PALEOCEANOGRAPHY, VOL. 13, NO.

5, PAGES 517-529, OCTOBER 1998

Global dominance of Gephyrocapsacoccolithsin the late

Pleistocene:Selectivedissolution,evolution, or global
environmental change?

J6rg Bollmann
GeologicalInstitute,Eidgen6ssische
TechnischeHochschule,Ziirich, Switzerland

Karl-Heinz Baumann

GeoscienceDepartment,Universityof Bremen,Bremen,Germany

Hans R. Thierstein

GeologicalInstitute,Eidgen6ssische
TechnischeHochschule,Ztirich, Switzerland

Abstract. Individual speciesof marine calcareousphytoplanktonare known to occur in bloomsin today'soceanand to

have dominated fossil assemblages.Interest in the study of marine phytoplankton-ocean-climateinteractionshas
increasedbecauseof the potentialinfluenceof phytoplanktonspecieslike Emiliania huxleyion globalclimateand on the
global carbon cycle. There is evidencethat Gephyrocapsa caribbeanica, which is closely related to E. huxleyi, was
globally dominantin the late Pleistocene(480-262 ka). Morphologicalanalysesof Gephyrocapsacoccolithsreveal that
only two of six Holocene morphologicalassociationsoccurredduring this time interval. We examine three potential
causesfor the dominance:preservation,environment,and evolutionary adaptation.We concludethat evolutionary
adaptationwas the likely processresponsiblefor Gephyrocapsadominance,althoughthere are indicationsin the mid-
Brunhesfor warmer climatesthan today.

1. Introduction of relatedtaxa,itsdensity
or itsbiomass
(e.g.,ce!l/Lor gC/m3).
The expression
"dominance"
is usedin this studyif onetaxo-
There is an increasinginterestin the studyof the relationship
nomic unit exceeds 50% relative abundance in a nannofossil as-
between marine phytoplanktonand climate in today's oceans.
Marine phytoplankton probably influences the formation of
semblage
of a sediment
sample.)It is theyoungest
globaldomi-
nanceintervalpriorto theongoingoneof E. huxleyito whichthe
cloudsthroughthe productionof dimethylsulfide(DMS) and may
play a major role in the global carboncycle {Holligan, 19921. species
of Gephyrocapsa
arecloselyrelatedasevidencedby the
Most researchin biological climate forcing has concentratedon morphologically
intermediate
speciesGephyrocapsa protohuxleyi
lMclntyre, 19701. This mid-Brunhes dominance interval of G.
Emiliania huxleyi,a prymnesiophyte belongingto the marinecal-
careous phytoplankton which it dominates in today's oceans
caribbeanicais especiallysuitablefor the studyof the relation-
[Westbroeket al., 19931.E. huxleyiforms bloomsin shelf edge,
shipbetweencalcareous marinephytoplankton andthe environ-
shelf sea, and open ocean environments.The causesfor these mentbecauseall formerlydominantmorphologicalassociations
bloomsof E. huxleyi are still not known [ Westbroeket al., 19931. of Gephyrocapsa arestillalivetodayandhigh-resolution oxygen
E. huxleyi may be one of the most importantcontributorsto the isotopestratigraphy
is availablefor manycoresduringthistime
interval.
carbonateof deep-seasediments.Furthermore,E. huxleyi is also
the geologicallymostrecentexampleof how a singlespeciescan In thisanalysiswe documentthe beginningandthe endof this
dominate the calcareousphytoplankton. It is known to have globaldominance interval.We relatethePleistocene
morpholog-
dominatedthe sedimentassemblagesin the Atlantic Ocean for icalassociations
withthoseof theHoloceneanddiscuss threehy-
the last 85 kyr [Thierstein et al., 19771.The study of the domi- potheticalcausesof the dominancewhich are (1) selectivedisso-
nanceof calcareousphytoplanktonspeciesin the pastmay pro- lution, (2) globalenvironmental
change,and (3) evolutionary
vide an insightinto the relationshipbetweenbloomsand local as adaptation.
well as global environmentalchanges. Gephyrocapsawas proposedas a genusby Kamptner119431
The global dominanceof Gephyrocapsacaribbeanicaduring and is believedto be the ancestorof E. huxleyi[e.g.,Mcintyre,
the mid-Brunhes(484-262 ka) hasnot yet beenwell documented. 1970; Young, 19891. The first frequent occurrence of
(Dominance is related to diversity measurementsand can be Gephyrocapsa coccolithsis usually observed in the lower
quantifiedasthe relativeabundance of onespecieswithina group P|iocene3.5 Myr ago in the ReticUtofenestra pseudoumbilicus
Zone (NeogeneNannoplanktonzone (NN) 15) in which it is as-
sumed to have evolved from its ancestor Reticulofenestra
Copyright
1998bytheAmerican
Geophysical
Union.
pseudoumbilicus IPirini-Radrizzani and Valleri, 1977;
Paper number 98PA00610. Samtleben,1980;Rio, 1982i- However,recently,thefirstappear-
0883-8 •05/98/98 PA-00610512.00 anceof Gephyrocapsacoccolithswasreportedto haveoccurred

517
 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
518 BOLLMANN ET AL.: LATE PLEISTOCENE GEPHYROCAPSA DOMINANCE

-180 - 140 - 100 -60 -20 20 60 100 140 180

b ß
6O GE•L 60

[] Go
4O GM ß 40
GC
. y .... i .... i ....... i .... ! .... i .... i ....

•;• '• • • • Length•m) 20

20 - • •' , '' -•-•-

-20
-20•'
-40 -40

C • Solid symbols: unimodal samples

Open symbols: dominant association within a polymodal assemblage
-80 •
-180 - 140 - 100 -60 -20 20 60 100 140 180
Cold: L: >2.4pm, BA: <27ø,< 21øC, &: Oligotroph:L: >3.1pm,BA: 270-56ø,22ø-25øC ,: Larger: L: >3.9pm,BA >56ø, 18ø-23øC
Transitional: L: 2.4-3.1pm,BA: 270-56ø, 19ø-20øCB: Equatorial:L:3.1-3.9pm,BA: > 56ø, 25ø-29.5øCI•h Minute: L:< 2.4pm, BA: 200-50ø, 19ø-27øC

Figure1. (a) Measurements

determined fromdigitized
scanningelectron
microscope(SEM)images of a singleplacolith:
BA,
Bridgeanglemeasured andL, Length.(b) Six morphological
associations
of Gephyrocapsadeterminedin Holocenesediment
assemblages.
(c) Biogeography
of thesixdifferentmorphological
associations
of genusGephyrocapsadetermined in Holocene
sediments
[afterBollmann1997].Thelinesatthesymbolsindicate
themeanorientation
ofthebridgewithinanassemblage.

