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Biological Selectivity of Extinction: A Link between Background and Mass Extinction

Author(s): Jennifer A. Kitchell, David L. Clark and Andrew M. Gombos, Jr.


Source: PALAIOS, Vol. 1, No. 5 (Oct., 1986), pp. 504-511
Published by: SEPM Society for Sedimentary Geology
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504 RESEARCHLETTERS

ofExtinction:
Selectivity
Biological ticularsignificance
highsurvival
is the unexpectedly
ofmarinediatoms,a pho-
A LinkBetweenBackground and toautotrophic group of major impor-
tanceforglobalprimary production. At
Mass Extinction present,quantitative estimatesof dif-
ferentialsurvival among plankton
groups,based on a comparison oflate
Maastrichtian and earlyDanianassem-
JENNIFERA. KITCHELL proximate biologicaladaptation and the blages,indicatea relatively severege-
Department ofGeological Sciencesand postulatedend-Cretaceousextinction neric extinctionof 73% for cocco-
MuseumofPaleontology mechanisms mayhaveplayeda major lithophorids, and even higherratesfor
UniversityofMichigan rolein theunexpectedly highsurvivalof radiolaria (85%) andforaminifera (92%)
AnnArbor,Michigan48109 planktonicdiatoms, documenting a clear (Thierstein,1982), comparedto only
exampleofcausal dependency between a 23% fordiatoms(Gombos,unpubl. ms.;
DAVID L. CLARK biologiccharacterselectedfor during Harwood,in press). Such differential-
Department ofGeology and timesofnormalorbackground extinction survivalestimatesare notpredicted by
Geophysics andmacroevolutionary survivorshipdur- the currently advancedcausal mecha-
UniversityofWisconsin ingtimesofcrisis. nism.Suppression ofphotosynthesis as
Madison,Wisconsin 53706 a consequenceoflargedustloadings has
ANDREWM. GOMBOS,JR. beensuggestedbyseverallinesofevi-
INTRODUCTION dence as the proximate cause forthe
ExxonProduction ResearchCo.
Houston,Texas77001 Understanding thebiological selectiv- Late Cretaceousextinction crisis(Al-
ityor non-random natureof majorex- varezet al., 1980; Pollacket al., 1983;
PALAIOS,1986, V. 1, p. 504-511 tinctioneventsis centralto unraveling Wolbachet al., 1985). The relativeim-
the problemof causalityof extinction munity of obligatephotoautotrophic di-
(Raup, 1986). Macroevolutionary pat- atomsto a cessationofphotosynthesis
The phenomenon of non-random or ternsof differential extinction among hasbeenconsidered bysome,however,
survival
selective acrossmajorextinction coevalgroupsin the geologicpast are evidenceto rejecttheglobal-
sufficient
boundaries in thegeologic past is poorly incompletely understood.A recurrent darkeninghypothesis (Thierstein,
understood but increasingly recognized issue,stemming from analysesofdiffer- 1982; Tappan, 1982; Ekdale and
areaforfutureresearch.
as a critical A entialsurvivalduringmajorextinction Bromley,1984).
currenthypothesis,developedfrom a com- events,is whethercommonbiological RecognizingthattheLateCretaceous
parisonofextinction patterns amongLate factorsseparate surviving fromnon- crisiswas particularly severe on sur-
Cretaceousmolluscs,is thatbiological surviving taxa(Padianet al., 1984).We face-dwellingmarineorganisms, partic-
adaptations oforganisms effectual dur- reporthereempirical evidenceofa life- ularlyplankton, Landman(1984) pro-
ing normaltimesof earthhistory are historycharacterwithin a Late Creta- posedthatbecauseofdifferences inlife
ineffectualduringtimesofmassextinc- ceous assemblageof marineplankton history, ammonites mayhave suffered
tion.Microsampling of laminatedbio- thatsupportsarguments thata biologi- extinction at the Cretaceous/Tertiary
siliceous
marine sediments fromthehigh- cal traitmaydetermine a macroevolu- boundarywhereas nautilidssurvived.
AlphaRidge(-86? N lat.)pro- tionary
latitutde patternof differentialsurvival. On thebasisofthesmallsize ofammo-
videsevidence ofan actively exploitedbio- These data contrastwith,but do not niteembryonic shellsin the Late Cre-
logicaladaptationbyLate Cretaceous exclude, the recent hypothesisthat taceous,Landmansuggestedthatam-
planktonic diatoms.Distinctive laminae mass extinctions are indifferentto bio- monites hada planktonic juvenilestage,
reveala life-history
strategy comprised of logicaladaptationsevolvedduring back- placingtheirearlylifehistoryin near-
alternating planktonic and non- groundtimes Jablonski, 1986). These surfacewaters,whereasmodernnautil-
planktonicstages.Contrary tothehypoth- datamayalso resolvetheapparentin- idsdo not(see alsoArnoldandCarlson,
esisthatmassextinctions are indifferent consistency betweentheobserveddif- 1986).It has also beenhypothesized by
to biologicaladaptations, thesedata in- ferentialsurvivalof photoautotrophic MilneandMcKay(1982) that,because
dicatea linkbetween selectionfortraits plankton and the global-darkening sce- somemodernphytoplankton formrest-
in normalpaleoenvironmental settings nario(e.g., Alvarezet al., 1980,1984) ing spores, resting-sporeformation
and differentialsurvivalduringan un- currently associatedwithexplanations might survival
facilitate amongplankton
expected episodeof paleoenvironmentaloftheLate Cretaceousmassextinction duringa globalatmospheric darkening
deterioration duringa mass extinction. event. suchas thathypothesized atthecloseof
Planktonicdiatoms,adaptedlocallyto Amongmarineorganisms,plankton theCretaceous.
survivingperiodsofstressbyleavingthe experiencedsevere losses duringthe We presentevidencethatmarinedi-
planktonenvironment, mayhavediffer- end-Cretaceous massextinction (Thier- atoms,althoughpartof the plankton,
survived
entially a globalcrisisat theend stein,1982; Raup, 1986). But among mayhavedifferentially survivedan un-
oftheCretaceous as a consequence ofa plankton, the magnitude of extinctions expectedglobaldarkening thatwas par-
trait.The interaction
life-history ofthis wasnotuniform (Tappan,1982).Ofpar- ticularlysevere on planktonic organ-

