (but,see Goldman
vitch-bandorbitalforcingmaybe inferred(Fischer,1986; et al., 1994,forcritiqueofthebentonitecorrelations), Cisne
DeDoer and Smith,1994). It may be possibleto use this and Rabe establishedgradientsof faunalchangeon a pa-
small scale cyclicityto inferratherpreciselythe timedu- leoslope. Quantitativeindices of fossilassociation(ordi-
rationsofparticularsedimentary packagesdownto 20,000 nation scores) werethen used to recognizeand correlate
year scales (Gale, 1989; House, 1985a; Elder et al., 1994; cyclesofdepth-related faunalchangeamongstratigraphic
DeBoer and Smith,1994,and references therein). sections,a procedurereferredto as coenocorrelation.In
subsequent studies Cisne and his colleagues have been
and Sequence Stratigraphy successfulin correlating minordepthfluctuations, perhaps
Ecostratigraphy
amountingto no morethana fewmetersofwater,recorded
Finally,it should be noted that a relatedapproachto in Ordovicianstrata(see particularly, Cisne et. al., 1984).
paleontologicalcorrelationhas been developedin thepast Once suchcyclicity is recognizable,
depth-related biofacies
severaldecades. Referredto as ecostratigraphy, it makes cycles become apparent even in seeminglymonotonous
use of the bathymetric distributionof benthicorganism deep waterfacies.Moreover,abruptjuxtapositionsofdis-
associationsor "communities"and the observationthat parate biofacies(or ordinationscores) pointto the pres-
these associationsappear to track cycles of relativesea enceofdiscontinuities, includingsequenceboundariesand
level fluctuation(see Ziegleret al., 1968; Boucot, 1975; surfacesof maximumstarvation.Thus, this methodpro-
Johnson,1987).For example,Johnsonet al. (1985) utilized vides a veryimportantecostratigraphic tool forsequence
detailedcyclesofrelativesea level,inferred partiallyfrom stratigraphy.
patternsof brachiopodcommunitychange,to correlate Overall, a refinedgraphic-basedbiostratigraphy and
Lower SilurianrocksbetweenNorthAmericaand China. ecostratigraphy, coupled with detailed sequence stratig-

raphymayproducean unprecedentedlevelofprecisionin bonate has the opportunity to accreteand formconcre-

correlationand in interpretation of absolute magnitudes tions,typicallynucleatedaroundorganicmaterialor cal-
of elapsed geologictime (Loutit et al., 1991; Armentrout,cium carbonateshells (see Waage, 1964; Elder, 1987,p.
1992). In turn,highprecisiontimeresolutionmaypermit 28). These horizons,althoughgeneratedbyunusualtaph-
muchbetterassessmentof ratesof bioticprocesses. onomicconditions,mayprovideextremely usefulmarkers
foreventstratigraphy.
Epiboles and Sequence Stratigraphy Ecological epiboles or "acme zones" (Buckman,1902)
representunusual abundances of normallyrare species.
Epiboles are thin stratigraphicintervalscharacterized They are confinedto thinintervalsbut maybe widespread
by the anomalousabundanceofspeciesthatare normally and even faciescross-cutting to a limitedextent.Factors
rareor absent (Buckman,1902; Brettand Baird, 1990,in responsibleforrapidproliferation ofotherwiseuncommon
press b). Epiboles wereoriginallyconceivedto represent formsare generallynot apparentfromthe stratigraphic
the hemerasor "acmes" of species undergoinga normal record.Nonetheless,sequence stratigraphy may provide
"lifespan"includinggradual buildup to a peak, followed clues as to the explanationofthese"bursts."Observation
bydecline(Buckman,1902).However,thisis an outmoded ofPaleozoicfaunasin theAppalachianbasinindicatesthat
and overlyrestrictive viewpoint.In fact,epiboles,as broad- such proliferations are most commonlyassociated with
lydefinedherein,representan empiricalphenomenonthat transgressive systemstracts;epiboles of a particularspe-
mayarise fromseveralfundamentally different processes. cies may recurin analogousparts of sedimentarycycles,
Epiboleshaverecently beenclassifiedgenetically intothree but not in each successivecycle. Perhaps unusual envi-
broad categories:taphonomic,ecological,and incursion ronmentalconditionsdevelop temporarilyin association
epiboles (see Brettet. al., 1990; Brettand Baird,in press withsedimentstarvationduringsea levelrise.These may
b). intermittently favora normallyrare,stenotopicspecies.
Taphonomic epiboles are an artifactof preservation. In contrastthe heavier sedimentationassociated with
Fragileskeletalremains,such as thoseof starfishand cri- highstandsand lowstandsmayfavorgeneralizedeurytopic
noids,are onlypreservableas intactspecimensundercon- organisms.
ditionsof exceptionalburialin an absence oflaterdistur- Likewiseincursionepiboles reflectthe abruptappear-
bance. Because these require geologicaccidents in for- ance of a normallyabsent species (as opposed to whole
mationone mightexpectthattaphonomicepiboleswould faunas) in a thinbut laterallyextensivestratigraphic in-
be quite erraticin occurrence.Nevertheless,conditions terval. They appear to representthe influx,temporary
whichare conduciveto appropriateburialand lack of re- proliferation, and abrupt extermination of "exotic" spe-
workingoccurcyclically.They are particularly character- cies. These speciesappear temporarily withina particular
istic of marginalenvironments of manyorganismswhich sedimentary basin as a resultofincursionoflarvaeorfree
are too deep to be disturbedbyfrequentstormwaves,but swimmingadults. Evidently,these organismswere only
stillshallowenoughto be occasionallyinundatedbypulses viable forrelativelyshortspans of geologictime (a few
of rapid storm-generated sedimentation.Hence, tapho- hundredsor thousandsofyears)beforebeinglocallyerad-
nomic epiboles may occur in predictableparts of sedi- icated. Episodic immigration eventsare particularlytyp-
mentarysequences. For example, infrequentpulses of ical of nektonicformssuch as ammonoids,and benthic
muddysedimentwhichflushedintothebasinduringtimes formswithplanktoniclarvae. Notable examples include
of relativedeepeningin the transgressivesystemstract widespreadammonoidincursionsthat are closelyassoci-
providedthe agent forabrupt destructionand burial of ated with markerbeds in the Jurassicof Germany(see
attached crinoidassemblages (Hagdorn, 1985). Because Bayer et al., 1985). Such incursionsmay reflectunusual
these weredevelopedwithina deepening-upward succes- sedimentaryconditions;thus, the developmentof wide-
siontheprobabilityoflaterdisturbancedecreasedthrough spread shell gravelsduringtransgressions in the Jurassic
timeas the sea bottomgraduallywas deepenedbelowthe basin seem to have permittedthe influxof hard substrate
zone of persistentwave winnowing. encrustersas well as certain nektobenthicammonoids
Anotherinstanceof taphonomicepiboles,of consider- (Bayer and McGhee, 1985; Bayer et al., 1985). Similarly,
able biostratigraphic significance, consistsof horizonsof Hagdorn(1982,1985) demonstrated thatwidespreadbeds
wellpreservedaragoniticfossils.These skeletonstypically of brachiopodsand crinoidstypicallyoccurat the caps of
occuras highlydeformedinternalmolds,butundercertain shallowing-up carbonatecycles(parasequences)withinthe
circumstancestheymay be well preservedas a resultof Triassic MuschelkalkLimestoneof Germany.Cementa-
earlymineralization, such as pyritizationor the develop- tion of the substrateassociatedwithmarinefloodingsur-
mentofearlydiageneticcarbonateconcretions (e.g.,Waage, faces developed hardgroundswhichpermittedextensive
1964). Once again,theseoccurpredictablyin sedimentary colonizationby species that requiredhard substratesfor
sequences. Conditionsparticularlyconduciveto the de- attachment. These specieswerenormally restricted
toshoal
velopmentof earlydiageneticnodules,forexample,typi- areas on the marginsofthe Tethyssea.
cally occur in the upper parts of parasequencesand in Changesin circulationpatternsand climaticameliora-
condensedsectionsare largersequences.Rapid burialmay tion associated withsea level rise may be the proximate
be followedby periodsofextensiveslowdownin sedimen- cause ofimmigration episodes.Kauffman(1986) attributes
tation and/orwinnowingduringwhichtimecalciumcar- the appearanceof a diversefaunawithina concretionary
I ?