in the middle to late Miocene [Jiangand Gartner, 1984; Pujos, and Medd, 1979|. Thompson[1989l describedthe dominanceof
1987; Raffi et al., 19931. several nannofossil species in annual laminae in Lower
At least27 specieshavesofar beendescribedwithinthe genus Cretaceousmarlsandexplainedthe succession of the dominant
Gephyrocapsaon the basisof variouscriteria [Bollmann,1997]. species withinonelaminaastheannualnannoplankton produc-
Br•h•ret |19781 and Samtleben| 19801demonstratedthe feasibil- tion in a seasonalcycle. The Upper Cretaceousof northern
ity of distinguishing
differentspeciesor morphotypesby using Germanyis, in part,dominatedbycalcifieddinoflagellates
called
easily measurableparameterssuchas size and bridge angle. calcispheresI Locker, 19671.The causesfor their dominanceare
Downcore studieshave revealedtime-progressivevariationsof thoughtto be a combination
of hydrodynamic
enrichmentand
relative abundanceas well as morphological variations of ecologicalcauses|Neuweilerand Bollmann, 1991,and references
Gephyrocapsa coccoliths.
Thesevariationshavebeeninterpreted thereinl. A geographicallyvariable successionof dominant
as indicatingeither evolution,environmentalchange,or both species(Thoracosphaera andBraarudosphaera) wasdeposited
[Gartner, 1972; Pujos-Lamy, 1976; BrJhEret, 1978; Samtleben, during the earliest Danian following the demise of most
1980; Rio, 1982; Pujos, 1987; Gard, 1988; Gartner, 1988; Cretaceoustaxa!e.g.,Thierstein,1981I. Severallayersof domi-
Matsuoka and Okada, 1989; Baumann, 1990; Gard and nantBraarudosphaera
rosaduringtheOligocene arethought
to
Backmann, 1990; Matsuoka and Okada, 1990; Gard and Crux, becausedbyenvironmental
stress
[Maxwell
etal., 1970;Tappan,
1991; Matsuokaand Fujioka, 1992; Raffi et al., 1993; Bollmann, 19801.Thewell-documentedsmallGephyrocapsa AcmeZone
1995; Su, 1996; Wells and Okada, 1997]. from0.93to 1.25Mawasinterpreted
byGartner [1988] asbeing
Morphometricmeasurements of Gephyrocapsacoccolithsin relatedto changesin theoceancirculation.
globally distributedHolocenesedimentsallowed the identifica- The Gephyrocapsa dominance intervalduringthe mid-
tion of six different morphologicalassociations(possibleeco- Brunhesis characterized
by somemajorchanges in theglobal
phenotypes) which appear to have distinct environmental environment.
Oneofthesechanges
istheonset
ofthepronounced
preferences with respect to temperature and productivity 100kyr climaticcyclicityin themid-Pleistocene
around620-650
|Bollmann,19971(Figure 1). ka |Mudelseeand Stattegger,1997,and references therein;
The dominanceof a singlespecies,like E. huxleyiin Holocene Raymo,19971.In addition, thebeginningof thegrowthof the
sediments,is not uniquein the geologicalpast.Dominanceinter- GreatBarrierReeffallswithinthebeginning of thisperiodat
vals of single specieswithin the calcareousphytoplanktonhave about0.5 Myr ago[DaviesandMcKenzie, 1993].Furthermore,
occurred since Late Jurassic time, but little is known about the intensifiedcarbonate
dissolution hasbeenwidelydocumented in
potential relationshipbetween the environmentand dominant lowlatitudes
during
thistime[Berger,
1968,1973;Thompson,
calcareousphytoplanktonspecies.The earliestknownexampleis 1976;Adelseck
andAnderson,
1978;Schiffelbein,
1984;Crowley,
Ellipsagelosphaerabritannicain the KimmeridgeClays [Gallois 1985;Petersonand Prell, 1985;Bergerand Vincent,1986;
 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
BOLLMANN ET AL.: LATE PLEISTOCENE GEPHYROCAPSADOMINANCE 519

Arrhehius, 1988; Farrell and Prell, 1989; Wu and Berger, 1989; -180 - 140 - 1O0 -60 -20 20 - 60 -1,00 140 180

Wu et al., 1990, Archer, 1991; Bassinot et al., 1994; Weber et al., ,

1995]. There is, however, still no agreementabout whether the
mid-Brunhes dissolutioncycle [Crowley, 1985] was caused by
kf' t '. -c,o
changesin bottom water chemistry [Berger, 1968; Fartel and 60 K708,•,7
Prell, 1989; Archer, 1991; Lea, 1995] or by changes of the
productionrate of calcite and organic matter [Arrhenius, 1988;
40 '.•-- _..• • •/•' - 40
• V23-100 f • • KH84-1,St.21
Archer, 1991]. The highestcarbonatesedimentationratesduring
0 + • .+ - 0
the last million years at high latitudesoccurredduring the time k ? a 3 •.. •ERDC•o•es
span of this dominance [Sancettaand Silvestri, 1984; Hodell,
1993; Henrich and Baumann, 1994] which may have been
balanced by enhanced dissolution at low latitudes [Luz and -40
-60
•, ' • ODP704
-DSDP593
DS•
•L-40
DP594-60
Shackleton, 1975]. Core Latitude Longitude Water Depth Stratigraphy
The interglacialsduring that time seemto have been warmer ODP 643A 67ø43'N 01ø02'E 2764 m Henrtch and Baumann 119941;Bleil 119891
K708-7 53ø56'N 24ø05'W 3502 m Ruddbnanet al. 119861
thanthe currentone [Burckle, 1993], and oxygenisotopestage11 V 16-205 15ø24'N 43ø24'W 4045 m van Donk [ 1976l; Glass et al. 11967l
at 362-420 ka, in the middleof the dominanceinterval, was prob- M 13519 05ø40'N 09ø5 !'W 2862 m Sarntheinet al. 119841;Bled et al. 119841
V28-239 03ø!5'N 159ø1!'E 3490 m Shackletonand Opdyke11976[
ably the warmest interglacial in the last million years ERDC92BX 02ø!3'S 156ø59'E 1598 m Wuand Berger 119891
[Shackleton, 1987; Howard, 1997]. From evidence from marine ERDC83BX2 01ø24'N 157ø!8'E 2342 m Wu and Berger 119891
ERDC!29BX 00ø00'S 161ø58'E 4169 m Wuand Berger 119891
and continentalrecords,Burckle [ 1993] identified stages11, 13,
and 15 as especially warm interglacials. The temperaturesin Figure2. Locationof coresusedin thisstudy.Crosses,
coreswherethe
theseinterglacialsmight have been up to 5øC warmer than today dominanceintervalof Gephyrocapsacaribbeanicawasshownandwhere
in north China, up to 3ø-6øC in the Himalaya, up to 3øC in morphologicalanalysisweredone(this study). Opensquares, cores
Siberia, and up to 4øC in the eastern Urals. Subtropical wherethedominanceintervalof Gephyrocapsawasreportedor shown;
OceanDrillingProgram(ODP) Hole704 [GardandCrux,1991,Figures
conditionswere also reportedfrom the Atlantic coastof France
3 and41;DeepSeaDrillingProject(DSDP)Site593 [DudleyandNelson,
(50øN) during stage 11 [Rousseauet al., 1992]. The end of the 1989,Figure31;DSDP Site594 [DudleyandNelson,1994,Figure31;
Gephyrocapsa dominance coincides with the mid-Brunhes KH84-1, Station 21 [Matsuoka and Okada, 1989, Figure 31. Solid
climatic event [Jansen et al., 1986]. squares,
coreswheretheendof thedominance
intervalwasreported
by
Thiersteinet al. [ 19771.Note thatthe stageboundaries
for coreV 16-205
wereidentifiedin this studyat the followingdepths:event7.0, 95 cm;
2. Materials and Methods event8.0, 127.5 cm; event9.0, 155 cm; event 10.0, 180 cm; event 11.0,
200 cm; event12.0,220 cm; event 13.0,232.5 cm;event14.0,252.5 cm;
Coccolith assemblagesof five deep-seacores from the event 15.0,272.5 cm; and event 16.0,293 cm.