Copyright? 1986, The Society of Economic Paleontologistsand Mineralogists 0883-1351/86/0001-0504/$1.50

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OF EXTINCTION
SELECTIVITY 505

0.

.
*~~K U'C A F )

,'# tafK.'inr -

ft~ ~ ~ ~ ~ ~ ~~~~~~~~~t

4 ' ~ ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~ 4

FIGUREI-Smear-slide samples of laminatedmarinesedimentsfromArcticcore Fl-437, showingalternatemodes inthe life-history of these


high-latitudeLate Cretaceous centricdiatoms. (a) and (c) are resting-sporeassemblages fromcore segment 11, lamina 6; magnification
400x and 100x, respectively;(b) and (d) show vegetative-cellassemblages fromcore segment 11, lamina 16 (b), and core segment 13,
lamina 33 (d); magnification400x and 100x, respectively.

isms,as an incidental consequenceor Maastrichtian/Danian boundary (Bukry, Microsampling ofthedistinctivelam-


effect strategy
ofa life-history adapted 1985). L. furcula,although commonin inae revealed(1) layersconsisting al-
locallyto survivingnot onlylongdark our core samples,disappearsbelow mostexclusively (max.93.3%, N>200
winterperiodsbutseasonalperiodsof (priorto) thecoresectionsexamined in per sample)of diatomrestingspores
low nutrient and highlight
availability detailforthisstudy. (Fig. la,c), and(2) layersnearlybarren
stress. We analyzedforcomparison 48 sam- ofrestingsporesand consisting ofthe
ples of individual laminaewithinthe vegetativecells (max. 96.4%, N>200
EVIDENCE OF AN ADAPTATION laminated portion(segments10-13) of per sample) of centricdiatoms(Fig.
A core fromthehigh-latitude Arctic Fl-437andanadditional 40 samplesfrom lb,d). A representative laminated sec-
site Fl-437(85.599?N,129.588?W)re- a mottled (orbioturbated)
portion ofthe tion,showing thealterna-
quantitatively
coveredan extremely well preserved core.Sampleswerenotsieved(thereis tion of vegetativecells and resting
and abundantdiatomassemblageof littlecontamnination withnon-siliceous, spores, is illustrated in Figure2. By
Late Cretaceousage. Morepreciseas- non-bioticmaterial),andwereexamined contrast,samplingmottledsections
signment on
of age is based presently at 400x and1000x magnification. Only yieldedassemblagesofcombined rest-
the disappearanceof the widespread wholeindividuals orfragments compris- ingsporesandvegetativecells.Sucha
species, Lyramulafur- ingmorethan50% ofanindividual
silicoflagellate were mixed-cell assemblageis alsocharacter-
cula,knowntobecomeextinct nearthe counted,at 400x power. isticof standardmacrosamples (e.g.,