602 BRETT
limestonein the Upper CretaceousGranerosShale to the imentarycyclesand thusin sequences.Hence,taphofacies,

influxof a warmsubtropicalwatermass intothe Western expressedas quantifiableindices of preservationalattri-
InteriorBasin. The lattereventis closelyassociatedwith butes in fossils,should show predictablerelationshipsto
peak eustatichighstand;and thusthe limestoneprobably positioninsedimentary sequencesand parasequences(Fig.
representsa condensedsection. Changes in water tem- 2).
peratureand biogeographicfactorsmay be importantin Skeletal preservation, on the whole,may be predicted
this incursion. to be poor in lowstandand earlytransgressive portionsof
The occurrenceofmultipleincursionepibolesofpartic- sequences. During these phases skeletonsaccumulate in
ular ammonoidgenerain the Appalachianbasin has been depositsthatrepresenttypicallyrathershallowwater,and
associatedwithmaximalhighstandsrepresentedbyblack commonly"highenergy"conditionsunderwhichratesof
shale tongues(see Kloc, 1986; Brett and Baird, in press skeletaldestructionwillbe high.Moreover,sedimentation
b). Hence, epibolescommonlyoccurin appropriatefacies rates in shelfor uprampareas will tend to be low during
followingmajorhighstands,recordedas condensedinter- sea level lowstandsas a resultof dynamicwinnowing and
vals ofdeep waterfacies,suchas blackshaletongues.Such bypass of fine grainedsediment.During transgressions
incursionsmay representthe temporaryconnectionbe- sedimentstarvationwill characterizeboth deeper water
tweennormallyisolatedbiogeographicprovinces.Connec- areas and much of the shallowshelf(Posamentieret al.,
tionsmightbe establishedduringmaximumflooding events. 1988; Posamentierand Vail, 1988; Loutit et al., 1988).
At such times,barriersbetweennormallyisolatedbasins Hence,skeletallag depositsdevelopedin shelfareaswithin
mightbe breached,allowingthe influxof pelagic adults these intervalsare typicallycomposedof highlybroken,
and larvalforms(Eicherand Diner,1985;Kauffman, 1986). abradedcoquinas (Loutitet al., 1988;Baum and Vail,1988;
Both ecologicaland incursionepibolesare the recordof Holland, 1993). Conversely,sedimentationrates during
short-livedbiotic "events" and thus theyhave utilityas lowstandmaybe highin downramporoffshelf areas,where
local biostratigraphicmarkers.Considerablymore study lowstandturbiditefansor wedgesaccumulate.The rapid
is requiredto fullydocumentthe relationshipsof these burialmayfavorpreservation ofpelagicmicrofossils;
how-
"marker horizons" to sequence stratigraphy.However, ever,benthicmacrofossilstend to be sparse in these de-
preliminaryobservationsuggestsassociationswith later posits due to dilutioneffectsand the adverselivingcon-
transgressiveand earlyhighstandphases. The abruptdis- ditionsin areas of highsedimentationand turbidity.
appearance of epibole taxa is enigmaticbut may reflect At maximummarineflooding,sedimentationrates in
abruptdeteriorationof keyenvironmental factorsduring offshoreareas should be at a minimum.As a resultof
rapid sea level change. relativelyrapid sea level rise,mostterrigenoussediments
will be impoundedin the nearshorealluvial areas, while
SEQUENCE STRATIGRAPHY manypartsof the sea bottomwillfallbelowthe euphotic
AND TAPHONOMY zone of importantcarbonateproduction.Consequently,
organismskeletonswillbe exposedforprolongedintervals
A second obviouslinkexistsbetweensequence stratig- oftimeon thesea floor(Loutitet al., 1988).Althoughthese
raphy and preservationmodes of fossils.As fossilsare skeletalremainsmayaccumulateunderlowerenergycon-
sedimentaryparticles(Seilacher,1973),theyprovidesen- ditionsthanthoseduringearlytransgressive phases,they
sitive gauges of many processes,particularlythose that may also be subjectto intensebio-geochemicalcorrosion
relateto dynamicsof sedimentationand geochemistry of (Fig. 2). Hence,thinlag depositsare characterizedbygeo-
the near surfacesediments(Kidwell and Behrensmeyer, chemicallystable particles,includingthe resistantcalcite
1993). Models of taphofaciesattemptto relateconsistent biocrystalsof echinodermplates,or phosphaticskeletons
patternsof fossilpreservation, such as the proportionof such as conodontelementsand/orvertebratebone (e.g.,
articulatedskeletons,fragmentation, abrasion,corrosion, bone beds, see Aepler and Reif, 1971; Seilacher,1982a;
bioerosion,as well as early diagenesisof fossilsto envi- Reif,1982;Kidwell,1989;Baird and Brett,1991).Evidence
ronmentalparameterssuch as turbulenceand sedimen- ofabrasionand physicalbreakagemaybe relativelylower
tation rate (Speyer and Brett,1986, 1988; Norris,1986; than in earlytransgressive deposits.However,microbor-
Davies et al., 1990; Meyeret al., 1989; Staffand Powell, ings,corrosionpitting,and otherphenomenaindicativeof
1990). biogeochemicaldissolutionprocessesmay be muchin ev-
It is criticalto distinguishbetweenfossilassemblages idence.Althoughmostcondensedbeds developunderlow
whichaccumulatebynearlyinstantaneousburialprocesses energy,offshore marineconditions,theystillmaypossess
and thosein any particularfacieswhichrepresent"back- evidence for physicalerosion as well as dissolution.In
groundconditions"(Speyer and Brett,1991). For back- carbonates,corrosivewaters developed duringtimes of
groundtypeaccumulations,fossilpreservation potentially earlyhighstand,low oxygenconditionsmay lead to dis-
can provideveryimportantinformation on ratesofburial solutionofoldercarbonates.Withinsiliciclasticsediments
and extent of processingby waves, currents,and other erosionis commonlyevidentat early highstandbound-
turbulentagencies. In turn,these parametersare con- aries.Deep stormwaves,bottomflowing contourcurrents,
trolledby aspects of relativewaterdepth,extentof sedi- and/or internal waves, propagated along water mass
mentinputand starvationand geochemistry (oxygenation, boundariesin stratifiedbasinsmayall be important agents
salinity,and alkalinity),whichvarypredictability in sed- of deep submarineerosionwhichservesto formlag de-

posits of chemicallyand mechanicallyresistantparticles A low

suchas thepyriticbeds reportedbyBaird and Brett(1986, ,.*v...:. t' : TST
........ ... ...
1991). Underappropriatecircumstances, beds ofreworked

sedimentary phosphorite(Baird, 1978;Loutitet al., 1988)
t^^TST= . ... .':'.'. ,
or pyrite,includingfossilsteinkerns,may also occur at

theseintervals.The mostimportantfactorresponsiblefor ..'-::--- LHS;SB/TS
5 & III r
r-tt
productionof such verythin geochemicallag depositsis scale S ) sr 7t LHS
sedimentstarvation.In the absence of a renewedsupply * ;
-7. -: -7
(cm)
40
0
a)
L .......
-T - scale
ofsediments,evengentlecurrents, whichserveto winnow .::~:i-!:---i:-
:- _
-20 uo 3 t
Y
and removesomefractionofpreviouslydepositedmaterial, I
ySB s-7.:v-._-..- [IM^^y'
-7-7- - rOP
may induce net erosionof sedimenton the sea floor. ;';--- PBOI V -
II .t{ T PB
Duringearlyhighstandphases preservationof skeletal EHS ? * EHS
materialsmaybe surprisingly poor (Fig. 2). The low rates 1+ 2