Norwegian-Greenland
Sea,theAtlanticOcean,andthe Pacific
Ocean were usedin this study(Figure 2). The corescover equa-
torial to northernsubpolarwater masses. Geometryand magnificationof the SEM were calibratedby
All coresstudied
havehigh-resolution
6180isotope
stratigra- measuringlatex spheresof a given meandiameterof 1.98ium +
phiesbackto at leastisotope
stage17 (references
in Figure2). 0.1. The averagesizeof 97 measured
sphereswas 1.89ium+ 0.09
OceanDrilling Program(ODP) Hole 643A and pistonCores which is within the statistical deviation.
V16-205, M13519, and V28-239 had magneticreversalsidenti- All morphometric measurementsand the subsequentdata
fied, and the isotopestratigraphies of all coreshavebeencali- treatmentwere conductedfollowing the proceduresdefined by
bratedby two synchronous nannofossil events:the lastoccur- Bollmann[19971 (Figure l a). Measurementsof the lengthtaken
renceof Pseudoemilianialacunosa(isotopestage 12) and the from Baumann[ 19901were dividedby 1.3 becauseof a constant
firstappearance of E. huxleyi(isotopestage7). The isotopestage sizeoffsetbetweenthe samesamplesmeasuredin this studyand
boundaries were taken from the literature for each core with the thosemeasuredby Baumann! 19901.
exception of V 16-205IvanDonk,19761.Thestageboundaries of Relative abundancecounts were done using the taxonomic
V 16-205 had not been identified and therefore had to be deter- conceptof Mcintyreet al. [ 19701to distinguishspecieswithin the
minedin thisstudy(seeFigures2 and3). The latestversionof .
genusGephyrocapsa.They useda bridgeangle greaterthan45ø
the variouspublishedisotopestratigraphies for Hole 643A by to define Gephyrocapsaoceanica (warm), a bridge angle less
Henrich and Baumann !19941 was used. They specifically than 45 ø to define G. caribbeanica (cold), and a small size (2.2-
changedthe isotopestage13 first publishedby Jansenet al. 1.9 •m) to recognizeGephyrocapsaericsonii. This taxonomic
[19891to stage 15. The age modelsof each core were created conceptwaschosenhere becausethereare alreadyseveralstudies
usingthe age estimatesof imbrie et al. [19841for the isotope in which this conceptwas applied,for example,Climate: Long-
stageboundaries up to isotopestage16. A linearsedimentation Range investigation,Mapping, and Prediction(CLIMAP) [1976,
ratewasassumedbetweenstageboundaries. 19811, Geitzenauer et al. !1976, 19771, Roche et al. [19751,
All sampleswereprepared asfollows:a smallamountof sedi- Ruddiman and Mcintyre 119761and Thiersteineta/. | 19771,and
mentwassuspended
inwateranddisaggreg
atedbyimmersion
in thedominance
intervalof G. caribbeanica
wasalreadypartly
an ultrasonicbath.A dropof highlydilutedsuspension
wasput documented/shown,for example, by Ruddiman and Mcintyre
on a cover slide and was dried. Then the cover slide was mounted 119761, Thierstein et al. 119771. However, the informal
on an aluminiumstubandsputteredwith goldfor subsequentob- nomenclature of Bollmann [19971 was used to define different
servation in a Hitachi S2300 scanning electron microscope morphologicalassociationsof Gephyrocapsacoccoliths(Figure
(SEM). 1). The comparison between Holocene morphological
 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
520 BOLLMANN ET AL.: LATE PLEISTOCENEGEPHYROCAPSA
DOMINANCE

ODP Hole 643a K708-7 V16-205 M13519 V28-239

G. caribbeanica (%) G. caribbeanica (%) G. caribbeanica (%) G. caribbeanica (%) G. caribbeanica (%)
0 20 40 60 80 100 0 20 40 60 80 100 0 20 40 60 80 lOO 0 20 40 60 80 lOO 0 2o 40 60 80 lOO
.,,i,,,i,,,i,,,i,,,
o
, , I , , I , ii• i I • , 1
ß
ß 3

500. ß

[]
7

7
? []
End
9

9 11

11
11
1500'
13
Beginning
ß
13 1:3
15

2000- 15

17 17 []
17 ß

2500

o. Cold:L: >2.4pm,BA:<27ø,< 21øC, A: Oligotroph:L: >3.1pm,BA:27o-56

ø, 22ø-25øCT: Larger:L: >3.9pm,BA >56ø,18ø-23øC
©' Transitional'
L: 2.4-3.1pm,
BA:270-56
ø,19ø-20øC
l: Equatorial'
L:3.1-3.9pm,
BA'> 56ø,25ø-29.5øC
I•: Minute:L:<2.4pm,BA'200-50
ø,19ø-27øC
Figure 3. Late Pleistocenestratigraphyof Gephyrocapsaspp. Solid lines connectingdots are relative abundancesof G.
caribbeanica(morphological associationsGT, GO andGM). The dashedlinesindicatethebeginningandthe endof the global
dominanceof Gephyrocapsacarribeanica.Symbolsindicatedominantmorphologicalassociations of Gephyrocapsafound(see
also Figure 1). Solid symbolsare unimodalassemblages whereasopen symbolsrepresentthe dominantassociationwithin a
polymodalassemblage. Note thatthe linesat the symbolsin Core K708-7 markthedepthrangeof samplesaddedto obtainat least
a samplesizeof 30 measurements.
For sourcesof dataseesection2.