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506 KITCHELL,CLARK& GOMBOS

Barron,1985)thatamalgamate laminae. RESTING SPORES gravesandFrench,1983). The forma-


0%
Restingsporesare hererecognized as 132.3 tionof restingsporesand consequent
100%

beingmoreheavilysilicified thanvege- rapidsinking outofthephoticzonepre-


tativecells,andhaving a reducedgirdle- ventthedestruction ofthediatomcell
bandregion.For comparison, vegeta- by metabolicimbalancesor photo-
tivecelltypesareillustrated inFigure3 0-
I
oxidativeeffectsor both(Davis et al.,
andresting-spore celltypesinFigure4. 1980). As a consequence, resting
Examplesof restingspores still at- a
=i==
spores sequesteror isolatesome por-
tachedto thevegetative cellwerealso 0 tion of the populatonfromselection
observed(e.g., Fig. 4a,b). Although whenthe ambientplanktonic environ-
heaviersilicificationintroduces a pres- 133.1
ment exceeds the physiological toler-
ervationalbias, preservation of both 100% 0% ances ofthevegetative cell.
vegetative-cell and resting-spore as- VEGETATIVE CELLS Restingsporeshavea reducedreac-
semblagesin adjacentlaminaeargues FIGURE 2-Quantitative counts of resting
tivesurfacearea andare specializedto
againstthelatterrepresenting a pres-
spores and vegetativecells ina sequence of persistin eithera benthicor deep-
ervationalartifact. eight light-dark laminae over a 0.8-cm core pelagicenvironment oflow to no light
Studies of Holocene laminated section. The horizontalblack lines desig- (FrenchandHargraves,1980; Fryxell,
diatomaceoussedimentsfromDSDP nate dark laminae. 1983). Low temperatures prolongsur-
Site 480 in the Gulfof California have vival times (Hargravesand French,
similarlyrevealedlayersdominated by 1983).Unlikedinoflagellate cysts(Dale,
restingspores(ofthediatomChaeoto- lifehistory. Mixed-cell assemblagesare 1983), therestingsporeis nota truly
ceros)andinterpreted torecordnutrientinterpreted torepresent physical mixing dormantstage and maygerminate al-
depletionduringseasonallyvariable ofthesetwobioticsedimentary compo- most immediately (French and Har-
coastalupwelling (Schrader,1982). A nents;mixing maybe post-depositional. graves, 1980; Garrison,1984). Rein-
majordifference in oceanographic Seasonalchangeinassemblagecompo- troduction
set- ofsporesto thephoticzone
tingevidencedbytheGulfofCalifornia sition,differential dissolution, and dif- maybe fromeitherthebenthicor pe-
recordandtheArcticrecordis thatthe ferential grazingmayalso havecontrib- lagicenvironment.
formerrepresentscoastal upwelling utedto thepreservedrecord.
whereas the latterrepresentsopen- Numerousstudieshave contributed
ocean upwelling; the Arcticsediments toan understanding oftheresting stage THE LOCAL LATE CRETACEOUS
are also notablydevoidof terrigenous amongmarine centric diatoms as a prox- ENVIRONMENT
input(KitchellandClark,1982).Resting imatesurvivalstrategy(Fryxell,1983; That the Late Cretaceous high-
sporesalsohavebeenreported inmod-Garrison, 1984). In themodernocean, latitude environment was quitedifferent
ernendemicspeciesfromtheAntarctic resting sporesappearimmediately after fromthatof todayis supportedby a
open-oceanupwelling zone (Hobanet a diatom bloomandtheonsetofnutrient greatdeal of paleontological evidence
al., 1980). depletion. In boththegeologicpastand (Axelrod,1984). In particular,theLate
the modernworldocean, the distribu- Cretaceouspoleswereicefree(Barron,
MEROPLANKTONICLIFE tionofdiatomaceous sediments is con- 1983). There is littlesupportfromei-
HISTORY: A PROXIMATE trolledlargelyby geographic zones of ther celestialmechanics(Harris and
SURVIVALSTRATEGY upwelling(Lisitsyn, 1972; Calvert, Ward,1982) or paleoclimatic modeling
A meroplanktonic includes 1983). The correlation
lifehistory is causal: sus- (Barron,1984) forreducedobliquity of
vegetative, resting,and sexualstages taineddiatomgrowth requirescontinual the earth'seclipticin the Late Creta-
(Smayda,1958)andis characteristic of replenishment ofsufficient dissolvedsil- ceous. Sincetheearth'seclipticserves
numerous modern diatomspeciesofthe icaandotheressentialnutrients tokeep as the majorcontrolon photoperiod,
order Centrales, class Bacillario- upwiththeexponential natureofdiatom thereis no reasonto suspectthatpho-
phyceae.Such species are pelagicas growth (Tilman,1982).Nutrient deple- toperiodsat highlatitudeswere differ-
vegetative cells,or onlyduringpartof tioncurtailsdiatomgrowthandensues entlyspacedintheLate Cretaceous.Of
theirlifecycle.Sedimentation ofbuoy- bothfromthe discontinuous natureof particularinterestare paleoclimatic
antvegetativecells is accomplished in upwelling and fromrapidgrowthrates modeling effortsthatshowthatopen-
themodernoceanbyfecal-pellet trans- thatexhaustlimiting nutrients. As a ocean upwellingwouldprevailin the
portwhereasincreasedsinkingrates consequence,planktonic diatomsmay ice-free centralArcticOceanduring the
removerestingspores,thought to be becomeentrained inhighlightlevelsin Maastrichtian winter,associatedwith
grazing resistant(Garrison,1981). an environment thatis increasingly nu- an atmospheric low-pressure zone and
We proposethatthevegetative-cell- trientdepleted(Garrison,1981; Holli- cyclonic (Kitchelland Clark,
circulation
dominatedlaminae of Arctic core baughet al., 1981;KieferandMitchell, 1982;ParrishandCurtis,1982;Barron,
F1-437recorda growth phaseofactive 1983).It is thecombination ofhighlight 1985). Hence,open-oceanupwelling at
nutrientuptakeandphotosynthesis. By levelsat lownutrient levelsthatforms F1-437, a site located on the Alpha
contrast,the resting-spore-dominated a particularly dangerousenvironment Ridge,woulddiminish andthencease at
laminaerecorda survivalphase in the forthemarinephytoplankton cell (Har- a timeof highlightintensity. Because