7
of burialin condensedportionsof sectionsare commonly 1 ~2 ~3 E 4 E; 5 E 6-E
recognizablein the developmentofpavementsofdisartic-
ulated and commonlyeven fragmented skeletalmaterial. Biostrotinomy Diogenesis
The exact agentsof fragmentation remainunclear,but it Disort Reori Froam Abros Tophofocies Sk diss
-I-'I-, ,
.... -I
- - Corros Erly dio
is likelythatthe processis aided by geochemicalsolution A A A A I v A HARD
GROUND
and thinningof shells.Even minorbottomcurrentsmay A A A V
then providesufficient V 11
energyto shatterthe weakened
C/PY
AV
shells. Best and Kidwell (1995, and pers. comm.) have A A III
foundthat the proportionof fragmentedshells may ac- V
A V-V IVA V-V C
tually be higherin deeper than shallowersettingsin a A-v
AV A
modernsiliciclasticshelf environment.They inferthat V V V IVB ?PH
much of this shell breakageprobablyresultedfrompre- T V V V PY

dation (pers.comm.). A A-A A VI A A--A
Early solutionof shells may also be an importantde-
structiveagentdependingupon the geochemistry of bot- A A V VII A--V A-V
tom and interstitialwaters (Aller,1982). In some cases, V low very high A
particularly thoseassociatedwithhighorganicproductiv- B V
very low high A
ity,low pH may be developed in the upper sediment,
partlyas a resultof oxidationof monosulfidesor pyrite FIGURE2-Predictedvariation oftaphonomic attributes andidealized
and the developmentof sulfuricacid. This processmay taphofacies forshallowing upwardcycles(parasequences)undercon-
ditionsof lowand highratesof siliciclastic sedimentation; notethat
produce substantialdissolutionof skeletalmaterialand becauseof sediment starvation, taphofacies
will commonlylead to the destructionof mostor all ara- VII transgression-associated
maybe rare.A) Generalized taphofaciesframework; columna,
goniticshells. Many mollusksmay be preservedonly as stratigraphic patternof upward-shallowing cyclewithlow ratesof
thin plasticallydeformedperiostracalfilmsor foils(Sei- sediment input, underwhichtaphofacies VI,IV,(rarely II)andI should
lacheret al., 1985). The plasticdeformation indicatesthat appear,inascendingorder;columnb,stratigraphic pattern forupward-
dissolutionoccurredquite early,priorto earlyphases of shallowing cyclewithhighratesof sedimentinput,underwhichta-
sedimentcompaction.Under appropriateconditionsof sides phofaciesVII,V, III,and I mayappear.Stratigraphic columnsalong
oftaphofacies chartrepresent shallowing successions(earlyto
moderatebackgroundsedimentationand anoxic,non-sul- late overlain bytransgressive systemstracts;patterned
highstands),
fidicsediment,however,earlydiageneticpyritemayinfill afterDevoniansequencesofmixedcarbonate-calcareous shalefacies
skeletalvoids priorto dissolutionof shells,thus yielding inwesternNewYork(leftcolumn)and centralNewYork,mudstone-
well preservedinternalmolds (Brettand Baird, 1986;see sandstonecoarseningupwardsequences (right column);thetapho-
herein). facies indicated along each sideof the middle chartwouldcharacterize
Fig. 7, levelsoftheadjacentstratigraphic columns;lithologic
Conditionsfavoring unalteredpreservation areprobably corresponding coral-
best developedduringtimesof mid highstandin the de- symbols keyedalongbase ofchartinclude:1) rugose-tabulate
richcalcareousshaleto limestone; 2) crinoidal wacke-to packstone;
positionofmarinesedimentary sequences(Fig. 2). During 3) blackshale;4) graymudstone; 5) graysilty mudstone; 6) finegrained
thisintervaloftime,largeareas ofshelfand epicontinental sandstone;7) calcareoussandstoneto sandylimestone withbrachio-
seas will be floodedto depths sufficient that theywill be pod-bivalve coquinas.B) Taphonomic attributes of thetaphofacies.
below wave base, and probablybelow all but the most Biostratinomic traitsof disarticulation (Disart),reorientation (Reori),
fragmentation (Fragm), and abrasion (Abras), vary largely as a function
As
severe stormwave effects. siliciclasticsedimentsare ofenvironmental relatedtodepthinthismodel),and
energy(inversely
increasingly suppliedto the depositionalarea,chancesfor exposuretime(relatedto sedimentation and frequency of reworking
burial of skeletalparticleswill increase,and, on average, events);earlydiagenetic features ofskeletaldissolution (Sk diss)and
exposuretimeoforganismremainson thesea floorwillbe corrosion (Corros)are a function ofseafloor/interstitial sediment pH,
lower.Relatively"time poor," rapidlyaccumulatedsedi- redoxconditions, and geochemical species present;earlydiagenetic
ment may be indicatedby the generallywell-preserved, mineralization (lastcolumn)mayincludetheformation ofhardground
cements, carbonateconcretions (C), pyrite nodules(PY),orphospho-
articulated, unworn nature of many of the fossilswithin ritenodules(PH). AdaptedfromSpeyerand Brett(1991).
these parts of sedimentarysequences (Kidwell,1991a,b;

604 BRETT
I
MINIMUM NET SEDIMENTATION RATE pertainto intensityof bioturbation,at least in environ-

ments characterizedby only minorskeletal production
Zero (Omission) Negative (Erosion) (Savrda, 1993). Sedimentationrateswillvarypredictably
LUi
in the courseoftransgressive-regressive
cycles(Fig. 3). As
Kidwell (1991a) has pointedout, relativelylow sediment
cc
I II accumulationratesmayoccureitherduringtimesoftrans-
z Slowdown c,cc-NCo 0 C-?' -
0 CYq CV:- CNCNC gressiveflooding,due to offshoresedimentstarvation,or
from + (C 5 duringrelativelowstands,due to winnowing and dynamic
to 0 or -
. : C . bypassingofsediments.In eithercase,thebuildupofskel-
z c)
. +. + etal gravelsmaypromotefurther skeletalaccumulationas
LU
a resultofpositivetaphonomicfeedback;i.e.,manyshelly
epibenthicorganismsmay colonize the hard substrates
Cl) 111 IV .
z Increase _, + ::: providedby skeletaldebris(Kidwelland Jablonski,1983;
Kidwell et al., 1986). Consequently,skeletal concentra-
LUJ from0 or- ::) CC" .. tionsoccurpredictablyat severallocationsin sedimentary
z to + N
- sequences or parasequences.In manymarinefaciesthese
CcCcNc
I shellbeds mayaid inrecognitionofkeycondensedsections
U
and erosionsurfaces(Fiirsich,1978).
FIGURE3-Model fordifferent patternsofskeletalaccumulationas-
sociatedwithdecreasing(upperrow)or increasing (lowerrow)rates Patternsof Skeletal and Trace Fossil
ofsedimentation;theleftcolumnshows patterns generatedbysedi-
mentstarvation,whilerightcolumndisplayspatterns associatedwith Accumulationin Parasequences
seafloorerosion.FromKidwell bypermission
(1986); reprinted ofthe
Paleontological
Society.
Where skeletal debris is sparse or lacking,condensed
intervalsand discontinuitiesat tops ofparasequencesmay
be markedbyintenselybioturbatedhorizons.Forexample,
Brettand Baird, 1993). Furthermore, the preservation of Mortimoreand Pomerol(1991) observedextremely wide-
local patchinessis a commonphenomenonwithinthese spread horizonsof trace fossilsin the Cretaceouschalks
of Europe that appear to recordomissionsurfacesasso-
expandedsectionsand indicatesa generalabsenceoftime- ciated withmarineflooding.Evidence forsedimentcon-
averagingand hencea rapidand perhapsmorenearlycon- solidationduringprolongedintervalsof minimaldeposi-
tinuous rate of sedimentation(Brett and Baird, 1993).
tionmayalso be recordedin complexcross-cutting ofbur-
Hence, mid to late highstanddepositsare at an opposite rowsshowingdiffering
extremefromearlyhighstandortransgressive bedsinterms degreesofsharpness(Savrda,1991;
of fossilpreservation. Savrda and Bottjer,1994).Even in nearshoresands,where
The laterphases of highstanddepositionshouldbe as- bodyfossilsaretypicallylacking,tracefossilconcentration
maypermitrecognition ofsedimentary condensation.For
sociatedwithincreasing rateofsedimentation up to a point,
followed,at least in shallowwaterareas, by development example,distinctive"piperock"horizons(bedsriddledwith
ofhigherenergyconditionsas sedimentsaggradeto levels Skolithos) markfloodingsurfacesin certainearlyPaleo-
wavebase zoic sandstones(Droserand Bottjer,1989,1993).
approachingaveragestormwavebase and finally A particularlyfavorablesituationfor developmentof
itself.This shouldlead to a predictablespectrumoffossil
skeletal concentrationsin siliciclastic-dominated parase-
preservation,beginningwith excellentpreservationto-
wardstheonsetofthelaterhighstandand witha decreas- quences occursduringtimes of initialfloodingfollowing
ing grade of preservationupward (Figs. 2, 7). Increasing shallowingepisodes.Thus, shell beds, commonlyshowing
amounts of disarticulation,skeletal fragmentation and some evidence of long term time-averaging, occur pre-
otherevidenceforreworking wouldbe predictedto appear dictablyat the bases of parasequences,overlyingmarine
in the laterpartsofthe regressive(or late highstand)sys- floodingsurfaces.These beds havebeenreferred to as BOP
temstractpreservation. As a lowstandis approached,skel- or base ofparasequenceshellbeds (Banerjee and Kidwell,
etalremainsaccumulatedin veryshallowwaterareasmight 1991). BOP shellbeds maybuildup duringtimesofinitial
be expected to be highlyfragmented,abraded and re- marinefloodingin siliciclastic-dominated parasequences.
worked. Such shellbedstypicallydisplayecologicaltimeaveraging;
fossilsrepresentingdistinctivesedimentary environments
may be commingledwithinshell beds a fewcentimeters
FOSSIL ACCUMULATIONS AND to severaldecimetersin thickness.Such beds may splay
DEPOSITIONAL SEQUENCES out intooverlying sedimentswitha seriesof interbedded
thinmud and shell-richlayers.
BackgroundAccumulations Similarprocessesmayoccurin offshore carbonatepara-
A keycontrolon the formationof fossilaccumulations sequences (or punctuatedaggradationalcycles,PACs of
on the seaflooris lack of dilutionby sedimentation(Fir- Goodwinand Anderson,1985),resultingin thinshellbeds
sich,1978;Fiirsichand Aberhan,1990;Kidwell,1988,1991a, whichrestabruptlyoverthecaps ofunderlying smallscale
b; Kidwelland Bosence,1991). A similarrelationshipmay cycles of more sparselyfossiliferous lime mudstonesor