associations
as definedby Bollmann
[1997]-and the excluded
fromallcounts.
Scatter
plotsofalldowncoremorpho-
Gephyrocapsaspeciesas defined by Mcintyre et al. [1970] metric
analyses
areshownonhttp://www.erdw.ethz.ch/-bolle/pa
revealedthat G. caribbeanicamost closely resemblesthe 196/pa
196.html.l
morphologicalassociationsGephyrocapsaTransitional(GT),
Gephyrocapsa Oligotroph(GO), andGephyrocapsaCold (GC),
3. Results
whereasG. oceanica
mostcloselyresemblesthemorphological
associations
Gephyrocapsa
Equatorial(GE) andGephyrocapsa
3.1. Downcore Abundance Data
Larger (GL), and Gephyrocapsa ericsonii resemblesthe
morphologicalassociationGephyrocapsaMinute (GM). The In all five analyzedcores,G. caribbeanicadominatedthe cal-
question,
whethertheHolocenemorphological
associations
each careousnannofossil
assemblage
duringisotopestages13-9
or in groups representbiological speciesor subspecies (Figure 3). During the dominanceinterval the relative abun-
(phenotypic
or ecotypic
variants),
is stillanopenquestion
andis dancesof G. caribbeanica usuallyvary from 70% to 99%. The
the subjectof ongoingresearch. riseto dominance in thedifferent
coresisdiachronous
(Table1).
The relative abundance counts of coccoliths for Core K708-7 It occurred
in latestage16to earlystage15at highlatitudes
in
are from Ruddimanand Mclntyre[ 1976].The relativeabundance the North Atlantic(in Hole 643a at 633 ka and in CoreK708-7 at
datafor ODP Hole643A andsomemorphometric measurements 613ka),whileat lowerlatitudes in theNorthAtlanticit began
for K708-7 andHole 643A are from Baumannl19901.The rela- only in theearlystage13 (in CoreV 16-205at 527 ka andin Core
tive abundance
countsin Core V28-239 wereconducted
by M1351 at 529 ka). In the westernequatorialPacific the
Thiersteinet al. 119771from stage11 to the Holocene.The rela- dominance beganin thelaterpartof stage13(in CoreV28-239at
tive abundance countsandmorphometric measurements werede- 484 ka). G. caribbeanica
abundance beganto decrease
in most
termined in this studyfor V 16-205, M 13519, and V28-239 and
partlyfor Hole 643A. At least200 specimens in everysample
werecounted to providea maximum errorof +7%at 95%proba- I Supporting
datatables,
rawcounts,
andscatterplots
areavailable
on
bility level.All countswereperformed at a magnification
of diskette
orviaAnonymous FTPfromkosmos.agu.org,
directory
APEND
(Username = anonymous,
Password= guest).
Diskette
maybeorderedby
6000xusingthesametaxonomic categoriesasCLIMAP[1981]. mailfromAGU,2000Florida Ave.,N. W.,Washington,
DC20009orby
Accordingly, theusuallyabundantFlorisphaera profundawas phoneat 800-966-2481;
$15.00.Paymentmustaccompanyorder.
 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
BOLLMANNET AL.:LATEPLEISTOCENE
GEPHYROCAPSA
DOMINANCE 521

Table 1. EstimatedAgesof the Beginningandthe End bridgeanglebetween27ø and56ø anda meanlengthsmallerthan

of the Dominance in All Five Cores Studied 3.1/•m. In Holocenesedimentsthis associationis only found in
areas with a mean sea surfacetemperaturebetween 19øC and
Core Beginning,Depthin Core, End, Depthin Core, 20øC. In Cores V16-205, M13519, and V28-239 the dominance
ka cm ka cm
interval is bracketed by assemblages consisting of two
643A 633 1740 269 837 morphologicalassociations (Figure3).
K708-7 613 1213 251 560 Core V16-205 showsthe lowest variability in morphology.
V16-205 527 255 279 146 Before the dominanceinterval, morphologicalassociationGT
M13519 529 746 251 375
was most abundantwithin the Gephyrocapsacomplex.During
V28-239 484 439 270 252
the dominanceintervalandafterthedominanceinterval,morpho-
logicalassociation
GO wasmostabundant(Figure3). The largest
morphologicalvariability during the last 620 kyr was found in
Core M13519. It is remarkablethat the sameassociation,GE, oc-
coresin latestage9, anditsdominance
wasdefinitelyoverby curredbeforeand after the dominanceinterval(Figure 3). The
early stage7. Thereforethe end of the global mid-Brunhes same is true for Core V28-239 (Figure 3). At higher latitudes
dominance intervaloccurredwithina shortertime period(<30 (Core K708-7 and ODP Hole 643A) a changefrom association
kyr) thanthe riseto dominance(4150 kyr). The endoccurredin
GT and GO to GC occurredbetweenstages8 and 6 at the end of
the Norwegian Sea (Hole 643A) at 269 ka and in the North
the dominanceinterval (Figure 3). From all data it is clear that
Atlantic(CoreK708-7)andtheeasternequatorial Atlantic(Core the morphologicalvariability during the dominanceinterval is
M13519)at 251 ka. In thewesternequatorialPacific(CoreV28-
very low comparedto the Holoceneand to the isotopestagesbe-
239) and North Atlanticgyre margin(core V16-205) the end
fore and after the dominanceinterval(Figure 3).
occurredat 279 and270 ka, respectively.
Triersteinet al. [1977,
Figure21showedin globallydistributedcoresthatthedecrease in
abundance of G. caribbeanicaoccurred duringoxygenisotope
stages9 and 8. There is no systematicgeographicpattern
observablefor theabundance decrease,although Gephyrocapsa • 7o- ODP643A a K708-7 b
spp.continued to dominate
in highlatitudesaftera breakduring 1•9 60
_

isotopestages
7 and6. Thereforetheendof theglobaldominance
interval
isassumed
tobesynchronous
atanaverage
agelevelof • so
262ka.Theduration
oftheglobal
dominance
interval
of G. • n0
caribbeanica is thereforeabout220 kyr. The dominanceof ':'
nn 30
Gephyrocapsa coccoliths can also be identified in additional
recordspublishedpreviously(Figure2)' at southernlatitudesin 20
ODPHole704A(46øS)
between
stage13and9 (GardandCrux .... i .... i,,, i .... i .... 1 .... i .... • .... .... i .... i,,. i .... i .... i .... i .... i ....

[19911their Figures3 and4, althoughthey lumpedall small 80

- C
placoliths
intoonecategory),
in theNorthPacific(20øN)from • 70- V16-205
-520 to -270 ka IMatsuoka
andOkada,1989,Figure31,andin
_

1•9 60
the SouthwestPacific(DSDP Site 593, 40øS;DSDP Site 594, _

45øS)fromisotope
stage14to isotope
stage
7 [Dudleyand • so
Nelson
' 1989,Figure3; Dudley
andNelson
' 1994,Figure31ø

nn 30
3.2. Morphometric Data
_

Morphological
analyses
of 2600Gephyrocapsa
coccoliths
of 20
late Pleistoceneage (650-10 ka) revealedthat all Pleistocene 10 .... 1 .... i,,, i .... i .... i .... i .... i ....
80
morphological associationsare readilyassignable to Holocene f
morphologicalassociations
definedby Bollmannl19971
(Figure o•,70i V28'239 • -Holocene
4). Theinformalnomenclature
of Bollmann!19971wasusedto •60 _
samples
describe
allmorphological
associations
ofgenus
Gephyrocapsa
• so
.