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SELECTIVITYOF EXTINCTION 507

FIGURE 3Vegetative cells ofLateCretaceousmarinecentricdiatoms,site FI-437.(a,b) Coscinodiscus;(c,g)Actinoptychus; (d-f,h-k,m)


Hemiaulus;(I,p)Gladius;(n)Stephanopyxis;(o) Trinacria.
(a = 18.98,udiam.;b= 24.9,udiam.;c = 2O.O8pu.
diam.;d= 21.09,ubasal length;
e= 1O.54,ubasal length;f= 6.02,ubasal length;g= 9.16,udiam.;h= 7.23,ubasal length;i= 23.7,ubasal length;
j= 17.11,ubasal length;
k= 9.94,ubasal length;I= 15.76,udiam.;m= 59.65,ubasal length;n= 13.86,ubasal length;o= 14.86,udiam.; p= 58.15,uvert.length).

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508 CLARK& GOMBOS
KITCHELL,

FIU 4Rsigsoe
E fLt rtcosmrn enti itm,st I4Taooi eiini rges yM oa,Clfri

Acdm icel iue n .( 101 egtb99p egtc=59p aa egh 181

ofSine.SMpooryJ
dim. e=1.0?da. =801 im;g .3?bsllnt;h 6.9?bsllnt 181 aa egh =281 im;k
I31 dim;1 *9?bsllnt;m 951 aa egh =681 aa egho 5 0?mx it;p 481 a.wdh
q=1.1?da. r= 1.71 upe valv dia .)..-

