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PALAIOS.
http://www.jstor.org
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-
0.
.
*~~K U'C A F )
,'# tafK.'inr -
ft~ ~ ~ ~ ~ ~ ~~~~~~~~~t
4 ' ~ ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~ 4
FIU 4Rsigsoe
E fLt rtcosmrn enti itm,st I4Taooi eiini rges yM oa,Clfri
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 .)..-
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
sis]: Universityof Wisconsin-Madison,
rineplankton. As such,thesedatadoc- StudytheAlphaRidge,ArcticOcean. Geo- 233 p.
logicalSurveyof Canada, Paper 84-22, p.
umentan incidental but causal depen- 125-135. HAJOS,M., 1975, Late Cretaceous ar-
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,
Geochemistry
Site 275, in KENNETT,J. P., HOUTZ,R. E.,
survivorship during an exceptional time York,AcademicPress, p. 143-186.
et al., eds., InitialReportsoftheDeep Sea
ofcrisisinearthhistory. DALE, B., 1983, Dinoflagellate restingcysts:
DrillingProject, v. 29: Washington(U.S.
"benthicplankton", in FRYXELL,G. A., ed.,
Govt. Printing Office),p. 913-1009.
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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