in the studiedcores(FiguresI and3). a}

'lJ 40
However,
during
thedominance
interval,
onlytwomorpho-'•
logical associations(GT and GO) out of six Holocene rn 30
associations
dominatethe coccolithassemblages
in all studied 20
cores(Figure 3). In Holocenesedimentsthesetwo associations
are restrictedto centralgyre and transitionalregions.The 10.0 I ....i....i....i....i....i....i....i....
1.5 2.0 2.5 3.0 3.5 4.0 4.5 5. 1. 1.5 2.0 2.5 3.0 3.5 4.0 4.5 5.0
morphologicalassociation
GO hasa meanbridgeanglebetween
Length (/jm) Length (prn)
27øand56øanda meanlengthgreater than3.1/•m.In Holocene
sedimentsthisassociationisonlyfoundin oligotrophicregions Figure4. Dominant morphological associations
in Pleistocene
(a-e)and
with a meansea surfacetemperaturebetween22ø and 25øC Holocene samples(f). Notethatall Pleistocene
morphologicalassocia-
tionsarereadilyassignableto Holocene morphological
associations
as
(centralgyre). The morphologicalassociationGT has a mean definedbyBollmannl19971 andindicated bydividing
lines.
 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
522 BOLLMANNET AL.: LATEPLEISTOCENE
GEPHYROCAPSA
DOMINANCE

1500 15ff
4. Interpretation and Discussion
There are severalpertinentquestionsto be addressed.Can en-
ERDC83B•(2
2000
ERDC92BX
'/ ' a ERDC92 200• b

vironmental processesand mechanismsbe identified that might •E2500 250• ERDC83BX2

have led to the rise to globaldominanceof two morphologicalas- •.3000 300•

sociationsof the genusGephyrocapsaabouthalf a million years

• Lysoc
3500 350•
ago? Why did only their dominanceand not their existenceend ysocline
4000
about a quarterof a million yearsago?What is the evidencefor ERDC129BX '.
4001
ERDC129BX
preservationalor environmentalchange?Must, largely by de- 4500 4501
fault, evolutionaryadaptationbe consideredas the main cause? 2.5 • 3'.5 4.5
Bridge Angle (o) Length (/zm)
8O 1500
4.1. Selective Preservation of G. caribbeanica
7O 2000 d ßß
Roth and Berger [ 1975] and Roth and Coulbourn [1982] • 60 GE GL
showed that G. oceanica is one of the more dissolution resistant
2500 %-
• 50
coccolithspeciesin Holocenesediments.Thus the dominanceof 3000

G. caribbeanica could be the result of dissolution, which could g 40 [• GT GO 3500

.'R_30
have eliminated most of the other species and also all
GephyrocapsamorphotypesotherthanGT or GO.
Enhanced carbonate dissolution between 500 and 300 ka in
20

10
:

1.5
GM

i i

2.5
GC
I i

3.5
I

4.5
4000

4500 •1
0
• Lysocli
i .... i ....

10 20
• .... i ....

30
i ....

40
, ....

50 60
, ....

70
low latitudesfrom all oceanbasinshasbeendocumentedin many
Length(/Jm) % Fraction>63/Jm
cores(see section1). This is supportedby a decreasein the per-
centageof the coarsefractiondominatedby plankticforaminifera Figure 5. Morphometricmeasurements of Holoceneassemblages along a
in the studied cores V28-239 [Shackletonand Opdyke, 1976, depthtransecton the OntongJava Plateau:(a) bridgeangle versusdepth,
(b) coccolithlengthversusdepth,(c) morphologyof threesamples(small
Figure 1], M13519 (data from Sarntheinet al. [1984], and V16- squares)and Holocenemorphologicalassociations (for a symbolexplana-
205 [Briskin and Berggren, 1975, Figure 12] during this time in- tion seeFigure 1), and (d) abundancefractionversusdepth(as a measure
terval. for dissolution,the relative abundanceof the fraction larger than 63 •um
However, if dissolution were the real cause of the dominance was used). All error bars show the 95% confidence intervals for the
means.
of only two morphologicalassociations,
one would expectthat
the morphologicalcompositionof Holoceneassemblages alonga
depth transect with demonstrateddissolution effects should
changetoward one of the dominantmorphologicalassociations,
[Bollmann, 1997]. Such correlationscould be usedto reconstruct
GT or GO, if the environmental conditions in the surface water
theseenvironmentalparameters duringthedominanceintervalto
were constant.Sucha well-knowndepthtransectis availableat
provideinformationaboutthe environmental parameters which
the Ontong Java Plateauin the westernequatorialPacific. The
mighthave led to the globallyobservedGephyrocapsadomi-
morphologyof Gephyrocapsacoccolithswas here analyzedin
nance.A successfulapplicationof transferfunctiontechniquesto
samples from water depths of 1598 (ERDC92BX1), 2342
reconstructpastpaleoenvironments, however,requiresthata few
(ERDC83BX2), and 4169 m (ERDC129BX). The local lysocline
crucialconditionsbe met [Imbrie and Kipp, 19711-Theseare (1)
lies at 3400 m depth [Gr6tschet al., 1991], and the mean seasur-
a close correlation of speciesabundancesor morphology in
face temperatureat each location exceeds 29øC at present Holocenemicrofossilassemblages with measuredoceanographic
[Levitus,1982]. In eachassemblage the associationGE dominates parameters, (2) no evolutionaryadaptation,(3) evensampledis-
the Gephyrocapsa complex, and there is no statisticallysig- tributionalongthe environmental gradientsto be reestimated,(4)
nificant change in morphology with depth of deposition exclusionof nonanalogassemblagecompositions,and (5) no or
(Figure5). minimal alterationof fossil assemblagesby differential dissolu-
In addition, Coccolithuspelagicusand Calcidiscusleptoporus tion, winnowingor reworking.
have been shown to be even more resistant to dissolution than
Conditions one and three were demonstrated by Bollmann
Gephyrocapsa spp. [Roth and Coulbourn, 1982]. They are [1997] and conditionfive wasdiscussedin section4.1. Condition
dominant in high northernlatitudesin plankton and Holocene four requiresthatonly thoseHoloceneassemblageswerechosen
sediment samples [Geitzenauer et al., 1977; Samtlebenand for the calibrationwhosemorphometricmeansand standardde-
Bickert, 1990]. Therefore a changeto high abundancesof G. viations were similar to thoseassemblagesencounteredin the
caribbeanica relative to the other two resistantspecieswould dominanceinterval (Figure 6a). These thirteenassemblagesare
morelikely indicateenhancedcarbonatepreservation,ratherthan locatedin subtropicalcentralgyre regionswith a mean seasur-
dissolution,at thesehigh latitudes.Furthermore,Gephyrocapsa face temperattire(SST) rangefrom 22ø to 25øC and in transi-
coccoliths are generally well preserved in the high-latitude tionalareaswith a meanSST rangefrom 19ø to 20øCas shownin
assemblagesof Hole 643A in the interglacial parts of the Figure6b.
dominance interval.
Standardlinear multipleregression
analysiswasappliedto the
Holocenedata set, linking the morphologicalparameters(mean
4.2. Global Environmental Change
bridgeangleA andmeanlengthL) to SST (Tmean, Tma x, Tmin)and
The morphologicalvariabilityof Gephyrocapsacoccolithsin to fertility(chlorophyllconcentration
C). The resultsindicatethat
Holocene sedimentassemblageshas been shown to be closely the environmental variability spannedby the Holocene as-
correlated with sea surface temperatureand fertility patterns semblagescan be describedby the variability observedin
 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
BOLLMANN ET AL.: LATE PLEISTOCENE GEPHYROCAPSA DOMINANCE 523