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SELECTIVITYOF EXTINCTION 509

diatomslacktheability to shutoffpho- monophyletic cladesviaoneoftwoproc- Such an exampleof enhancedsurvi-


tosynthesis (Fryxell,1983), highlight esses, effect macroevolution or species vorshipofa majorextinction eventas a
is a critical
intensity stressfactor,par- selection(Stanley,1979; Vrba, 1983, consequenceof a biologicaladaptation
in concertwithnutrient
ticularly deple- 1984). We interpret ourcase as an ex- contrastswiththe recentlydescribed
tion.Rapidformation ofrestingspores ampleof effectmacroevolution rather ineffectual survivorship valueof plank-
wouldremovethe endangered popula- thanspeciesselectionforthefollowing totrophiclarval developmentwithin
tionfroma now perilousphoticzone. reasons: the propertyof a mero- molluscsduringthesame mass extinc-
These atmospheric, oceanic,and bio- planktonic life historyis a collective tion,despiteits enhancement of sur-
logicmechanisms wouldoperateon a characterofthespeciesemergent atthe vivorshipduringtimesof background
seasonalbasis. We are notconcluding, leveloftheindividual ratherthanat the extinction (ablonski,1986). This mol-
however,thatadjacentpreservedlami- levelofthespecies,andtheinteraction luscanexamplehas led Gould(1984) to
nae necessarilyrecordinter-seasonal betweencharacter and environment at generalize that"massextinction is prob-
events,becauseoftheproblems ofsuf- the among-organism level causally ablyblindto the exquisiteadaptations
accurateage resolution
ficiently andthe determines differentialextinction (Vrba, evolvedforpreviousenvironments of
forcurrent
potential scouranda discon- 1983). A meroplanktonic life history normal times."The diatomexampledis-
tinuoussedimentrecord.That these benefitstheindividual inparticular types cussedinthispaperprovidesa counter
LateCretaceoussediments areoverlain of marineenvironments and, as we case. Moreover,amongmolluscs,geo-
byoniya thinveneerofLate Cenozoic propose,has benefitedclades during graphicrangeofa specieswas a good
sediments(Kitchelland Clark,1982) exceptional timesofcrisis.In thiscase, predictorof clade survivalduringthe
supportsthiscaveat. differential survivalis reducibleto Late Cretaceousmass extinction Ua-
selection upon individualsand the blonski,1986). Amongdiatoms,fluctu-
MACROEVOLUTIONARY macroevolutionary phenomenon of dif- ationin stressconditions, and notbio-
SIGNIFICANCEOF A BIOLOGICAL ferentialsurvivalis a consequenceof geography, is the betterpredictorof
ADAPTATION upwardcausation, theessentialcharac- spore formation (Hargraves and
The biologicalpropertyof forming teristics ofeffect macroevolution (Vrba, French,1983).Geographic rangewould
restingspores as a stage in the life- 1984). be a predictor but not the postulated
historycycleof marinediatomsis an The adaptation of a meroplanktonic causal factorof a life-history trait
adaptation in the currentevolutionary lifecycleamongpopulations of centric adaptedto environments ofdiscontinu-
usage(Sober,1984).We are proposing diatomsin the Late Cretaceousin- ous upwelling.
thatthisadaptation resultedina fortu- creasedtheprobability of survivorship The production of restingspores is
itousbenefit to individualsand,hence, duringnormaltimesof local environ- considered a primitiveevolutionary trait
to species.Diatomsmayhavedifferen- mentaldeterioration, as it does today. (Simonsen,1979; Fryxell,1983). Fam-
tiallysurviveda mass extinction event Thisinnovation ofa meroplanktonic life iliesthatevolvedmorerecently thanthe
thatwas particularly severe on plank- historymay have been differentiallyLate Cretaceous-theMiocenerepre-
tonicorganisms as a consequence(ef- enhancedduringthe Late Cretaceous sentingan approximate timeofdivision
fect)ofa life-history cycleadaptedlo- mass-extinction regime.As evidence, betweenprimitive and advancedfami-
callyto surviving stressbyleavingthe all taxa presentin the Fl-437assem- lies withregardto thisbiological adap-
plankton environment. The observation blageat thegenuslevelrangeintothe tation(Simonsen,1979)-forma phys-
thatmarineplanktonic diatomssuffered Tertiary,withthe exceptionof Glad- iologically restingcell. This physiologi-
proportionately less extinctionacross ius, a ubiquitousformknownworld- calshutdown doesnotentailmorpholog-
theCretaceous/Tertiary boundarythan wide to be restrictedto the Creta- ical change, and could not, conse-
othermarineplanktongroups,rather ceous. A surveyof the literature (see quently,be detected in the fossil
thanremaining an enigma(e.g., Thier- reviewby Fenner,1985; also Prosh- record.Retention oftheprimitive trait
stein,1982; Tappan,1982; Ekdaleand kina-Lavrenko, 1974; Schrader and in modernspecies has been explained
Bromley,1984),is consistent withthe Fenner,1976; Gombos,1976, 1984; by a possiblegeneticconstraint and a
hypothesisthat global atmospheric Hajos, 1975, 1976; Benda, 1972; knownselectiveadvantageundercer-
darkening mayhavebeena causalfactor Mukhina,1976; Jouse,1978) indicates tainconditions. Sporeformation permits
intheLateCretaceousextinction event. that all of the spore-generaknown a morerapidandimmediate responseto
However,the evidencereportedhere fromthe Cretaceoussurvivedintothe nutrientdepletion (Hargraves and
does not addressthe global-darkeningTertiary. Interestingly, all are foundat French,1983).
hypothesis per se, and is consistent high-latitude sites and feware known
withvariousscenariosthatpostulate fromeitherlow-or mid-latitude sites. Biological SelectivityandRecognition of
a stressed planktonhabitat (e.g., Only two spore-generaare foundin Mass Extinctions
McLean, 1982; Zoller et al., 1983; both the Maastrichtian-age Moreno
Toon,1984). Shale of California (paleolatitude The currentprocedurefor distin-
It is currentlyhypothesized thatbe- -40?N) and the Late Maastrichtian- guishing timesofbackground extinction
tween-lineage differences may fortu- age assemblagefromDSDP Site216 in from timesofmassextinction restson a
itouslydetermine an evolutionarypat- theIndianOcean (paleolatitude -30?S) criterion of magnitude (Raup and Sep-
ternor macroevolutionary trendwithin (Gresham,1985). koski,1984), a divisionshownto be