SEQUENCE STRATIGRAPHYIN PALEONTOLOGY
SEQUENCE STRATIGRAPHY PALEONTOLOGY 605
m
Approx.
A
IWater Depth
1 i 2': E :*,j;e Pyr. ^^* ^..'* ^
SSS/PSB
:.-..
25 m 50m Pyr.
MFS,?'3 + C + -t
CB
7
1? ?I?,??-?:
s
I .I:L
I6 ?4,~'
L CI
r
Z?c:e Q
!
:c.
" '- '- .'J-:
..... 4
E
:Y =Tr ic
r r,- Lr L? ........... Y i3,
4P .sx;
Zlr i:LI'?l'??;
"""'-?---?
Cn?nc;r.
rrryinrr
-??...
tE: 3..1.I Lt.?:?-?-"?
c
??.?-i......?-???
IC.CI _ L?c.ccrr=Tr?-,?-?:
1(
::::,, ;iltl'LI;?* ,L1 r*CI_? LJLI?. =L,,-?Cr\r

: ;..,:??:;??;;rc.;;;"-irl??=??..
rc
:;;." -;;';;
/r;_S_ce,rr
,,,r?X?i; rr -i_c
?:
?:r ~-'~""~e"-
--t,..
cn
i: C c,,;Y rn
.Nlrrrrr_r'
kU?- rz
r u ??\-rc?:f
"-?n7,-
A
r
?, :'?
:Ili clrr,
z (
:::-
,..... r C L c : -..
.7 ????-?:
I jt
s I'-?'? ?-.
.r
:i71 i
..^;r. , i - i^r-- ^! -d ^
?-?--
?rr..i:5
cr3rft i ??-- B 77^7^'!^^~~~~r '
.N'f:c^
r :c
i .:'r7 1
js^
^^^^ ? f^^^/
FS ,?.? B .'Y1-.L:_;PCL?L?;L
rAAAIAAIA
ICIY..._-?-,_
r v ,,,13
rr--r
I_Jlc-
rle
cu C a
.
?-???
....
?14
4rr L_f A C rrh CI C
:: I C A AAAAnAA_ CLIAAIIAA IIL
r
-C,v
c
r, :i
It e a z
T sl i:-: 7, 5??;i ?? ?? "-u.rr--, ??
-*.?L.C
C :?i?1 ?-... +
1"
i ' Ay)ll YrYI' -?l-.rljL1I --;7? --?
: Lnji , C?LLIAI
:sr:
+ t t c tf_ PYt? I;;'se,ppppppPb 1Pia _ SF-CC
."- .1
I )' I: ' '' 1. r??
A i'
rr7 r2r_7
F.nAU
MFS 'r tr rr
i:
i"' BP:,iE
:?:1: ??r
CB jLL
?f ?? ?.,
?r??IIICCA?ICI o
.c.c??-? +++*++*+?
Yr:'f1Y:.i PYI? +t:=;;sL=-?r:?*?:*i:r.-;-::-+-,-:o:
fr "~'-?
L
'-? ?: ;r-.?t.
D
I .
FIGURE4-Variationofshellbedstratinomy, taphonomy within

andskeletalaccumulation representativemeter-scale
carbonateparasequences
fromtheMiddleOrdovician Trenton GroupincentralNewYork.Thefollowing features A) base ofparasequence,characterized
are indicated:
bya seriesofshalesandcalcisiltitesthatrepresent distalstormlayers(tempestites);
well-preserved andtrilobite
crinoid remainsindicaterapid
eventburial;B) similarbutmoreproximal stormeventbeds,also displaying obrution
features;C) gradedtempestites displayinglaminatedto
burrowed ("lam-scram") fossilsless common;D) amalgamated
articulated
fabrics, skeletalpack-andgrainstonesofTOP (topofparasequence)
bed; notescouredbases; E) condensednodularwackestonesnearcycletopdisplayabundantdisarticulated skeletaldebrisas wellas whole
fossils;nautiloids
are typically
concentrated forming condensedBOP shellbeds (CB); notecorrosion surfaceat topoverlain bya thinlag of
fragmented, corrodedand reworked crinoidand shelldebrisat base ofnextcycle;Pyr= pyrite-rich
sediment.Alsonoteminor surface
flooding
ofsmallscale cyclemarkedFS. SSS = surfaceofsediment starvation;PSB = parasequenceboundary.
wackestones(Fig. 4). In such instances,theshellsare typ- scale biohermsmay occuron floodingsurfaces(Hagdorn,

ically embedded in a finegrainedsiliciclasticrich mud, 1982; West et al., in press). Withinthe peritidalzone,
and may include abundant small corals,brachiopods,or widespreadhorizonsofstromatolites or thrombolites
may
mollusks.These somewhatcondensedskeletalconcentra- also recordmarinefloodingevents.Stromatolitehorizons
tions appear to result fromtemporarydrowningof the have been tracedforover100 kmparallelto strikein lower
carbonateplatformby risingsea level.In otherwords,the Proterozoiccarbonatesof the Great Slave Lake area in
organismswhichproducefinecarbonatesediment,suchas Canada (Grotzinger,1986). Eagan and Liddell (in press)
algae, are temporarilydiminishedduringthese times of traced beds of columnarstromatolitesinto thrombolitic
deepeningand thus siliciclasticmud accumulatesas the biostromesin intertidalto subtidalfaciesin theCambrian
mainmatrixaroundlargerbioclasts,notbecauseit is being ofthe GreatBasin. These authorsattributedthe buildup
supplied in greaterconcentration, but because of the ab- of microbialmoundsto relativelyrapid deepening.Such
sence of "dilution"by fine-grained carbonatesediment. featuresmayprovideusefulregionalmarkersand permit
In shallowerwater,the firming of sedimentand abrupt delineationof subtleparasequenceboundaries.
minorriseofsea levelmayfacilitatecolonizationbythick- In the higherportionsofsmallscale shallowing-upward
ets orbanksofcorals,stromatoporoids, bryozoansorother parasequences,shell beds may again appear. Such shell
organisms.Thus, laterallyextensivebiostromesor small beds havebeenreferred to as TOP, ortopofparasequence,

*_,g:
606 BRETT
lower
Muschelkalk
Aalenlan beds. These are typicallydiscontinuousand show local,
within-habitat timeaveraging.Such beds may occuras a
Fl
resultofthe shallowingofthe sea floor(by sedimentpro-
gradationor minorsea level fall) into areas which are
affectedmorefrequentlyby the winnowingand scouring
effectsof stormwaves. Near the tops of some parase-
quences skeletal accumulations,consistingof physically
r MrS
SB
reworked,typicallydisarticulatedand fragmentary
etal material,mayagain becomecommonand build up to
skel-
thicknessesof severalcentimetersor decimeters.