-180 - 140 - 100 -60 -20 20 60 100 140 180

80 - - 80
Sample
ID Latitude Water Depth, • •r• ' 60 '
E48-36 -30.93
Longitude
Depth,
mcm •
87.78 1366 2-4cm
55 a
N---•"•I
I •'•x7-17.6• 3'-•- 0---•'•'-• 50
INMD127Bx5-29.88
-33.982509 0-1cm•/ ø • 45
RCll-22 -20.10 -32.414167 TVVT• I• I '•'
-,o.oo o/ 't
RC8-91 -33.42-111.902723 1-5cm -- , z•' .,rl' I 35
6O RC8'94 '2728'102
083074 2cm •1••1 •,• '" 6O
RG9-104 -27.30 -123.30 3632 TVVT ... .,]1,{, •_•-
V19-55 -17.00-114.183177 4cm O ••' [ ,-•
V20-103 33.59-177.50
3442 TWT •;• .•J 0325
4O V20-171 -21.6068.29
2833 2cm l'J,.,,,•• • 20 4O
V20-176 -22.40 60.18 3946 TVVT ', %,,,./•,,F •_•,,,•
V29-64 -27.2074.523404 TWT _• • 15
2O
. - ---• .... •' ' 10
• 2.53 3.54 4.5
_i
Length(/.,m)
2O

-2O -2O

-4O -4O

ß Holocene assemblages
-6O oPleistocene
• • assemblages_
• -60
- 180 - 140 - 100 -60 -20 20 60 100 140 180

Figure 6. Comparison between the morphologyof Holocene assemblagesand assemblagesfrom the Pleistocenedominance
interval. (a) Samplemeansof unimodalmorphologicalassociations in the Holocene(diamonds,GT and triangles,GO) and in the
Pleistocene(circles). Error bars equal 1 standarddeviation. (b) Locations of Holocene assemblagesthat are morphologically
comparableto assemblages of the dominanceinterval.

Gephyrocapsacoccoliths(Figure 7). The mean SST comparison interglacials(21.1ø + 0.3øC; 0.071-0.081 /•g/L chlorophyll). In
shows
thehighest
multiple
regression
coefficient
(r2 = 0.76).The general,the estimatesof the chlorophyllconcentrationssuggest
temperaturerange coveredis restrictedto 19ø-27øC(Figure 7a) lower productivityduring the dominanceinterval in the northern
and the standarderror of estimatesis 1.2øC. Slightly different and eastern Atlantic and higher productivity in the western
valuesapply for the minimumand maximumtemperatures(Table equatorialPacific than today(Table 3).
2). The fertility transferfunctionshowsa multipleregressionco- These rather puzzling resultsare not likely to be causedby a
efficient
of r2 = 0.66forthecovered
chlorophyll
rangeof0.045- violation of the crucial conditionsthree and four given above.
0.095/• g chlorophyll/L(Figure 7b). This leavestwo alternatives:eitherthe Gephyrocapsadominance
Application of these transfer-functionsto the assemblages intervalrepresents a globaloceanography very unliketoday's,or
from the dominance interval in these various areas reveals some the secondassumptiondoes not apply; that is, Gephyrocapsa
spectacularresults(Figure8 andTable 3). Interglacialpaleotem- populationshave evolvedin the past0.5 Myr.
peratureestimatesfor stages9, 11, and 13 comparedto modem As discussed in the introduction, there is some evidence for
valuessuggesta 11ø-14øCwarmer SST at high latitudes,a 4ø- markedly warmer climatesduringthe mid-Brunhes.Warmer SST
9øC coolerSST in the equatorialareasof the Atlantic,and a 5ø- than today at high latitudesof all oceansare also suggestedby
11øC cooler SST in the Pacific. Glacial paleotemperatureesti- high carbonatecontentsas documentedin North Atlantic Hole
matesduringthe dominanceinterval(stages10 and 12) in eastern 643A (67øN) [Henrich and Baumann, 1994, Figure 21, North
equatorialAtlantic Core M13519 are 23.8øCand 23.6øC(Table Pacific Core V20-119 IHays and Shackleton,1976; Sancettaand
3) and thusare comparableto the 24.2øC estimatedby CLIMAP Silvestri, 1984, Figure 31 and South Atlantic Hole 704 (40øS)
[1981] for the last glacialmaximum(Table 3). However, in Core IHodell,1993,Figures5 and61.In addition,
the{5i80isotope
V28-239 in the westernequatorialPacific the •leotemperatures recordof Neogtoboquadrinapachydermain South Atlantic ODP
for the glacial stages10 and 12 are 6ø-7øCcoolerthan the 18 ka Hole 704A [Hodell 1993, Figure61 showsa trendtoward warmer
estimateof 27.2øCby CLIMAP [ 19811(Table 3). In easternequa- SST than today in this region.Additional paleontologicalevi-
torial Atlantic Core M13519 the derived,morphology-based pa- dencealso pointsto warm interglacialsduringthe dominancein-
leotemperature estimatesfor the dominanceintervalshowan ad- terval. The increasedabundanceof plankticforaminiferain ODP
ditional oddity (Table 3). Glacials(23.7ø + 0.1øC; 0.055 + 0.003 Hole 643A during stages 15-11 [Spiegler and Jansen, 1989,
/• g/L chlorophyll)seemto have beenwarmerand lessfertile than Figure 61 are thoughtto be the resultof higher oceanicheat flux
 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
524 BOLLMANN
ETAL.:LATEPLEISTOCENE
GEPHYROCAPSA
DOMINANCE

to the northernlatitudes[Berger et al., 1994]. Paleotemperature

o]b r^2=0'66 estimatesderivedfrom plankticorganismtransferfunctionssug-
(3 27-
o... 5 gestSST up to 24.8øCin nearbyODP Hole 646 (68øN) in inter-
i- 25- • glacial stage5e [Aksuet al., 19921and up to 11øC(4øC higher
U)23- • than today) for the interglacialstages 11 and 9 [Aksuet al.,
19921.Transfer functionsof foraminifera (FTF) in core K708-7
ß•, 21
16
t • show interglacialsummerSST up to 19øCfrom 700 to 410 ka
.E [Ruddimanet al., 1986,Figure2] thatis 5. IøC warmerthantoday
U.117-t
15
15
..............
i
17
i
19
=16+014A+474L
i
21 23
i
'•
,
25
i
27
,
29
• :•t
0'037t/
/....C:
0.03
,0'2
....
-0'0004A
O.OS
,.....
- ,L and 4.4øC lower than mean SST derived from Gephyrocapsa
morphology.If Gephyrocapsawere to have bloomedduringa
0.07 0.09

Mean SST (øC) Chl. a Concentration (pg/L) very shorttime periodof 2-4 weeksin summer,possiblyduring
maximumwarmingof the surfacewaters,the discrepancy would
29:
C rn2=ø'42
•
O 27-

• 23-
25-

•
dr^2=0'71 not be as large. Yet, thereis little known aboutGephyrocapsa
bloomsin today'soceans.Temperatureestimatesbasedon radio-
larian transfer functionsin South Atlantic Core RC13-229 (25øS)
31 .Z" ß
suggestSST up to 20øCduringstage13 and 11, i.e., 6øCwarmer
• 21 than today [Morley and Hays, 1981, Figure9]. Howard and Prell
E19-
....
• [ 1992, Figures8 and91 usingFTF-derivedSST up to 3øChigher
than in the Holocenefor stages11 and 9 in severalcoresin the
• 17- t -129+007A+277L
southernIndian Ocean (42ø-48øS). Currently, there are no mor-
15 • • • • ,
............... , 5y ...,,.., ...,...,...,...,,..,,
• .....