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510 KITCHELL,CLARK& GOMBOS

equivocal(Stigler,in press). We pro- gions: Palaeogeography, Palaeoclimato- phytoplankton. II. Restingspore cycles in
coherent
posethata biotically patternof logy,Palaeoecology,v. 45, p. 105-147. coastaldiatompopulations: Journal ofPlank-
onlargescalebe con-
selectivesurvival BARRON, E. J., 1983, A warm,equable Creta- tonResearch,v. 3, p. 137-156.
ceous: the natureof the problem:Earth GARRISON,D. L., 1984, Planktonic diatoms,in
sideredin conjunction withmagnitude. Science Reviews,v. 19, p. 305-338. STEIDINGER, K., and WALKER, L. M., eds.,
A mass-extinction regimeshouldbe BARRON, E. J., 1984, Climaticimplications of Marine PlanktonLife Cycle Strategies:
supportedby evidenceof predictable the variableobliquity explanation of Creta- Boca Raton,Florida,CRC Press, p. 1-17.
patterns survival
ofdifferential toa suite ceous-Paleogenehighlatitudefloras:Geol- GOMBOS,A. M., JR.,1976,PaleogeneandNeo-
ofcloselyrelatedcausalforces.Times ogy,v. 12, p. 595-598. gene diatomsfromtheFalklandPlateauand
of background extinction,by contrast, BARRON, E. J., 1985, Numericalclimatemod- MalvinasOuter Basin, Leg 36, Deep Sea
shouldnot. Background extinction is eling,a frontier in petroleumsource rock Drilling Project,in BARKER,P. R., DALZIEL,
prediction:results based on Cretaceous I. W. D., et al., eds., InitialReportsofthe
best characterizedas thesumofinde- simulations: American AssociationofPetro- Deep Sea DrillingProject,v. 36: Washing-
pendent causalevents(Hoffman andKit- leum Geologists Bulletin, v. 69, p. ton (U. S. Govt. Printing Office), p.
chell,1984). 448-459. 575-687.
BARRON, J. A., 1985, Diatombiostratigraphy of GOMBOS,A. M., JR., 1984, Late Paleocene
SUMMARY theCesar 6 core,AlphaRidge,inJACKSON, diatomsintheCape Basin,in Hsu, K. J.,LA
We concludethatrecognition of the H. R., MUDIE,P. J.,and BLASCO,S. M., BRECQUE, J.L., et al., eds., InitialReports
eds., InitialGeologicalReporton Cesar- of the Deep Sea DrillingProject, v. 73:
exploitation ofa meroplanktonic lifecy- The CanadianExpedition to StudytheAlpha Washington (U.S. Govt.Printing Office),p.
cle bycentric diatoms mayhelpresolve Ridge,ArcticOcean: GeologicalSurveyof 495-512.
whathas beentermedan enigmatic im- Canada, Paper 84-22, p. 137-148. GOMBOS, A. M., JR.,A reviewoftherecordof
munity ofa majorgroupofmarine plank- BENDA, L., 1972, The diatomsof the Moler Late Cretaceousdiatomextinctions: unpub-
tonto a globalregimeofmass extinc- Formationof Denmark(lowerEocene). A lishedmanuscript.
tion.The interaction ofthislife-history preliminary report:Nova Hedwigia,v. 39, GOULD,S. J., 1984, The cosmicdance ofSiva:
p. 251-266.