TOP shell beds are typicallywell developedin carbon-
ate-dominated parasequences(Fig.4) wheretheymayform
extensiveshell-richcaps up to severalmetersthick(Kid-
well and Aigner,1985; Aigner,1985). These beds are the
m, productoferosionalwinnowing or dynamicbypassoffiner
grainedsedimentsas the sea bottomapproachesand then
exceedsa thresholdofhigherenergyconditionsassociated
withaveragestormwave base.
. .
Patternsof Skeletal and Trace Fossil
A Accumulationin Sequences
,,KLP FEL,,
Somewhatmoretime-significant skeletalaccumulations
ci
proximal _distal occur again in a predictablearrayin largerscale cycles
_ _ representedby fourthor thirdordersequences (Kidwell,
= e M FS
L-
ci' ,,-
M?. 1991a,b; Kidwelland Behrensmeyer, 1993;Brettand Baird,
1993,in pressb). Mixed carbonateand siliciclasticdepo-
4)
sitionalsequences typicallyshow a concentrationof car-

a
4)
>:-.? SB
_-
,F
_
_
-
_
,
- ;
bonateswithinthe transgressive systemstract,as thisin-
tervalis characterizedby offshoresiliciclasticsediment
-; } *- *..
C, ...--.- _
starvation(Fig. 5). Duringtimeswhenthe sea flooris still

-_-_-__--
---_
*. _
_
_ _ =
relativelyshallow,and whensea level rise eventsare not

_
sufficientlyrapid to drownthe platform,skeletal accu-
=_~ mulationsmay build up and formprimarycarbonates.
Excellentexamplesof thickersuccessionsof skeletalcar-
bonate associated with transgressivetracts include the
_
widespreadencriniteor crinoidallimestonedepositstyp-
B ical of many Paleozoic sedimentarysequences (Ausich,
1990; Kidwell and Brenchley,1994). Such deposits are
typicallyrathertexturallymature,skeletal grainstones
condensation -s .
composedofsand-to finegravel-sizedand typicallyrather

...
--_.. I II well sorted,abraded and rounded pelmatozoanossicles
non deposition
4-
ofthe"roofbed" is hereininterpreted as a sequenceboundary, top

ofbed as a condensedinterval; Bayeret al. (1985)con-
alternatively,
sideredtheroofbedto represent theregressive maximum (lowstand).
c T ~- 2 ~ "1F_---..
.~~c B) Comparison of Klupfelcyclealonga proximal (upramp)to distal
gradientnotethattheshallowing upwardportion ofthecycleis largely
truncatedatan erosionsurface(hereinterpretedas a sequencebound-
.77~~~~~~~?' arySB) beneathammonite-rich limestone bedinleftfigure;conversely,
thisportionof the cycleis wellrepresented in thicker,moredistal
sections;themaximum surface(MFS,orsurfaceofmaximum
flooding
sedimentstarvation) is shownas the datum(horizontal line)in the
FIGURE5-Comparisonofan idealizedasymmetrical transgressive-cross section.C) Inferredtimerelationshipswithinthecycleat each
shallowingcycles(depositionalsequence;leftcolumn)withobserved of the threesectionsshownabove; blackareas indicatehiatuses
patternsofsequencesincarbonates(TriassicMuschelkalk Limestone, producedbyerosion,whiteregionsarehiatusesduetosediment star-
southern Germany) (MiddleJuras- vation(non-deposition).
andmixedcarbonates-siliciclastics Figuremodified and adaptedfromBayeret
sic,Aalenian),
"Klupfel" base al. (1985).
cycles,Lorainearea,France).Erosional

(Figs. 5, 7). Comparable beds, composed of abraded or - /L Hb orC E Hs

-_- - - -
corrodedshell fragments, occur in many Mesozoic and
Cenozoicsequences.For example,complexshell-rich grav- COQUNA Lag (L) Composite(C)
or Hialal
Events (E) Hiatal
starved (Hs)
els, oolitic sands and reworkedconcretionscharacterize A TYPE:
bypassed (Hb)
the so-called"roofbeds" thatcap (or overlie?)largescale,
shallowing-upward cyclesin theTriassicMuschelkalkand UDEfLY1NG Shoreface Nearshore Open Shelf Shelf/Basin
Middle Jurassicof the Loraine area of France (Kliipfel, CAUSE: Erosion Storm Sediment Distal
1917; Aigner,1985; Bayer et al., 1985;Fig. 5, herein).The Accretion Dilution Starvation
bases ofsuch beds are typicallysharpand commonlypre- BASIN AXIS

servecasts ofsharplydefinedburrows,evidenceofa firm-
groundendofaunalassociationthatis otherwiseunrecord- E '
ed (see Kidwell and Aigner,1985,forexamplesfromthe Hsa ?

Miocene of Maryland). Landing and Brett (1987) de- E
scribedone such firmground ichnofaunafromthe Middle C
DevonianofOntarioand inferred thatovercompacted muds B Maryand Miocene
MarylandMiocene
had been exposed on the seaflooras a resultof erosional
truncationof overlyingsedimentduringa sea level low-
stand.
Most researchershave interpreted the roofbeds as "re-
gressivemaxima";however,the factthatthesebeds over-
lap erosion surfacestowardbasin margins,and display x tbUgHs
max. transgr. _._ L/Hs
E
fining-upward successionsinternally,suggeststhat they

Lorraine Jurassic
representcomponentsoftransgressive systemstracts(Figs.
5, 6). This distinctionunderscoresthe difference in em- C
phasis betweenthetraditionalapproachofdividingstrata FIGURE6-Distribution of shellbeds (coquinas)concentrated as a
into T-R cyclesand that of sequence stratigraphy. Roof resultoferosionlagformation (L),hiatalsediment starvation
(Hs),hiatal
beds may,indeed,representthe shallowestwaterdeposits bypassing(Hb),sedimentation-erosion events(E), orcombinations of
in the stratigraphiccolumn,but discontinuitiesbeneath theseprocesses(C), inshallowing-upward cycles(parasequences).A)
idealizedonshore-offshore spectrum ofdepositional processes.B, C)
themwereproducedby stillshallowerconditions,not re- seriesofshellbeds intransgressive-regressive cyclesoflocalsections
cordedby deposits. alongonshore-offshore transects.B) siliciclastic-dominated
shelfset-
Kidwell and Brenchley(1994) have documenteda gen- ting,based ontheMiocenethird-order sequencesfrom Maryland(Kid-
eral increasein the thicknessand complexityoflaterMe- well,1989);C) mixedsiliciclastic-carbonatecyclesbasedonthird-order
sozoic and Cenozoic shell beds, as comparedto Paleozoic "Klupfel cycles"fromtheJurassicofGermany and France(Bayeret
examples.They attributethistrendto the increasedpro- al., 1985). FigureadaptedfromKidwell (1991a).
ductivityand robustnessof molluscanshells.Brachiopod
shells are relativelythinnerand moreproneto fragmen-
tationdue to theirmicroarchitecture; withrareexceptions, 1917;Firsich,1979;Bayeret al., 1985;Fig. 5,herein).Early
theydo not appear to have formedshellbeds thickerthan cementationof carbonatesmayhave been morefrequent
a fewcentimeters. at certaingeologicalintervals,such as the Ordovician,as
Transgressiveskeletal limestones commonlydisplay a resultof differences in oceanic chemistry(Walkerand
hardgroundsat theirupper surfaces,markingsurfacesof Diehl, 1985).
maximumsedimentstarvationduringwhichthe skeletal In deeper waterareas, skeletalhash limestonesof this
deposits became stabilized and sedimentation rates type may be replaced in the transgressive systemstract
droppedsufficiently lowthatinterstitial cementsbeganto by pelagic limestones (e.g., "Cephalopodenkalk,"con-
formin the pore spaces ofthe skeletaldebris(Wilsonand densed cephalopodlimestonesof Paleozoic and Mesozoic
Palmer,1992). The presenceofcementedepibenthos,e.g., age). Outstandingexamplesoccur in the Silurian of the
oysters,or borings,e.g., Trypanites,may provideimpor- PraguebasinofBohemia(Kriz,1992),and in theDevonian
tant evidence of such discontinuities.Commonly,hard- redto blackcephalopodlimestonesofMoroccoand central
groundsoccurat shale-on-limestone contacts,and in some Europe (Wendt, 1988; Wendt and Aigner,1982, 1985;
cases,thelast generationofencrusting organismshas been Wendtet al., 1984). Some belemnite"battlefields"(Doyle
rapidlysmotheredby pulses of mud deposition(Fig. 7). and Macdonald, 1993) also fall in this category.Pelagic
Excellent examples of hardgroundsoccur at the upper tests and small skeletonsof pelagic organismsmay form
contactsof a numberof transgressive marinelimestones a dominantcomponentof these offshoreskeletal lime-
in the Middle Ordovicianof the MississippiValley and stones. In the Paleozoic, nautiloids,goniatitesand sty-
Ontario(see Palmer and Palmer,1977; Brettand Brook- liolinidsor nowakiidsmaybe common(Wendtet al., 1984;
field,1984; Walkerand Diehl, 1985), in the Triassic Mu- Walliser,1990), whereas,in later Mesozoic and Cenozoic
schelkalkLimestonein southcentralGermany(see Aigner, timesforaminiferal, coccolith,calpionellid,and pteropod
1985;Hagdorn,1982,1985),and at thetops of"roofbeds" remainsmaydominatetheseoffshore limestones(Hallam,
in JurassicKlipfel cyclesfromthe Loraine area (Klipfel, 1975; Loutit et al., 1988; Sarg, 1988; Aberhan,1994; Bra-