5 17 19 21 23 25 27 29 15 17 19 21 23 25 27 29 phometricdataavailablefrom the southernhemisphere, but it ap-

Maximum SST (øC} Minimum SST (øC) pearslikely that the sameGephyrocapsa morphological associa-
tionswerepresentduringthedominanceintervalas in thenorth.
Figure 7. Plotsof measuredenvironmentalparametersversustheir esti-
There is little independentevidencefor coolerSST thantoday
mated values in thirteen Holocene assemblagesbasedon transfer func-
tions using Gephyrocapsa morphology (mean bridge angle A, mean at low latitudes duringthe dominance interval.The meanSST of
lengthL): (a) multiple linear regressionagainstmeanSST (residualstan- 23øC derivedfrom Gephyrocapsamorphologyin Core V16-205
darderror,1.232on10degrees of freedom; multiple r2,0.7676;f statistic, (subtropicalAtlantic)duringthe dominanceintervalis consider-
16.51 on 2 and 10 degreesof freedom; and p value, 0.0006782); (b) ably lessthanthe foraminiferapaleotemperatures of 26øC mean
multiple linear regressionagainst chlorophyll-concentration(residual
standard error,0.01126on10degrees of freedom; multiple r2,0.6689;f SST [Briskinand Berggren,1975, Figure 12], which is the same
statistic,10.1 on 2 and 10 degreesof freedom;and p value,0.003977); (c) temperature as today. In eastern equatorial Atlantic Core
multiple linear regressionagainstmaximumSST (residualstandarderror, M13519, the mean Gephyrocapsa SST estimatesin glacials
1.515on i0 degrees of freedom; multiple r2,0.421;f statistic, 3.635on2 (23.6 ø and 23.8øC) are close to the foraminifera estimates for
and 10 degreesof freedom;and p value,0.06508); and(d) multiplelinear
regressionagainst minimum SST (residual standarderror, 1.747 on 10
glacial stages 10 and 12 and for the last glacial maximum
degrees of freedom; multiple r2,0.7126;f statistic, 12.4on2 and10de- (24.3øC) [Pfiaumann, 1986]. However, the mean Gephyrocapsa
greesof freedom:and p value = 0.001961). SST estimatesduringinterglacialsat this locationare up to 7øC

Table 2. ComparisonBetweenthe Modem Maximum,Minimum,Mean Sea SurfaceTemperaturesand Chlorophyll

Concentrationand the EstimatedValues Derived From a Multiple RegressionAnalysis

Core Mean Standard Mean Standard Estimated Residual, Estimated Residual, Estimated Residual, Estimated Residual,
Bridge Deviation,Length,Deviation, Mean Maximum Minimum Chlorophylla
Angle, Temperature, Temperature, Temperature, Concentration,
deg deg /• m /• m øC øC øC oC oC oC /•gL'l /•gL-]

E48-36 38.9 15.9 2.6 0.6 19.2 0.0 23.4 -0.5 16.6 -0.4 0.095 -0.007
INMD115Bx7 35.2 11.6 3.4 0.6 23.7 -0.9 25.1 -0.1 22.7 -2.2 0.065 -0.007
INMD127Bx5 33.2 10.7 3.7 0.4 21.9 2.0 23.7 2.0 19.0 2.7 0.065 -0.017
RC 11-22 43.2 14.2 3.7 0.6 25.7 -0.6 27.5 - 1.2 23.3 0.1 0.055 -0.010
RC 13-17 43.1 13.3 3.6 0.6 26.2 - 1.4 27.4 - 1.3 24.9 - 1.9 0.045 0.002
RC8-91 40.5 12.6 2.8 0.6 19.2 1.5 21.1 2.7 16.3 1.7 0.08 -0.002
RC8-94 41.2 11.0 3.4 0.5 22.3 1.3 23.8 1.7 19.7 1.9 0.045 0.010
RC9-104 41.7 11.1 3.6 0.4 22.9 1.4 25.0 0.9 21.2 1.2 0.055 -0.005
V 19-55 40.2 11.7 3.5 0.4 24.4 -0.7 25.5 0.0 23.5 - 1.9 0.055 -0.001
V20-103 32.5 8.3 2.7 0.4 19.5 -0.5 23.9 - 1.1 15.8 0.0 0.095 -0.009
V20-171 38.0 14.0 3.3 0.6 23.7 - 1.0 26.7 - 1.7 21.8 - 1.4 0.065 -0.005
V20-176 41.4 14.1 3.5 0.6 24.3 -0.4 26.5 -0.9 22.5 -0.6 0.045 0.008
V 29-64 30.9 14.4 3.2 0.5 21.7 -0.6 24.6 -0.5 19.4 - 1.0 0.045 0.022

Residualis the difference betweenestimatedand measuredtemperaturesand chlorophyll concentrations.

 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
BOLLMANN ET AL.: LATE PLEISTOCENE GEPHYROCAPSADOMINANCE 2525

-180 -1 40 -1 O0 -60 -20 20 60 100 140 180

180 80

OD• 643A

60
60

K708 -7

40
40

20 '"',, V16-205 20
• • 'V28-239
0 ' M13519
• • 0

-20 ' '- •1 -20

-4 0 -4 0

-6O • ' • • • ' • 60

-1 80 -1 40 -1 O0 -60 -20 20 60 100 140 180

Figure 8. Comparisonof today'smeanSSTswith paleotemperature estimatesbasedon Gephyrocapsa morphology.The upper

lines within boxesindicatetoday'smeanSST obtainedfrom Levitus [19821(left) and the SST duringthe last glacialmaximum
(LGM, 18 ka) takenfrom Climate: Long-RangeInvestigation,Mapping,and Prediction(CLIMAP) 119811(right). The lower lines
showthe morphology basedSST estimates
duringinterglacials
(left) andglacials(fight)of thedominance
interval.SeealsoTable
3.