property and thepostulated Late Cre- BUKRY, D., 1985, Correlationof Late Creta-
NaturalHistory,v. 8, p. 14-19.
taceous extinctionmechanismsmay ceous Arctic silicoflagellatesfromAlpha
GRESHAM, C. W., 1985, Cretaceous and Pa-
haveplayeda majorroleintheobserved leocene siliceous phytoplankton assem-
Ridge,inJACKSON,H. R., MUDIE, P. J.,and blages fromDSDP sites 216, 214 and 208
evolutionary patternofbiological selec- BLASCO, S. M., eds., InitialGeologicalRe-
in the Pacificand IndianOceans [M.S. the-
tivityor differential survival amongma- porton Cesar-The CanadianExpedition to
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rineplankton. As such,thesedatadoc- StudytheAlphaRidge,ArcticOcean. Geo- 233 p.
logicalSurveyof Canada, Paper 84-22, p.
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dencybetweena biologicalcharacter, CALVERT,S. E., 1983, Sedimentarygeochem- chaeomonadaceae, diatomacae and
selectedforinnormal background times istryofsilicon,in ASTON,S. R., ed., Silicon
silicoflagellataefrom the South Pacific
of geologichistory,and evolutionary and Biogeochemistry: New Ocean, Deep Sea DrillingProject,Leg 29,
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ACKNOWLEDGMENTS Survival Strategies of the Algae: Cam-
bridge, CambridgeUniversityPress, p. HAJos,M., 1976, Upper Eocene and lower
We thankS. Stanley,H. Tappan,H. 69-136. Oligocene diatomaceae, archaeomon-
Thierstein, J. Barron,H. Schrader,G. DAVIS, C. O., HOLLIBAUGH,J. T., SEIBERT,
adaceae, and silicoflagellatae in southwest-
Fryxell,M. Hoban,J. Fenner,and S. D. L. R., THOMAS,W. H., and HARRISON, ern Pacificsediments,DSDP Leg 29, in
HOLLISTER, C. D., CRADDOCK,C., et al.,
and P. Kilhamforreadingthe manu- P. J., 1980, Formation ofrestingspores by
eds., InitialReportsoftheDeep Sea Drilling
script.We thankE. Barron,J. Kutz- Leptocylindrus danicus (Bacillariophyceae)
in a controlledexperimentalecosystem: Project,v. 35: (U.S. Govt.Printing Office),
bach,F. J.R. Taylor,andJ.P. Bujakfor Journalof Phycology, v. 16, p. 296-302. p. 817-884.
discussion. Thisresearchwasfunded by EKDALE, A. A., and BROMLEY, R. G., 1984, HARGRAVES, P. E., and FRENCH,F. W., 1983,
NSF grantsDPP-7926251and BSR- Sedimentology and ichnology of the Creta- Diatom resting spores: significanceand
8307099toJ. A. K. ceous-Tertiary boundaryin Denmark:im- strategies,in FRYXELL,G. A., ed., Survival
plicationsfor the causes of the terminal Strategiesof the Algae: Cambridge,Cam-
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Ithas been justlysaid, thatthegreaterthecircleoflight,the greaterthe boundaryofdarkness bywhich


itis surrounded.

-Charles Lyell

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