_ _ .
608 BRETT
i ~
O E,.' MFS
_2 F SB/SSB F
:::
.5. :-l~~TFS
'.E '%o,. -,." ....'>
'
'?
7-'"-I -
r_- . . )C3. . . C
-r
.
PB H
SWB~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?~i''''''
WB ~`~?- :
I- SB/SSB
SWaterDepth .-.---
FIGURE7-Taphofaciesandskeletalaccumulation withina 10 meterthickmudrock-dominated sequenceintheMiddleDevonianofNewYork.

InsetsA, B representearlyand laterportions
oftransgressive systemstract(TST); A) sequencebounding unconformities
(SB), are overlain
bytransgressiveskeletalgravel;despiteamalgamation, remnants ofsingleburialeventsare rarely condensedupper
preserved.B) relatively
partofTST withstackedobrution horizons;notethinlagofreworked fossilsand phosphatic nodulesmantlingthemaximum floodingsurface
(MFS). C-D) early(maximum) highstand; lowenergy, dysoxicmudfacies;noteobliquely-oriented nautiloid
and insitunuculidbivalveswith
pyritic
overgrowths.D) mid-highstand condensedbed ("precursor bed" of Brettand Baird,inpressa) marksinitiationofrapidsea levelfall;
erosionofsubjacentdysoxicmudsproduceslag ofreworked concretionsandfossils.E) regressive
or latehighstandphase characterizedby
rapidlyaggradedbutheavilybioturbated (noteZoophycos)sediment; fossilsscatteredbuttypicallywellpreserved,storm-winnowing events
becomeincreasingly numerous upward.F) "regressive cap" ofcycle;seafloorapproaching normalwave base; noteshellysandstoneswith
hummocky and latediagenetic
crossstratification lenticular
septarialconcretion; shelllags (coquinites)
mayoccurdue to seafloorwinnowing
eventsand sediment bypass.Modified fromBrettand Baird(1993).
sier, 1995). Typically,these relativelythin successions The biohermsextendupwardintotheoverlying highstand

(decimetersto a fewmetersin thickness)containnumer- systemstractand interfinger withargillaceousor micritic
ous internalcorrosionsurfacesormineralizedhardgrounds "background"facies.In some cases, the deepeningevent
(Sellwood and Jenkyns,1975; Hallam, 1975). was sufficiently
rapid that these biohermsfailedto keep
Among carbonate-dominatedsequences, biohermsor up withdepositionand becamedrownedand buriedinfine
reefsseemparticularly characteristic
ofthetransitionfrom grain carbonateor siliciclasticsediment.As noted,this
transgressivesystemstractto earlyhighstands(Sarg,1988; patternalso holds true formany smallerbiohermsand
Buchbinderet al., 1993). Many Paleozoic bioherms,for biostromesthat developedat floodingsurfacesof parase-
example,appear to have been establishedon firmor hard quences.
substratesassociatedwiththe stabilizationof olderskel- Biohermsare generallyabsentat thesesurfacesin deep
etal sand depositsand coincidentwithsurfacesof maxi- waterareas. Instead,surfacesof maximumstarvationare
mumsedimentstarvation(Walkerand Alberstadt,1975). most commonlymarkedby corrosionsurfaces,typically

withthinskeletallags of phosphaticmaterialsuch as fish that are interpretedas true sequence boundaries.They