lower than today (27.6øC) and than foraminifera paleo- Pullentiatinaobliquiloculata)are alsodiscerniblein manyother
temperatures(27øC) (Figure 8 and Table 3). The mean SST coresof this area [Schiffelbein,1984;Bergeret al. 1993, Figure
estimatesobtainedfrom foraminiferashow both the highestand 5]. The interpretationof thesetrendsis difficult becausethey
lowest estimatesof the last 750 kyr during stages 14-9 (21 ø- could be influenced by temperature, salinity, or increased
27øC), and temperatures are generallyhigherin the interglacials carbonatedissolutionI Wu and Berger, 1989; Wu et al., 19901.
than in the glacials [Pflaumann, 19861. The reversed Alternatively,thesetrendsmightreflectchangesin the depthof
glacial/interglacialtemperatures
obtainedfrom the Gephyrocapsa the mixed layer, e.g., a risingthermocline[Bergeret al., 1993I.
morphologyindicateenhancedupwellingduringinterglacials,re- All of theseobservations castsomedoubtson the validity of
sulting in lower temperaturesand higher productivity, which is the ratherdramaticallylow, morphology-based paleotemperature
contraryto whathasbeeninferred fromincreased Corgcontents estimatesfor low latitudes.Independentevidenceclearly would
duringglacialperiodsin thiscore [Miiller et al.,1983, Figure41. be desirable.Alternatively,assumptiontwo for applicationsof
Equally problematicare the morphology-basedmean paleo- transferfunctionsmay havebeenviolated;that is, evolutionary
temperatureestimatesof 20-21.4øC for the dominanceinterval adaptationsmay have occurred.
(Table 3) in westernequatorialPacific Core V28-239 because
they are up to 9øC lower thantoday'smeantemperature(29.2øC) 4_3. Evolution
and 7øC lower than the foraminifera paleotemperatureestimates
for the last glacial maximum(Table 3). There are no foraminifera Certain morphological associationsof Gephyrocapsa
paleotemperatureestimatesavailable for the dominanceinterval (specificallyGT and GO) might have becomedominantbecause
of this core, but an indicationof lower temperaturesduring the they evolved especially successful,new traits around 633 ka.
dominance interval might be the abundancedecreaseof the Morphologicalchangewithin the Gephyrocapsacomplexhasof-
planktic foraminifer Globigerinoidesruber, a warm temperature ten been assumedto be the result of evolution [Samtleben, 1980;
indicator [Thompson1976, Figure 61. However, G. ruber is also Matsuoka and Okada, 1990; Gard and Crux, 1991; Matsuoka
one of the most solubleplankticforaminifera [Berger, 19681and and Fujioka, 19921.
thus could be rarer because of differential dissolution.Although the first appearances
Lower of morphologicalassociations
temperatures thantodayaresupported by thedecreasing 6180 GO and GC havebeenidentifiedin isotopestage14 in V 16-205
values of Globigerinoides sacculifer since isotopestage 12 in and in stage5 in core K708-7 and Hole 643A, respectively,they
western equatorial Pacific Core V28-239 |Shackleton and cannotbe inferred to have evolved then becauseno assemblages
Opdyke,1976,Figures I and21.Identicallong-term 6J80trends older than 633 ka have been analyzedyet in thesecoresand the
of the <38 /,tm fraction and some planktic foraminifera (e.g., two morphologicalassociations are likely to haveexistedbefore.
19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
0
BOLLMANN ET AL.: LATE PLEISTOCENE GEPHYROCAPSADOMINANCE

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526
 19449186, 1998, 5, Downloaded from https://agupubs.onlinelibrary.wiley.com/doi/10.1029/98PA00610 by Algeria Hinari NPL, Wiley Online Library on [18/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
BOLLMANN ET AL.: LATE PLEISTOCENE GEPHYROCAPSA DOMINANCE 527

The occurrenceof the other four Gephyrocapsamorphological were warmer than today duringthe dominanceinterval, suchas
associations(GT, GL, GE, and GM) before and after the domi- paleotemperature estimatesderivedfrom plankticforaminifera,
nanceintervalsuggests that no evolutionarychangeoccurredin high carbonatecontents,and temperatureestimatesfrom
their morphology.
Thusit is mostlythe failureof all attemptsto continentalrecords.However,coolertemperatures thantodayat
identifyenvironmental or preservationalevidencefor conceivable low latitudesaresuggestedby ourtemperatureestimatesderived
processes responsiblefor the observedmid-Brunhesdominance fromGephyrocapsa morphology, butarenotsupported by other
of Gephyrocapsamorphologicalassociations which leavesevo- paleotemperature indicators.
Coolertemperaturesthantodayin
lutionarychanges,leadingto increased, thoughtransitory,fitness, the westernequatorialPacificmightbe inferredfrom heavier
as a likely cause. Ongoing analysesof the morphological 6180values
inplanktic
foraminifera,
butthe6180signature
of
developmentof gephyrocapsids prior to the dominanceinterval planktic foraminiferamay also have been influencedby
may revealwhetherthe dominantmorphologicalassociations had dissolution
andsalinitychanges.
evolved alreadyearlier or representan evolutionarynovelty that Unrealisticallylow temperature
estimatesobtainedfrom trans-
originatedat the beginningof thedominanceinterval. fer functions
basedonthemorphology
of Gephyrocapsa
suggest
thatthe morphological associations GT andGO maybothhave
5. Summary and Conclusions responded to environmental parameters differentfromthosethey
seemto respondto today.The possibilityof morphological de-
The global dominanceof two morphologicalassociationsof pendence from environmental parameters otherthansurfacewa-
Gephyrocapsa(GO and GT) duringoxygen isotopestages13-9, tertemperature andfertilityis currentlybeinginvestigated. There
which constitute70%-99% of all placolithsin the fine fractionof are someindications that Gephyrocapsa morphological associa-
all deep-seacoresanalyzed,can be documented.The earliestrise tionsmayhaveresponded to globalchanges in thedepthof the
to dominanceof Gephyrocapsahas been observedat 633 ka in thermocline. The far off morphology-based paleotemperature
the Norwegian-Greenland Sea and the youngest has been estimates,
however, makeit appearlikelythatthemorphological
observed150,000 yearslater in the westernPacific. The end of associationsof Gephyrocapsa havechangedtheirenvironmental
the Gephyrocapsadominanceintervalwas synchronous at about dependencies andpreferences through evolutionary processes.
262 ka, and the minimum duration of the global dominance Progressin our understanding of theeffectsof a globaltake-
interval of G. caribbeanica was approximately 220 kyr. The overby a singlecalcareous phytoplankton species on theglobal
morphology of Gephyrocapsa assemblagesin single samples carboncycle will requirebudgetaryconsiderations for which
shows bimodality before and after the dominanceinterval and accumulation ratesof crucialcarbonate components in a global
unimodalityduring the dominanceinterval. Selectivepreserva- andtemporalframeworkwill be needed.
tion as a majorcausefor the dominanceof the two morphological
associationscan be excludedbecausein Holoceneassemblages Acknowledgments.We wouldlike to thankA. Mcintyreand B.
Gephyrocapsamorphologyis not influencedby dissolution,al- Molfino for providingsamplesfrom North Atlantic Core K708-7; M.
thoughthere is no doubt that during the dominanceintervalen- Sarntheinfor samplesfrom equatorialAtlantic Core M13519; and N.
hanced carbonate dissolution occurred in low latitudes. The ob- Shackletonfor providingsamples
fromequatorialPacificCoreV28-239.
K. yon Sails Perch-Nielsen and P. Westbroek made useful comments on
served dominanceof Gephyrocapsain nannofossilassemblages an earlyversionof thispaper.The manuscript profitedfromthe reviews
and the carbonatedissolutionmore likely representa globally of J. Backman,
M. Delaney andM. Leckie.Thisworkwassupported by
changedenvironment.There is evidencethat global climates grantsfrom the Swiss National ScienceFondation.

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