bone beds (Kaufmann,1986;Kidwell,1989) or,in the Pa- overlierelativelymonotonousdeep waterfaciesoftheearly
leozoic,conodontbeds (Heckel, 1977,1986;see Baird and highstand.Thus, ratherthanrepresenting maximumlow-
Brett,1986 forreview).Similarbut thinnerand less per- stand,these featuresappear to have developedduringan
sistentbone beds mayoccurat smallerscale floodingsur- initialrapid phase of sea level fall.
faceson the tops of minorparasequences. It is unclearwhysedimentary starvationorcondensation
Siliceous deposits,composedof spongespicules or ra- should occur duringinitialregressiveevents.One possi-
diolarians,may typifyearly highstanddeposits in areas bilityis that during"forcedregressions"(i.e., these pro-
whicharelackingin carbonateorsiliciclasticdilution(Hal- ducedbyactualsea-levelfall,as opposedto progradation),
lam,1975).These maybe manifestas chert-rich beds.They shallowingbringsportionsof the seafloorwithinstorm
will be particularlycharacteristicof earlyhighstandsin wavebase,causingsubmarineerosion.Initially,in offshore
areas ofoceanicupwellingwhichprovidenutrients to nour- areas,thiserosionis notbalancedbyinputofnewsediment
ish radiolariansor, in post-Jurassicsediments,diatoms due to a time-lageffectbetweensea-levelfalland progra-
(see Wignall,1994,and referencestherein). dation.Later,continuedloweringofbase-levelmaycause
Manycondensed,transgressive depositsin thelowerPa- erosionin coastalareas and progradation ofa clasticwedge
leozoicconsistofthin,tabularto nodulardiageneticlime- basinwardover the precursorbed. Consequently,in sili-
stones and/orconcretionhorizons,typicallyrich in the ciclastic-dominated sequences,theprecursorbed typically
remainsof trilobites,as well as certainbrachiopods,pel- lies at the base of a coarseningupwardparasequenceset
matozoandebris,and sponges.For example,trilobitehash whichformsthe late highstand(regressive)systemstract
and spongespicularbeds occur in the transgressive sys- (Fig. 7).
temstractsof sequencesand as thinBOP beds ofparase- The midto laterhighstanddepositsofmostsiliciclastic-
quencesintheMiddleCambrianofwesternNorthAmerica dominated depositional sequences display fewerthick
(Westrop,1989; Allisonand Brett,1995). skeletalconcentrations, althoughthin and typicallydis-
A smoothtransitionbetweenearlyand laterhighstand continuousshell beds may occur at or near the caps of
phases would be predicted by most sequence models. componentparasequences (Fig. 7). Depending upon lo-
Moreover,gradually increasingrates of sedimentation cation,themidhighstandmaybe reflected in dysoxiclam-
would also be anticipated.However,carefulexamination inatedshale facies,or graybioturbatedmudrockfacies.In
of many sequences, at least in the Paleozoic of the Ap- nearshoreareas,hummocky crossbeddedsandstones,typ-
palachianBasin, revealsthedevelopmentofa distinctdis- icallyinterbedded withmudstone,mayprevailwithinthese
continuity and condensedshellbedbetweentheearlier(ag- settings.Fossils occuras skeletallags at the bases ofthese
gradational)and later (regressive)partsof the highstand stormbeds, or as scatteredindividualswithinthe back-
systemstract.This changeis markedin manycases by a groundmudrocks.Mud to silt faciesare characterizedby
thin (decimeters) condensed bed which occurs at the abundantbioturbation.
boundarybetweendeeper and abruptlyshallowerwater In themiddlePaleozoic,thetracefossilZoophycoscom-
deposits(Fig. 7). These beds commonlycontainreworked monlydominatesthe bioturbatedsilty mudrockfacies
concretions,derivedfromerosionof underlyingdysoxic (Bottjerand Droser,1994). However,in pre-Siluriantime
sedimentaryfacieswhereconcretionsare typical(see for theZoophycostrace-maker was veryuncommonorabsent
example,Bayer,Altheimer, and Deutchle,1985).Corroded in offshore facies,occurringmorerarelyin nearshoresands
fossilsof mixed communitytypesprovideadditionalevi- (Thayer et al., 1990). The result of this is that storm-
dence for ecological-scalecondensation.Such beds, re- dominated,siltyheterolithicfaciesof Cambrianand Or-
ferredto as "precursorbeds" by Brettand Baird (in press doviciantime are apparentlyreplaced by moremassive,
a), occurpredictablyin the manyPaleozoic sequences in non-bedded,heavily-churned, siltymudstonefaciesin lat-
theAppalachianforelandbasinand reconnaissance studies er times. In the Mesozoic, the Zoophycos trace-makers
indicatetheirpresenceelsewhere(Brettand Baird,inpress appear to have retreatedfartheroffshore, and theirsprei-
b). For example,Heckel (1977) commentson the abrupt ten occuron beddingplanes withinrelativelydysoxicoff-
appearanceofshellrichbeds abovedarkgray"coreshales" shore facies (Bottjer and Droser, 1994). However,Zoo-
(maximumhighstand)and belowregressivelimestonesin phycos appears to be replaced in shallow,siltymudrock
Pennsylvaniancyclothems. faciesby othertypesof bioturbatingtrace markerssuch
It mightbe suggestedthat such "precursorbeds" ac- as Teichichnus,Rhizocorallium,and Planolites (Bottjer
tuallyoverliesequence boundariesand representthe bot- and Droser,1994).
toms of lowstand systemstracts (S. M. Holland, pers. The extentofbioturbation generallyappearsto increase
comm.,1995). However,in contrastto lowstanduncon- upwardto a pointwithinmost shallowing-upward cycles
formities, the discontinuitiesbeneaththese beds become (Sellwood,1970; Savrda, 1991). It may be diminishedin
less pronouncedin uprampor nearshoreareas. The beds higher,regressiveportionsof some cycles as a resultof
themselvesappear mostdistinctand condensedin offshore frequentphysicalreworking of the sediments(Fig. 7).
areas,as do condensedhighstanddeposits;theyappear to Many carbonate-dominated sequences display similar
"splay-out"into continuoussectionstowardsiliciclastic fabrics.These showscatteredfossilsofvariedtypesimbed-
shorelines.Moreover,precursorbeds generallylie wellbe- ded in a sparselyfossiliferous, heavilybioturbatedmatrix
low, but may be cut out by, moremajor disconformities (Figs. 4, 6). Thus, burrowed,stylonodular wacke-to pack-

610 BRETT
stones tend to be characteristicof the upper portionsof transgressions (see Elder,1987). For example,manyofthe
manyscale sedimentary cycles,especiallyin the earlyPa- famoustrilobiteand crinoidbeds in the Paleozoic ofeast-
leozoic (Droser and Bottjer,1988, 1989). However,as is ernNorthAmericaoccurin thinbeds ofargillaceous,con-
true near the tops of parasequences,the late highstand cretionarylimestone,in condensed upper transgressive
systemstractsin some carbonatefaciesmay be typified systemstracts(Speyerand Brett,1985;Brettand Taylor,
by skeletalpack- or grainstones.For example,the regres- in press; Hickerson,in press).
sive limestonesofPennsylvaniancyclothems maybe mas- In relativelydeep-wateroffshore settingstheearlyhigh-
sive,skeletalhash dominatedbeds, withabundantdebris stand intervalalso maydisplayunique conditionsforthe
ofshallowwaterorganisms, suchas phylloidalgae (Heckel, preservationof extraordinaryfossils or "lagerstatten."
1977). Many offshoresettingswill be deep enoughthat distur-
Erosionat sequenceboundariescommonly producesme- bance ofthe seafloorbywavesor currentswillbe minimal.
chanicallags ofcoarse,robustand geochemically resistant The frequentdevelopmentof dysoxicto anoxic bottom
particles,such as phosphatizedsteinkernsof shells and watersduringearlyhighstands(see Hallam and Bradshaw,
prefossilizedbone and wood (Baum and Vail, 1988; Kid- 1979; Wignalland Hallam, 1991; Wignall,1994) mayalso
bioclastsmay be incor-
well, 1991a). These pre-fossilized inhibitscavengingand favorarticulatedpreservationof
porated into basal transgressivelags of the overlyingse- carcassesthatsettleintothisenvironment evenunderrel-
quence; theseremanieclasts are commonlyboredand en- ativelylow ratesofburial(so-called"stagnationdeposits"
crustedbyshallowmarineorganisms(see Webb,1994,and of Seilacher and Hemleben,1966; Seilacher et al., 1985).
Bryan,1992, forexcellentCarboniferousand Cretaceous Articulatedskeletonsof fish,epiplanktoniccrinoids,and
examples,respectively).Savrda (1991) also identified
bored marinereptilesin the Early Jurassic(Lias) black shales
"woodgounds"(Terredolitesichnofacies)overlying trans- of Germanyand the Dorset Coast of Britainprovideex-
gressivesurfacesin the PaleogeneofAlabama,and argued cellentexamples(Seilacher,1982b; Kauffman,1981; Sei-
that the drowningof coastal woodlandsduringtransgres- lacheret al., 1985). However,in certaindeeperwaterset-
sion providedabundantwoodysubstrates. tings,the sea bottommayremaintoo distalto sourceareas
Wherelowstanddepositsare developedin shelfand epi- to receive sedimentblankets requisite for preservation
continentalsea areas,theyare typicallysiliciclasticfacies, (Speyerand Brett,1991).
displayingscatteredfossilsof marginalmarineor non- Giventheincreasedratesofoffshore sedimentation typ-
marine types. Shell beds are generallylacking (see Ar- ical of mid to later highstand,one mightpredictthe oc-
mentrout,1992). currenceof numeroussmotheredbottomassemblages,or
eventbeds. This is particularlytruebecause,duringparts
of thisdeposition,the sea floorin manyareas willremain
DistributionofFossil Event Deposits belowaveragestormwavebase and henceeventlayerswill
(Lagerstdtten)in Depositional Sequences be protectedfromreworking. Thus, even in fullyoxygen-
ated settingsthe potentialforpreservationof obrution
In additionto thebackgroundaccumulationsofskeletal deposits should be quite high duringmid-highstand in-
debris,certainbeds occur as the resultof nearlyinstan- tervals.The famousMazon Creek Essex fauna fromthe
taneous sedimentationprocesses.These includethe well Pennsylvanianof Illinois (Baird et al., 1986) providesan
known obrutionbeds or smotheredsea bottomassem- excellentexample. Extraordinarily rapid burial and en-
blages,as wellas transportedassemblagesofskeletalma- tombmentin earlydiageneticsideritenodules preserved
terialin turbidites,and stormwinnowedcoquinadeposits. even softbodied organismsduringmid-highstand progra-
Such beds mayoccurthroughoutsedimentarycyclesand dation of a clasticwedgeinto a shallowbasin.
sequences,althoughtheytend to occurmorepredictably Nonetheless,in many sedimentarysuccessions,fewer
at certainpositions(Fig. 7). eventdepositsare recognizedwithinthemidto laterhigh-
Obrution("smotheredbottom")depositsare mostfre- stand intervalsthan in more condensedsections.Those
quentlyrecognizedin the late transgressive to earlyhigh- eventbeds that are discernedtend to be sparselyfossil-
stand depositsof manysuccessions(Brettand Seilacher, iferous.Presumably,many burial layers go unnoticed.
1991). This is an empiricalrelationshipthat seemssome- Scatteredcompletelyarticulatedfossilsare not generally
whatparadoxical.Because thesesectionsare relatively the registeredas partsofburialhorizons,althoughtheirpres-
most sedimentstarved,one mightpredictfewerburial ervationis probablytheresultofsingledepositionalevents.
events.However,manycondensedsuccessionsmayaccu- Hence, the low densityand patchydistributionof fossils
mulate as a stack of veryinfrequentlarge depositional mitigatesagainst the recognitionof obrutionhorizons
events.Major stormsmay carrysiliciclasticor allodapic withinthisportionofthesection.Furthermore, manybur-
carbonatesedimentsintobasinsthatare otherwisealmost rowingorganisms, suchas theproducersofZoophycos,are
completelysedimentstarved.These are protectedfrom favoredby moderateratesof sedimentationwhichsupply
later reworking because of the deep (and deepening)po- the substratewithabundantfinearticulateorganicmatter
sitionofthesea floor.The recognition ofobrutiondeposits uponwhichtheburrowers feed(Scott,1978;Fiirsich,1978;
is facilitatedby the burial of abundantskeletonizedepi- Rhoads et al., 1972). Hence, rates of bioturbationmay
benthicorganismsthat inhabit shallow seafloorsduring actuallyincreaseup to a pointas sedimentaccumulation
these times of relativelylow turbidityassociated with rates increase,thoroughlyhomogenizingsediments,and

eradicatingtracesofformereventburiallayers(Sellwood, vationmay provideinformation criticalto the recog-

1970; Goldring,1995; Fig. 7, herein). nitionof different portionsof sedimentarycyclesand
sequences.Fossil debrisbeds or intenselybioturbated
layerstypicallypermitus to recognizesurfacesofmax-
CONCLUSIONS imum starvation,for example. Preservationalattri-
There are numerousimportantconnectionsbetweense- butes also provideone of the fewavenuesof approach
to assessmentoftimerichnesswithinsedimentary rocks.
and
quence stratigraphy paleontology, many of which are
6) Sequence stratigraphy provides a heuristic paradigm
only just being explored.The presentpaper providesa forinvestigating
crosssectionof some empiricaland hypotheticalcorrela- patternsand processesin biostratig-
tionsbetweensequence stratigraphy and biostratigraphy, raphy and taphonomy. Increasedintegrationof pale-
ontology and sequence stratigraphy will surelyyield
taphonomyand stratinomy(fossilbed genesis). Most of into the interaction of biotic and
the generalizationsshould be viewedas preliminary and important insights
abiotic processes in earth history.
tentative.Nearlyall ofthesuggestedrelationships require
further documentationand rigoroustesting.Nonetheless,
integration ofstratigraphicand paleontologicdata should
ACKNOWLEDGMENTS
pave the way towarda much improvedunderstanding of
earthand lifehistory.Reviewofthe connectionsbetween I gratefully acknowledgethe ongoingsupportand en-
sequence stratigraphybiostratigraphyand taphonomy couragementof mycolleagueGordonC. Baird in studies
suggeststhe followinggeneralnotions. of sequence stratigraphyand paleontologyin the field;
numerous studentshave also aided in these endeavors,
1) Integrationofimprovedbiostratigraphic zonation-and
including, especially, Bill Goodman,Dave Lehmann,Steve
physical sequence-based physical stratigraphywill Keith
LoDuca, Miller, Steve Speyer, and Charles Ver
greatlyimprovethe precisionof correlation.Sequence Straeten.Lars Olsen in preparationoffigures.
helped The
biostratigraphy permitsextensionof biostratigraphic was reviewedby PeterAllison,Steve Holland,
dating from condensed offshore sections,enriched in manuscript
strata. Brad Sageman,and Charles Ver Straeten,who provided
diagnosticfossils,to shorewardless fossiliferous usefuland constructive My researchon
Epiboles ("abundancezones") oftaphonomic,ecologic, many suggestions.
and sequencestratigraphy
and immigration be used to refine
local cor- taphonomy has beensupported
originmay
relations.Mixed zonal fossils permitrecognitionof bythreegrantsfromthePetroleumResearchFund,Amer-
highlytime-richbiostratigraphically condensedbeds. ican ChemicalSociety,and by NSF GrantEAR-9219807.
2) Taphofacies(faciesdefinedon the basis of fossilpres-
ervationalfeatures)shouldcloselyparalleldepositional
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Sequence Stratigraphy, Biostratigraphy, and Taphonomy in Shallow Marine Environments

Author(s): Carlton E. Brett

This content downloaded from 134.129.115.40 on Sat, 10 Aug 2013 22:05:32 PM

DepartmentofEarth and EnvironmentalSciences, UniversityofRochester,Rochester,N.Y. 14627

PALAIOS, 1995, V. 10, p. 597-616 marinefloodingsurfacesoccurat bases ofparasequences,

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oped fromseismicrecordsofcontinentalshelfsedimentary tributionand preservationof ancientorganisms.Manyof

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not have sufficient

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FirstOccurrences WaterDepth(m) LastOccurrences

theorderofa fewhundredsofthousandsofyears(Goodwin The most detailed use of ecostratigraphy as a tool for

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raphymayproducean unprecedentedlevelofprecisionin bonate has the opportunity to accreteand formconcre-

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limestonein the Upper CretaceousGranerosShale to the imentarycyclesand thusin sequences.Hence,taphofacies,

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posits of chemicallyand mechanicallyresistantparticles A low

1991). Underappropriatecircumstances, beds ofreworked

or pyrite,includingfossilsteinkerns,may also occur at

Duringearlyhighstandphases preservationof skeletal EHS ? * EHS

materialsmaybe surprisingly poor (Fig. 2). The low rates 1+ 2

much of this shell breakageprobablyresultedfrompre- T V V V PY

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MINIMUM NET SEDIMENTATION RATE pertainto intensityof bioturbation,at least in environ-

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::::,, ;iltl'LI;?* ,L1 r*CI_? LJLI?. =L,,-?Cr\r

FIGURE4-Variationofshellbedstratinomy, taphonomy within

wackestones(Fig. 4). In such instances,theshellsare typ- scale biohermsmay occuron floodingsurfaces(Hagdorn,

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thicknessesof severalcentimetersor decimeters.

sitionalsequences typicallyshow a concentrationof car-

starvation(Fig. 5). Duringtimeswhenthe sea flooris still

relativelyshallow,and whensea level rise eventsare not

composedofsand-to finegravel-sizedand typicallyrather

ofthe"roofbed" is hereininterpreted as a sequenceboundary, top

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(Figs. 5, 7). Comparable beds, composed of abraded or - /L Hb orC E Hs

bases ofsuch beds are typicallysharpand commonlypre- BASIN AXIS

ed (see Kidwell and Aigner,1985,forexamplesfromthe Hsa ?

fining-upward successionsinternally,suggeststhat they

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'.E '%o,. -,." ....'>

FIGURE7-Taphofaciesandskeletalaccumulation withina 10 meterthickmudrock-dominated sequenceintheMiddleDevonianofNewYork.

sier, 1995). Typically,these relativelythin successions The biohermsextendupwardintotheoverlying highstand

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withthinskeletallags of phosphaticmaterialsuch as fish that are interpretedas true sequence boundaries.They

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eradicatingtracesofformereventburiallayers(Sellwood, vationmay provideinformation criticalto the recog-

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This content downloaded from 134.129.115.40 on Sat, 10 Aug 2013 22:05:32 PM

EINSELE, G.,RICKEN, W.,andSEILACHER, A. 1991,Cyclesand Events HAQ,B.U., 1991,Sequencestratigraphy, sea-levelchange,and sig-

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KIDWELL, S.M., 1986,Modelsforfossilconcentrations: Paleobiologic phonomyof echinoderms in carbonatefacies:Fort PayneFor-

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This content downloaded from 134.129.115.40 on Sat, 10 Aug 2013 22:05:32 PM

This content downloaded from 134.129.115.40 on Sat, 10 Aug 2013 22:05:32 PM

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